Ceratopogonidae, Borkent, 2014

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, Zootaxa 3879 (1), pp. 1-327 : 24-37

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)

Figures

FIGURE 3. Habitus of pupae of Culicoides sonorensis (from Borkent 2012). A. Female, in dorsal view. B. Male, in lateral view. C. Female, in ventral view. A and C have the abdominal segments separated by expanded membrane, not shown in B. Shagreen not shown.

FIGURE 8. Habitus. A. Austroconops mcmillani, male, in lateral view (from Borkent & Craig 2004). B. Leptoconops spinosifrons, male, in lateral view (from Laurence & Mathias 1972). C. Leptoconops gallicus, female, in dorsal and ventral view (from Clastrier 1972). D. Forcipomyia (Euprojoannisia) spatulifera, female, dorsal view (from Saunders 1957). E. Forcipomyia (Euprojoannisia) wirthi, female, in dorsal view (from Saunders 1957). F. Forcipomyia (Euprojoannisia) bromeliae, male, in dorsal view (from Saunders 1957). G. Forcipomyia (Caloforcipomyia) caerulea, female, in dorsal view (from Saunders 1957). H. Forcipomyia (Caloforcipomyia) varicolor, female, in dorsal view (from Saunders 1957).

FIGURE 9. Habitus. A. Forcipomyia (Schizoforcipomyia) petersoni, female, in dorsal view (from Chan & LeRoux 1971b). B. Forcipomyia (Metaforcipomyia) cerifera, female, in dorsal view (from Saunders 1957). C. Forcipomyia (Warmkea) lesliei, male, with female segment 9 separate, in dorsal view (from Saunders 1957). D. Forcipomyia (Warmkea) tuberculata, male, in dorsal view (from Saunders 1957). E. Forcipomyia (Microhelea) intonsa, female, with male segment 9 separate, in dorsal view (from Chan & LeRoux 1971a). F. Forcipomyia (Forcipomyia) brevipennis, female, with male segment 9 separate, in dorsal view (from Saunders 1924). G. Forcipomyia (Forcipomyia) ciliata, female, with male segment 9 separate, in dorsal view (from Saunders 1924). H. Forcipomyia (Forcipomyia) pulchrithorax, male, with female segment 9 separate, in dorsal view (from Saunders 1924).

FIGURE 10. Habitus. A. Forcipomyia (Forcipomyia) costata, female, with male segment 9 separate, in dorsal view (from Saunders 1924). B. Forcipomyia (Forcipomyia) costata, female, with male segment 9 separate, in ventral view (from Saunders 1924). C. Forcipomyia (Forcipomyia) costata, female, in lateral view (from Saunders 1924). D. Forcipomyia (Lepidohelea) bicolor, male, in dorsal view (from Saunders 1924). E. Forcipomyia (Phytohelea) grandis, male, in dorsal view (from Chan & LeRoux 1971a). F. Forcipomyia (Trichohelea) eques, female, in dorsal view (from Chan & LeRoux 1965). G. Forcipomyia (Trichohelea) crinita, male, with female segment 9 separate, in dorsal view (from Saunders 1964). H. Forcipomyia (Synthyridomyia) acidicola, male, in dorsal view (from Saunders 1957).

FIGURE 11. Habitus. A. Forcipomyia (Thyridomyia) rugosa, male, in dorsal view (from Chan & LeRoux 1970). B. Forcipomyia (Pterobosca) crinume, female, in ventral view (from Tokunaga 1932). C. Forcipomyia (Lasiohelea) rugosa, male, in dorsal view (from Saunders 1964). D. Atrichopogon hexastichus, male, in dorsal view (from Nielsen 1951). E. Atrichopogon wirthi, female, in dorsal view (from Chan & Linley 1988). F. Atrichopogon muelleri, sex unknown, in dorsal view, with larval exuviae on posterior of abdomen (from Szadziewski et al. 1997). G. Dasyhelea vericolor, sex unknown, in dorsal view (from Szadziewski et al. 1997). H. Schizonyxhelea bulla, sex unknown, in dorsal view (from Thomsen 1937).

FIGURE 12. Habitus. A. Pellucidomyia leei, male, in dorsal view (from Debenham 1970). B. Dibezzia prominens, sex unknown, in dorsal view (form Mayer 1934c). C. Macropeza albitarsis, male, in dorsal view (from Szadziewski & Dominiak, 2007, modified). D. Mallochohelea munda, male, in ventral view (from Szadziewski et al. 1997). E. Sphaeromias fasciata, sex unknown, in dorsal view (from Mayer 1934a). F. Bezzia glabra, female, in ventral view (from Thomsen 1937). G. Palpomyia sp., male, in dorsal view (from Grogan & Wirth 1979). H. Paryphoconus oliveirai, female, in ventral view (modified from Ronderos et al. 2007).

