Worldwide Generic Key to Pupae of Ceratopogonidae This is the first attempt at keying pupae of all known genera of Ceratopogonidae of the world. Generally, regional keys are more useful for identification but the material I examined was seriously limited and often included reared species representing genera which were otherwise unknown within a given region as immatures. The inclusion of distributions for each keyed genus, based on all stages, will help to exclude some taxa for those keying material from a given area. A key to the known subgenera of Forcipomyia is also included. Each of the families of Culicomorpha was keyed by Borkent (2012). Ceratopogonidae are the only Diptera with a curved S-shaped hind leg lying under the wing, with the terminal process at the apex of the abdomen either lobe-shaped or tapering to a point, without abdominal spiracles on segments 5–7 (or elsewhere on the abdomen) and a respiratory organ with at least some apical pores (many with additional subapical pores). Some ceratopogonid pupae are superficially quite similar to those of some Psychodidae but can be distinguished by their S-shaped hind leg. In the following key I have attempted to key out common taxa first and to use features that can be seen in material preserved in alcohol or glycerin. Regrettably, for some genera, slide mounting is required and in such instances, readers are strongly urged to read the materials and methods section—poorly mounted specimens and especially those that are crushed by coverslips, are often miserable to identify. Well prepared material is always of benefit in using keys to insects but is particularly important with ceratopogonid pupae. If pupae, and especially pupal exuviae, are compressed by coverslips, this can make identification hellish but in most instances, yet possible, if the reader is experienced in recognizing morphological features. So too, dirty specimens often obscure important sensilla and can make their detection nearly impossible. Features of segment 4 are generally also present on segments 5–7, so that if segment 4 is obscured the specimens may yet be identified using these other segments as surrogates. Readers will see that some genera, such as Palpomyia, key out in more than one place in the key. Some genera could not be diagnosed as separate from all other genera and this may indicate the need for further research. Either some of these genera are paraphyletic, I may have missed some important features, been fooled by features that are markedly homoplastic (i.e. using highly variable features as part of the key), was misled by misassociations and/or misidentifications, or any combination of these potential problems. If there are differences between males and females noted in the key, males can be distinguished from females by the presence of two well-developed genital lobes, nearly always situated ventromedially (some Forcipomyia males have the lobes placed dorsally). The relative development of the male genital lobes reflects the general length of the developing gonocoxites and gonostyli. Readers should be particularly aware that this key is undoubtedly flawed and seriously incomplete, with only 45 of 111 genera known, and of those known, for many genera, only a small percentage have been reared (Table 3). Of these known genera, Amerohelea is too poorly understood to be included. A key to the subgenera of Forcipomyia is included below but only 14 of 35 subgenera are known as pupae. For further challenges, readers are urged to read "Warning to Readers" above. Properly prepared specimens are important and sometimes vital to identification (see Materials and Methods). Overall, pupal exuviae in good condition are easier to identify than whole pupae (which need to be carefully cleared to see sensilla). 1. Mouthparts without mandibles or maxillae and the two halves of the labium either broadly abutting medially or slightly separated (Figs. 23F–G); abdominal segment 4 with or without the following combination of features: posterior sensilla on short, bilobed or shelf-like tubercles, all more or less the same size and evenly spaced along posterior margin of abdominal segment (Fig. 56C); terminal process either tapering from narrow (Fig. 73E) to broad base or with secondary divisions, with pair of short to moderately elongate setae arising from or near its apex (Figs. 72K–L, 73A–D, F–G), some with basal elongate seta (Fig. 73E)....... DASYHELEINAE....................................................... Dasyhelea (pg. 45) (worldwide) - Mouthparts with at least mandibles present (e.g. Figs. 23A, 24A–E) or if not evident then the posterior margin of the labium forming a continuous (not divided medially) structure (Fig. 24F) or separated medially by posterior extension of labrum and/or hypopharynx (Figs. 23C, 23E); abdominal segments with posterior sensilla of variable sizes and on differing sizes of tubercles (or not on tubercles) but not arranged in the combination of being on short, bilobed or shelf-like tubercles and all more or less the same size and evenly spaced along posterior margin of abdominal segment (e.g. Figs. 55A–C, 56A–B, 57A–C); terminal process simple, either short and apically rounded (Figs. 72B–E) or tapering from base (e.g. Figs. 72A, 72F–H, 73H–L), with, at most, a single short to elongate seta (some Forcipomyia (Fig. 72I) and Dibezzia (Fig. 75I))..........2 2. Dorsolateral cephalic sclerites abutting medially (Fig. 13J); hind leg not visible at lateral margin of wing in lateral view (Figs. 32A–B); terminal process either rounded (Figs. 72B–E) or pointed (Fig. 72A) apically (the latter only in western Australian specimens)................ LEPTOCONOPINAE............ 3 (worldwide) - Dorsolateral cephalic sclerites separated medially by dorsal apotome (which reaches the anterior margin of the mesonotum); hind leg visible at lateral margin of wing (Figs. 32C–L, 33A–L); terminal process tapering and apically pointed (Figs. 72F–L, 73A–L)...................................................4 3. Body with short thick, undivided setae, with many on abdomen (D-5-IV, D-8-IV, L-2-IV, L-3-IV, V-5-IV, V- 7-IV) directed posteriorly (Fig. 8B) or hooked anteriorly (Figs. 8C, 55B); apex of palpus posterior to the posterior margin of labium (Fig. 23B); terminal process short and rounded apically (Figs. 72B–E)............................................................................. Leptoconops (pg. 38) (worldwide but with marked habitat restrictions, see generic description) - Body with elongate setae (bifurcating in at least A. mcmillani, only species known as pupa) (Figs. 8A, 55A); apex of palpus anterior to posterior margin of labium (Fig. 23A); terminal process tapering and pointed apically (Fig. 72A)........................................................ Austroconops (pg. 37) (southwestern Australia) 4. Mesonotum nearly smooth or with at least one pair of elongate tubercles of various shapes from short (about as long was wide) (Figs. 8D–E, 8H, 10E–G) to very elongate (Figs. 9A–H, 10A, 10C–D); halter well separate from hind leg (Figs. 32C–D); prothoracic extension not present (Figs. 23C–E); with, at most, 1 anterolateral seta on the mesonotum; mesonotum with 2 dorsal setae and 1–2 campaniform sensilla (Figs. 29D–F); male genital lobes dorsal (Figs. 9C–H) or ventral to terminal processes; abdomen free or partially encased by larval exuviae (Fig. 11F).................................... FORCIPOMYIINAE............ 