Craspedochaeta, Czerny, 1903

Lonsdale, Owen & Marshall, Stephen A., 2006, Revision of the New World species of Craspedochaeta (Diptera: Clusiidae), Zootaxa 1291 (1), pp. 1-101 : 13-18

Key to the New World species of Craspedochaeta

1. Posterior fronto­orbital bristle 2/3 length of anterior fronto­orbitals ( Fig. 9 ). Anterior dorsocentral bristle 2/3 length of posterior dorsocentral. Clypeus yellow with dorsallateral corners brown. Ecuador, Bolivia (Map 14) ( C. biseta species group) ............... ................................................................................................... C. argoniae spec. nov.

­ All fronto­orbital bristles subequal in length. Anterior dorsocentral bristle half length of posterior bristle. Clypeus yellow or brown; if bicoloured, yellow with entire dorsal margin ( C. phaios ) or lateral margins brown (some specimens of C. annulipes )........ 2

2. Five pairs of fronto­orbital bristles; two reclinate bristles in front of inclinate frontoorbital ( Fig. 8 ). Bristles along basal section of costa usually lighter in colour. Acrostichal bristles almost always present. Ventral receptacle with long subterminal flagellum ( Figs. 67–77 ) ( C. transversa species group)......................................................... 3

­ Four pairs of fronto­orbital bristles; one reclinate bristle in front of inclinate frontoorbital ( Fig. 7 ). All costal bristles black. Acrostichal bristles almost always absent. Ventral receptacle simple, without subterminal flagellum ( Figs. 106–109 ) ( C. concinna species group) ................................................................................................. 19

3. Notum predominantly brown, at most with yellow arch ( Fig. 17 ) or one pair of anterolateral spots ( Fig. 18 ), but never with cruciate pattern. Back of head entirely dark brown or almost entirely dark on dorsal half. Male cerci shorter than surstylus and variably coloured. Surstylus twisted so that inner face is visible posteriorly ( Fig. 27 ), or surstyli with basal process on dorsal or ventral margin ( Figs. 40 & 44 ). Distiphallus large and sac­like........................................................................................................... 4

­ Notum largely yellow, or with brown to dark brown pattern; pattern often cruciate with one transverse stripe behind notopleuron (sometimes converging upon it) and one median longitudinal stripe; stripes sometimes interrupted centrally ( Figs. 16, 19–23 ). Back of head sometimes yellow, but almost always with one pair of stripes radiating from central foramen. Male cerci white, united, large and plate­like, often as long as epandrium ( Fig. 47 ). Surstylus usually small and smooth ( Fig. 47 ). Distiphallus absent or small without heavily sclerotized regions............................................................... 11

4. Pleuron black with lower half of katepisternum fading to yellow. First flagellomere variably pigmented. Face and clypeus usually dark brown to orange, or yellow with venter darker. Mid and hind femora sometimes brown, at least basally ...................... 5

­ Pleuron yellow; anepisternum sometimes with brown spot or stripe. First flagellomere yellow. Face and clypeus yellow. Legs white to yellow (sometimes hind tibia with median brown band)...................................................................................................... 9

5. Legs entirely yellow. Face light yellow. First flagellomere sometimes infuscated along anterior margin. Surstylus strongly lobate posteriorly and with anterobasal thumb ( Fig. 40 ). Cerci thin basally and wide and emarginate apically ( Fig. 41 ). Brazil (Map 4)...... ............................................................................................. C. amazonensis spec. nov.

­ Legs predominantly brown, or at least with brown median band on hind tibia. Female face orange to dark brown; male face usually yellow (venter sometimes darker). First flagellomere brown on ventral or anterior half. Surstylus not strongly lobate, but if thumb present, situated on posterior margin ( Fig. 44 ). Cerci variable, but not as above ....................................................................................................................................... 6

6. Face entirely dark brown. Female frons dark brown excluding lateral margins. Female mid tibia yellow. Female mid and hind femora yellow with apices brown. Male femora yellow with base of mid femur brown. Brazil, Costa Rica, Ecuador, Mexico, Peru (Map 3)....................................................................................... C. protomis spec. nov.

­ Face yellow, at least on upper half. Female frons yellow laterally and on anterior half (sometimes tinted with orange). Female mid tibia light brown to brown medially. Female mid and hind femora usually brown apically. Male femora yellow (brown basally in Costa Rican males) ...................................................................................... 7

7. Mid coxa yellow. Posterior half of postpronotum yellow. Female femora yellow. Proepisternum yellow. Back of head dark brown with yellow dorsomedial stripe. Basal segment of spermatheca approximately 6 times length. Median segment of spermatheca smooth and globose. Brazil (Map 4)..................................... C. feminea spec. nov.