FIGURE 13. Anterior portion of heads and thoraces, in anterior or anterodorsal view. A. Leptoconops kerteszi, ecdysial sutures of head enhanced to show position. B. Culicoides denticulatus. C. Ceratopogon nr. abstrusus. D. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). E. Alluaudomyia bella, dorsal apotome absent. F. Clinohelea curriei. G. Pellucidomyia leei. H. Mallochohelea caudellii. I. Bezzia sp. (from Bolean Lake, BC, Canada).

FIGURE 19. Dorsal apotomes, anterior view. A. Culicoides circumscriptus (from Kettle & Lawson 1952). B. Culicoides cubitalis (from Kettle & Lawson 1952). C. Culicoides truncorum (from Kettle & Lawson 1952). D. Culicoides chiopterus (from Jamnback 1965). E. Paradasyhelea minuta (from Kettle & Elson 1975a). F. Paradasyhelea albipunctata (from Elson- Harris & Kettle 1985a). G. Paradasyhelea reyei (from Elson-Harris & Kettle 1985a). H. Ceratopogon boomerangus (from Borkent & Grogan 1995). I. Ceratopogon abstrusus (from Borkent & Grogan 1995). J. Ceratopogon arcanus (from Borkent & Grogan 1995). K. Brachypogon artemis (from Debenham 1991). L. Alluaudomyia appendiculata (from Elson-Harris & Kettle 1985b). M. Alluaudomyia fumosipennis (from Elson-Harris & Kettle 1985b). N. Baeodasymyia michaeli (from Borkent & Craig 1999). O. Austrohelea campbellensis. P. Stilobezzia rabelloi (from Borkent & Craig 2001).

FIGURE 20. Dorsal apotomes, anterior view. A. Stilobezzia soror (from Mayer 1934c). B. Stilobezzia lutea (from Wirth & Grogan 1981). C. Stilobezzia antennalis (from Wirth & Grogan 1981). D. Stilobezzia pallidiventris (from Wirth & Grogan 1981). E. Schizonyxhelea forattinii (from Borkent 2000a). F. Schizonyxhelea bulla (from Wirth & Grogan 1981, modified). G. Serromyia sp.. H. Allohelea japonica. I. Monohelea obscura. J. Atyphohelea macroneura (from Borkent 1998). K. Parabezzia alexanderi male (from Grogan & Wirth 1977a, modified). L. Parabezzia alexanderi female (from Grogan & Wirth 1977a, modified). M. Clinohelea dryas (from Elson-Harris & Kettle 1986). N. Clinohelea tasmaniensis (from Elson-Harris & Kettle 1986). O. Heteromyia wokei (from Wirth & Grogan 1977, modified). P. Pellucidomyia leei (from Elson-Harris & Kettle 1986).

FIGURE 21. Dorsal apotomes, anterior view. A. Dibezzia prominens (from Mayer 1934c). B. Hebetula tonnoiri (from Elson- Harris 1987). C. Jenkinshelea albaria (from Wirth & Grogan 1979). D. Macropeza albitarsis (from Szadziewski & Dominiak, 2007). E. Mallochohelea termophila (from Spinelli 1984, modified). F. Probezzia pallida (from Wirth & Grogan 1979). G. Neobezzia fittkaui (from Ronderos et al. 2011, modified). H. Nilobezzia basispinigera (from Elson-Harris 1987). I. Nilobezzia virago (from Elson-Harris 1987). J. Johannsenomyia argentata (from Wirth & Grogan 1979). K. Lanatomyia electra (from Elson-Harris 1987). L. Anebomyia atripes (from Wirth & Grogan 1979, modified).

FIGURE 22. Dorsal apotomes, anterior view. A. Sphaeromias longipennis (from Wirth & Grogan 1979). B. Amerohelea sordidipes (from Lane et al. 1955). C. Bezzia mollis (from Mayer 1934c). D. Bezzia roldani (from Spinelli & Wirth 1989). E. Bezzia brevicornis (from Spinelli 1983a). F. Clastrieromyia dycei (from Spinelli & Grogan 1986). G. Pachyhelea pachymera (from Spinelli 1983a). H. Palpomyia lineata (from Grogan & Wirth 1979). I. Palpomyia belkini (from Grogan & Wirth 1979, modified). J. Palpomyia flaviceps (from Grogan & Wirth 1979). K. Phaenobezzia opaca (from Wirth & Grogan 1982). L. Paryphoconus oliveirai (from Ronderos et al. 2007).

