Dasyhelea

Grogan, Willliam L., Díaz, Florentina, Spinelli, Gustavo R. & Ronderos, Maria M., 2016, The biting and predaceous midges of Guadeloupe (Diptera: Ceratopogonidae). II. Species of the subfamily Dasyheleinae, Zootaxa 4184 (2), pp. 201-254 : 203-205

Key to adult Dasyhelea of Guadeloupe

(females are unknown for D. brachystyla , huertai , patrycjae , quadrifurca , thomasi and sp. nr. thomsenae )

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

- Females........................................................................................... 27

2 Aedeagus covered with hyaline envelope ( Figs. 1 F, 2E); gonocoxal apodemes and paramere symmetrical ( Fig. 2 D) ( cincta group).............................................................................................. 3

- Aedeagus without hyaline envelope ( Figs. 4 A, 12A, 22A, 28A, 30A); gonocoxal apodemes and paramere symmetrical or asymmetrical ( Figs. 4 C, 12C, 22C, 28C, 30C).............................................................. 4

3 Paramere fused to gonocoxal apodemes; large species, wing length 1.39–1.49 mm .................. D. cincta (Coquillett)

- Paramere not fused to gonocoxal apodemes; small species, wing length 0.98–1.06 mm ...... D. caribbeana Spinelli & Wirth

4 Frontal sclerite broad elliptical, bilobate or heart-shaped ( Figs. 5 B, 9A); antennal flagellomere 13 with tapered slender apex ( Figs. 3 A, 5B, 7A, 9A); tergite 9 usually with long apicolateral processes ( Figs. 4 A, 6A, 10A); gonocoxite often with mesal hook-like, pointed or blunt spur ( Figs. 4 A, 6A, 10A) ( grisea group)............................................. 5

- Frontal sclerite slender elliptical ( Figs. 11 A, 17B); antennal flagellomere 13 usually without tapered slender apex ( Figs. 11 A,19A); tergite 9 with long or short apicolateral processes ( Figs. 12 A, 14A, 22A, 28A); gonostylus without mesal hook-like, pointed or blunt spur ( Figs. 12 A, 14A, 22A, 30A).......................................................... 15

5 Posterolateral arms of aedeagus with serrate inner-most margins ( Figs. 4 D, 8D)................................... 6

- Posterolateral arms of aedeagus with smooth inner-most margins ( Figs. 6 D, 10D)................................... 7

6 Tergite 9 with tapered distal portion that extends beyond apices of gonocoxites and elongate tubular, non-divergent apicolateral processes ( Fig. 4 A); paramere with slender distal portion with a subapical sharply pointed tubercle and short recurved apex ( Fig. 4 C); apices of posterolateral processes of aedeagus abruptly recurved anteroventrally with slender pointed tips ( Fig. 4 D)....................................................................................... D. hamula n. sp.

- Tergite 9 with elongate, quadrangular distal portion that extends well beyond apices of gonocoxites and short, triangular, slightly divergent apicolateral processes ( Fig. 8 A); paramere with distal portion greatly expanded, flat, triangular, spatula- shaped with recurved tip ( Fig. 8 C); apices of posterolateral processes of aedeagus straight with rounded tips ( Fig. 8 D)............................................................................................. D. spatula n. sp.

7 Gonocoxite with mesal hook-like, pointed or blunt spur...................................................... 8

- Gonocoxite without mesal hook-like, pointed or blunt spur.................................................. 14

8 Gonocoxite with mesal hook-like spur.................................................................... 9

- Gonocoxite with mesal pointed or blunt spur............................................................... 11

9 Body and genitalia except gonostylus yellowish................................... D. luteogrisea Wirth & Williams

- Body and genitalia brown............................................................................. 10

10 Gonocoxite with large mesal spur; paramere fused to both gonocoxal apodemes................... D. grisea (Coquillett)

- Gonocoxite with small mesal spur; parameres fused to right gonocoxal apodeme.. D. flavifrons (Guérin-Méneville) (in part)

11 Apex of gonostylus with notched tip.......................................... D. pseudoincisurata Waugh & Wirth

- Apex of gonostylus rounded or slightly pointed, not notched................................................. 12

12 Aedeagus with a four-pronged fork-like distal portion ( Fig. 6 D)................................ D. quadrifurca n. sp.

