Leptoconops (Proleptoconops) werneri Wirth and Atchley

Phillips, Robert A., 2022, Culicoides Latreille and Leptoconops Skuse biting midges of the southwestern United States with emphasis on the Canyonlands of southeastern Utah (Diptera: Ceratopogonidae), Insecta Mundi 2022 (907), pp. 1-214 : 29-45

Leptoconops (Proleptoconops) werneri Wirth and Atchley

( Fig. 13, 14 )

Leptoconops (Leptoconops) werneri Wirth and Atchley, 1973: 34 (key; female, male; fig. female head, wing, genitalia,

hind tibial comb, antenna, spermathecae, palpus, hind tarsomere 5, male genitalia, antenna; California). Leptoconops (Proleptoconops) werneri: Clastrier 1974: 231 (assignment to subgenus Proleptoconops ). Borkent and Spi-

nelli 2000: 10 (in Neotropical catalog).

Diagnosis. ( Table 13) Entirely dark brown to black; stigma indistinct; eyes bare; palpal segment 3 sensilla in superficial excavation wider than deep (female as in Fig. 12 L . carteri); clypeus with four setae, median pair farther apart from each other than from corresponding lateral setae; hind tibial comb with four spines. Female: antenna with 12 flagellomeres; mandible with 10–13 teeth; hind tarsomeres 3 and 5 subequal; claw with prominent basal tooth; two large and one small elongate ovoid spermathecae; cerci>3× longer than wide (as in Fig. 6 L . torrens). Male: flagellomere 13 ~1.4× longer than 12; tergite 9 evenly tapering to base of fingerlike widely separated apicolateral processes, which are ~4× longer than wide and separated by a distance about equal to their length; aedeagal sclerites separate, apically expanded, blunt; distal sclerite of paramere ~2× longer than wide, apical half greatly expanded quadrate with blunt ventromedian tooth.

Distribution. California, Arizona, New Mexico, Kansas, Missouri, Texas, Baja California, Baja California Sur, Sonora, Zacatecas, Morelos.

Adult behavior. Females have been collected in a quail-baited trap and while biting a human ( Wirth and Atchley 1973). In addition, Weinmann et al. (1979) collected blood-engorged L. werneri from California quail ( Callipepla californica [Shaw], Odontophoridae )-baited traps; however, they found no L. werneri infected with quail heartworm ( Splendidofilaria californiensis [Wehr and Herman] [Nematoda: Filarioidea]) or other evidence it can transmit the parasite.

Key to Culicoides Males

(Measurement and ratio definitions are in the “Morphological terms and data” section of Materials and Methods)

1. Wing without pattern of pale spots (often false pale areas caused by paucity of macrotrichia in base of wing and from r-m crossvein posterior along r 1 and r 2 to around end of costa, but veins not pale)(as in Fig. 3, 4 )............................................................................ 2

— Wing with pale spots at least at tip of costa and on r-m crossvein ( Fig. 50, 51 , 57 , 117 –169)........ 14

2(1). Distal portion of parameres fused ( Fig. 34–40 ); SCo absent from flagellomeres 9–13.................................................................................... C. ( Selfia Khalaf ) 3

— Parameres separate ( Fig. 41–43 ) or rarely fused only at base by a narrow bridge ( C. saundersi Wirth and Blanton ); SCo often on some of flagellomeres 9–13....................................... 9

3(2). Brown or yellowish to reddish yellow ( Fig. 33 female); scutellum with four setae; hind tarsomeres without apical spines; posterior margin of sternite 9 convex over basal portion of aedeagus, without median lobes; aedeagus apically trifurcate ( Fig. 34, 35 ).................................... 4

— Variously medium to dark brown or gray ( Fig. 33 female); scutellum with six to eight setae; hind tarsomeres with apical spines (Fig. 281); posterior margin of sternite 9 variously lobular, cleft, concave, or straight, not simply convex; apical portion of aedeagus entire, if appearing trifurcate, then not into separate lobes ( Fig. 36–40 )........................................................ 5

4(3). Brown; aedeagus about as long as wide, lateral arms in the form of boomerang-shaped sclerites ( Fig. 34 )......................................................... C. (Selfia) brookmani Wirth — Yellowish to reddish yellow ( Fig. 33 female); aedeagus ~2× longer than wide, lateral arms not boomerang-shaped sclerites ( Fig. 35 )................................. C. (Selfia) moabensis Phillips

5(3). Sternite 9 with pair of prominent caudomedial lobes ( Fig. 36 , 37 ); gonocoxal apodeme without a basal process............................................................................. 6

— Sternite 9 without caudomedial lobes; gonocoxal apodeme with a small posterior hooklike or pointed toothlike basal process ( Fig. 38–40 ).................................................... 7

6(5). Caudomedial lobes of sternite 9 not much longer than wide, apex of gonostylus rounded clublike ( Fig. 36 ).................................................. C. (Selfia) denningi Foote and Pratt

— Caudomedial lobes of sternite 9 ~4× longer than wide; apex of gonostylus expanded foot-shaped ( Fig. 37 )................................................... C. (Selfia) hieroglyphicus Malloch

7(5). Apex of gonostylus markedly expanded and foot-shaped; aedeagus elongate, almost parallel-sided except on basal 0.1 ( Fig. 38 )......................................... C. (Selfia) jamesi Fox

— Apex of gonostylus at most only slightly expanded, blunt; aedeagus Y-shaped ( Fig. 39, 40 )......... 8

8(7). Posterior margin of sternite 9 concave or cleft; parameres with heavily sclerotized knoblike shoulders at ~0.5, posterior median process elongate parallel-sided, fingerlike ( Fig. 39 )................................................................................ C. ( Selfia ) j acksoni Atchley

— Posterior margin of sternite 9 nearly straight; parameres without distinct shoulders, posterior median portion tapering blunt triangular ( Fig. 40 )...................... C. (Selfia) tenuistylus Wirth

9(2). Head black, thorax mostly shiny black; aedeagus V-shaped, apex bifurcate; paramere with conspicuous submedian blunt triangular process and truncate tip......... (unplaced) C. monoensis Wirth

— Light to dark brown; aedeagus Y-shaped, apex entire; paramere without submedian process, apex either a simple straight, slightly curved, or strongly hooked point ( Fig. 41, 42 ) or a long curved serrated saberform blade ( Fig. 43 )............................................................. 10

10(9). Distal portion of paramere conspicuous as a long curved heavily sclerotized serrated saberform blade ( Fig. 43 )............................................... C. (Wirthomyia) bottimeri Wirth

— Paramere relatively discreet, distal portion simple, pointed, straight to hooklike ( Fig. 41, 42 )..... 11

11(10). Eyes with interommatidial pubescence (as in Fig. 18 L . carteri, but more distinct); ventro-posterior membrane of sternite 9 bare; distal portion of paramere slender, straight to slightly curved; hind tibial comb with five or six (occasionally 4) spines........................................ 12

— Eyes bare; ventro-posterior membrane of sternite 9 spiculate; distal portion of paramere a stout hook ( Fig. 41, 42 ); hind tibial comb with four spines.......................................... 13

12(11). Parameres separate, distal tip not extending beyond tip of aedeagus...................................................................... (Saundersi group) C. atchleyi Wirth and Blanton

— Parameres fused at base by a narrow bridge, distal tip extending beyond tip of aedeagus................................................... (Saundersi group) C. saundersi Wirth and Blanton

13(11). Dark to medium brown, including legs; median process of aedeagus truncate ( Fig. 42 ).............

............................................ (Stonei group) C. werneri Wirth and Blanton — Variously brown to yellowish brown, legs yellow to straw-colored; median process of aedeagus blunt

( Fig. 41 )................... (Stonei group) C. stonei James , C. mortivallis Wirth and Blanton,

C. owyheensis Jones and Wirth

14(1). Flagellomeres 9–10 each only half as large as 8 ( Fig. 59 ); SCo absent from flagellomeres 2–13; eyes with interommatidial pubescence (as in Fig. 18 L . carteri); minute, wing length 0.7 mm ....................................................................... (Leoni group) C. reevesi Wirth

— Flagellomeres 9–10 normal, each larger than 8 (Fig. 251, 252); SCo on some of flagellomeres 2–13; eyes bare (except sometimes on C. boydi Wirth and Mullens and C. chiopterus [Meigen], which have normal antennae and SCo on flagellomeres 11–13); wing length usually> 1 mm .............. 15

15(14). At least distal portion of cell r 2 within a pale spot ( Fig. 117–125 ); tergite 9 posterior margin either convex with or without tiny apicolateral processes not extending beyond median portion ( Fig. 65–72 ) or broadly concave with square shoulders ( Fig. 63 ) or with short apicolateral processes ( Fig. 64 ); hind tibial comb with 5–7 spines; SCo absent from 2–9, often 10...................... 16

— Cell r 2 entirely dark ( Fig. 50, 51 , 126–169); tergite 9 posterior margin concave, with distinct apicolateral processes extending beyond medial portion ( Fig. 73 –116); hind tibial comb often with four spines; SCo often present on some of 2–9..................................................... 27

16(15). Mesal surface of gonocoxite clothed with short pale inconspicuous setae ( Fig. 63–65 ); ventral apodeme of gonocoxite ~2× longer than dorsal apodeme, slender, 5–7× longer than basal width ( Fig. 63–65 ); tergite 9 broadly convex without apicolateral processes ( Fig. 65 ), or broadly concave with square corners ( Fig. 63 ) or short triangular apicolateral processes ( Fig. 64 ); wing pattern of pale spots faint to absent from distal third ( Fig. 117, 118 ); hind tibial comb with five spines....................................................................................... C. ( Avaritia Fox ) 17

— Mesal surface of gonocoxite clothed with short dark conspicuous setae or spinules ( Fig. 66–72 ); ventral apodeme of gonocoxite not longer than dorsal apodeme, strongly tapered, pointed, 1–2× as long as basal width; tergite 9 tapering convex with tiny apicolateral processes (rarely absent) not extending past mesal portion of tergite ( Fig. 66–72 ); wing pattern of pale spots extensive into distal portion of wing ( Fig. 119–125 ); hind tibial comb with six or seven spines (Fig. 286).................................................... C. ( Silvicola Mirzaeva and Isaev ) 20