FIGURE 23. Posterior portion of heads, in ventral view. A. Austroconops mcmillani. B. Leptoconops kerteszi. C. Forcipomyia (Euprojoannisia) sp. (from Peru). D. Forcipomyia (Phytohelea) n. sp. near antiguensis. E. Atrichopogon sp. (from St. Pierre de Wakefield, Quebec, Canada). F. Dasyhelea paracincta. G. Dasyhelea pseudoincisurata.

FIGURE 24. Posterior portion of heads, in ventral view. A. Culicoides guttipennis. B. Paradasyhelea albipunctata. C. Ceratopogon arcanus. D. Brachypogon sp. (Flag Glade, Pocahontas Co., West Virginia, USA). E. Alluaudomyia needhami. F. Baeodasymyia michaeli. G. Austrohelea campbellensis.

FIGURE 25. Posterior portion of heads, in ventral view. A. Stilobezzia antennalis. B. Stilobezzia rabelloi (from Borkent & Craig 2001). C. Schizonyxhelea bulla. D. Serromyia sp.. E. Allohelea japonica. F. Monohelea obscura (reconstructed from distorted specimen). G. Atyphohelea macroneura. H. Parabezzia huberti.

FIGURE 26. Posterior portion of heads, in ventral view. A. Echinohelea lanei (reconstructed from distorted specimen). B. Clinohelea curriei. C. Heteromyia wokei. D. Pellucidomyia leei. E. Dibezzia debenhamae (reconstructed from distorted specimen). F. Hebetula tonnoiri. G. Jenkinshelea magnipennis H. Macropeza bayeri.

FIGURE 27. Posterior portion of heads, in ventral view. A. Mallochohelea caudellii. B. Probezzia sabroskyi. C. Neobezzia fittkaui. D. Nilobezzia curticornis. E. Johannsenomyia argentata. F. Lanatomyia electra. G. Anebomyia atripes. H. Homohelea delanoe.

FIGURE 28. Posterior portion of heads, in ventral view. A. Leehelea hispida. B. Sphaeromias longipennis. C. Xenohelea galatea. D. Bezzia nobilis. E. Clastrieromyia dycei. F. Pachyhelea pachymera. G. Palpomyia subasper. H. Phaenobezzia opaca. I. Paryphoconus grandis.

FIGURE 29. Dorsal setae from left side of thoraces. A. Austroconops mcmillani. B. Leptoconops kerteszi. C. Leptoconops linleyi. D. Forcipomyia (Euprojoannisia) sp. (from Peru). E. Forcipomyia (Forcipomyia) bipunctata. F. Forcipomyia (Phytohelea) n. sp. nr. antiguensis. G. Atrichopogon sp. (from Lake Opinicon, Ontario, Canada). H. Dasyhelea paracincta. I. Dasyhelea atlantis. J. Dasyhelea pseudoincisurata. K. Culicoides haematopotus. L. Culicoides phlebotomus. M. Paradasyhelea albipunctata. N. Ceratopogon boomerangus. O. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). P. Alluaudomyia needhami. Q. Baeodasymyia christopheri (from Borkent & Craig 1999). R. Austrohelea campbellensis. S. Stilobezzia antennalis. T. Schizonyxhelea forattinii (from Borkent 2000a).

FIGURE 30. Dorsal setae from left side of thoraces. A. Serromyia sp. B. Allohelea japonica. C. Monohelea obscura. D. Atyphohelea macroneura. E. Parabezzia huberti. F. Echinohelea lanei. G. Clinohelea curriei. H. Heteromyia clavata. I. Pellucidomyia leei. J. Dibezzia debenhamae (dirty specimen, D-5-T perhaps broken setal socket). K. Hebetula tonnoiri. L. Jenkinshelea magnipennis. M. Macropeza natalensis. N. Mallochohelea caudellii. O. Probezzia sabroskyi. P. Neobezzia fittkaui (from Ronderos et al. 2011, modified). Q. Nilobezzia curticornis. R. Johannsenomyia argentata. S. Lanatomyia miles. T. Anebomyia atripes.

FIGURE 31. Dorsal setae from left side of thoraces; anteromedial setae. A. Homohelea delanoe. B. Leehelea hispida. C. Sphaeromias longipennis. D. Xenohelea galatea. E. Bezzia nobilis. F. Clastrieromyia dycei. G. Pachyhelea pachymera. H. Palpomyia jonesi. I. Phaenobezzia mashonensis. J. Paryphoconus oliveirai. K. Echinohelea lanei, partial lateral view (distorted slide specimen). L. Palpomyia (JAD685), dorsal view. M. Palpomyia (JAD685), dorsolateral view. N. Probezzia seminigra, dorsal view of left side.