- Aedeagus with undivided pair of posterolateral processes ( Fig. 10 D)............................................ 13

13 Paramere with broad scalpel-shaped apical portion ( Fig. 10 C); tergite 9 apically rounded with short triangular apicolateral pro- cesses ( Fig. 10 A); posterior margin of sternite 9 with narrow deep posteromedian excavation ( Fig. 10 B)... D. scalpela n. sp.

- Paramere with slender apical portion; tergite 9 apically truncate with elongate tubular apicolateral processes; posterior margin

of sternite 9 with broad, shallow posteromedian excavation................... D. flavifrons (Guérin-Méneville) (in part) 14 Tergite 9 with elongate, tubular apicolateral processes; posterolateral processes of aedeagus without mesally directed arm........................................................................ D. flavifrons (Guérin-Méneville) (in part)

- Tergite 9 with short rounded apicolateral processes; posterolateral processes of aedeagus with mesally directed arm............................................................................................... D. griseola Wirth

15 Frontal sclerite elongate elliptical with lower marginal ear-like lobes ( Fig. 15D ); gonocoxal apodemes and paramere asymmetrical, paramere elongate, fused to right gonocoxal apodeme ( Figs. 12 C, 14C, 16C,18C) ( leptobranchia group).......... 16

- Frontal sclerite elongate elliptical without lower marginal ear-like lobes ( Fig. 17 B); gonocoxal apodemes and paramere asymmetrical or symmetrical, fused to both gonocoxal apodemes ( Figs. 20 C, 22C, 24C, 26D)............................ 19

16 Palpal segment 3 extremely elongate ( Fig. 13 D), palpal ratio 7.13; sternite 9 with posteromedian projection bifid ( Fig. 14 B); posterolateral arms of aedeagus stout, sub-triangular ( Fig. 14 D).................................. D. patrycjae n. sp.

- Palpal segment 3 short to moderately long, palpal ratio 2.25–4.85; sternite 9 with posteromedian projection entire, not bifid; posterolateral arms of aedeagus slender to stout, but not sub-triangular......................................... 17

17 Posterolateral arms of aedeagus short, stout ( Fig. 16 D); gonostylus with distal 1/2 abruptly tapered, slender and greatly curved ( Fig. 16 A); posterior margin of sternite 9 greatly elongate, slender ( Fig. 16 B)......................... D. waughi n. sp.

- Posterolateral arms of aedeagus elongate, slender; gonostylus with distal 1/2 not abruptly tapered or greatly curved; posterior margin of sternite 9 moderately short, broad............................................................... 18

18 Tergite 9 with greatly divergent apicolateral processes ( Fig. 12 A); posterolateral arms of aedeagus broadly separated ( Fig. 12 D); distal portion of paramere stout, apex curved 75º ( Fig. 12 C)................................. D. juanae n. sp.

- Tergite 9 with parallel apicolateral processes ( Fig. 18 A); posterolateral arms of aedeagus narrowly separated ( Fig. 18 D); distal portion of paramere slender, apex recurved 180º ( Fig. 18 C)....................................... D. thomasi n. sp.

19 Aedeagus with long to moderately long posteromedian projection and posterolateral arms divided or single; gonocoxal apodemes and paramere usually asymmetrical; tergite 9 with well developed conical or elongate cylindrical apicolateral processes ( mutabilis group)................................................................................... 20

- Aedeagus with or without posteromedian projection, posterolateral arms single; gonocoxal apodemes and paramere symmetrical; tergite 9 with very short to moderately elongate apicolateral processes (species unplaced to group)............... 26

20 Gonostylus with deeply bifid apex......................................................... D. scissurae Macfie

- Gonostylus with pointed or rounded apex, not deeply bifid.................................................. 21

21 Gonocoxal apodemes and paramere symmetrical ( Figs. 22 C, 24C)............................................. 22

- Gonocoxal apodemes and paramere asymmetrical ( Figs. 20 C, 26D)............................................ 23

22 Gonostylus very short, stout with an abruptly tapered apex and sharply pointed tip ( Fig. 24 A); paramere moderately broad, the apex extends beyond apex of aedeagus ( Fig. 24 A); aedeagus with broad crescent-shaped basal arch, posteromedian projection conical, apex entire ( Fig. 24 D); tergite 9 with elongate cylindrical apicolateral processes ( Fig. 24 A).... D. brachystyla n. sp.