17(16). Sternite 9 posterior margin with broad caudomedian excavation; tergite 9 posterior margin concave, with blunt apicolateral processes ( Fig. 64 ) or square lateral corners ( Fig. 63 ); aedeagus with sclerotized membrane between arms and sclerotized anterior-directed point at base of median process; aedeagal ratio <0.5 ( Fig. 63, 64 )....................................................... 18

— Sternite 9 posterior margin notched; tergite 9 posterior margin convex, without apicolateral processes; aedeagus without sclerotized membrane between basal arms and without sclerotized anterior-directed point at base of median process; aedeagal ratio>0.5 ( Fig. 65 ).............. 19

18(17). Basal arms of aedeagus forming pointed lateral shoulders adjacent to median process; tergite 9 with square apicolateral corners ( Fig. 63 ).................. C. (Avaritia) boydi Wirth and Mullens

— Aedeagus without lateral shoulders; tergite 9 with short broad apicolateral processes ( Fig. 64 )............................................................... C. (Avaritia) chiopterus (Meigen)

19(17). Aedeagal ratio>0.7; tip of aedeagus convex, sides bare ( Fig. 56 ); paramere tip with minute hairs, which may require an oil-immersion lens to see.................... C. (Avaritia) obsoletus (Meigen)

— Aedeagal ratio <0.7; tip of aedeagus concave, sides usually spinulose; paramere tip simple............................................................... C. (Avaritia) sanguisuga (Coquillett)

20(16). Major bend of paramere near its middle; aedeagal ratio ~0.3; coastal tidelands.......................................................................... C. (Silvicola) tristriatulus Hoffman

— Major bend of paramere at a fifth to a third of length from basal end ( Fig. 66–72 ); aedeagal ratio often 0.4–0.6; inland..................................................................... 21

21(20). Hind tarsomeres with apical spines (Fig. 282) (species D and E likely key this path, but males unknown)................................................................................... 22

— Hind tarsomeres without apical spines.................................................... 24

22(21). Hind tibiae with distinct basal pale band ~2× longer than wide (Fig. 286)....................................................................... C. (Silvicola) sierrensis Wirth and Blanton

— Hind tibiae without pale band, or sometimes with indistinct band not longer than wide......... 23

23(22). Distal wing pattern indistinct, hourglass-shaped distal dark spot in r 3 usually not darker than other distal dark patches ( Fig. 123 ); gonocoxite with moderately strong black setae on mesal surface; aedeagal ratio <0.4 ( Fig. 70 )............................. C. (Silvicola) neomontanus Wirth

— Distal wing pattern distinct, hourglass-shaped distal dark spot in r 3 darker than other distal dark patches ( Fig. 119 ); gonocoxite with fine brown setae on mesal surface; aedeagal ratio>0.4 ( Fig. 66 )................................................. C. (Silvicola) cockerellii (Coquillett)

24(21). Fore tarsomeres 1 or 2 with apical spines (as in Fig. 285 C. calexicanus ); dark spot over portions of r 1 and r 2 not broadly connected to dark spot over M fork ( Fig. 120 )............................................................................ C. (Silvicola) freeborni Wirth and Blanton

— Fore tarsomeres without apical spines; dark spot over r 1 and r 2 broadly connected to dark spot over M fork ( Fig. 121, 122, 124 ).............................................................. 25

25(24). Median process of aedeagus broadly triangular, tapering to broad blunt tip ( Fig. 71 ) ; tergite 9 often without apicolateral processes; palpal segment 3 with patch of scattered sensilla instead of pit; known only from the low southern deserts of California and Nevada ... C. (Silvicola) saltonensis Wirth

— Median process of aedeagus more abruptly narrowing on distal half to slender fingerlike tip ( Fig. 68, 69 ); tergite 9 with apicolateral processes; palpal segment 3 with distinct sensory pit (Fig. 241); widespread............................................................................. 26

26(25). Hind tibiae with distinct basal pale band as in Fig. 286 C. sierrensis ; distal pale wing spots with distinct borders ( Fig. 122 )............................ C. (Silvicola) neofagineus Wirth and Blanton

— Hind tibiae without pale band; distal pale wing spots with indistinct borders ( Fig. 121 )............................................................ C. (Silvicola) lahontan Wirth and Blanton

27(15). Parameres fused at base ( Fig. 46, 48 ); hind tibial comb with six or seven spines; fore and hind tarsomeres 1, 2 with apical spines................................ C. ( Monoculicoides Khalaf ) 28

— Parameres separate ( Fig. 73 –116); hind tibial comb with four or five spines (rarely six on C. californiensis and C. kettlei ); fore and hind tarsomeres without apical spines (except on C. usingeri, Palmerae group species, and unplaced species F)........................................ 30

28(27). Aedeagus apex entire; wing with faint pattern (as in Fig. 173 female); scutum yellowish brown without dark spots; legs pale brown without apparent banding........................................................................... C. (Monoculicoides) grandensis Grogan and Phillips

— Aedeagus apex deeply bifurcate ( Fig. 47, 49 ); wing with distinct pattern ( Fig. 50, 51 ); scutum brown with dark spots at setal bases (as in Fig. 55 ); legs brown with distinct pale banding............... 29

29(28). Ventral surface of aedeagus spiculate ( Fig. 47 ).... C. (Monoculicoides) sonorensis Wirth and Jones

— Aedeagus bare ( Fig. 49 )....................... C. (Monoculicoides) occidentalis Wirth and Jones

30(27). Aedeagal arms each with sclerotized spurlike process on posterior margin ( Fig. 73–78 )......................................................................... C. ( Diphaomyia Vargas ) 31

— Aedeagal arms simple ( Fig. 79 –116)....................................................... 35

31(30). Paramere without submedian lobe ( Fig. 77 ); posterior process of footlike ventral apodeme of gonocox- ite difficult to see, often unapparent ( Fig. 78 )............................................ 32

— Paramere with submedian lobe; posterior process of footlike ventral apodeme of gonocoxite distinct ( Fig. 73–76 )........................................................................ 33

32(31). Wing pattern extensive with distinct distal pale spots in r 3, m 1, m 2 (as in Fig. 194 female)................................................................... C. (Diphaomyia) bergi Cochrane

— Wing pattern without distal pale spots in r 3, m 1, m 2 (Fig. 130)....... C. (Diphaomyia) salihi Khalaf

33(31). Distal half of r 3 mostly dark, with small distinct pale spot entirely within distal 0.2 of cell (Fig. 126, 127)..... C. (Diphaomyia) defoliarti Atchley and Wirth , C. (Diphaomyia) haematopotus Malloch

— Distal pale spot in r 3 central in distal half or filling most of distal third of cell, but may be faint or absent (Fig. 128, 129)....................................................................... 3 4

34(33). Wing pattern extensive with diffuse pale spots in r 3, m 1, m 2 (Fig. 128); SCo pattern 1, (6), (7), 8, 9................................................ C. (Diphaomyia) inyoensis Wirth and Blanton

— Distal pales spots absent from r 3, m 1, m 2 (Fig. 129); SCo pattern 1, (10), 11–13.................................................................... C. (Diphaomyia) erikae Atchley and Wirth

35(30). Ventral apodeme of gonocoxite complex, with two widely divergent processes, footlike ( Fig. 79–87 )..................................................................................... 36

— Ventral apodeme of gonocoxite simple ( Fig. 88 –116)......................................... 45 36(35). Parameres with submedian lobe ( Fig. 79, 80 ); SCo absent from flagellomeres 11–13.............. 37

— Parameres without submedian lobe ( Fig. 81–87 ); SCo present on 12–13........................ 40

37(36). Hind tibial comb with four spines........................................................ 38

— Hind tibial comb with five or six spines........................ C. ( Haematomyidium Goeldi ) 39

38(37). Aedeagus tip truncate, spinose ( Fig. 79 ); inland................ (Mohave group) C. mohave Wirth

— Aedeagus tip rounded, without spines; coastal....... (Mohave group) C. hoguei Wirth and Moraes

39(37). Two separate pale spots in m 1 (Fig. 133).. C. (Haematomyidium) kettlei Breidenbaugh and Mullens

— Three separate pale spots in m 1 (Fig. 134, 135)......... C. (Haematomyidium) stellifer (Coquillett)

40(36). Distal pale spots distinct in r 3, m 1, m 2 (Fig. 136–139)......................................... 41

— Distal pale spots faint or absent from r 3, m 1, m 2 (Fig. 140, or as in Fig. 233, 234 females).......... 42

41(40). Pale band or patch along posteromedian margin of anal cell (Fig. 136, 137); sternite 9 caudomedian excavation ~0.3 as deep as wide, about evenly curved; heavily sclerotized basal arms of aedeagus with basal ends sharply bent laterally, posteromedial ends connected only by unsclerotized or barely sclerotized membrane, aedeagal ratio ~0.7 ( Fig. 81–83 )........................................................................... (Piliferus group) C. cavaticus Wirth and Jones

— No pale band or patch on posteromedian margin of anal cell (Fig. 138, 139); sternite 9 caudomedian excavation ~0.5 as deep as wide, about three-sided; heavily sclerotized basal arms of aedeagus with basal ends bent more ventrally, posteromedial ends fused by moderate sclerotization, aedeagal ratio 0.45–0.6 ( Fig. 84, 85 ) ( C. lophortygis Atchley and Wirth may key here too, but male unknown)........................................ (Piliferus group) C. doeringae Atchley

42(40). Hind tibial comb with five spines...................... (Piliferus group) C. unicolor (Coquillett)

— Hind tibial comb with four spines..................................................... 43

43(42). Pale yellowish-brown............................................. (Piliferus group) species A

— Dark brown............................................................................ 44

44(43). Faint distal pale spots in r 3, m 1, m 2, straddling midportions of M 1 and M 2 (as in Fig. 233 female)............................................. (Piliferus group) likely species B (male unknown)

— Distal pale spots absent from r 3, m 1, m 2, sometimes from midportions of M 1 and M 2 (as in Fig. 234 female) (Piliferus group) likely C. chewaclae Glick and Mullen (male unknown)

45(35). Distal half of paramere with elongate curved pointed thornlike process and distal fringe of spines ( Fig. 88 ); SCo pattern 1, (7), 8–10...................... (Limai group) C. luglani Jones and Wirth