FIGURE 32. Cephalothoraces and abdominal segments 1–3, in lateral view. A. Austroconops mcmillani. B. Leptoconops linleyi. C. Forcipomyia (Phytohelea) sp. (Florida, USA). D. Atrichopogon sp. (from 6.5 km NW Enderby, BC, Canada). E. Dasyhelea sp., in anteroventral view (Herman Lake, 4 km SE Eagle Bay, BC, Canada). F. Dasyhelea oppresa. G. Culicoides denticulatus. H. Ceratopogon nr. abstrusus. I. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). J. Alluaudomyia bella. K. Baeodasymyia michaeli. L. Stilobezzia antennalis.

FIGURE 33. Cephalothoraces and abdominal segments 1–3, in lateral view. A. Parabezzia sp. (5 km. E. Danby, Vermont, USA). B. Clinohelea curriei. C. Pellucidomyia leei. D. Hebetula tonnoiri. E. Jenkinshelea sp. (from Ottawa, ON, Canada). F. Mallochohelea caudellii. G. Probezzia seminigra. H. Nilobezzia sp. (from Kruger NP, South Africa). I. Sphaeromias longipennis. J. Bezzia sp. (6 km E. Salmon Arm, BC, Canada) K. Palpomyia sp. (Black Lake, Quebec, Canada). L. Phaenobezzia sp. (from Skulsuza, South Africa).

FIGURE 35. Legs, wings and apices of the antennae, in ventral view. A. Paradasyhelea sp. (from Borlase Stream, Lake Rotoiti, New Zealand). B. Ceratopogon sp. nr. abstrusus. C. Brachypogon sp. (from Spanish Lake, 6 km E. Falkland, BC, Canada). D. Alluaudomyia needhami. E. Baeodasymyia michaeli. F. Austrohelea campbellensis (reconstructed from distorted specimen).

FIGURE 36. Legs, wings and apices of the antennae, in ventral view. A. Stilobezzia antennalis. B. Stilobezzia lutea. C. Schizonyxhelea forattinii. D. Serromyia sp.. E. Allohelea japonica (reconstructed from distorted specimen). F. Monohelea obscura (reconstructed from distorted specimen).

FIGURE 37. Legs, wings and apices of the antennae, in ventral view. A. Atyphohelea macroneura. B. Parabezzia huberti. C. Echinohelea lanei (reconstructed from distorted specimen). D. Clinohelea currei. E. Heteromyia clavata. F. Pellucidomyia leei.

FIGURE 38. Legs, wings and apices of the antennae, in ventral view. A. Dibezzia debenhamae (reconstructed from distorted specimen). B. Hebetula tonnoiri (reconstructed from distorted specimen). C. Jenkinshelea sp. (from Ottawa, ON, Canada). D. Macropeza natalensis. E. Macropeza sp. (from Laos). F. Mallochohelea caudellii.

FIGURE 39. Legs, wings and apices of the antennae, in ventral view. A. Probezzia sabroskyi. B. Neobezzia fittkaui. C. Nilobezzia sp. (from Australia). D. Johannsenomyia argentata. E. Lanatomyia electra. F. Anebomyia atripes.

FIGURE 40. Legs, wings and apices of the antennae, in ventral view. A. Homohelea delanoe, male (reconstructed from distorted specimen). B. Homohelea albitudinis, female (drawn from photograph). C. Leehelea hispida, female. D. Sphaeromias longipennis, male. E. Sphaeromias longipennis, female. F. Xenohelea galatea, female (reconstructed from distorted specimen).

FIGURE 41. Legs, wings and apices of the antennae, in ventral view. A. Bezzia nobilis. B. Clastrieromyia dycei (reconstructed from distorted specimen). C. Pachyhelea pachymera. D. Palpomyia occidentalis. E. Phaenobezzia opaca. F. Paryphoconus oliveirai (reconstructed from distorted specimen).