5 (worldwide) - Mesonotum smooth, with bumps, or with apically rounded tubercles at most about as long as wide (Fig. 33A); halter and hind leg closely approximated (Figs. G–I) or abutting (Figs. 32J–L, 33A–L); prothoracic extension well-developed (Figs. 24A–C, F–G), although in some it is restricted to abutting either the antenna (Figs. 24D–E) or palpus (Figs. 26C–E) (difficult to see in some slide preparations of Culicoides and Paradasyhelea but these with 2 well-developed anterolateral setae on the mesonotum and mesonotum with 4 dorsal setae and a campaniform sensillum (Figs. 29K–M)); mesonotum with at least 3 dorsal setae and 1–2 campaniform sensilla (Figs. 29N–T, 30A–T); male genital lobes ventral to terminal processes; abdomen without attached larval exuviae........................... CERATOPOGONINAE............ 17 (worldwide) 5. Terminal process with elongate and well-developed lateral seta (Figs. 10E, 72I); mesonotum smooth or with very small, short tubercles (Figs. 10E, 29F)............. Forcipomyia (in part) (F. (Phytohelea)) (pg. 40) (worldwide except for Nearctic Region) - Terminal process with, at most, a short lateral seta; mesonotum with tubercles variable, from very short (Figs. 8D–E, 8H, 10E–G) to very elongate (Figs. 9A–H, 10A, 10C–D).............................6 6. Terminal processes each tapering from thick base to pointed or somewhat rounded apex, with bases abutting medially, each either bare or with spicules directed posteriorly or with very fine, anteriorly directed spicules restricted to the very base of the terminal process (latter only in a few species in the subgenus F. (Forcipomyia) (Figs. 72F–G)................................. Forcipomyia (in part) (pg. 40)............ 7 (worldwide) - Terminal processes each nearly cylindrical for most of its length in dorsoventral view, with the bases well separated medially (with a medial truncate, nearly or completely transverse area between the bases) (Figs. 11D–E), each with strong, anteriorly directed spicules at least at apex (Fig. 72J)..... Atrichopogon (pg. 43) (worldwide) 7. Dorsum of head and mesonotum with very elongate slender setae arising directly from cuticle surface (no tubercles) (Figs. 9E).......................................... Forcipomyia (Microhelea) (pg. 40) (worldwide) - Dorsum of head and mesonotum with or without elongate tubercles but never with very elongate slender setae arising directly from cuticle surface (e.g. Figs. 9A, 9F, 10A–D)..............................8 8. Respiratory organ with elongate pores arranged in a round semicircle at its apex (Figs. 42E–F)................................................................. Forcipomyia (Euprojoannisia) (pg. 40) (worldwide) - Respiratory organ with in either a single row of round or somewhat oblong pores or with these undulating somewhat (Figs. 42G–M, O–R)............................................................9 9. Larval exuviae not retained on pupal abdomen...............................................10 - Larval exuviae retained on pupal abdomen..................................................12 10. Posteromedial protrusion of mesonotum extending posteriorly to tergite 2 (Figs. 9C –D); male genital lobes dorsal to terminal processes (Figs. 9C–D).......................... Forcipomyia (Warmkea) (pg. 40) (worldwide) - Posteromedial protrusion of mesonotum not extending posteriorly to tergite 2 (Figs. 10H, 11A); male genital lobes ventral to terminal processes (Figs. 10H, 11A)...........................................11 11. Abdomen with elongate lateral setae (Fig. 11B); terminal process elongate, relatively slender (Fig. 11B)............................................................. Forcipomyia (Pterobosca) (pg. 40) (worldwide except for Australia) - Abdomen with, at most, short lateral setae (Figs. 10H, 11A); terminal process short, relatively blunt (Figs. 10H, 11A)..................... Forcipomyia (Thyridomyia) or Forcipomyia (Synthyridomyia) (pg. 40) (both worldwide) 12. Terminal process short and directed laterally to posterolaterally (Figs. 10F–G, 11C); male genital lobes ventral to terminal processes................................................................13 - Terminal process elongate and directed posteriorly (Figs. 8G–H, 9A–B, F–H, 10A–D); male genital lobes dorsal to terminal processes (not known for F. (Metaforcipomyia))...............................14 13. Abdomen with at most, very low tubercles (Fig. 10G).............. Forcipomyia (Trichohelea) (pg. 40) (worldwide) - Abdomen with well-developed but short tubercles (Fig. 11C).......... Forcipomyia (Lasiohelea) (pg. 40) (worldwide) 14. Mesonotum with 6–7 pairs of elongate tubercles (Figs. 9A, 10D)............................................................... Forcipomyia (Lepidohelea) + Forcipomyia (Schizoforcipomyia) (pg. 40) (both worldwide) - Mesonotum with no more than five pairs of elongate tubercles (Figs. 8G–H, 9B, 9F–H, 10A, 10C)......15 15. Respiratory organ more or less slender (Figs. 42K–L) to markedly swollen apically (Fig. 42M); mesonotum with 0–4 pairs of elongate tubercles, if only 1–2 present, then the base of each relatively slender, not broadly conical (Figs. 9F–H, 10A, 10C); abdominal segments with very short to elongate tubercles (Figs. 9F–H, 10A–C).................................................. Forcipomyia (Forcipomyia) (pg. 40) (worldwide) - Respiratory organ more or less slender (Fig. 42J) to slightly swollen apically (Figs. 42H–I); mesonotum with 1–2 pairs of elongate tubercles, each with a broadly conical broad base and an elongate seta; abdominal segments with at most very short tubercles (Figs. 8G–H, 9B)......................................16 16. Mesonotum with, at most, 2 pairs of tubercles (Figs. 8G–H); respiratory organ with pores restricted to apical 0.29–0.35 of its length (Figs. 42H–I)........................ Forcipomyia (Caloforcipomyia) (pg. 40) (worldwide) - Mesonotum with 4 pairs of tubercles (Fig. 9B); respiratory organ with pores extending basally to basal 0.50 of its length (Fig. 42J).................................... Forcipomyia (Metaforcipomyia) (pg. 40) (worldwide) 17. With prothoracic extension extending from palpus to antenna (Figs. 24A–B) (difficult to see in some slide preparations), apex of halter narrowly separated from or barely touching hind leg in lateral view (Fig. 32G) and cephalic apotome lacking central dome bearing sensilla (Figs. 19A–G).........................18 - With prothoracic extension either extending from palpus to antenna (e.g. Figs. 24C, F–G) or only extending for part of this area (e.g. Figs. 24D–E, 28A–C), apex of halter broadly touching hind leg in lateral view (Figs. 32J–L, 33A–L), or if narrowly separated from or barely touching hind leg in lateral view (Fig. 32H) then cephalic apotome with central dome bearing sensilla (Figs. 19H–J)...........................19 18. Dorsal apotome with or without spicules but if with lateral row of stout, pointed spicules then also with further stout spicules more medially (Figs. 19A–D); respiratory organ with or without spicules or shagreen, with or without annulations at near midlength, 0–4 pores scattered along length with the remainder grouped apically (Figs. 43H–N).................................................... Culicoides (pg. 49) (worldwide, except for New Zealand) - Dorsal apotome with stout, pointed spicules restricted to a dorsolateral row, in some this row short, with