— Distal half of paramere simple or with only apical spines ( Fig. 89 –116); SCo pattern 1, (2), (3), (4), (5), (11), 12, (13)........................................................................ 46

46(45). Ventral apodeme of gonocoxite inconspicuous as an obtuse antero-medially directed toothlike bump ( Fig. 89 ); distal pale wing spots ovoid, no isolated pale spots straddling M 1 or M 2 (Fig. 142, 143)............................................. C. (Beltranmyia) crepuscularis Malloch

— Ventral apodeme of gonocoxite slender elongate, 3–10× longer than median width,>0.5 as long as dorsal apodeme ( Fig. 90 –116); distal pale wing spots often irregular or absent, isolated pale spots often straddling M 1 or M 2 (Fig. 144–169).................................................... 47

47(46). Antennae, palpi, and wings feminized (Fig. 147, 251); genitalia normal ( Fig. 93 )................. 48

— Antennae, palpi, and wings normal (Fig. 252).............................................. 49

48(47). Wing pattern extensive with distal pale spots (Fig. 147); tergite 9 apicolateral processes long and slender, ~5× longer than wide, projecting nearly parallel caudally ( Fig. 93 ).................................................................................. (Palmerae group) C. utahensis Fox

— Wing pattern greatly reduced, with pale spots only at tip of costa and on r-m crossvein; tergite 9 apicolateral processes stout, ~2× longer than wide, broadly flaring laterally ~30°........................................................................... (Palmerae group) C. leechi Wirth

49(47). Hind tibial comb with five spines; distal pale spots present in cua 1 and anal cell, absent from or barely discernable in r 3, m 1, m 2; no pale spots straddling M 1 or M 2; basal arms of aedeagus slightly curved, slender, each ~10× longer than wide, distal process simple, tapering to narrow blunt tip; parameres without apical spines........................... (unplaced) C. nanellus Wirth and Blanton

— Hind tibial comb often with four spines; distal pale spots sometimes absent from or often present and distinct in in r 3, m 1, m 2, cua 1, anal cells; pale spots often straddling M 1 or M 2; basal arms of aedeagus often more robust ( Fig. 90–92 , 94 –116), distal process often broad or truncate ( Fig. 90–92 , 104 –116); parameres sometimes with apical spines ( Fig. 90 , 94–96 )................................. 50

50(49). Fore tarsomeres with apical spines (Fig.285)(uncertain for C. novamexicanus and C. oregonensis );mid and hindfemorawithoutpalebands;nopalespotstraddlingmidportionofM 1 (Fig.144–146);SCooftenabsent from flagellomere 13; gonostylus often abruptly bent medially ~90° at ~0.2, L-shaped ( Fig. 92 )... ................................................................................... 51

— Fore tarsomeres without apical spines; mid or hind femora often with basal or subapical pale band; pale spot often straddling at ~0.5 on M 1 (Fig. 154–167); SCo on flagellomere 13; gonostylus more or less evenly curved, not L-shaped.............................................. 55

51(50). Median process of aedeagus with wide, usually sharp, shoulders abruptly constricting to a narrow tip ( Fig. 92 )............................................ (Palmerae group) C. palmerae James

— Median process of aedeagus simple, without shoulders ( Fig. 90, 91 )............................ 52

52(51). Aedeagus broadly Y-shaped; median process slender, nearly parallel-sided, <0.1 as wide as basal arm spread, ~5× longer than median width.... (Palmerae group) C. oregonensis Wirth and Rowley

— Aedeagus tapering with almost a V-shape; median process of aedeagus broad, tapering to narrow or broad tip ( Fig. 90, 91 )................................................................ 53

53(52). Aedeagus tapering to broad tip ~0.4 as wide as basal arm spread ( Fig. 91 ); distal pale wing spots faint or absent (Fig. 145)............................ (Palmerae group) C. hawsi Wirth and Rowley

— Aedeagus tapering to narrow tip ~0.2 as wide as basal arm spread ( Fig. 90 ); wing with extensive distal pale spots, but may be diffuse (Fig. 144)................................................ 54

54(53). Gonostylus abruptly bent medially ~90° at ~0.2, L-shaped (as in Fig. 92 C . palmerae); scutellum about same brown as scutum (as in Fig. 221 C. utahensis )....... (Palmerae group) C. novamexicanus Atchley

— Gonostylus evenly curved, not abruptly bent ( Fig. 90 ); scutellum more yellowish than scutum (Fig. 221).................................. (Palmerae group) C. calexicanus Wirth and Rowley

55(50). Distal portion of paramere with 3–6 spines ( Fig. 94–96 )..................................... 56

— Distal portion of paramere simple, tapering to filamentous tip ( Fig. 97 –116).................... 59

56(55). Wing pattern extensive (as in Fig. 158, 159, 161, 162)............ C. (Drymodesmyia) bakeri Vargas

— Wing pattern reduced, distal pale spots absent from r 3, m 1, often from m 2, cua 1, anal cell (Fig. 148– 150).................................................. C. ( Silvaticulicoides Glukhova ) 57

57(56). Distal pale spots present in cua 1, anal cell, often m 2 (Fig. 148); paramere apex with 3–4 spines ( Fig. 94 )................................................... C. (Silvaticulicoides) sublettei Atchley

— Distal pale spots absent (Fig. 149, 150); paramere apex with 4–6 spines ( Fig. 95, 96 ).............. 58

58(57). Hind tarsomeres with apical spines (as in Fig. 283 C. palmerae ); ventral apodeme of gonocoxite slender, 2× longer than dorsal apodeme; paramere apex with five or six spines ( Fig. 95 ).................................................... C. (Silvaticulicoides) usingeri Wirth

— Hind tarsomeres without apical spines; ventral apodeme of gonocoxite triangular tapering, about as long as dorsal apodeme; paramere apex with four spines ( Fig. 96 )............................................................. C. (Silvaticulicoides) vetustus Breidenbaugh and Mullens

59(55). Distal pale spots absent from r 3, m 1, m 2, or small, faint, and only in apices of cells; pale spots not straddling M 1 or M 2 (Fig. 151–153)......................................................... 60

— Distal pale spots extensive, often prominent, in r 3, m 1, m 2; pale spots straddling M 2, often M 1 (Fig. 154–169)........................................................................... 62

60(59). Pale spots absent from above CuA fork, distally from r 3, m 1, m 2 (Fig. 151)....................................................................... (unplaced) C. posoensis Wirth and Blanton

— Distinct pale spot anterior to CuA fork, sometimes faint distally in r 3, m 1, m 2 (Fig. 152, 153)............................................................... C. ( Sensiculicoides Shevchenko ) 61

61(60). Aedeagal ratio ~0.5; median process stout at base, tapering to blunt tip ( Fig. 98 )................................................................... C. (Sensiculicoides) kibunensis Tokunaga

— Aedeagal ratio ~0.4; median process narrow, parallel-sided on distal half ( Fig. 99 )....................................................................... C. (Sensiculicoides) travisi Vargas

62(59). Hind tibial comb with five, occasionally six, spines; hind tibiae with prominent subapical pale band; distinct pale spots straddling at ~0.3 on M 1; distal pale spot in r 3 double, bilobed about C- or 8-shaped, oriented perpendicular to M 1, central in distal half of cell (Fig. 154–156)................................................................................. C. ( Amossovia Glukhova ) 63

— Hind tibial comb with four spines; subapical pale band often absent from or indistinct on hind tibiae, but if distinct then no pale spot straddling M 1; distal pale spot in r 3 often single or U-shaped extending into apex of cell (Fig. 157–169)..................................................... 66

63(62). Yellowish coppery-brown; fore and mid tibiae with subapical pale band; aedeagus nearly V-shaped, median process broad triangular with lateral caudally directed spines on midsection, tapering to small simple blunt tip ( Fig. 100 )............. C. (Amossovia) californiensis Wirth and Blanton

— Dark brown; fore and mid tibiae without subapical pale band; aedeagus Y-shaped, median process slen- der with truncate or pointed tip or with lateral filaments ( Fig. 101, 102 ).................... 64

64(63). Hind femora with prominent subapical pale band; median process of aedeagus narrow parallel-sided with truncate tip........................................ C. (Amossovia) pecosensis Wirth

— Hind femora entirely dark or with only faint pale band; median process of aedeagus with pointed tip ( Fig. 101, 102 )....................................................................... 65

65(64). Hind femora unbanded; median process of aedeagus simple ( Fig. 101 )....................................................................... C. (Amossovia) cochisensis Wirth and Blanton

— Hind femora with faint basal and subapical pale bands; median process of aedeagus with lateral pair of subapical filaments ( Fig. 102 ).......................... C. (Amossovia) oklahomensis Khalaf

66(62). Ventro-posterior membrane of sternite 9 spiculate (Fig. 110–112); distal pale spot in r 3 simple (Fig. 165) or if irregular then not U- or distinctly 8-shaped (Fig. 164, 166).................................................................................. C. ( Drymodesmyia Vargas ) (part) 67

— Ventro-posterior membrane of sternite 9 bare ( Fig. 103–109 , 113–116); distal pale spot in r 3 irregular (Fig. 157–163) or simple (Fig. 167–169)................................................. 69

67(66). SCo pattern 1, 11–13; tip of median process of aedeagus papilliform, <0.1 as wide as arm spread (Fig. 110)..................................... C. (Drymodesmyia) insolatus Wirth and Hubert

— SCo pattern 1, 12–13; tip of median process of aedeagus rounded blunt or truncate,>0.1 as wide as arm spread (Fig. 111, 112)................................................................ 68

68(67). Ventro-posterior membrane of sternite 9 densely spiculate; ventral apodeme of gonocoxite ~0.5 as long as dorsal apodeme; aedeagal ratio ~0.6 (Fig. 111).................................................................................. C. (Drymodesmyia) arizonensis Wirth and Hubert

— Ventro-posterior membrane of sternite 9 with a few spicules; ventral apodeme of gonocoxite about as long as dorsal apodeme; aedeagal ratio 0.3–0.4 (Fig. 112)............................................................................. C. (Drymodesmyia) copiosus Root and Hoffman