FIGURE 42. Respiratory organs, most in dorsal view, Fig. 42B in lateral view, pattern on tracheal tubes shown only in some. A. Austroconops mcmillani (from Borkent & Craig 2004). B. Austroconops mcmillani (from Borkent & Craig 2004). C. Leptoconops americanus (from Clastrier & Wirth 1978). D. Leptoconops asilomar (from Clastrier & Wirth 1978). E. Forcipomyia (Euprojoannisia) wirthi (from Saunders 1957). F. Forcipomyia (Euprojoannisia) longispina (from Saunders 1957). G. Forcipomyia (Warmkea) malayae (from Saunders 1957). H. Forcipomyia (Caloforcipomyia) caerulea (from Saunders 1957). I. Forcipomyia (Caloforcipomyia) varicolor (from Saunders 1957). J. Forcipomyia (Metaforcipomyia) cerifera (from Saunders 1957). K. Forcipomyia (Forcipomyia) genualis (from Saunders 1957, as raleighi). L. Forcipomyia (Forcipomyia) nigra (from Saunders 1924). M. Forcipomyia (Forcipomyia) ciliata (from Saunders 1924). N. Forcipomyia (Phytohelea) bromelicola (from Saunders 1925, (as an Apelma)). O. Forcipomyia (Synthyridomyia) acidicola (Synthyridomyia) (from Saunders 1924). P. Forcipomyia (Trichohelea) crinita (from Saunders 1964). Q. Forcipomyia (Lasiohelea) quasicornuta (from Saunders 1964). R. Forcipomyia (Lasiohelea) cornuta (from Saunders 1964). S. Atrichopogon wirthi (from Chan & Linley 1988). T. Atrichopogon thienemanni (from Lenz 1934).

FIGURE 43. Respiratory organs, most in dorsal view, Fig. 43T in lateral view, pattern on tracheal tubes shown only in some. A. Dasyhelea pseudoincisurata (from Waugh & Wirth 1976). B. Dasyhelea grisea (from Waugh & Wirth 1976). C. Dasyhelea traverae (from Waugh & Wirth 1976). D. Dasyhelea pseudocincta (from Waugh & Wirth 1976). E. Dasyhelea leptobranchia (from Waugh & Wirth 1976). F. Dasyhelea brevicornis (from Waugh & Wirth 1976). G. Dasyhelea hitchcocki (from Wirth 1976a). H. Culicoides chiopterus (from Jamnback 1965). I. Culicoides furens (from Jamnback 1965). J. Culicoides guttipennis (from Jamnback 1965). K. Culicoides dewulfi (from Kettle & Lawson 1952, as C. pseudochiopterus). L. Culicoides punctatus (from Kettle & Lawson 1952). M. Culicoides sanguisuga (from Jamnback 1965). N. Culicoides variipennis (from Jamnback 1965). O. Paradasyhelea albipunctata (from Elson-Harris & Kettle 1985a). P. Ceratopogon inverecundus (from Borkent & Grogan 1995). Q. Ceratopogon willisi (from Borkent & Grogan 1995). R. Brachypogon sp. (from Virginia, USA). S. Alluaudomyia needhami (from Wirth & Grogan 1981). T. Alluaudomyia fumosipennis (from Debenham 1971).

FIGURE 44. Respiratory organs, in dorsal view, pattern on tracheal tubes shown only in some. A. Alluaudomyia latipennis (from Debenham 1971). B. Alluaudomyia footei (from Wirth & Grogan 1981). C. Baeodasymyia christopheri (from Borkent & Craig 1999). D. Austrohelea campbellensis. E. Stilobezzia antennalis (from Wirth & Grogan 1981, modified). F. Stilobezzia glauca (from Wirth & Grogan 1981). G. Stilobezzia ochracea (from Kettle & Lawson 1952). H. Stilobezzia diversa (from Wirth & Grogan 1981). I. Stilobezzia rabelloi (from Borkent & Craig 2001). J. Stilobezzia navaiae (from Wirth & Grogan 1981). K. Stilobezzia lutea (from Wirth & Grogan 1981). L. Schizonyxhelea bulla (from Wirth & Grogan 1981, as a Stilobezzia). M. Schizonyxhelea forattinii (from Borkent 2000a). N. Serromyia femorata (from Kettle & Lawson 1952). O. Allohelea japonica. P.Monohelea hirsuta (from Wirth & Grogan 1981, modified). Q. Atyphohelea macroneura (from Borkent 1998). R. Parabezzia alexanderi (from Grogan & Wirth 1977a). S. Parabezzia huberti (from Grogan & Wirth 1977a). T. Echinohelea lanei (from Wirth 1994b).