only 3 spicules (Figs. 19E–G); respiratory organ bare, without spicules or shagreen (not to be confused with annulations along its length), with annulations at least near midlength (some with more extensive annulations), 1–3 pores scattered along length with the remainder grouped apically (Fig. 43O).................................................................................... Paradasyhelea (pg. 52) (Australia, New Caledonia, New Zealand, southern South America, Olympic Peninsula in Washington, USA) 19. Abdominal segment 8 with D-3-VIII (Fig. 12H); female with wings abutting posteroventrally (Figs. 12H, 41F)............................................................... Paryphoconus (pg. 112) (Neotropical and southern Nearctic Regions) - Abdominal segment 8 without D-3-VIII (Macropeza with anterior L-1-VIII, Fig. 12C); female with wings separated posteroventrally by forelegs (e.g. Figs. 12A, 12D, 12F, 41A–E)..........................20 20. With at least abdominal sternites 6 and 7 each with very thin membranous disc (Fig. 12D).............21 - Abdominal sternites without any thin membranous areas (e.g. Figs. 3C, 12F, 12H)...................37 21. Sternites 4–8 each with a narrow membranous disc; dorsolateral cephalic sclerite seta very short; one very short anteromedial setae and an associated campaniform sensillum; one clypeal-labral seta present as a very small, short, blunt peg (Fig. 25H)................ Parabezzia (in part, alexanderi species group) (pg. 72) (New World, Palaearctic (only in north Africa) and Afrotropical Regions) - With, at most, sternites 4–7 each with a narrow or broad membranous disc; dorsolateral cephalic sclerite seta elongate; one short and one very elongate anteromedial seta, with the elongate seta extending to or toward respiratory organ (Figs. 31N), in some with an additional associated campaniform sensillum; with two thick, stout clypeal labral setae (Figs. 26G–H, 27A–C)..............................................22 22. Abdominal sternites 5–7 each with membranous disc bearing a posteromedial dark pigment spot (present only in N. schwarzii); mesonotum with D-1-T, D-2-T elongate and slender (Fig. 30Q); respiratory organ elongate (Figs. 45O–P)............................................. Nilobezzia (in part) (pg. 88) (worldwide) - Abdominal sternites 4–7, 5–7, or 6–7 each with membranous disc and without a posteromedial dark pigment spot on the membrane, or if spot present, only abdominal sternites 6–7 with membranous discs; mesonotum with D-1-T, D-2-T short and blunt (Figs. 30L–P); respiratory organ squat to elongate (Figs. 45I–N).....23 23. Abdominal sternites 4–7 or 5–7 each with membranous disc or, if only sternites 6–7 with membrane then these each with a posteromedial dark pigment spot............................................24 - With each of abdominal sternites 6–7 with membranous disc and without a posteromedial dark pigment spot on membrane.........................................................................27 24. With only abdominal sternites 6–7 each with membranous disc.............. Probezzia (in part) (pg. 85) (Holarctic Region) - Abdominal sternites 4–7 or 5–7 each with membranous disc....................................25 25. Abdominal sternite 4 with at least V-5-IV and V-6-IV on well-developed stout, pointed tubercles (V-6-IV shorter than V-5-IV); V-7-IV on a very short rounded to small pointed tubercle (Fig. 66B)............................................................................... Probezzia (in part) (pg. 85) (Holarctic Region) - Abdominal sternite 4 with V-5-IV on well-developed stout tubercle, contrasting with very short tubercles bearing each of V-6-IV and V-7-IV (Figs. 65A, 66A)..........................................26 26. With either abdominal sternites 4–7 or 5–7 each with membranous disc but with membrane on 4 and/or 5 not as thinly developed as in 6–7 and with arrangement of V-5, V-6, V-7 setae of sternites 4–5 well-developed, not clumped and reduced as they are on sternites 6–7 (Fig. 65A)................................................................................... Jenkinshelea (in part; J. magnipennis) (pg. 81) (Nearctic) - With abdominal segments 5–7 each with membranous disc and arrangement of V-5, V-6, V-7 setae similar on each sternite (Fig. 66A)................ Mallochohelea (in part; M. inermis, M. nr. caudelli) (pg. 84) (Nearctic) 27. Nearctic taxa..........................................................................28 - From all regions (including Nearctic).......................................................31 28. Campaniform sensillum D-4-IV situated between (Fig. 65C) or medial to D-8-IV and D-9-IV(Fig. 65B); L- 1-VIII present (Fig. 12C) (based on species outside of Nearctic Region)............. Macropeza (pg. 82) (eastern Nearctic) - Campaniform sensillum D-4-IV medial to D-8-IV and D-9-IV (e.g. Figs. 65A, 66A); L-1-VIII absent (e.g. Fig. 12G).............................................................................29 29. Abdominal tergite 1 with anterior campaniform sensillum D-7-I situated near lateral tubercle (Fig. 51C)................................................................ Jenkinshelea (in part) (pg. 81) - Abdominal tergite 1 with anterior campaniform sensillum D-7-I situated near D-2-I and D-3-I (Figs. 51E–F)....................................................................................30 30. Abdominal sternite 4 with V-5-IV on well-developed stout tubercle, contrasting with very short tubercles bearing each of V-6-IVand V-7-IV (Fig. 66A)........................ Mallochohelea (in part) (pg. 84) - Abdominal sternite 4 with each of V-5-IV, V-6-IV and V-7-IV on well-developed stout, pointed tubercles (of increasing size) (similar to Fig. 66B)....................... Probezzia (in part; P. fuscipennis) (pg. 85) (eastern Nearctic) 31. Abdominal sternite 4 with V-5-IV on well-developed, stout tubercle, contrasting with very short tubercles bearing each of V-6-IV and V-7-IV (Fig. 65A)...............................................32 - Abdominal sternite 4 with each of V-5-IV, V-6-IVand V-7-IV on well-developed stout, pointed tubercles (e.g. Figs. 66 B-C)......................................................................34 32. Abdominal sternite 4 with V-5-IV on bifid tubercle (as in Fig. 65C)...... Macropeza (M. bayeri) (pg. 82) (Afrotropical) - Abdominal sternite 4 with V-5-IV on pointed tubercle (Figs. 65A, 66A)...........................33 33. Abdominal tergite 1 with campaniform sensillum D-7-I situated near L-1-I and L-2-I (Fig. 51C)....................................................................... Jenkinshelea (in part) (pg. 81) (Nearctic species) - Abdominal tergite 1 with campaniform sensillum D-7-I situated near D-2-I and D-3-I (Fig. 51E)..................................................................... Mallochohelea (in part) (pg. 84) (worldwide) 34. Abdominal tergites 1–7 with D-2 and D-5 on short, sclerotized tubercles, each of these tubercles arising directly from surrounding cuticle (Fig. 66C).................................................35 - Abdominal tergites 1–7 with D-2 and D-5 each on a well-developed, sclerotized protuberance arising from large, swollen, unpigmented base (Figs. 65B)................................................36 35. Abdominal segment 4 with L-1-IV an elongate seta on a moderately sized, pointed tubercle (Fig. 66C)........................................................................... Neobezzia (pg. 87) (Neotropical Region) - Abdominal segment 4 with L-1-IV a short, thick seta on a small, apically rounded tubercle (as in Fig. 66B).......................................... Probezzia (in part; P. fuscipennis, one paratype) (pg. 85) (eastern Nearctic) 36. Swollen base at each of D-2 and D-5 on each of abdominal tergites 1–7 smooth, without spicules; abdominal segment 4 with L-2-IV on bifid tubercle; abdominal tergite 1 with campaniform sensillum D-7-I situated near lateral tubercle (Figs. 51D)................................ Macropeza (M. natalensis) (pg. 82) (Afrotropical) - Swollen base at each of D-2 and D-5 on abdominal tergites 1-7 with well-developed, rounded spicules (making them look bumpy); abdominal segment 4 with L-2-IV on pointed tubercle; abdominal tergite 1 with anterior campaniform sensillum situated near D-2-I, D-3-1 (as in Fig. 51E)....................................................................................... Jenkinshelea (J. polyxenae) (pg. 81) (Afrotropical) 37. 0–2 anteromedial setae on anterior margin of mesonotum very short, none extending to or long enough to extend to respiratory organ; exuviae with dorsolateral cephalic sclerite either detached from anterior margin of mesonotum (Figs. 13C–E) or fused with it (Figs. 13F–I).....................................38 - 2 anteromedial setae on anterior margin of mesonotum with either one or both elongate, extending to respiratory organ (Figs. 31K–N); exuviae with dorsolateral cephalic sclerite fused to anterior margin of mesonotum (Figs. 13F–I)......................................................................50 38. Mesonotum with at least one pair of large, rounded tubercles (Fig. 33A)...........................39 - Mesonotum with, at most, low, relatively flat bumps (Figs. 32K, 33D–G)..........................42 39. Lateral margin of foreleg with angular bend posterolaterally from tibial apex (Fig. 37B); antenna extending to or posterior to apex of anterior portion of midleg (Figs. 37B); abdominal sternite 4 with only 2 ventral posterior setae (V-5-IV, V-6-IV) (Figs. 62B–C)........................................................................ Parabezzia (in part: those other than in the alexanderi species group) (pg. 72) (worldwide except for Palaearctic and Australasian Regions) - Lateral margin of foreleg evenly curved posterolaterally from tibial apex (Figs. 36E–F, 37A); antenna apex anterior to or extending to apex of anterior portion of midleg (Figs. 36E–F, 37A); abdominal sternite 4 with 3 ventral posterior setae (V-5-IV, V-6-IV, V-7-IV) (Figs. 61A–C, 62A)............................40 40. Respiratory organ with separate pores near midlength, in addition to those at apex, most of surface of internal tracheal tube appearing as abutting, lumpy scales (Fig. 44O).................... Allohelea (pg. 68) (worldwide) - Respiratory organ with all pores grouped at apex (Figs. 44P–Q), surface of internal tracheal tube smooth between spiraled base and pore tubes (Fig. 44P) or with spirals extending ¾ length of tracheal tube.....41 41. Abdominal segment 4 with L-1-IV dorsal to L-3-IV (Figs. 61 B-C)................. Monohelea (pg. 69) (worldwide) - Abdominal segment 4 with L-1-IV anterior to L-3-IV (Figs. 61 B-C)............... Atyphohelea (pg. 71) (Nearctic) 42. Head with prothoracic extension restricted to dorsolateral area and abutting antenna, not extending to palp (Figs. 24D–E).........................................................................43 - Head with prothoracic extension extending from antenna to palp (Figs. 24C, 24F–G, 25A–D)..........44 43. Mesonotum with dorsal setae D-1-T, D-2-T, D-4-T, D-5-T clumped on single raised area (Fig. 29O); respiratory organ with many annulations, with single pore near base and with remaining pores tightly grouped at very apex in a semicircle (Fig. 43R); halter and hind leg slightly separate and nearly abutting at anterior margin of tergite 2 (Fig. 32I)................................................ Brachypogon (pg.55) (worldwide) - Mesonotum with dorsal setae D-1-T, D-2-T, D-4-T, D-5-T separate from one another and especially seta D- 5-T well posterior of D-1-T, D-2-T, at most with each seta on a separate, very short tubercle (Fig. 29P); respiratory organ without annulations, with or without separate pores along length and with remaining pores either tightly grouped at apex (as above) or, in most species, with a distinctive pair of tightly appressed rows of wide pores, with these rows extending from the apex more basally along length of respiratory organ (Figs. 43S–T, 44A–B); halter and hind leg broadly abutting, well posterior of anterior margin of tergite 2 (Fig. 32J).................................................................... Alluaudomyia (pg. 57) (worldwide) 44. Dorsal apotome with central dome bearing both dorsal apotome setae (Figs. 19H–J); respiratory organ with apex of tracheal tube J-shaped (Figs. 43P–Q); with most abdominal setae on bifid tubercles, in some with these tubercles greatly extending beyond length of seta, forming a pair of slender, elongate spicules (Figs. 57C, 58A); halter and hind leg separate (Fig. 32H)............................ Ceratopogon (pg. 54) (Holarctic Region) - Dorsal apotome without central dome, each dorsal apotome seta arising from separate low or well-developed tubercle (Figs. 19N–P, 20A–G, 20M–N); respiratory organ with apex of tracheal tube more or less straight or divided equally into pore tubes (Figs. 44C–N, 45A–B); with or without most abdominal setae on bifid tubercles (Figs. 59B, 60A), never with tubercles forming a pair of elongate spicules; halter and hind leg broadly abutting (Figs. 32K–L, 33B)..............................................................45 45. Respiratory organ with numerous annulations and a separate, short, expanded apical portion bearing pores in a single plane (a single pore is also near the base of the respiratory organ) (Fig. 44C); without mandible (Fig. 24F); apex of foreleg far anterior to apex of midleg (Fig. 35E); total length = 0.91–1.19 mm............................................................................ Baeodasymyia (pg. 59) (Neotropical Region) - Respiratory organ without annulations and surface smooth, forming a single uniform structure (Figs. 44 D- N); with mandible (Figs. 24G, 25 A-D); apex of foreleg close to that of midleg or, at most, moderately anterior (Figs. 35F, 36 A-D); total length = 1.19-4.47..............................................46 46. Exuviae with dorsolateral cephalic sclerite fused to anterior margin of mesonotum (Fig. 13F); mesonotum with 4 dorsal setae and campaniform sensillum (D-3-T) (Figs. 30A, 30G); respiratory organ with all pores grouped at apex (Figs. 44N, 45 A-B).......................................................47 - Exuviae with dorsolateral cephalic sclerite detached from anterior margin of mesonotum (as in Figs. 13 B-E); mesonotum with 3 dorsal setae (D-5-T absent) and campaniform sensillum (D-3-T) (Figs. 29R–T); respiratory organ with all pores grouped at apex or with additional pores scattered along length (Figs. 44D–M).......................................................................................48 47. Dorsal apotome with seta very short, barely longer than width of basal socket (Fig. 20G); abdominal segment 4 with short setae on rounded or, at most, very slightly bilobed tubercles with each lobe rounded (Fig. 60C); metathorax with 1 small seta (M-1-T) and 2 campaniform sensilla (M-2-T, M-3-T) (Fig. 49D); face without clypeal-labral setae (Fig. 25D)........................................ Serromyia (pg. 67) (worldwide except Neotropical Region) - Dorsal apotome with seta elongate, longer than distance from base of seta to lateral margin of dorsal apotome (Figs. 20 M-N); abdominal segment 4 with at least setae D-5-IV, L-2-IV, and V-5-IV more elongate and on pointed, bifid tubercles (Fig. 63B); metathorax with only 2 campaniform sensilla (M-2-T, M-3-T) (Fig. 50D); face with two elongate clypeal-labral setae (Fig. 26B)............... Clinohelea (in part) (pg. 75) (worldwide) 48. Apex of hind leg ventral to apex of midleg (Fig. 35F) (perhaps applying only female, male unknown); apex of antenna far anterior to apex of medial portion of midleg (Fig. 35F).............. Austrohelea (pg. 60) (Australia, New Zealand) - Apex of hind leg lateral to apex of midleg (Figs. 36A–C); apex of antenna just anterior, equal, or posterior to apex of medial portion of midleg (Figs. 36A–C)..............................................49 49. Mesonotum with dorsal setae D-1-T, D-2-T, D-4-T and campaniform sensillum D-3-T close to one another, arising directly from cuticle or on very small, round tubercles (Fig. 29S); abdominal segment 4 with each dorsal sensillum on a tubercle with a single rounded, pointed or bifid apex (Figs. 59C, 60A)................................................................................... Stilobezzia (pg. 61) (worldwide) - Mesonotum with dorsal setae D-1-T, D-2-T, D-4-T and campaniform sensillum D-3-T distant from one another, D-1-T much thicker than others and on moderately sized tubercle larger than those of D-2-T and D- 4-T (Fig. 29T); abdominal segment 4 with each dorsal sensillum on a comb-like tubercle (Fig. 60B)......................................................................... Schizonyxhelea (pg. 64) (New World and Oriental Region) 50. Mesonotum projecting posteromedially over metathorax (Fig. 50C); abdominal segment 4 with most sensilla on strongly bifid tubercles with each half slender and elongate (Fig. 63A)........... Echinohelea (pg. 74) (worldwide except for the Palaearctic Region) - Mesonotum not projecting posteromedially over metathorax (Figs. 50E–F, 51A–F); abdominal segment 4 with sensilla on, at most, short, shallowly bifid tubercles (e.g. Figs. 63B, 65C)......................51 51. Cephalothorax much wider than slender abdominal segments 3–4 (Fig. 12A); apex of respiratory organ with pores arranged in a nearly perfect circle and separated by a circular, cuticular disc arising from the apicomedial margin of the respiratory organ (Fig. 45F); shape of abdominal segment 9 distinctive, with terminal processes moderately elongate, pointing nearly posteriorly and with their bases separated by broad, nearly transverse area (Figs. 75G–H)........................................... Pellucidomyia (pg. 77) (Neotropical, Afrotropical and Australasian Regions) - Cephalothorax not much wider than abdominal segments 3 and 4 (Figs. 12B–G); apex of respiratory organ with pores arranged either in a linear group or, at most, in a semicircular arrangement and with an oblong cuticular area arising from the apicomedial area of the respiratory organ (Figs. 45G–M, 45O–T, 46A–F, 46H–S); abdominal segment 9 with terminal processes pointing laterally to nearly posteriorly and with their bases separated by narrow v- or u-shaped area, if with bases separated by broad, nearly transverse area then terminal process very slender, elongate and directed posterolaterally or short and directed laterally (Figs. 75D–F, 75I–L, 76A–E, 76G–L, 77A–L)....................................................52 52. Segment 9 with terminal process very slender, nearly equal in diameter for most of its length and with dorsobasal seta extending nearly to the apex of the terminal process (Fig. 75I); surface of respiratory organ with very fine spicules arising from very fine ridges (giving the appearance of a somewhat reticulated surface) (Fig. 45G)................................................................ Dibezzia (pg. 80) (Afrotropical and Oriental Regions) - Segment 9 with terminal process very slender and nearly equal in diameter for most of its length to tapering from thick base, without seta (Figs. 75D–F, 75J–L, 76A–L, 77A–L); surface of respiratory organ without any spicules, either smooth or with, at most, light wrinkles (Figs. 45H–T, 46A–F, 46H–S).............53 53. Abdominal segment 8 with L-1-VIII seta on pointed or bifid tubercle, in addition to transverse arrangement of more posteriorly placed setae (so in dorsoventral view there are two laterally projecting tubercles, one anterior to the other) (Fig. 12C); terminal process directed laterally (Figs. 76A–B); abdominal segment 4 with campaniform sensillum D-4-IV between or just anterior to space between D-8-IV and D-9-IV (Figs. 65C).................. Macropeza (in part; M. albitarsis and two unnamed species from Laos) (pg. 82) (Palaearctic and Oriental Regions) - Abdominal segment 8 without L-1-VIII seta, with only transverse arrangement of posterior setae (so in dorsoventral view there is one laterally projecting tubercle or dorsoventrally arranged group of tubercles) (Figs. 12E–G); terminal process directed laterally to posteriorly (Figs. 76G–L, 77A–L); abdominal segment 4 with campaniform sensillum D-4-IV medial to D-8-IV and D-9-IV (e.g. Figs. 63B, 67A–C, 68A–C)........54 54. Dorsal apotome with two pairs of tubercles, well separated dorsoventrally (Fig. 21L); abdominal segment 4 with L-1-IV and L-3-IV more or less dorsoventral to one another so that laterally there are two pairs of setae, an anterior pair (L-1-IV and L-3-IV) and a posterior pair (L-2-IV and L-4-IV) (Fig. 68A)..................................................................................... Anebomyia (pg. 92) (Nearctic, Afrotropical and Oriental Regions) - Dorsal apotome with only 1 pair of tubercles (Figs. 21H–K, 22A–K); abdominal segment 4 with L-1-IV anterior to group of L-2-IV, L-3-IV and L-4-IV (Figs. 63B, 69 A-C, 70A, 70C) or with L-1-IV and L-3-IV more or less dorsoventral to one another so that laterally there are two pairs of setae, an anterior pair (L-1-IV and L-3-IV) and a posterior pair (L-2-IV and L-4-IV) but if so, the anterior sensilla are situated dorsoventrally to posterior pair (Fig. 70B)..........................................................55 55. Mesonotum with anteromedial setae (those directly anterior to base of respiratory organ) with one very elongate and the other very short, less than 0.20 the length of the long seta and, in some a peg (Fig. 31N); mesonotum with each of D1-T, D-2-T, D-5-T either a peg, a short thick seta, or elongate (Figs. 30G–H, 30O, 30Q, 30L)............................................................................56 - Mesonotum with anteromedial setae both very elongate, generally equal in length but in some with one shorter than the other but if so, the shorter is more than 0.75 the length of the long seta (Figs. 31L–M); mesonotum with each of D1-T, D-2-T, D-5-T an elongate seta (Figs. 30K, 30R–S, 31A–I).............60 56. Abdominal segment 9 with terminal process very long, slender, directed either posteriorly or very nearly posteriorly, with apex hooked somewhat medially (Figs. 75D–F); width of forefemur very wide compared to width of foretibia; exuviae