69(66). Distal pale spot in r 3 distinctly double, bilobed, or mushroom-shaped, oriented perpendicular to M 1, extending to anterior margin of wing, not extending into distal 0.2 of cell; apices of M 1, M 2 pale, forming small pale spots at wing margin; distinct isolated pale spot straddling at ~0.3 on M 1 (Fig. 158–163)............................................ C. ( Drymodesmyia Vargas ) (part) 70

— Distal pale spot in r 3 single, ovoid, or quadrate (Fig. 167–169) to irregular U-shaped extending into distal 0.3 of cell (Fig. 157); apices of M 1, M 2 dark or no more pale than interior portions of veins and not forming apical spots; distinct isolated pale spot often absent from M 1 (Fig. 167–169) 74

70(69). Aedeagus V-shaped, straight-sided until broad flaring truncate tip ~0.25 width of arm spread; aedeagal ratio ~0.3 ( Fig. 107, 108 )............................................................. 71

— Aedeagus more Y-shaped, constricting at base of median process, tapering to truncate or rounded tip <0.2 width of arm spread; aedeagal ratio 0.4–0.5 ( Fig. 104–106, 109 )....................... 72

71(70). Smaller, mean wing length 0.86 mm; pale wing markings more extensive (Fig. 161); Chihuahuan Desert from southeastern Arizona to Texas and probably Mexico............................................................................... C. (Drymodesmyia) jonesi Wirth and Hubert

— Larger, mean wing length 0.98 mm; pale wing markings less extensive (Fig. 162); Sonoran Desert and Colorado Plateau from Baja California to California, Utah, Arizona, and New Mexico ..................................................... C. (Drymodesmyia) sitiens Wirth and Hubert

72(70). Two pale spots of about same intensity in distal half of anal cell, often merged with broad pale area along wing margin; pale apices of M 1 and M 2 broadly flaring (Fig. 158, 159); aedeagal ratio>0.43; narrowest diameter of paramere before first ~90° bend in apical half> 0.0027 mm ( Fig. 104, 105 )....................................... C. (Drymodesmyia) butleri Wirth and Hubert

— Only one distinct pale spot in distal half of anal cell, usually not merging with fainter pale area along wing margin; pale apices of M 1 and M 2 not flaring (Fig. 160, 163); either aedeagal ratio <0.42 or narrowest diameter of paramere before first ~90° bend in apical half <0.0026 mm ( Fig. 106, 109 ).. 73

73(72). Aedeagal ratio>0.43; narrowest diameter of paramere before first ~90° bend in apical half <0.0026 mm ( Fig. 106 )................................ C. (Drymodesmyia) cacticola Wirth and Hubert

— Aedeagal ratio <0.42; narrowest diameter of paramere before first ~90° bend in apical half> 0.0027 mm ( Fig. 109 )................................. C. (Drymodesmyia) torridus Wirth and Hubert

74(69). Median process of aedeagus broad, tapering abruptly at ~0.6 to slender blunt fingerlike tip ( Fig. 103 ); pale spot in distal half of r 3 broadly U-shaped, extending from near anterior wing margin through midportion of distal half of cell along M 1 to apex of cell (Fig. 157)................................................ (Daedalus group) C. pampoikilus Macfie

— Median process of aedeagus evenly narrowing to broad truncate or blunt tip (Fig. 113–116); distal pale spot in r 3 simple, ovoid or quadrate to slightly C-shaped, not broadly U-shaped or extending to apex of cell (Fig. 167–169, or as in Fig. 195 female)............................................ 75

75(74). Ventral apodeme of gonocoxite much more slender and often shorter than the broad, pointed dorsal apodeme (Fig. 113); distal pale spot in r 3 elongate ovoid, tapering distally, oriented parallel with M 1 (Fig. 167)................................ C. (Drymodesmyia) ryckmani Wirth and Hubert

— Ventral apodeme of gonocoxite as long as or longer and only a little more slender than dorsal apodeme (Fig. 114–116); distal pale spot in r 3 ovoid or quadrate to slightly C-shaped, oriented oblique or perpendicular to M 1 (Fig. 168, 169, or as in Fig. 195 female)............. 76

76(75). Aedeagus Y-shaped, aedeagal ratio ~0.6, median process narrow, nearly parallel-sided (Fig. 116); southern, rare........................................... (Daedalus group) C. daedalus Macfie

— Aedeagus more V-shaped, aedeagal ratio 0.3–0.4, median process more tapered (Fig. 114, 115); widespread treehole species............................................................... 77

77(76). Ventral apodeme of gonocoxite about as long as dorsal apodeme (Fig. 115); yellow (in alcohol); distal spot in r 3 oriented perpendicular to M 1; pale spot at tip of costa bent back beneath r 2 (Fig. 169); pale leg-banding subapical on mid femora, basal and subapical on hind femora, subapical on hind tibiae (Fig. 250)........................................... C. (Drymodesmyia) hinmani Khalaf

— Ventral apodeme of gonocoxite up to 2× longer than dorsal apodeme (Fig. 114); brown (in alcohol); distal spot in r 3 oriented oblique to M 1; pale spot at tip of costa not bent back beneath r 2 (Fig. 168); pale leg-banding reduced: absent subapically from mid femora, basally and subapically from hind femora, subapically from hind tibiae (Fig. 250)............. C. (Drymodesmyia) byersi Atchley

Key to Culicoides Females

(Measurement and ratio definitions are in the “Morphological terms and data” section of Materials and Methods)

1. Wing without pattern of pale spots(often false pale areas caused by paucity of macrotrichia in base of wing and from r-m crossvein posterior along r 1 and r 2 to around end of costa, but veins not pale) ( Fig. 3, 4 ).................................................................................... 2

— Wing with pale spots at least at tip of costa and on r-m crossvein ( Fig. 2 , 56 , 170–240)........... 14

2(1). Three long unsclerotized or lightly sclerotized fingerlike spermathecae, usually unrecognizable even in slide preparations.................................................... C. ( Selfia Khalaf ) 3

— Two distinctly sclerotized ovoid or pyriform spermathecae, with vestigial third ( Fig. 44, 45 )....... 7

3(2). Brown or yellowish to reddish yellow ( Fig. 33 ); scutellum with four setae if brown, eight setae if yellow; hind tarsomeres without apical spines.................................................. 4

— Variously medium to dark brown or gray ( Fig. 33 ); scutellum with 8–14 setae; hind tarsomeres with apical spines (Fig. 281)................................................................ 5

4(3). Brown; scutellum with four setae.................................. C. (Selfia) brookmani Wirth

— Yellowish to reddish yellow ( Fig. 33 ); scutellum with eight setae....... C. (Selfia) moabensis Phillips

5(3). SCo pattern 1, 5–8........................................ C. (Selfia) denningi Foote and Pratt

— SCo pattern 1, (2), (3), 4–8................................................................ 6

6(5). SCo pattern 1, 4–8................................... C. (Selfia) hieroglyphicus Malloch (part)

— SCo pattern 1, (2), 3–8...... C. (Selfia) hieroglyphicus Malloch (part), C. (Selfia) jacksoni Atchley , C. (Selfia) jamesi Fox , C. (Selfia) tenuistylus Wirth

7(2). Head black, thorax mostly shiny black; SCo pattern 1, 12–13...... (unplaced) C. monoensis Wirth

— Light to dark brown; SCo on some of flagellomeres 2–11....................................... 8

8(7). Eyes with prominent interommatidial pubescence (as in Fig. 18 L . carteri, but more distinct); man- dible either with 14–20 teeth or without teeth; anterior wing veins dark and thick ( Fig. 4 )..... 9

— Eyes bare; 9–13 mandibular teeth; anterior wing veins more normal ( Fig. 3 ).................... 10

9(8). Eyes separated by <1 ommatidium diameter; mandibular teeth vestigial.................................................................. (Saundersi group) C. atchleyi Wirth and Blanton

— Eyes separated by ~2 ommatidium diameters; mandible with 14–20 teeth................................................................ (Saundersi group) C. saundersi Wirth and Blanton

10(8). Hind tibial comb with five or six spines; eyes separated by <1 ommatidium diameter; SCo pattern 1, (2), 3, (4), 5–8; posterior portion of sternite 8 medially concave but not cleft, with pair of pointed submedian posterior projections ( Fig. 44 ).................. C. (Wirthomyia) bottimeri Wirth

— Hind tibial comb with four spines; eyes separated by>1 ommatidium diameter; SCo on at least 8–12; posterior portion of sternite 8 medially cleft, with blunt submedian posterior projections ( Fig. 45 ) ................................................................................... 11

11(10). SCo pattern 1, 8–12, sometimes also on flagellomere 5 or 7.................................................................................. (Stonei group) C. werneri Wirth and Blanton

— SCo pattern 1, 5–12 or 1, (2), 3–12......................................................... 12

12(11). SCo pattern 1, (2), 3–12......................................... (Stonei group) C. stonei James

— SCo pattern 1, 5–12..................................................................... 13

13(12). Wing length <1.06 mm; length of flagellomeres 7+8 divided by length of flagellomere 9 <1.50; length of palpal segment 3 divided by length of flagellomere 9 <1.90; length of proboscis divided by length of flagellomere 9 <3.95....................... (Stonei group) C. mortivallis Wirth and Blanton

— Wing length> 1.06 mm; length of flagellomeres 7+8 divided by length of flagellomere 9>1.50; length of palpal segment 3 divided by length of flagellomere 9>1.90; length of proboscis divided by length of flagellomere 9>3.95......................... (Stonei group) C. owyheensis Jones and Wirth

14(1). Flagellomeres 9–10 much shorter and narrower than any others ( Fig. 58 ); eyes with interommatidial pubescence (as in Fig. 18 L . carteri); one fully developed spermatheca with vestigial second; minute, wing length ≤ 0.8 mm ...................................... (Leoni group) C. reevesi Wirth

— Flagellomeres 9–10 normal, each as large as or larger than any of 2–8 (Fig. 253); eyes bare (except sometimes on C. boydi and C. chiopterus , which have normal antennae and two fully developed spermathecae); either one developed spermatheca rarely with vestigial second ( Fig. 52–54 ) or two developed spermathecae with vestigial third (Fig. 255–278); wing length usually> 1 mm ..... 15