FIGURE 45. Respiratory organs, in dorsal view, pattern on tracheal tubes shown only in some. A. Clinohelea tasmaniensis (from Elson-Harris & Kettle 1986). B. Clinohelea bimaculata (from Wirth & Grogan 1979, modified). C. Heteromyia wokei (from Wirth & Grogan 1977). D. Heteromyia clavata (from Wirth & Grogan 1977). E. Pellucidomyia leei (from Elson-Harris & Kettle 1986). F. Pellucidomyia geari (from de Meillon & Wirth 1981). G. Dibezzia prominens (from Mayer 1934c). H. Hebetula tonnoiri (from Debenham 1974). I. Jenkinshelea albaria (from Wirth & Grogan 1979). J. Macropeza albitarsis (from Szadziewski & Dominiak, 2007). K. Mallochohelea albihalter (from Wirth & Grogan 1979). L. Probezzia pallida (from Wirth & Grogan 1979). M. Probezzia xanthogaster (from Wirth & Grogan 1979). N. Neobezzia fittkaui (from Ronderos et al. 2011). O. Nilobezzia aranea (from Elson-Harris 1987). P. Nilobezzia virago (from Elson-Harris 1987). Q. Johannsenomyia argentata (from Wirth & Grogan 1979). R. Lanatomyia electra (from Debenham 1974). S. Lanatomyia miles (from Debenham 1974). T. Anebomyia atripes (from Wirth & Grogan 1979).

FIGURE 46. Respiratory organs, most in dorsal view, Figs. 46G. in lateral view, pattern on tracheal tubes shown only in some. A. Homohelea albitudinis (from de Meillon & Wirth 1981). B. Homohelea delanoe (from de Meillon & Wirth 1981). C. Leehelea wasselli (from Elson-Harris 1987). D. Leehelea hispida (from Elson-Harris 1987). E. Sphaeromias longipennis (from Wirth & Grogan 1979). F. Xenohelea galatea. G. Amerohelea sordidipes (from Lane et al. 1955). H. Bezzia glabra (from Wirth 1983b). I. Bezzia obelisca (from Wirth 1983a). J. Bezzia dorsasetula (from Wirth 1983a). K. Bezzia turkmenica (from Glukhova 1979b). L. Clastrieromyia dycei (from Spinelli & Grogan 1986). M. Pachyhelea pachymera (from Spinelli 1983a). N. Palpomyia lineata (from Grogan & Wirth 1979). O. Palpomyia jonesi (from Grogan & Wirth 1979). P. Palpomyia belkini (from Grogan & Wirth 1979). Q. Palpomyia rufa (from Grogan & Wirth 1979). R. Palpomyia flaviceps (from Grogan & Wirth 1979). S. Phaenobezzia opaca (from Wirth & Grogan 1982). T. Paryphoconus oliveirai (from Ronderos et al. 2007).

FIGURE 49. Metathoraces and tergites 1, in dorsal view. A. Austrohelea campbellensis. B. Stilobezzia antennalis. C. Schizonyxhelea forattinii. D. Serromyia sp.. E. Allohelea japonica. F. Monohelea obscura.

FIGURE 50. Metathoraces and tergites 1, in dorsal view. A. Atyphohelea macroneura. B. Parabezzia huberti. C. Echinohelea lanei. D. Clinohelea currei. E. Heteromyia wokei. F. Pellucidomyia leei.

FIGURE 51. Metathoraces and tergites 1, in dorsal view. A. Dibezzia debenhamae. B. Hebetula tonnoiri (reconstructed from distorted specimen). C. Jenkinshelea magnipennis. D. Macropeza natalensis. E. Mallochohelea caudellii. F. Probezzia sabroskyi.

FIGURE 55. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Austroconops mcmillani (from Borkent & Craig 2004). B. Leptoconops kerteszi. C. Forcipomyia (Euprojoannisia) sp. (from Peru).

FIGURE 56. Abdominal segment four, in dorsal and ventral view (shagreen not shown). A. Forcipomyia dominicana. B. Atrichopogon sp. (from Lake Opinicon, Ontario, Canada). C. Dasyhelea paracincta.

FIGURE 57. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Culicoides haematopotus (from Borkent 2012). B. Paradasyhelea albipunctata. C. Ceratopogon boomerangus (from Borkent & Grogan 1995, compressed dorsoventrally).

FIGURE 58. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Ceratopogon nr. inverecundus. B. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). C. Alluaudomyia needhami.

FIGURE 59. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Baeodasymyia christopheri (from Borkent & Craig 1999). B. Austrohelea campbellensis. C. Stilobezzia antennalis.

FIGURE 60. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Stilobezzia rabelloi (from Borkent & Craig 2001). B. Schizonyxhelea forattinii (from Borkent 2000a, modified). C. Serromyia sp.

FIGURE 61. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Allohelea japonica (reconstructed from distorted specimen). B. Monohelea obscura (reconstructed from distorted specimen). C. Monohelea hirsuta.

FIGURE 62. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Atyphohelea macroneura (from Borkent 1998). B. Parabezzia huberti. C. Parabezzia alexanderi.