- Gonostylus long, slender with gradually tapered apex and curved tip ( Fig. 22 A); paramere very slender, the apex extends to apex of aedeagus ( Figs. 22 A&C); aedeagus with very shallow basal arch, posteromedian projection broad proximally, apex deeply bifurcate; tergite 9 with very short conical apicolateral processes ( Fig. 22 A).................. D. megatheca n. sp.

23 Aedeagus moderately short with broadly spaced posterolateral arms............................................ 24

- Aedeagus moderately long with narrowly spaced posterolateral arms............................................ 25

24 Gonostylus bifurcate, short inner gonostylus with subapical prong ( Fig. 26 C); aedeagus covered with thin hyaline envelope, posterolateral arms apically divided ( Fig. 26 E); posterior margin of sternite 9 with elongate greatly tapered distal portion and slender apex ( Fig. 26 B)................................................................... .. D. nelidae n. sp.

- Gonostylus entire, not bifurcate; aedeagus not covered with hyaline envelope with single posterolateral arms; posterior margin of sternite 9 with moderately short non-tapered distal portion with broad apex......... D. guadeloupensis Delécolle & Rieb

25 Aedeagus with tubular posteromedian portion, apex extending to or near apices of gonocoxites, tip flared with recurved apicolateral extensions; tergite 9 with short apicolateral processes; gonocoxal apodemes slightly asymmetrical, barely fused or narrowly separated, paramere small triangular, lightly sclerotized …....................... D. bermudae Wirth & Williams

- Aedeagus with short tubular posteromedian process extending near apices of posterolateral processes, apex narrow, rounded ( Fig. 20 D); tergite 9 with elongate apicolateral processes ( Fig. 20 A); gonocoxal apodemes asymmetrical, fused to base of paramere, paramere elongate, heavily sclerotized, distal portion J-shaped ( Fig. 20 C)............ D. sp. nr. thomsenae Wirth

26 Aedeagus short, broad with short broadly separated posterolateral arms and shallow basal arch ( Fig. 30 D); paramere short with truncate apex ( Fig. 30 C); posterior extension of sternite 9 narrow with heart-shaped apex ( Fig. 30 B); tergite 9 with long apicolateral processes ( Fig. 30 A)............................................................... D. turnbowi n. sp.

- Aedeagus elongate with slender long narrowly separated posterolateral arms and deep basal arch ( Fig. 28 D); paramere moderately long with arrow-head shaped apex ( Fig. 28 C); posterior extension of sternite 9 triangular with rounded apex ( Fig. 28 B); tergite 9 with very short apicolateral processes ( Fig. 28 A).......................................... D. huertai n. sp.

27 Two spermathecae ( cincta group)........................................................................ 28

- One spermatheca................................................................................... 29

28 Large species, wing length 1.18–1.47 mm; spermathecae small (48 x 42 µm), spherical to ovoid with numerous hyaline punctations; abdominal pleurae with numerous dark streaks........................................ D. cincta (Coquillett)

- Smaller species, wing length 0.99–1.18 mm; spermathecae large (68 x 48 µm), elongate ovoid to sub-quadrangular with sparse hyaline punctations; abdominal pleura without dark streaks........................... D. caribbeana Spinelli & Wirth

29 Frontal sclerite broad elliptical, bilobate or heart-shaped ( Figs. 3 F, 7F, 9E) ( grisea group)........................... 30

- Frontal sclerite slender elliptical ( Figs. 11 E, 15E, 21F, 25E, 29G).............................................. 37

30 Femora, tibiae with broad dark bands; large species, wing length 1.20–1.47 mm ...... D. pseudoincisurata Waugh & Wirth

- Femora, tibiae without dark bands or with poorly developed band on fore and hind femora; smaller species, wing length 0.84– 1.23 mm ........................................................................................... 31

31 Subgenital plate with arrowhead-shaped anterior portion ( Figs. 3 I, 7I, 9H)...................................... 32

- Subgenital plate with slender anterior portion ( Figs. 11 H, 15I)................................................ 36

32 Spermatheca with straight to slightly oblique neck ( Fig. 3 I).................................................. 33

- Spermatheca with oblique or retort-shaped neck ( Figs. 7 I, 9H)................................................ 35

33 Body yellowish............................................................. D. luteogrisea Wirth & Williams

- Body brown to dark brown............................................................................ 34

34 Spermatheca pyriform without distinct neck or very short neck................................ D. grisea (Coquillett)

- Spermatheca ovoid with broad tapered neck ( Fig. 3 I)............................................ D. hamula n. sp.