15(14). One fully developed spermatheca ( Fig. 52–54 ), sometimes with vestigial second; r 2 entirely dark; distal wing pattern extensive and either with one isolated ovoid distal pale spot in r 3 and two isolated ovoid pale spots in m 1 and no spots straddling M 1 or M 2 (Fig. 170–172) or of dark irregular curves, extensively interconnected pale streaks, and zigzags (Fig. 173, 174)........................ 16

— Two fully developed spermathecae with vestigial third (Fig. 255–278); r 2 often in a pale spot (Fig. 175–186); only one or no isolated ovoid pale spot within m 1 not straddling a vein (Fig. 175–186, 190–215, 217–240) or isolated distal spots in r 3 irregular, not ovoid (Fig. 187–203, 207–210, 216). .... .... ... .... .... .... .... .... .... ... .... .... .... .... .... ... .... .... .... .... ..... 19

16(15). Distal wing pattern of ovoid pale spots (Fig. 170–172); eyes separated by ~1 ommatidium diameter; hind tarsomeres without apical spines; spermatheca ovoid, opening ~0.1 the diameter of the spermatheca ( Fig. 52 ).......................................... C. (Beltranmyia) crepuscularis Malloch

— Distal wing pattern more of dark irregular curves, pale streaks, and zigzags than ovoid spots (Fig. 173, 174); eyes widely separated by>2 ommatidium diameters; hind tarsomeres with apical spines (as in Fig. 281–284); spermatheca with opening>0.5 the diameter of the spermatheca, often U-shaped ( Fig. 53, 54 ).............................................. C. ( Monoculicoides Khalaf ) 17

17(16). Spermatheca ovoid ( Fig. 53 ); wing with faint pattern (Fig. 173); scutum yellowish brown without dark spots; legs pale brown without apparent banding; mandibular teeth vestigial....................................................... C. (Monoculicoides) grandensis Grogan and Phillips

— Spermatheca U-shaped ( Fig. 54 ); wing with prominent pattern (Fig. 174); scutum brown with dark spots at seta bases ( Fig. 55 ); legs brown with prominent pale banding; mandibular teeth well-developed................................................................................... 18

18(17). Immatures in aquatic alkaline or highly organic soils (females morphologically indistinguishable)......................................... C. (Monoculicoides) sonorensis Wirth and Jones

— Immatures in aquatic highly saline soils (females morphologically indistinguishable)....................................................... C. (Monoculicoides) occidentalis Wirth and Jones

19(15). At least distal portion of cell r 2 within a pale spot (Fig. 175–186); hind tibial comb with 5–7 spines (Fig. 286); eyes contiguous (except for C. tristriatulus ); SCo on at least 1, 10–13.................. 20

— Cell r 2 entirely dark or with only the vein at distal tip of r 2 in a pale spot (Fig. 187–240); hind tibial comb often with four spines; if with five spines, then: eyes separated, or SCo absent from 11–13, or pale wing spots distinct and extensive (Fig. 187–193) or greatly reduced (Fig. 227)............... 33

20(19). Wing pattern of pale spots faint to absent from distal third (Fig. 175–177); palpal segment 3 with distinct round sensory pit ≤0.5 width of segment; hind tibial comb with five spines.. C. ( Avaritia Fox ) 21

— Wing pattern of pale spots extensive into apex (Fig. 178–186); palpal segment 3 with scattered sensilla or broad shallow pit>0.5 width of segment; hind tibial comb with six or seven spines (Fig. 286).................................................... C. ( Silvicola Mirzaeva and Isaev ) 24

21(20). Wing length usually <0.9 mm; pale spot overlapping only distal tip of r 2 (Fig. 175); five or six scutellar setae; SCo pattern 1, (9), 10–13; known from deserts of Southern California and Baja California ..................................................... C. (Avaritia) boydi Wirth and Mullens

— Wing length usually> 0.9 mm; pale spot overlapping distal half of r 2 (Fig. 176, 177); four scutellar setae; SCo pattern 1, 9–13; rare or absent from arid warm areas of southwestern North America... 22

22(21). Length of palpal segment 3 <1.5× that of palpal segment 5; mandible with 6–10 teeth; proboscis ratio <0.75.................................................. C. (Avaritia) chiopterus (Meigen)

— Length of palpal segment3>1.5×that of palpal segment5;mandible with11–17teeth;proboscis ratio>0.75................................................................................... 23

23(22). Vein CuA 1 with 7–14 macrotrichia; cell cua 1 with>4 macrotrichia, including those near the margin........................................................ C. (Avaritia) obsoletus (Meigen)

— Vein CuA 1 with 1–6 macrotrichia; cell cua 1 with <4 macrotrichia, including those near the margin.................................................... C. (Avaritia) sanguisuga (Coquillett)

24(20). Eyes separated by>1 ommatidium diameter; palpal ratio>3.5; coastal tidelands........................................................................ C. (Silvicola) tristriatulus Hoffman

— Eyes contiguous; palpal ratio <3.5; inland.................................................. 25

25(24). Hind tarsomeres with apical spines (Fig. 282).............................................. 26

— Hind tarsomeres without apical spines.................................................... 30

26(25). Palpal segment 3 with scattered sensilla instead of pit (Fig. 245)............................... 27

— Palpal segment 3 with regular roundish pit (as in Fig. 242 C. lahontan ) or irregular subdivided pit (Fig. 246).......................................................................... 29

27(26). Distal wing pattern faint (Fig. 182); eyes contiguous for>0.3 ommatidium diameter; SCo on some of flagellomeres 2–8...................................... C. (Silvicola) neomontanus Wirth

— Distal wing pattern relatively distinct (Fig. 178, 184); eyes contiguous for <0.3 ommatidium diameter (Fig. 287); SCo rarely on any of flagellomeres 2–8....................................... 28

28(27). Hind tibiae with distinct basal pale band ~2× longer than wide (Fig. 286)....................................................................... C. (Silvicola) sierrensis Wirth and Blanton

— Hind tibiae without basal pale band, sometimes with indistinctly pale base........................................................................... C. (Silvicola) cockerellii (Coquillett)

29(26). SCo pattern 1, (5), (6), 7, (8), 9–13; sensory pit on palpal segment 3 irregular, subdivided (Fig. 246)................................................................... C. ( Silvicola ) species D

— SCo pattern 1, 9–13; sensory pit on palpal segment 3 regular roundish (as in Fig. 242 C. lahontan )..................................................................... C. ( Silvicola ) species E

30(25). Fore tarsomeres 1 or 2 with apical spines (as in Fig. 285 C. calexicanus ); distal hourglass-shaped dark spot in r 3 with narrow waist (Fig. 179)............. C. (Silvicola) freeborni Wirth and Blanton

— Fore tarsomeres without apical spines; distal hourglass-shaped dark spot in r 3 with wider waist (Fig. 180, 181, 183)....................................................................... 31

31(30). Palpal segment 3 with patch of scattered sensilla instead of pit; known only from the low southern deserts of California and Nevada .............................. C. (Silvicola) saltonensis Wirth

— Palpal segment 3 with distinct wide sensory pit, sometimes irregular with scattered sensilla (Fig. 242, 244); more widespread............................................................... 32

32(31). Hind tibiae with basal pale band; distal pale wing spots with sharply defined borders (Fig. 181); palpal segment 3 ratio ~2.8, with irregular pit and sometimes scattered sensilla (Fig. 244)......................................................... C. (Silvicola) neofagineus Wirth and Blanton

— Hind tibiae without pale band; distal pale wing spots with indistinct borders (Fig. 180); palpal segment 3 ratio ~2.1, with distinct wide sensory pit (Fig. 242).. C. (Silvicola) lahontan Wirth and Blanton

33(19). Hind tibial comb with five or six spines; hind tibia with prominent subapical pale band; wing pattern prominent with distal pale spots in r 3, m 1, m 2; distal spot in r 3 irregular C-shaped or 8-shaped centered in distal half of cell, sometimes extending along wing margin into apex of cell to form an arch (Fig. 187–193); pit in palpal segment 3 not broadening internally (Fig. 247)................. 34

— Hind tibial comb with four (occasionally five) spines; if comb with five spines, then distal pale spot in r 3 regular ovoid (Fig. 211, 212, 225, 228) or absent (Fig. 227, 231–239) or hind tibia without prominent subapical pale band; if hind tibia with pale band or distal pale spot in r 3 irregular (Fig. 197, 202), then comb with four spines or pit in palpal segment 3 broadening internally to ~2× diameter of pore (Fig. 248).............................................................. 39

34(33). SCo pattern 1, (3), (4), (5), 6–8; M 1 and M 2 dark, not straddled by distinct pale spots (Fig. 187–189)....................................................... C. ( Haematomyidium Goeldi ) 35

— SCo pattern 1, (2), 3, (4), 5, (6), 7, 9–13; isolated pale spots straddling at 0.2–0.3 on M 1, 0.4–0.5 on M 2 (Fig. 190–193).............................................. C. ( Amossovia Glukhova ) 36

35(34). Two pale spots in m 1, distal-most not at wing margin (Fig. 187, 188); SCo pattern 1, (4), 5–8.......................................... C. (Haematomyidium) kettlei Breidenbaugh and Mullens

— Three pale spots in m 1, distal-most at wing margin, sometimes conjoined narrowly with second back along posterior margin of cell to form a U (Fig. 189); SCo pattern 1, (3), (4), (5), 6–8........................................................... C. (Haematomyidium) stellifer (Coquillett)

36(34). Yellowish coppery-brown; fore and mid tibiae with subapical pale band................................................................... C. (Amossovia) californiensis Wirth and Blanton

— Dark brown; fore and mid tibiae without subapical pale bands................................ 37

37(36). M 1, M 2 barely pale at wing margin (Fig. 191); SCo pattern 1–7, 9–13; hind femora without pale bands........................................... C. (Amossovia) cochisensis Wirth and Blanton

— M 1, M 2 broadly pale at wing margin (Fig. 192, 193); SCo often absent from flagellomeres 2, 4, 6; hind femora with pale bands.............................................................. 38

38(37). SCo pattern 1, (2), 3, (4), 5, (6), 7, 9–13; proboscis ratio ~1.0; 14–20 mandibular teeth; ~22 scutellar setae....................................................... C. (Amossovia) pecosensis Wirth

— SCo pattern 1, 3, 5, 7, 9–13; proboscis ratio ~0.8; 11–14 mandibular teeth; ~16 scutellar setae............................................................. C. (Amossovia) oklahomensis Khalaf