FIGURE 63. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Echinohelea lanei (reconstructed from distorted specimen). B. Clinohelea unimaculata. C. Heteromyia wokei.

FIGURE 64. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Pellucidomyia leei. B. Dibezzia debenhamae (reconstructed from distorted and dirty specimen). C. Hebetula tonnoiri.

FIGURE 65. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Jenkinshelea magnipennis. B. Macropeza natalensis. C. Macropeza sp. (from Laos).

FIGURE 66. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Mallochohelea nr. caudellii (from Hope, British Columbia, Canada). B. Probezzia sabroskyi. C. Neobezzia fittkaui (from Ronderos et al. 2011, modified).

FIGURE 67. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Nilobezzia curticornis. B. Johannsenomyia argentata. C. Lanatomyia miles.

FIGURE 68. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Anebomyia atripes. B. Homohelea delanoe. C. Leehelea hispida (reconstructed from distorted specimen).

FIGURE 69. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Sphaeromias longipennis. B. Xenohelea galatea. C. Bezzia nobilis.

FIGURE 70. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Clastrieromyia dycei. B. Pachyhelea pachymera. C. Palpomyia jonesi.

FIGURE 71. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Palpomyia occidentalis. B. Phaenobezzia opaca. C. Paryphoconus oliveirai.

FIGURE 72. Abdominal segments 9. A. Austroconops mcmillani, male, in dorsal and ventral view (from Borkent & Craig 2004). B. Leptoconops whitseli, male, in dorsal and ventral view (from Clastrier & Wirth 1978). C. Leptoconops whitseli, female, in dorsal and ventral view (from Clastrier & Wirth 1978). D. Leptoconops asilomar, male, in dorsal and ventral view (modified from Clastrier & Wirth 1978). E. Leptoconops asilomar, female, in dorsal and ventral view (modified from Clastrier & Wirth 1978). F. Forcipomyia anabaenae, female, in dorsal view (from Chan & Saunders 1965). G. Forcipomyia eques, male, in ventral view (from Chan & LeRoux 1965). H. Forcipomyia rugosa, female, in dorsal view (from Chan & LeRoux 1970). I. Forcipomyia brevis, female, in dorsal view (from Mayer 1934a). J. Atrichopogon wirthi, male, in dorsal view, arrow indicating anteriorly directed spines (from Chan & Linley 1988). K. Dasyhelea contigua, female, in ventral view (from Mayer 1934c). L. Dasyhelea grata, male, in ventral view (from Mayer 1934c).

FIGURE 73. Abdominal segments 9. A. Dasyhelea laeta, female, in ventral view (from Mayer 1934c). B. Dasyhelea assimilis, male, in lateral view with ventral to the right (from Mayer 1934c). C. Dasyhelea assimilis, male, in lateral view with ventral to the right (from Mayer 1934c). D. Dasyhelea grisea, male, in dorsal view (from Waugh & Wirth 1976). E. Dasyhelea chani, female, in dorsal view (from Wirth & Linley 1990). F. Dasyhelea atlantis, female, in ventral view (from Waugh & Wirth 1976). G. Dasyhelea gluchovae, female, in ventral view (from Brodskaya 1996). H. Culicoides haematopotus, female, in dorsal and ventral view (shagreen not shown). I. Paradasyhelea albipunctata, female, in dorsal view (Elson-Harris & Kettle 1985a). J. Ceratopogon boomerangus, female, in ventral view (modified from Borkent & Grogan 1995). K. Brachypogon artemis, female, in dorsal view (from Debenham 1991). L. Alluaudomyia appendiculata, female, in dorsal view (from Elson-Harris & Kettle 1985b).

FIGURE 75. Abdominal segments 9. A. Parabezzia huberti, female, in ventral view (from Grogan & Wirth 1977a). B. Echinohelea lanei (from Wirth 1994b). C. Clinohelea bimaculata, female, in ventral view (from Wirth & Grogan 1979). D. Heteromyia wokei, male, in dorsal and ventral view (from Wirth & Grogan 1977). E. Heteromyia wokei, female, in dorsal and ventral view (from Wirth & Grogan 1977). F. Heteromyia clavata, female, in dorsal and ventral view (from Wirth & Grogan 1977). G. Pellucidomyia leei, male, in dorsal view (from Elson-Harris & Kettle 1986). H. Pellucidomyia geari, male, in ventral view (from de Meillon & Wirth 1981). I. Dibezzia debenhamae, male, in dorsal and ventral view (shagreen not shown). J. Hebetula tonnoiri, female, in dorsal view (from Elson-Harris 1987). K. Jenkinshelea albaria, male, in dorsal and ventral view (from Wirth & Grogan 1979). L. Jenkinshelea albaria, female, in dorsal and ventral view (from Wirth & Grogan 1979).