with ecdysial split extending from ventrolateral margin of dorsal apotome posteriorly as a strongly sinuous curve a short distance on eye and with separation of lateral portion of mouthparts and prothoracic extension from foreleg, so that face remains connected to rest of exuviae only at the base of antenna, the labium, and medial surface of the palp (Fig. 79G).................... Heteromyia (pg. 76) (New World) - Abdominal segment 9 with terminal process at most moderately elongate, directed laterally to posterolaterally, with, for those with moderately elongate terminal process, apex evenly curved posteriorly or curved somewhat laterally (Figs. 75K–L, 76G); width of forefemur not much wider than width of foretibia; exuviae with ecdysial split extending from ventrolateral margin of dorsal apotome posteriorly as a sinuous curve far on eye and with remaining portion of face yet connected to the rest of the exuviae (Fig. 79I), with the ecdysial split extending from the dorsal apotome just to the base of the antenna (Fig. 79E) or with the ecdysial split extending from ventrolateral margin of dorsal apotome posterolaterally around base of antenna, laterally along face and, in some, including mouthparts, so that the entire face remains attached to the rest of the exuviae only through a connection at the mouthparts (Figs. 79H).................................57 57. Dorsal apotome elongate, narrow (Figs. 20M–N); metathorax with two campaniform sensilla near its anterior margin (M-2-T, M-3-T); exuviae with ecdysial split extending from the dorsal apotome just to the base of the antenna (Fig. 79E)........................................... Clinohelea (in part) (pg. 75) (worldwide) - Dorsal apotome wide (Figs. 21C, F, H–I); metathorax with one campaniform sensillum well posterior from its anterior margin (M-2-T, M-3-T); exuviae with ecdysial split extending from ventrolateral margin of dorsal apotome posteriorly as a sinuous curve far on eye and with remaining portion of face yet connected to the rest of the exuviae (Fig. 79I)or with the ecdysial split extending from ventrolateral margin of dorsal apotome posterolaterally around base of antenna, laterally along face and to and, in some, including mouthparts, so that the entire face remains attached to the rest of the exuviae only through a connection at the mouthparts (Figs. 79H)...........................................................................58 58. Respiratory organ with basal annulations (some species with these only at the very base) (Fig. 45P); male with very short genital lobes, each slightly longer than wide and apex anterior or to level of base of terminal process (as in Figs. 78I–J); exuviae with the ecdysial split extending from ventrolateral margin of dorsal apotome posterolaterally around base of antenna, laterally along face and to and, in some, including mouthparts, so that the entire face remains attached to the rest of the exuviae through a connection at the mouthparts (consequently, the anterior margin of the face has a characteristic and bilaterally symmetrical margin, with a concave anterolateral excavation where the base of the antenna otherwise fits) (Fig. 79H)......................................................................... Nilobezzia (in part) (pg. 88) (worldwide) - Respiratory organ without basal annulations (Figs. 45I, 45L–M); male with elongate genital lobes, each longer than wide and apex extending posteriorly to near posteromedial gap between terminal processes (Figs. 75K, 76E); exuviae with ecdysial split from ventrolateral margin of dorsal apotome extending posteriorly as a sinuous curve far on eye and with remaining portion face yet connected to the rest of the exuviae (Figs. 79I).................................................................................59 59. Apex of antenna anterior, equal or just slightly posterior to apex of middle portion of midleg (Fig. 39A).................................................................... Probezzia (in part) (pg. 85) (Holarctic Region) - Apex of antennal well posterior to apex of middle portion of midleg (as in Fig. 38E)......................................................... Jenkinshelea (in part; J. rhodesiensis, J. tokunagai) (pg. 81) (Afrotropical and Oriental Regions) 60. Nearctic taxa..........................................................................61 - Worldwide taxa (including Nearctic)...................................................... 77 61. Apex of foreleg extending to or posterior to apex of wing and ventral to apex of midleg (Fig. 40E)...................................................................... female Sphaeromias (pg. 97) - Apex of foreleg well anterior to apex of wing and apex of midleg (Figs. 41A, 41C–E)................62 62. Apex of halter lateral or anterior to anterior margin of tergite 2 (as in Figs. 33B, 33D–E); abdominal segment 4 with all lateral setae (L-1-IV, L-2-IV, L-3-IV, L-4-IV) in a transverse row at about anterior 0.3-0.4 of segment and anterior to D-8-IV, D-9-IV, V-7-IV, giving the appearance of two sets of lateral tubercles in dorsoventral view, one anterior to the other (Fig. 67B).......................... Johannsenomyia (pg. 90) - Apex of halter posterior to anterior margin of tergite 2 (Figs. 33I–L); abdominal segment 4 with L-1-IV seta anterior to L-2-IV, L-3-IV, L-4-IV, which are positioned at or posterior to midlength of segment and more or less anteroventral to D-8-IV, D-9-IV and anterodorsal to V-7-IV, giving the appearance of one set of lateral tubercles in dorsoventral view (Figs. 69A, 69C, 70B–C)........................................63 63. Very large body, with total length = 6.63–8.04 mm; prothoracic extension restricted to medial portion, not extending laterally to antenna (Figs. 28B, 28F); clypeal-labral setae well anterior to level of rounded area joining mandible and palp (Figs. 28B, 28F); abdominal segments 3–7 without ventral creeping welt or, if slightly present, then covered with spicules in a pattern continuous with surrounding cuticle...........64 - Small to large body, with total length = 2.00– 7.03 mm; prothoracic extension extending laterally to antenna, in some only as a narrow band (Figs. 28D, 28G–H); clypeal-labral setae slightly anterior to posterior to level of rounded area joining mandible and palp (Figs. 28D, 28G–H), well anterior only in Palpomyia novitibialis which has V-5 and V-6 on a transverse, narrow, raised welt on each of segments 3–7 and which is lacking spicules..............................................................................65 64. Palp short, not protruding much beyond posterolateral margin of labium (Fig. 28B); V-5-IV, V-6-IV, V-7-IV grouped closely together (Fig. 69A)................................... male Sphaeromias (pg. 97) - Palp elongate, protruding well beyond posterolateral margin of labium (Fig. 28F); V-7-IV closer to L-4-IV than V-5-IV and V-6-IV, which are closely abutting (Fig. 70B)................... Pachyhelea (pg. 105) 65. Segment 4 with D-3-IV lateral to D-2-IV (Fig. 71A)..................... Palpomyia (in part) (pg. 106) - Segment 4 with D-3-IV posterolateral to D-2-IV (Figs. 69C, 71B)................................66 66. Male................................................................................67 - Female...............................................................................72 67. Genital lobes very short, each slightly longer than wide and apex extending posteriorly to about level of lateral base of terminal processes (Figs. 78I–J)................................ Phaenobezzia (pg. 110) - Genital lobes elongate, each longer than wide and apex extending posteriorly to near posteromedial gap between terminal processes (Figs. 77E, 78A, 78C, 78E, 78G)...................................68 68. Dorsal apotome with at least one seta and two campaniform sensilla (Figs. 22C–D, 22J)..............69 - Dorsal apotome with only one seta and one campaniform sensillum (Figs. 22E, 22H–I)...............71 69. Segment 4 with L-2-IV, L-3-IV, L-4-IV each on a single tubercle (Fig. 69C); V-7-IV situated far lateral to the closely paired V-5-IV and V-6-IV (Fig. 69C)....................... Bezzia (Bezzia) (in part) (pg. 100) - Segment 4 with L-3-IV situated basally or subbasally on same tubercle as L-2-IV (so there are only two prominent pointed posterolateral tubercles) (Fig. 70C); V-5-IV, V-6-IV and V-7-IV forming a single closely gathered group (Fig. 70C)................................................................70 70. Dorsal apotome with two setae (and additional campaniform sensilla)..................................................................................... Palpomyia (in part; P. jonesi) (pg. 106) - Dorsal apotome with one seta (and additional campaniform sensilla) (Figs. 22D, 22J)................................. Bezzia (Homobezzia) (in part; B. (Bezzia) gibbera, Palpomyia flaviceps) ( pgs. 100, 106) 71. Abdominal segment 4 with D-5-IV arising directly from surrounding cuticle; each tubercle bearing setae D- 8-IV and D-9-IV elongate, thick, with strong apical point (Fig. 70C)...................................................................................... Palpomyia (in part; P. lineata) (pg. 106) - Abdominal segment 4 with D-5-IV on a short, round tubercle; each tubercle bearing setae D-8-IV and D-9- IV similar in size, each short, apically rounded or, at most, with tubercle bearing D-9-IV with very short apical point (Fig. 69C)...................................... Bezzia (in part; B. dorsasetula) (pg. 100) 72. Dorsal apotome with at least one seta and two campaniform sensilla (Figs. 22C–D, 22J)..............73 - Dorsal apotome with only one seta and one campaniform sensillum (Figs. 22E, 22H–I)...............75 73. Segment 4 with L-2-IV, L-3-IV, L-4-IV each on a single tubercle; V-7-IV situated far lateral to the closely paired V-5-IV and V-6-IV...................................... Bezzia (Bezzia) (in part) (pg. 100) - Segment 4 with L-3-IV situated basally or subbasally on same tubercle as L-2-IV (so there are only two prominent pointed posterolateral tubercles); V-5-IV, V-6-IV and V-7-IV forming a single closely gathered group................................................................................74 74. Dorsal apotome with two setae (and additional campaniform sensilla)..................................................................................... Palpomyia (in part; P. jonesi) (pg. 106) - Dorsal apotome with one seta (and additional campaniform sensilla) (Figs. 22E, 22H–I)............................... Bezzia (Homobezzia) (in part: B. (Bezzia) gibbera , Palpomyia flaviceps) (pgs. 100, 106) 75. Area between terminal processes truncated or nearly so (Fig. 78K).............. Phaenobezzia (pg. 110) - Area between terminal processes u-shaped (Fig. 78B).........................................76 76. Abdominal segment 4 with setae D-2-IV, D-3-IV, D-8-IV and D-9-IV on large pointed tubercles, D-5-IV arising directly from surrounding cuticle (as in Fig. 70C)......... Palpomyia (in part; P. lineata) (pg. 106) - Abdominal segment 4 with setae D-2-IV, D-3-IV, D-5-IV,D-8-IV and D-9-IV on small, rounded tubercles (as in Fig. 69C)............................................ Bezzia (in part; B. dorsasetula) (pg. 100) 77. Apex of foreleg ventral to apex of midleg or, at most, with just a wider than long portion of the midleg visible in ventral view (Fig. 40E).............................................................78 - Apex of foreleg well anterior to apex of midleg so that visible portion of midleg longer than wide in ventral view (Figs. 39D–E, 41A–E) (The presently unknown males of Leehelea and Xenohelea are keyed through couplet 81, assuming they have the same sexual dimorphism as is present in Sphaeromias, in which females have long and males short forelegs. If these two genera are not dimorphic they will likely successfully key to their females below)....................................................................81 78. Abdominal segment 4 with lateral setae (L-1-IV, L-2-IV, L-3-IV, L-4-IV) on tubercles on anterior half of segment and with D-8-IV and D-9-IV situated more laterally, giving the appearance of two sets of lateral tubercles in dorsoventral view (Fig. 67B)................... female Johannsenomyia (in part) (pg. 90) (worldwide but absent from most of the Palaearctic (known only in north Africa)) - Abdominal segment 4 with lateral setae (L-1-IV, L-2-IV, L-3-IV, L-4-IV) on tubercles on posterior half of segment and with D-8-IV and D-9-IV situated more mediodorsally, giving the appearance of one transverse set of lateral tubercles in dorsoventral view (Figs. 67C, 68B–C, 69A–C)...........................79 79. Abdominal segment 3–8 with projecting tubercles apically pointed (Figs. 68C, 69A).............................................................. female Leehelea + female Sphaeromias (pgs. 96, 97) (Leehelea: Oriental and Australasian Regions; Sphaeromias: worldwide except Neotropical Region) - Abdominal segments 3–8 with projecting tubercles broadened apically, appearing rounded apically in dorsoventral view (Figs. 68B, 69B)............................................................80 80. Abdominal segments 3–7 with D-8-IV and D-9-IV on a single rounded tubercle (Fig. 69B)........ female...................................................................... Xenohelea (pg. 99) (Afrotropical and Oriental Regions) - Abdominal segment 3–7 with D-8-IV and D-9-IV on a bilobed tubercle (Fig. 68B)....................................................................................female Homohelea (pg. 95) (Palaearctic, Afrotropical and Oriental Regions) 81. Apex of halter just anterior or at level of anterior margin of tergite 2 (Fig. 33D).....................82 - Apex of halter well posterior to level of anterior margin of tergite 2 (Figs. 33J–L)...................85 82. Abdominal segment 3–6 with lateral setae (L-1, L-2, L-3, L-4) on tubercles on anterior half of segment and with D-8-IV and D-9-IV situated more laterally, giving the appearance of two sets of lateral tubercles in dorsoventral view, one more anterior than the other (Figs. 67B–C)..................................83 - Abdominal segment 3-6 with lateral setae (L-1, L-2, L-3, L-4) on tubercles on posterior half of segment and with D-8-IV and D-9-IV situated more mediodorsally, giving the appearance of one set of lateral tubercles in dorsoventral view (Figs. 64C, 70A)........................................................84 83. Abdominal segments 2–7 with D-2 placed nearly directly anterior of D-5 (so that D-2 and D-5 on both halves of a segment form a nearly perfect square) (Fig. 67B); apex of antenna posterior to apex of midportion of midleg (Fig. 39D); exuviae light brown, with pigmented patches on abdomen only somewhat darker (not strongly contrasting); terminal process directed