39(33). Distal pale spot in r 3 irregular: bilobed, 8-shaped, C-shaped, or quadrate, centered in distal half of cell, sometimes with additional tiny pale spot in apex of cell; pale spot straddling at ~0.5 on M 2 (Fig. 194–203, 208–210).................................................................. 40

— Distal pale spot in r 3 simple, single, convex, ovoid, diffuse, faint, or absent; not bilobed, medially constricted, or divided (Fig. 204–207, 211–240); pale spot often absent from M 2 ................. 50

40(39). SCo absent from 10–13; at least midportion of CuA 1 within pale area (Fig. 194, 208–210), often entire CuA 2 within pale stripe (Fig. 194); pit in palpal segment 3 not broadening internally 41

— SCo present on 10–13; CuA 1 and CuA 2 nearly to entirely dark (Fig. 195–203); pit in palpal segment 3 broadening internally to ~2× diameter of pore (Fig. 248)................................. 42

41(40). CuA 2 entirely within a pale stripe; pale spots relatively small and isolated (Fig. 194); 11–13 scutellar setae.................................................. C. (Diphaomyia) bergi Cochrane

— CuA 2 entirely dark; pale spots relatively large and somewhat interconnected (Fig. 208–210); 6–8 scutellar setae........................................ (Mohave group) C. mohave Wirth (part)

42(40). Sclerotized ring absent from spermathecal duct; either SCo absent from flagellomeres 2–8 or small third distal pale spot in apex of r 3 (Fig. 196)............................................. 43

— Sclerotized ring present on spermathecal duct; SCo on some of flagellomeres 2–8; no pale spot in apex of r 3 (Fig. 197–203)................................... C. ( Drymodesmyia Vargas ) (part) 44

43(42). Distal pale spot in m 1 not extending to wing margin (Fig. 195); SCo pattern 1, 9–13.................................................................... (Daedalus group) C. daedalus Macfie

— Distal pale spot in m 1 at wing margin (Fig. 196); SCo pattern 1, (2), 3, (4), 5, (6), 7, 9–13............................................................. (Daedalus group) C. pampoikilus Macfie

44(42). Two distinct pale spots in distal half of anal cell (Fig. 197, 198, 201, 202); spermathecae saclike with broad neckless opening ~0.3 width of spermatheca (Fig. 255–257) or subspherical with narrow sclerotized neck as long as wide (Fig. 259).......................................... 45

— One distinct pale spot in distal half of anal cell, not reaching wing margin (Fig. 199, 200, 203); spermathecae subspherical with narrow neckless opening <0.2 width of spermatheca (Fig. 258)...... 4 8

45(44). Spermathecae subspherical to slightly pyriform, with sclerotized necks reducing to <0.2 width of spermatheca (Fig. 259)..................................... C. (Drymodesmyia) bakeri Vargas

— Spermathecae pyriform to elongate saclike, opening ~0.3 width of spermatheca, without sclerotized necks (Fig. 255–257)................................................................. 46

46(45). Spermathecae <1.5× longer than wide (Fig. 255); antennal ratio>1.1, combined distal five flagellomeres (including intersegmental spaces) longer than combined proximal eight; wing length> 1.3 mm ........................................... C. (Drymodesmyia) sitiens Wirth and Hubert

— Spermathecae>1.5× longer than wide (Fig. 256, 257); antennal ratio ~1.0, combined distal five flagellomeres(including intersegmental spaces) shorter than combined proximal eight;wing length<1.3 mm ................................................................................... 47

47(46). Spermathecae <2× longer than wide (Fig. 256)....... C. (Drymodesmyia) jonesi Wirth and Hubert

— Spermathecae>2× longer than wide (Fig. 257)...... C. (Drymodesmyia) butleri Wirth and Hubert

48(44). Apices of M 1, M 2, CuA 1 dark; anterior distal pale spot in r 3 usually attenuated; pale spot at ~0.3 of M 1 sometimes reduced to only touching anterior side of (not straddling) vein (Fig. 200); pale bands absent from mid femora apically, hind femora entirely......................................................................... C. (Drymodesmyia) insolatus Wirth and Hubert (part)

— Apices of M 1, M 2, sometimes CuA 1 pale; anterior distal pale spot in r 3 distinct and separate from or conjoined with posterior spot; pale spot at ~0.3 of M 1 straddling vein (Fig. 199, 203); faint pale bands apical on mid femora, basal on hind femora............................................ 49

49(48). Distal pale spot in r 3 not divided into two separate spots (Fig. 199)........................................................................ C. (Drymodesmyia) cacticola Wirth and Hubert

— Distal pale spot in r 3 distinctly divided into two separate spots (Fig. 203).................................................................... C. (Drymodesmyia) torridus Wirth and Hubert

50(39). Distal half of r 3 dark except for small distinct pale spot entirely within distal 0.2 of cell; distinct isolated pale spots straddling at ~0.3 on M 1 and ~0.5 on M 2 (Fig. 204, 205)......................... 51

— Distal pale spot in r 3 central in distal half or filling most of distal third, but may be faint or absent (Fig. 206–240); pale spots often absent from M 1 or M 2 ........................................ 52

51(50). SCo pattern 1, (5), (7), (8), 9–13; antennal ratio <1.4; distal pale spots in r 3, m 1 usually not touching wing margin (Fig. 204)........................... C. (Diphaomyia) defoliarti Atchley and Wirth

— SCo pattern 1, 3, (4), 5, (6), 7–13; antennal ratio>1.4; distal pale spots in r 3, m 1 usually at wing margin (Fig. 205)....................................... C. (Diphaomyia) haematopotus Malloch

52(50). SCo absent from at least 10–13; sclerotized spermathecal necks 2–3× longer than wide (Fig. 269, or as in Fig. 268 C. haematopotus )............................................................ 53

— SCo present on at least 11–13; sclerotized spermathecal necks absent to about as long as wide (Fig. 260–266, 270–278).................................................................. 57

53(52). Prominent subapical pale band on all femora; distal pale spot pattern distinct (Fig. 206)........................................................... (Limai group) C. luglani Jones and Wirth

— Subapical pale bands absent from mid and hind femora, at most faint on fore femora; distal pale spot pattern more diffuse or absent (Fig. 207, 211–213)....................................... 54

54(53). Distal pale spot in r 3 present but not filling into apical 0.2 of cell (Fig. 207); eyes separated>0.4 ommatidium diameter; SCo pattern 1, (4), (5), 6–8............................................ 55

— Either distal pale spot in r 3 extending into apex of cell (Fig. 211, 212) or absent (Fig. 213); eyes contiguous to separated <0.4 ommatidium diameter; SCo pattern 1, (2), 3–8, (9)................ 56

55(54). Eyes narrowly separated ~0.5 ommatidium diameter; palpal ratio 2.1; inland...................................................................... (Mohave group) C. mohave Wirth (part)

— Eyes widely separated ~2 ommatidium diameters; palpal ratio 2.5; coastal.................................................................... (Mohave group) C. hoguei Wirth and Moraes

56(54). Indistinct distal pale spots extensive into r 3, m 1, m 2 (Fig. 211, 212)........................................................................... C. (Diphaomyia) inyoensis Wirth and Blanton — Distal pale spots absent from r 3, m 1, m 2 (Fig. 213)................. C. (Diphaomyia) salihi Khalaf

57(52). Hind tibial comb with five spines; SCo pattern 1–5, 7, 9–13; distal pale spots faint or absent from r 3, m 1, m 2; pale spots absent from M 1, M 2; palpal segment 3 fusiform, with a deep sensory pit 0.4 as wide as segment (similar to pit in Fig. 249 C. hinmani )...... (unplaced) C. nanellus Wirth and Blanton

— Hind tibial comb often with four spines; SCo patterns from 1, 11–13 to 1–13, not 1–5, 7, 9–13; distal pale spots often distinct in r 3, m 1, m 2; pale spots often straddling M 1, M 2; palpal sensory pit often more broad than deep, wider than 0.5 diameter of segment (as in Fig. 242, 247, 251).............. 58

58(57). Palpal segment 3 sensory pit widening internally, distinctly wider and deeper than pore diameter; pore diameter <0.4 the diameter of segment (Fig. 249, or similar to 248 C. sitiens ); distal pale spots present in r 3, m 1, m 2 (Fig. 200, 214–220); hind tibial comb with four spines; SCo often absent from flagellomeres 2–8; pale spot at tip of costa bi-lobed, often extending back beneath r 2 (Fig. 200, 215–220); if distal pale spot in anal cell double then no pale spots straddling M 1 or M 2 (Fig. 214, 216)................................................ C. ( Drymodesmyia Vargas ) (part) 59

— Palpal segment 3 sensory pit not or slightly widening internally; pore diameter>0.5 the diameter of segment (as in Fig. 242, 244, 251); distal pale spots often absent from r 3, m 1, m 2; hind tibial comb sometimes with five spines; if SCo absent from flagellomeres 2–8, then no distal pale spots in r 3, m 1, m 2; pale spot at tip of costa often ovoid, not distinctly extending back beneath r 2; if distal pale spot in anal cell double, then distinct pale spots straddling M 1 and M 2 (Fig. 229, 230)............ 64

59(58). SCo on some of flagellomeres 2–7......................................................... 60

— SCo absent from flagellomeres 2–7........................................................ 62

60(59). Pale spot at tip of costa not bent back under r 2 (Fig. 214); SCo pattern 1, 3, 5, 7, 9–13...................................................................... C. (Drymodesmyia) byersi Atchley

— Pale spot at tip of costa bent back under entire r 2 (Fig. 200, 215); SCo pattern 1–3, (4), (5), (6), (7), 9–13................................................................................... 61

61(60). Spermathecal ratio ~1.5; SCo pattern 1–3, (4), (5), 9–13; pale spot at ~0.3 of M 1 straddling vein (Fig. 215)................................... C. (Drymodesmyia) arizonensis Wirth and Hubert

— Spermathecal ratio ~1.2; SCo pattern 1–5, (6), (7), 9–13; pale spot at ~0.3 of M 1 sometimes reduced to only touching anterior side of (not straddling) vein (Fig. 200).................................................................... C. (Drymodesmyia) insolatus Wirth and Hubert (part)