35 Spermatheca with retort-shaped neck ( Fig. 9 H)................................................ D. scalpela n. sp.

- Spermatheca with straight or slightly oblique neck................................. D. flavifrons (Guérin-Méneville)

36 Spermatheca spherical with slender oblique neck............................................... D. griseola Wirth

- Spermatheca ovoid with stout elongate, retort-shaped neck ( Fig. 7 I)................................. D. spatula n. sp.

37 Frontal sclerite with lower marginal ear-like lobes ( Figs. 11 E, 15D) ( leptobranchia group).......................... 38

- Frontal sclerite without lower marginal ear-like lobes ( Figs. 21 F, 25E, 29G)...................................... 39

38 Anterior portion of subgenital plate club-shaped, posterior portion elongate, ovoid, with large central lumen; spermatheca large spherical, diameter 86 µm; neck moderately long, slender ( Fig. 11 H)............................... D. juanae n. sp.

- Anterior portion of subgenital plate triangular, posterior portion somewhat rectangular without central lumen; spermatheca heavily sclerotized, slightly ovoid, diameter 40 µm, neck short, moderately broad ( Fig. 15I )............. D. waughi n. sp.

39 Subgenital plate chalice-shaped ( Fig. 29 K) (unplaced to species group)............................. D. turnbowi n. sp.

- Subgenital plate circular to ovoid with a central lumen ( Fig. 21 J) ( mutabilis group)............................... 40

40 Spermatheca very large (length 118 µm), elongate ovoid with a moderately long, very slender neck ( Fig. 21 I)................................................................................................. D. megatheca n. sp.

- Spermatheca smaller (length 34–71 µm), globular to ovoid with short broad neck................................ 41

41 Spermatheca very small (length 34 µm), ovoid with very short broad neck; wing membrane infuscated; subgenital plate with a circular very lightly sclerotized anterior portion, a slightly rectangular heavily sclerotized posterior portion and a small lumen ( Fig. 25 I)............................................................................... D. nelidae n. sp.

- Spermatheca larger (length 44–71 µm), globular or ovoid with short neck; wing membrane not infuscated; subgenital plate cir- cular to ovoidal with a large well defined lumen........................................................... 42

42 Spermatheca spherical with slender retort shaped neck........................... D. guadeloupensis Delécolle & Rieb

- Spermatheca spherical or ovoid with short straight neck..................................................... 43

43 Spermatheca large (length 64–71 µm), globular with moderately broad tubular neck; subgenital plate circular, anterior and pos- terior portions broad with moderately broad lumen.................................. D. bermudae Wirth & Williams

- Spermatheca smaller (length 48 µm), ovoid with short narrow neck; subgenital plate with broad triangular anterior portion, narrow posterior portion and small triangular lumen........................................... D. scissurae Macfie

Figures

FIGURE 1. Dasyhelea caribbeana Spinelli & Wirth. Male. A. head. B. palpus. C. thorax. D. scutellum. E. wing. F. genitalia.

FIGURE 2. Dasyhelea caribbeana Spinelli & Wirth. Male. A. genitalia. B. sternite 9. C. gonostylus. D. gonocoxal apodemes and paramere. E. aedeagus. Scale bar: 0.05 mm.

FIGURE 3. Dasyhelea hamula n. sp. Holotype male, A – D, Allotype female, E – I. A, E. head. B, G. palpus. C, H. wing. D. genitalia. F. frontal sclerite. I. subgenital plate and spermatheca.

FIGURE 4. Dasyhelea hamula n. sp. Holotype male. A. genitalia. B. sternite 9. C. gonocoxal apodemes and paramere. D. aedeagus. Scale bar: 0.05 mm.

FIGURE 5. Dasyhelea quadrifurca n. sp. Holotype male. A. head. B. frontal sclerite. C. palpus. D. wing. E. genitalia.

FIGURE 6. Dasyhelea quadrifurca n. sp. Holotype male. A. genitalia. B. sternite 9. C. gonocoxal apodemes and paramere. D. aedeagus. Scale bar: 0.05 mm.

FIGURE 7. Dasyhelea spatula n. sp. Holotype male, A – E, Allotype female, F – I. A, F. head. B, G. clypeus. C, G. palpus. D, H. wing. E. genitalia. I. subgenital plate and spermatheca.