nearly straight laterally (or only slightly posterior to lateral) (Figs. 76H–I)........................................... Johannsenomyia (in part) (pg. 90) (worldwide but absent from most of the Palaearctic (known only in north Africa)) - Abdominal segments 2–7 with D-2 placed more laterally than D-5 (so that D-2 and D-5 on both halves of a segment form an isosceles trapezoid) (Fig. 67C); apex of antenna anterior to apex of midportion of midleg (Fig. 39E); exuviae dark brown, with pigmented patches on abdomen very dark and clearly evident; terminal process directed posterolaterally (Fig. 76J)................................... Lanatomyia (pg. 91) (Oriental and Australasian Regions) 84. Abdominal segment 4 with V-5-IV, V-6-IV and V-7-IV in a close group and all setae elongate and slender (Fig. 64C); L-3-IV without tubercle (Fig. 64C); exuvial face with cuticular connection to base of antenna (Fig. 79G)............................................................... Hebetula (pg. 79) (Afrotropical, Oriental and Australasian Regions) - Abdominal segment 4 with V-6-IV and V-7-IV close together and V-5-IV more lateral, closer to L-4-IV than to V-6-IV and V-7-IV (Fig. 70A); seta D-5-IV, D-8-IV, L-1-IV, L-2-IV, L-4-IV, and V-5-IV all short pegs, other setae elongate and slender; L-3-IV on tubercle (Fig. 70A); exuvial face separate anteriorly and laterally from remainder of exuviae (without any connection to base of antenna) (Fig. 79H) Clastrieromyia (pg. 104) (Neotropical Region) 85. Segment 4 with D-3-IV posterolateral to D-2-IV (Fig. 71B) or if D-3-IV lateral to D-2-IV then the two are closely approximated...................................................................86 - Segment 4 with D-3-IV lateral to and distant from D-2-IV (Figs. 68B–C, 69A–B, 70B)...............91 86. Dorsal apotome with at least one seta and two campaniform sensilla (Figs. 22C–D, 22J)..............87 - Dorsal apotome with only one seta and one campaniform sensillum (Figs. 22E, 22H–I)...............89 87. Segment 4 with L-2-IV, L-3-IV, L-4-IV each on a single tubercle; V-7-IV placed laterally, closer to L-4-IV than to closely paired V-5-IV and V-6-IV (Fig. 69C) or about halfway between L-4-IV and V-6-IV.................................................................. Bezzia (Bezzia) (in part) (pg. 100) (worldwide) - Segment 4 with L-3-IV situated basally or subbasally on same tubercle as L-2-IV (so there are only two prominent pointed posterolateral tubercles) (Fig. 70C); V-5-IV, V-6-IV, and V-7-IV forming a closely approximated group (Fig. 70C)...........................................................88 88. Dorsal apotome with two setae (and additional campaniform sensilla)..................................................................................... Palpomyia (in part; P. jonesi) (pg. 106) (Nearctic Region) - Dorsal apotome with one seta (and additional campaniform sensilla) (Figs. 22E, 22H–I)........................ Bezzia (Homobezzia) (in part; B. (Homobezzia) annulipes , B. (Bezzia) gibbera), Palpomyia (in part; P. flaviceps) (pgs. 100,106) (Holarctic and Neotropical Region) 89. Area between terminal processes truncated or nearly so (Fig.78K); male with genital lobes very short, each slightly longer than wide and apex extending posteriorly to about level of lateral base of terminal processes (Figs. 78I–J)......................................................... Phaenobezzia (pg. 110) (worldwide except for Australasian Region) - Area between terminal processes u-shaped (Fig. 78B); genital lobes elongate, each longer than wide and apex extending posteriorly to near posteromedial gap between terminal processes (Figs. 77E, 78A, 78C, 78E, 78G)............................................................................90 90. Abdominal segment 4 with setae D-2-IV, D-3-IV, D-8-IV and D-9-IV each on large pointed tubercles, D-5- IV arising directly from surrounding cuticle (Fig. 70C)........... Palpomyia (in part; P. lineata) (pg. 106) (Holarctic Region) - Abdominal segment 4 with setae D-2-IV, D-3-IV, D-5-IV, D-8-IV and D-9-IV on small, rounded tubercles (Fig. 69C)............................................ Bezzia (in part; B. dorsasetula) (pg. 100) (Nearctic Region) 91. Abdominal segment 4 with V-7-IV placed laterally, closer to L-4-IV than to closely paired V-5-IV and V-6- IV (Fig. 69C).........................................................................92 - Abdominal segment 4 with V-5-IV, V-6-IV, and V-7-IV forming a closely gathered group (Figs. 70C, 71A).....................................................................................93 92. Abdominal segments 3-7 each with V-5 and V-6 on both sides of segment on a raised transverse welt, bare of spicules (P. tibialis is more generally bare); segments 3–7 each with D-5 on a short tubercle (as in Fig. 69C)............................. Palpomyia (in part, P. novitibilialis, P. subaspera, P. tibialis) (pg. 106) (Holarctic and Neotropical Region) - Abdominal segments 3–7 each with V-5 and V-6 arising from very short tubercles, without a transverse welt and with spicules present between the two V-5 on either side of the segment; segments 3-7 each with D-5 situated on flat cuticle (not on a tubercle) (Fig. 70B)............................. Pachyhelea (pg. 105) (New World) 93. Prothoracic extension clearly extending laterally to and abutting with a truncated margin against antenna (as in Fig. 28H)..................................................... Palpomyia (in part) (pg. 106) (worldwide) - Prothoracic extension not extending laterally to antenna or, at most, as a very thin band (Fig. 28G)......94 94. Apex of antenna posterior to apex of midportion of midleg (as in Fig. 41E)........................................ Palpomyia (in part; P. altispina, P. jamnbacki, P. nemorosa, P. rufipes, P. stonei) (pg. 106) (Holarctic Region) - Apex of antenna anterior to apex of midportion of midleg (Figs. 40A, 40D, 41D)...................95 95. Abdominal tubercles all apically pointed (Figs. 68C, 69A)........................................................... male Leehelea (unknown but likely keying here) + male Sphaeromias (pgs. 96, 97) (Leehelea: Oriental and Australasian Regions; Sphaeromias: worldwide except Neotropical Region) - Abdominal tubercles all apically rounded (Figs. 68B, 69B).....................................96 96. Palp extending well beyond posterolateral margin of labium (Fig. 28G).................................................................................... Palpomyia (in part; P. tuvae) (pg. 106) (Palaearctic Region) - Palp extending just beyond posterolateral margin of labium (Figs. 27H, 28C).......................97 97. Abdominal segments 3–7 with D-8 and D-9 on a single rounded tubercle (Fig. 69B)..........................................................male Xenohelea (unknown but likely keying here) (pg. 99) (Afrotropical and Oriental Region) - Abdominal segment 3–7 with D-8 and D-9 on a bilobed tubercle (Fig. 68B)........................................................................................... male Homohelea (pg. 95) (Palaearctic, Afrotropical and Oriental Regions)

Published as part of Borkent, Art, 2014, The Pupae of the Biting Midges of the World (Diptera: Ceratopogonidae), With a Generic Key and Analysis of the Phylogenetic Relationships Between Genera, pp. 1-327 in Zootaxa 3879 (1) on pages 24-37, DOI: 10.11646/zootaxa.3879.1.1, http://zenodo.org/record/4949051