62(59). Yellow (in alcohol); distal pale spot in anal cell double, reaching wing margin; no pale spots straddling M 1 or M 2 (Fig. 216); eyes contiguous; sensory pit on palpal segment 3 little widening internally (Fig. 249); pale leg-banding basal and subapical on mid and hind femora, subapical on hind tibiae (Fig. 250); dendrophilic................................... C. (Drymodesmyia) hinmani Khalaf

— Pale brown (in alcohol); distal pale spot in anal cell single, not reaching wing margin; pale spots straddling M 2, often M 1 (Fig. 217–220); eyes separated; sensory pit on palpal segment 3 greatly widening internally (as in Fig. 248 C. sitiens ); pale leg-banding reduced: absent from mid and hind femora, subapically from hind tibiae (as in Fig. 250 C. byersi ); cactiphilic.......................... 63

63(62). Spermathecae unequal by ~1.2×, pyriform, largest is larger than flagellomere 1 (Fig. 260, 261); distal pale spot in r 3 circular to ovoid, 1–1.5× longer than wide (Fig. 217, 218); SCo on flagellomeres 9–10..................................... C. (Drymodesmyia) copiosus Root and Hoffman

— Spermathecae subequal, subspherical, smaller than flagellomere 1 (Fig. 262); distal pale spot in r 3 elongate parallel with M 1, ~2× longer than wide (Fig. 219, 220); SCo often absent from flagellomeres 9–10..................................... C. (Drymodesmyia) ryckmani Wirth and Hubert

64(58). Fore tarsomeres and often hind tarsomeres with apical spines (Fig. 283–285) (uncertain for C. novamexicanus and C. oregonensis ); eyes separated>1 ommatidium diameter (except for species F); no pale spot straddling midportion of M 1 (except for species F);SCo on at least flagellomeres 1, 3, 5, 7, 9,11–13. .... .... ... .... .... .... .... .... .... ... .... .... .... .... .... ... .... .... .... .... ..... 65

— Fore tarsomeres and hind tarsomeres without apical spines (except for hind tarsomere spines on C. usingeri ); eyes contiguous to separated <1 ommatidium diameter (except for C. cavaticus and C. posoensis ); pale spot often straddling at ~0.5 on M 1; SCo often absent from some of flagellomeres 3, 5, 7, or 9

....................................................................................... 73

65(64). Eyes separated ~0.5 ommatidium diameter; isolated pale spots straddling at ~0.3 on M 1 and ~0.5 on M 2 (Fig. 226).......................................................... (unplaced) species F

— Eyes widely separated>1 ommatidium diameter; no isolated pale spots straddling M 1; if pale spot apparently straddling M 2, then contiguous with larger proximal pale area in m 2 (Fig. 222–225) ................................................................................... 66

66(65). Mandible with eight teeth (Fig. 254); hind tarsomeres without apical spines................................................................................... (Palmerae group) species C

— Mandible with 13–16 teeth; hind tarsomeres often with apical spines (as in Fig. 283 male)........ 67

67(66). Spermathecal ratio ≥1.3; sclerotized ring on spermathecal duct (Fig. 276); antennal ratio>1.4; distal pale spots faint in r 3, m 1, m 2, or absent (Fig. 224)....................................................................................... (Palmerae group) C. hawsi Wirth and Rowley

— Spermathecae subequal (Fig. 275, 277); sclerotized ring often absent; antennal ratio <1.4; distal pale spots often more distinct in r 3, m 1, m 2 (Fig. 223, 225, or as in Fig. 147 male)................. 68

68(67). SCo pattern 1–13; spermathecae without sclerotized necks (Fig. 277).......................... 69

— SCo often absent from some of flagellomeres 2, 4, 6, 8, 10; spermathecae with short, sclerotized necks (Fig. 275).......................................................................... 70

69(68). Wing with distal pale spots in r 3, m 1, m 2, cua 1 ............... (Palmerae group) C. palmerae James

— Wing without distal pale spots.............. (Palmerae group) C. oregonensis Wirth and Rowley

70(68). SCo pattern 1–5, (6), 7, (8), 9–13; scutellum about same brown as scutum (Fig. 221).............. 71

— SCo pattern 1, (2), 3, (4), 5, 7, 9, (10), 11–13; scutellum often lighter and more yellowish than scutum (Fig. 221)............................................................................... 72

71(70). Wing with extensive pale spots, but often faint on distal half (as in Fig. 147 male); SCo often absent from 6, 8, 10............................................... (Palmerae group) C. utahensis Fox

— Wing pattern greatly reduced to pale spots only at tip of costa and on r-m crossvein; SCo pattern 1–13.................................................. (Palmerae group) C. leechi Wirth

72(70). Proboscis ratio <0.8; SCo pattern 1, 3, 5, 7, 9, 11–13................................................................................... (Palmerae group) C. calexicanus Wirth and Rowley

— Proboscis ratio>0.9; SCo pattern 1, (2), 3, (4), 5, 7, 9, (10), 11–13.............................................................................. (Palmerae group) C. novamexicanus Atchley

73(64). Eyes contiguous; SCo pattern 1, 11–13; apex of labrum with small distinct median lightly sclerotized grayish tonguelike projection (Fig. 279); 11 or fewer mandibular teeth; hind tibial comb with five spines; pale bands absent from all tibiae, apically from all femora; wing with only proximal pale spots (Fig. 227), no distal pale spots in r 3, m 1, m 2, cua 1, or anal cell............................................................................ (Piliferus group) C. unicolor (Coquillett)

— Eyes separated; SCo usually on at least some of flagellomeres 2–10; labral projection absent; 11 or more mandibular teeth; hind tibial comb often with four spines; pale bands on at least fore femora or basally on tibiae; distal pale spots distinct to faint in cua 1, anal cell, often in r 3, m 1, m 2 (Fig. 228–240).. 74

74(73). Distinct pale spots distal in r 3, m 1, m 2, straddling at ~0.3 on M 1 and ~0.5 on M 2 (Fig. 228–230).... 75

— Distal pale spots faint or absent from r 3, m 1, m 2; spots faint or absent from M 1, M 2 (Fig. 231–240).. 77

75(74). SCo pattern 1–13; antennal ratio 1.0; scutellum with 26–30 setae; spermathecal ratio ~1.1 (Fig. 263)........................................... (Piliferus group) C. cavaticus Wirth and Jones

— SCo pattern 1, 3, (5), (7), (9), (10), 11–13; antennal ratio>1.2; scutellum with 8–12 setae; spermathecal ratio 1.3–1.4 (Fig. 264)............................................................... 76

76(75). Eyes separated <1.0 ommatidium diameter; SCo pattern 1, 3, 5, 7, (9), (10), 11–13..................................................................... (Piliferus group) C. doeringae Atchley

— Eyes separated>1.0 ommatidium diameter; SCo pattern 1, 3, (5), (7), 11–13............................................................... (Piliferus group) C. lophortygis Atchley and Wirth

77(74). SCo always absent from some of flagellomeres 2–10 from both antennae, never forming a 1, 3, 5, 7 SCo pattern; antennal ratio <1.3; spermathecae subequal, necks as long as wide (Fig. 270, 271); sclerotized ring often absent from spermathecal duct; pale spots never straddling M 1 or M 2 (Fig. 236–240). .... .... ... .... .... .... .... .... .... ... .... .... .... .... .... ... .... .... .... .... ..... 78

— SCo pattern 1–13 or 1, 3, 5, 7, (8), (9), (10), 11–13, rarely also with SCo also on 2, 4, or 6 of one antenna; antennal ratio>1.2; spermathecae subequal to unequal by 1.6×, necks absent to half as long as wide (Fig. 265, 266, 273, 274); sclerotized ring on spermathecal duct; faint pale spots often straddling at ~0.3 on M 1 and ~0.5 on M 2 (Fig. 233–235).............................................. 82

78(77). Eyes widely separated ~2 ommatidium diameters; SCo pattern 1, 9–13; pale spots only at tip of costa and over r-m crossvein (Fig. 236); sclerotized ring on spermathecal duct (Fig. 272)............................................................ (unplaced) C. posoensis Wirth and Blanton

— Eyes separated ≤1 ommatidium diameter; SCo often on some of flagellomeres 2–8; pale spots often distally in cua 1 and anal cell, basally in m 1, m 2, anal cell (Fig. 237–240); ring often absent..... 79

79(78). Distal pale spots present in cua 1, anal cell, often m 2 (Fig. 237); eyes narrowly separated ~0.2 omma- tidium diameter.................................... C. (Silvaticulicoides) sublettei Atchley

— Distal pale spots absent (Fig. 238–240); eyes moderately separated ≥0.4 ommatidium diameter... 80

80(79). SCo on 1, 6, 8–13, often others; antennal ratio>1.15; pale bands basal on all tibiae, apical on fore tibiae; sclerotized ring on spermathecal duct (Fig. 271).... C. (Diphaomyia) erikae Atchley and Wirth

— SCo absent from some or all of 2–10; antennal ratio <1.15; pale bands often absent from tibiae; sclerotized ring often absent from spermathecal duct (Fig. 270)................................ 81

81(80). SCo pattern 1, (5), (6), (7), (8), 9–13; proboscis ratio <0.9; hind tarsomeres with apical spines on the specimen examined (as in Fig. 283 C. palmerae ).......... C. (Silvaticulicoides) usingeri Wirth

— SCo pattern 1, (9), (10), 11–13; proboscis ratio>0.9; hind tarsomeres without apical spines............................................ C. (Silvaticulicoides) vetustus Breidenbaugh and Mullens

82(77). SCo pattern 1–13; no pale spots straddling M 1 or M 2 (Fig.231, 232); spermathecal ratio <1.2 (Fig. 273, 274)...................................................... C. ( Sensiculicoides Shevchenko ) 83

— SCo pattern 1, 3, 5, 7, (9), (10), 11–13, rarely also with SCo on 2, 4, or 6 of one antenna; faint pale spots straddling at ~0.3 on M 1 and ~0.5 on M 2 (Fig. 233–235); spermathecal ratio>1.2 (Fig. 265, 266)... .... .... ... .... .... .... .... .... .... ... .... .... .... .... .... ... .... .... .... .... ..... 84