FIGURE 8. Dasyhelea spatula n. sp. Holotype male. A. genitalia. B. sternite 9. C. gonocoxal apodemes and paramere. D. aedeagus. Scale bar: 0.05 mm.

FIGURE 9. Dasyhelea scalpela n. sp. Holotype male, A – D, Allotype female, E – H. A, E. head. B, F. palpus. C, G. wing. D. genitalia. H. subgenital plate and spermatheca.

FIGURE 10. Dasyhelea scalpela n. sp. Holotype male. A. genitalia. B. sternite 9. C. gonocoxal apodemes and paramere. D. aedeagus. Scale bar: 0.05 mm.

FIGURE 11. Dasyhelea juanae n. sp. Holotype male, A – D, Allotype female, E – H. A, E. head. B, F. palpus. C, G. wing. D. genitalia. H, subgenital plate and spermatheca.

FIGURE 12. Dasyhelea juanae n. sp. Holotype male. A. genitalia. B. sternite 9. C. gonocoxal apodemes and paramere. D. aedeagus. Scale bar: 0.05 mm.

FIGURE 13. Dasyhelea patrycjae n. sp. Holotype male. A. head. B. antennal flagellum. C. Clypeus. D. palpus. E. wing. F. genitalia.

FIGURE 14. Dasyhelea patrycjae n. sp. Holotype male. A. genitalia. B. sternite 9. C. gonocoxal apodemes and paramere. D. aedeagus. Scale bar: 0.05 mm.

FIGURE 16. Dasyhelea waughi n. sp. Holotype male. A. genitalia. B. sternite 9. C. gonocoxal apodemes and paramere. D. aedeagus. Scale bar: 0.05 mm.

FIGURE 17. Dasyhelea thomasi n. sp. Holotype male. A. head. B. frontal sclerite. C. palpus. D. wing. E. genitalia.

FIGURE 18. Dasyhelea thomasi n. sp. Holotype male. A. genitalia. B. sternite 9. C. gonocoxal apodemes and paramere. D. aedeagus. Scale bar: 0.05 mm.

FIGURE 20. Dasyhelea sp. nr. thomsenae Wirth. Male. A. genitalia. B. sternite 9. C. gonocoxal apodemes and paramere. D. aedeagus. Scale bar: 0.05 mm.

FIGURE 21. Dasyhelea megatheca n. sp. Holotype male, A – D, Allotype female, E – J. A, E. head. B, G. palpus. C, H. Wing. D. genitalia. F. frontal sclerite and clypeus. I. abdomen and spermatheca. J. subgenital plate.

FIGURE 22. Dasyhelea megatheca n. sp. Holotype male. A. genitalia. B. sternite 9. C. gonocoxal apodemes and paramere. D. aedeagus. Scale bar: 0.05 mm.

FIGURE 24. Dasyhelea brachystyla n. sp. Holotype male. A. genitalia. B. sternite 9. C. gonocoxal apodemes and paramere. D. aedeagus. Scale bar: 0.05 mm.

FIGURE 25. Dasyhelea nelidae n. sp. Holotype male, A – D, Allotype female, E – I. A, E. head. B, G. palpus. C, H. wing. D. genitalia. F. clypeus. I. subgenital plate and spermatheca.

FIGURE 26. Dasyhelea nelidae n. sp. Holotype male. A. genitalia. B. sternite 9. C. gonostylus. D. gonocoxal apodemes and paramere. E. aedeagus. Scale bar: 0.05 mm.

FIGURE 28. Dasyhelea huertai n. sp. Holotype male. A. genitalia. B. sternite 9. C. gonocoxal apodemes and paramere. D. aedeagus. Scale bar: 0.05 mm.

FIGURE 29. Dasyhelea turnbowi n. sp. Holotype male, A – E, Allotype female, F – K. A, F. head. B, G. frontal sclerite and clypeus. C, H. palpus. D, J. wing. E. genitalia. I. thorax. K. subgenital plate and spermatheca.

FIGURE 30. Dasyhelea turnbowi n. sp. Holotype male. A. genitalia. B. sternite 9. C. gonocoxal apodemes and paramere. D. aedeagus. Scale bar: 0.05 mm.

FIGURE 15. A – I. Dasyhelea waughi n. sp. Holotype male, A – C, Allotype female, D – I. A, D. head. B, H. wing. C. genitalia. E. frontal sclerite. F. clypeus. G. palpus. I. subgenital plate and spermatheca.