­ Mid coxa brown, at least in part. Posterior half of postpronotum rarely yellow. Female femora brown, at least in part on hind femur. Proepisternum almost always brown. Back of head dark brown. Basal segment of spermatheca much less than 6 times width. Median segment of spermatheca squat and annulated or otherwise textured .... 8

8. Posterior half of occiput brown. First flagellomere brown on ventral half or with variable apical infuscation. Male hind coxa yellow. Fore femur yellow. Female face entirely flat. Surstylus rounded and convex with anterodistal angle produced internally. Surstylus without basal process. Cerci broadly rounded with height half width and with wide shallow emargination ( Fig. 31 ). Central and South America to Bolivia (Map 2). ................................................................................. C. nigrithorax (Hennig)

­ Occiput entirely brown. First flagellomere brown on anterior half. Male hind coxa brown, at least faintly. Fore femur brown. Female face sharply angulate ventrally. Surstylus rounded with apex concave and slightly tapered. Surstylus with basal process directed medially ( Fig. 44 ). Cerci small, truncate, narrow basally and strongly sunken. Brazil (Map 4). ................................................................... C. melanosoma spec. nov.

9. Notum black with large yellow arch ( Fig. 17 ). Tip of scutellum sometimes yellow. Back of head with dorsal half largely black. Bolivia, Ecuador, Dominican Republic (Map 1)...................................................................................... C. apsilutea spec. nov.

­ Notum black, sometimes with yellow spot on notopleuron ( Fig. 18 ). Scutellum entirely black. Back of head black, or with ventral margin yellow. ........................... 10

10. Scutum almost always with one pair of yellow spots on the anterolateral margin ( Fig. 18 ). Hind tibia with a brown median band. Meron and (usually) anepisternum brown. Back of head often entirely brown. Surstylus produced anteriorly. Bolivia, Brazil (Map 3)........................................................................................... C. ornata (Hennig)

­ Scutum brown. Legs and pleuron entirely yellow. Back of head brown with ventral margin yellow. Surstylus with ventrobasal process. Costa Rica, Ecuador (Map 4). ...... ............................................................................... C. claripennis (Melander & Argo)

11. Notum largely yellow with one small pair of spots or one quadrate spot on anterior margin of notum and one small pair of supra­alar spots; notum with thin small brown semicircle (or less) in front of scutellum. Laterotergites yellow in part .................... 12

­ Notum with cruciate pattern including one transverse stripe behind notopleuron (sometimes converging upon notopleuron) and one central longitudinal stripe (sometimes interrupted centrally) ( Figs. 19–23 ). Laterotergites entirely brown to black ... 14

12. Anterior margin of scutum with one pair of small median spots ( Fig. 14 ). Scutellum entirely brown. Laterotergites brown with one pair of yellow stripes lateral to scutellum. M 1+2 ratio more than 7.0. Jamaica, Florida, Mexico, Peru (Map 5) ....................... ...................................................................................................... C. weemsi spec. nov.

­ Anterior margin of scutum with large median spot. Scutellum yellow medially, at least on distal 2/3. Laterotergites yellow with one pair of brown stripes lateral to scutellum or entirely brown lateral to scutellum. M 1+2 ratio distinctly less than 7.0. Brazil (Map 3) ..................................................................................................................................... 13

13. Lateral margin of scutum with one pair of minute supra­alar spots and sometimes with one pair of wider subbasal spots ( Fig. 15 ). Anterior margin of scutum with large quadrate spot. Occiput shiny. Frons with orange tint ..................... C. pacaraima spec. nov.

­ Lateral margins of scutum yellow ( Fig. 11 ). Anterior margin of scutum brown. Anterior margin of occiput silvery tomentose. Frons yellow with posterior half brown ....... ............................................................................................................ C. festiva (Sóos)

14. Legs brown on at least hind tibia (sometimes very faint); if legs entirely yellow (very uncommon) then frons evenly brown; palpus sometimes light brown. Pleural stripe usually extending from dorsal margin of anepisternum to anterior face of katepisternum. Frons sometimes darkest on anterior margin ..................................................... 15

­ Legs yellow to white and palpus always yellow. Pleural stripe restricted to dorsal spot on anepisternum, or (rarely) with small additional faded spot on anterior face of katepisternum. Frons never darkest anteriorly ............................................................ 16

15. Frons darkest along anterior margin or entirely dark (excluding lateral margins; Fig. 23 ), with clypeus, and ventral and dorsal margins of face black. Male with first flagellomere, buccal cavity, palpus and gena brown. All tibiae brown excluding apex and base. Fore femur brown excluding tip. Mid and hind femora brown excluding yellow tip and wide white base. Surstylus 2/3 length of cerci and pointed distally ( Fig. 49 ). Costa Rica, Ecuador, Guatemala, Peru (Map 5) ..................... C. chela spec. nov.

­ Frons yellow or darkest medially or posteriorly, but if frons evenly light brown, face yellow. Male with first flagellomere ( Fig. 22 ), palpus, gena, and buccal cavity yellow. Only hind tibia brown; often brown on basal 5/6, but sometimes reduced to faded band on basal 1/3; remainder of legs yellow to white. Surstylus half length of cerci and rounded distally ( Fig. 47 ). Bolivia, Brazil, Costa Rica, Ecuador, Mexico, Panama, Peru, Venezuela (Map 7) ........................................................... C. pollostos spec. nov.