83(82). Spermathecae subequal (Fig. 273)...................... C. (Sensiculicoides) kibunensis Tokunaga

— Spermathecal ratio ~1.2 (Fig. 274)............................ C. (Sensiculicoides) travisi Vargas

84(82). SCo pattern 1, 3, 5, 7–13, occasionally also with SCo on 2, 4, or 6 of one antenna; eyes separated <0.2 ommatidium diameter........................................ (Piliferus group) species B

— SCo pattern 1, 3, 5, 7, (9), (10), 11–13; eyes separated ≥0.5 ommatidium diameter................ 85

85(84). Dark brown; combined length of flagellomeres 7+8 greater than 9, antennal ratio <1.45; proboscis ratio <0.70.................................... (Piliferus group) C. chewaclae Glick and Mullen

— Pale yellowish brown; combined length of flagellomeres 7+8 less than 9, antennal ratio>1.45; proboscis ratio>0.70................................................... (Piliferus group) species A

Systematics of Culicoides

Diptera : Ceratopogonidae

Subfamily Ceratopogoninae

Tribe Culicoidini

Figures

Figures 9–15. 9) Leptoconops americanus female clypeus, setae (s). 10) L. knowltoni female clypeus. 11) L. knowltoni female palpal segment 3. 12) L. carteri female palpal segment 3 (Yolo County, CA). 13) L. werneri female tarsomeres 4–5 and claw (paratype, Imperial County, CA [UCRC]). 14) L. werneri male genitalia, ventral view, tergite 9 apicolateral processes (T9alp), meso-posterior lobe setae (s) (Riverside County, CA [UCRC]). 15) L. belkini male genitalia, ventral view, tergite 9 sclerotized lobe (L) (Riverside County, CA [UCRC]).

Figures 3–8. Female wings with cells (lower case) and veins (upper case) labeled. 3) Culicoides stonei. 4) C. atchleyi (Bonneville County, ID). 5) Leptoconops torrens. Female abdomen, ventral view. 6) L. torrens. 7) C. stonei. 8) L. californiensis, dorsal habitus (in alcohol, Yuma County, AZ [Carl Olson, with permission]).

Figures 50–55. 50) Culicoides sonorensis male wing. 51) C. occidentalis male wing. Spermathecae. 52) C. crepuscularis. 53) C. grandensis (paratype). 54) C. sonorensis. 55) C. sonorensis, lateral view (anterior to left) of female mesonotum showing prescutal pit (presct pit) and dark patches around seta bases.

Figures 56–62. Culicoides reevesi. 56) Female wing. 57) Male wing. 58) Female antennal flagellomeres 9 (f9) and 10 (f10). 59) Male antennal flagellomeres 9 (f9) and 10 (f10). 60) Female head. 61) Male genitalia, ventral view, focused on left paramere (P). 62) Focused on aedeagus, tip (At), subapical processes (Asp) (Lake County, CA).

Figures 117–125. Male wings of Culicoides subgenera Avaritia and Silvicola. 117) C. boydi (in grayscale) (Riverside County, CA [BM]). 118) C. obsoletus (MD [FSCA]). 119) C. cockerellii (Bonneville County, ID). 120) C. freeborni (San Diego County, CA [BM]). 121) C. lahontan. 122) C. neofagineus (in grayscale) (paratype, Mendocino County, CA [FSCA]). 123) C. neomontanus. 124) C. saltonensis (Imperial County, CA [FSCA]). 125) C. sierrensis (paratype, Modoc County, CA [FSCA]).

Figures 33–36. 33) Culicoides moabensis female (top) and unidentified brown C. subgenus Selfia female (bottom), lateral habitus (in alcohol). Male genitalia, ventral view focused on aedeagus. 34) C. brookmani, basal arm of paramere (Pa), distal tip of median process of paramere delineated (Pt), basal arm of aedeagus (Aa), tip of median process of aedeagus (At), gonocoxite (Gc), gonostylus (Gs). 35) C. moabensis, median process of paramere (Pmp) (paratype). 36) C. denningi, sternite 9 caudal lobes (S9cl).

Figures 37–40. Male genitalia of Culicoides, ventral view focused on aedeagus. 37) C. hieroglyphicus, basal arms of aedeagus (Aa), tip of median process of aedeagus (At), gonostylus (Gs), sternite 9 caudal lobes (S9cl). 38) C. jamesi. 39) C. jacksoni, paramere shoulder (Ps), apodeme of gonocoxite (Gca) tooth (t), tip of median process of paramere (Pt). 40) C. tenuistylus, parameres (P) (Ventura County, CA).

Figures 41–45. Male genitalia of Culicoides, ventral view, gonocoxite (Gc), gonostylus (Gs), paramere (P), aedeagus (A), apicolateral processes of tergite 9 (T9alp), sternite 9 (S9). 41) C. mortivallis. 42) C. werneri (paratype, Pima County, AZ [FSCA]). 43) C. bottimeri. Spermathecae, rudimentary third spermatheca (rsp), sternite 8 (S8), sternite 9 (S9). 44) C. bottimeri. 45) C. stonei.

Figures 16–19. Leptoconops carteri (Yolo County, CA). 16) Female terminalia with normal spermathecae. 17) Variant with spermathecae similar to those more typical of L. torrens. 18) Lateral view of lower portion of eye showing interommatidial pubescence.19) Male genitalia, ventral view, proximal sclerites and distal shieldlike fusion of the aedeagus (A), proximal and distal sclerites of a paramere (P), tergite 9 apicolateral processes (T9alp).

Figures 63–65. Male genitalia of Culicoides subgenus Avaritia, ventral view. Upper images focused on aedeagus (A). Lower insets focused on caudal margin of tergite 9. Ventral apodeme of gonocoxite labeled (Gcva). Anteriordirected median process of aedeagus circled. 63) C. boydi (Riverside County, CA [BM]). 64) C. chiopterus (MD [FSCA]). 65) C. obsoletus (MD [FSCA]).

Figures 66–72. Male genitalia of Culicoides subgenus Silvicola, ventral view. 66) C. cockerellii (Bonneville County, ID). 67) C. freeborni (San Diego County, CA [BM]). 68) C. lahontan. 69) C. neofagineus (paratype, Mendocino County, CA [FSCA]). 70) C. neomontanus, basal and apical tips of aedeagus triangulated. 71) C. saltonensis (Imperial County, CA [FSCA]). 72) C. sierrensis (paratype, Modoc County, CA [FSCA]).

Figures 73–78. Male genitalia of Culicoides subgenus Diphaomyia, ventral view. 73) C. defoliarti-haematopotus, focused on aedeagus (A), paramere (P). 74) C. defoliarti-haematopotus, focused on ventral apodemes of gonocoxites (Gcva) and median lobes on parameres (Pml). 75) C. inyoensis, posterior projections on basal arms of aedeagus circled.76) C. erikae, median lobes on parameres circled.77) C. salihi, focused on aedeagus and parameres, posterior projections on basal arms of aedeagus (App). 78) C. salihi, focused on posterior portion of ventral apodemes of gonocoxites, circled.

Figures 46–49. Male genitalia of Culicoides subgenus Monoculicoides, ventral view. 46) C. sonorensis, focused on fused base of parameres (P). 47) C. sonorensis, focused on aedeagus (not to same scale), spicules circled. 48) C. occidentalis, focused on fused base of parameres (P). 49) C. occidentalis, focused on aedeagus (not to same scale).

Figures 79–80. Male genitalia of Culicoides, ventral view, basal arms of aedeagus (Aa), submedian lobe on paramere (Pml), ventral apodeme of gonocoxite (Gcva), distal spines of paramere (Pds). 79) C. mohave, apex of aedeagus circled (San Bernardino County, CA [FSCA]). 80) C. kettlei (Riverside County, CA [BM]).

Figures 88–93. Male genitalia of Culicoides, ventral view. 88) C. luglani, submedian processes of parameres circled. 89) C. crepuscularis, ventral apodeme of gonocoxite circled. 90) C. calexicanus, apices of parameres circled, apex of aedeagus delineated. 91) C. hawsi (paratype, Modoc County, CA [FSCA]). 92) C. palmerae, sharp lateral shoulders on median process of aedeagus indicated (A). 93) C. utahensis.

Figures 81–87. Male genitalia of Culicoides Piliferus group, ventral view. 81) C. cavaticus, focused on aedeagus. 82) C. cavaticus, focused on aedeagus arm base (Aab), basal head of paramere (Ph), ventral apodeme of gonocoxite (Gcva) (paratype, Sacramento County, CA [FSCA]). 83) C. cavaticus (Grand County, UT). 84) C. doeringae, focused on aedeagus. 85) C. doeringae, focused on basal head of paramere (Ph), ventral apodeme of gonocoxite (Gcva). 86, 87) C. Piliferus group species A variations.

Figures 94–99. Male genitalia of Culicoides, ventral view. Ventral apodeme of gonocoxite and apex of paramere circled on Fig. 94–96. 94) C. sublettei (Gillespie County, TX [FSCA]). 95) C. usingeri. 96) C. vetustus (San Bernardino County, CA [BM]). 97) C. posoensis (paratype, Kern County, CA [FSCA]). 98) C. kibunensis. 99) C. travisi (VA [FSCA]).

Figures 104–109. Male genitalia of Culicoides subgenus Drymodesmyia, ventral view. 104) C. butleri (Greenlee County, AZ). 105) C. butleri (Gila County, AZ [FSCA]). 106) C. cacticola, bottom of basal arch of aedeagus delineated (San Diego County, CA [BM]). 107) C. jonesi (Greenlee County, AZ). 108) C. sitiens. 109) C. torridus, height (h) of basal arch of aedeagus indicated (San Diego County, CA [BM]).

Figures 100–103. Male genitalia of Culicoides, ventral view. 100) C. californiensis, left edge of paramere (P), right edge of aedeagus (A), lateral aedeagal spines (As). 101) C. cochisensis (Cochise County, AZ [UCRC]). 102) C. oklahomensis, aedeagus filaments circled (LA [FSCA]). 103) C. pampoikilus (Cochise County, AZ [UCRC]).

Figures 1–2. Female lateral habitus (in alcohol). 1) Leptoconops (Holoconops) Kerteszi group. 2) Culicoides (Monoculicoides) sonorensis.