FIGURE 76. Abdominal segments 9. A. Macropeza albitarsis, male, in ventral view (modified from Szadziewski & Dominiak, 2007). B. Macropeza albitarsis, female, in ventral view (from Szadziewski & Dominiak, 2007). C. Mallochohelea termophila, male, in ventral view (from Spinelli 1984). D. Mallochohelea termophila, female, in dorsal view (from Spinelli 1984). E. Probezzia pallida, male, in dorsal and ventral view (from Wirth & Grogan 1979). F. Neobezzia fittkaui, male, in dorsal view (from Ronderos, et al. 2011). G. Nilobezzia virago, female, in dorsal view (from Elson-Harris 1987). H. Johannsenomyia argentata, male, in dorsal and ventral view (from Wirth & Grogan 1979). I. Johannsenomyia argentata, female, in dorsal and ventral view (from Wirth & Grogan 1979). J. Lanatomyia miles, female, in dorsal view (from Elson- Harris 1987). K. Anebomyia atripes, male, in dorsal and ventral view (from Wirth & Grogan 1979). L. Anebomyia atripes, female, in dorsal and ventral view (from Wirth & Grogan 1979).

FIGURE 77. Abdominal segments 9. A. Homohelea delanoe, female, in dorsal view (from de Meillon & Wirth 1981). B. Leehelea wasselli, female, in dorsal view (from Elson-Harris 1987). C. Sphaeromias longipennis, male, in dorsal and ventral view (from Wirth & Grogan 1979). D. Xenohelea galatea, female, in dorsal and ventral view (shagreen not shown). E. Bezzia roldani, male, in dorsal view (from Spinelli & Wirth 1989). F. Bezzia roldani, female, in dorsal view (from Spinelli & Wirth 1989). G. Bezzia brevicornis, male, in dorsal view (from Spinelli 1983a). H. Bezzia acanthodes, female, in dorsal view (from Spinelli 1983a). I. Clastrieromyia dycei, male, in ventral view (from Spinelli & Grogan 1986). J. Clastrieromyia dycei, female, in ventral view (from Spinelli & Grogan 1986). K. Pachyhelea pachymera, male, in ventral view (from Spinelli 1983a). L. Pachyhelea pachymera, female, in dorsal view (from Spinelli 1983a).

FIGURE 78. Abdominal segments 9. A. Palpomyia lineata, male, in dorsal and ventral view (from Grogan & Wirth 1979). B. Palpomyia lineata, female, in dorsal and ventral view (from Grogan & Wirth 1979). C. Palpomyia jonesi, male, in dorsal and ventral view (from Grogan & Wirth 1979. D. Palpomyia jonesi, female, in dorsal and ventral view (from Grogan & Wirth 1979). E. Palpomyia belkini, male, in dorsal and ventral view (from Grogan & Wirth 1979). F. Palpomyia belkini, female, in dorsal and ventral view (from Grogan & Wirth 1979). G. Palpomyia rufa, male, in dorsal and ventral view (from Grogan & Wirth 1979). H. Palpomyia flaviceps, female, in dorsal and ventral view (from Grogan & Wirth 1979). I. Phaenobezzia vacunae, male, in ventral view (from Haeselbarth 1965). J. Phaenobezzia mashonensis, male, in ventral view (from Haeselbarth 1965). K. Phaenobezzia opaca, female, in dorsal and ventral view (from Wirth & Grogan 1982). L. Paryphoconus oliveirai, female, in dorsal view (from Ronderos et al. 2007).

FIGURE 79. Evolution of ecdysial sutures and splits in pupal exuviae of Ceratopogonidae, head and anterior portion of thorax in ventral view. Numbers refer to character states of characters 2 and 8 in text. Red indicates ecdysial splits. A. Leptoconopinae. B. Forcipomyiinae. C. Dasyheleinae. D. Most Ceratopogonini. E. Most Ceratopogonini, Clinohelea. F. Allohelea, Atyphohelea, Parabezzia. G. Heteromyia, Pellucidomyia, Hebetula. H. most Sphaeromiini s. lat., most Palpomyiini, Paryphoconus; some Bezzia are nearly completed fused (as in Fig. 79I), with only the dorsal apotome separate, along with an ecdysial split along the base of the antenna and the dorsal thoracic split. I. Jenkinshelea, Macropeza, Mallochohelea, Probezzia, Neobezzia.