16. Veins M 1 and R 4+5 slightly convergent distally ( Fig. 26 ). Distance between crossveins not greater than length of posterior cross­vein. Surstylus with numerous minute setulae and thin distal process ( Fig. 53 ). Notum as in Fig. 19 . Argentina, Bolivia, Brazil, Costa Rica, Paraguay, Peru (Map 6) ........................................... C. transversa Czerny

­ Veins M 1 and R 4+5 parallel ( Figs. 24 & 25 ). Distance between crossveins variable. Surstylus bare on outer face ............................................................................................. 17

17. Antenna (excluding arista) white, in striking contrast to frons. Frons orange, with posterior half fading to brown ( Fig. 16 ). Pleuron with vertical stripe along posterior margin of anepisternum. Surstylus broadly rounded with triangular process on posterior margin ( Fig. 45 ). Katepisternum sometimes with small faded spot. Bolivia, Brazil (Map 5) ....................................................................................... C. candida spec. nov.

­ Antenna (excluding arista) light yellow to yellow. Frons yellow, at least on anterior half. Anepisternum with short dorsal stripe not touching posterior margin. Surstylus thin, acute and without triangular process. Katepisternum entirely light yellow ....... 18

18. Distance between crossveins rarely greater than length of posterior cross­vein ( Fig. 25 ). Wing sometimes darkly clouded on distal 1/3, fading posteriorly, but usually restricted to cloud around apex of R 2+3. Face sometimes brown on upper 2/3. Surstylus half length of cerci and subtriangular ( Fig. 55 ). Notum as in Fig. 20 . Bolivia, Brazil, Costa Rica, El Salvador, Panama, Trinidad (Map 9) ................ C. synneura (Hennig)

­ Distance between crossveins distinctly greater than length of posterior cross­vein ( Fig. 24 ). Wing never darkly clouded on entire distal 1/3. Face sometimes light brown on lower half. Surstylus 2/3 length of cerci and distinctly arched; thin just past base, then wider medially ( Fig. 57 ). Notum as in Fig. 21 . Mexico to Bolivia (Map 8) .................. ............................................................................... C. pleuralis (Williston) comb. nov

19. Notum yellow to brownish­yellow, or yellow with brown pattern ............................ 20

­ Notum and laterotergites entirely brown to black; several small yellow spots sometimes present on scutum .............................................................................................. 22

20. Notum entirely light yellow ( Fig. 10 ). Laterotergites yellow below scutellum. Peru (Map 10) ............................................................................ C. xanthonotum spec. nov.

­ Notum yellow with brown pattern. Laterotergites entirely brown ............................. 21

21. Laterotergites brown; oblique supra­alar stripes approaching median longitudinal stripe on scutum posteriorly ( Fig. 13 ). Legs yellow. Brazil (Map 10) ........................... ......................................................................................................... C. trivittata (Sóos)

­ Laterotergites largely yellow beneath scutellum ( Fig. 12 ); oblique supra­alar stripes approaching median longitudinal stripe anteriorly. Legs yellow with mid and hind tibiae brown basally. Peru (Map 10) ................................................... C. loreto spec. nov.

22. Wing evenly clouded, with base lighter or costal margin slightly darker .................. 23

­ Wing ornamented with spots or cross bands .............................................................. 24 23. Wing evenly dusky, excluding base. Frons yellowish­orange on anterior 1/3, occasionally entirely dark. Acrostichal bristle present or absent. Bristles yellow or dark brown to black. Pregonite straight and relatively thick with apex broadly rounded and irregular in outline. Common. Mexico to Bolivia, Florida and New Mexico (Map 11) .......... ................................................................................................. C. concinna (Williston)

­ First and second radial cells much darker than remainder of wing. Frons yellowishorange on anterior margin. Acrostichal bristle present. Bristles brown. Pregonite thin, arched medially and bilobed apically. Uncommon. Bolivia (Map 10) .......................... ....................................................................................................... C. phaios spec. nov.

24. Head (excluding mouthparts and antenna) dark brown; sometimes anterior margin of frons orange. Pleuron, fore coxa, and anterior half of mid coxa dark brown. Mexico to Bolivia (Map 12) ................................................. C. fascipennis (Melander & Argo)

­ Head (excluding mouthparts and antenna) yellow in part. Coxae yellow to white. Pleuron entirely yellow or only yellow ventrally .............................................................. 25

25. Three dorsocentral bristles (each half length of preceding bristle) and mid and hind tibiae brown excluding base and apex. Pleuron entirely brown, or yellow with anepisternum and anterior margin of anepimeron brown. Female with gena silvery­tomentose, excluding ventral margin. Peru (Map 10) ................................. C. basalis Czerny

­ Two dorsocentral bristles with minute anterior bristle no more than 1/3 length of anterior dorsocentral; if additional bristle approaching half length of anterior dorsocentral (occasionally attaining), anepisternum predominantly yellow and legs not as above. Pleuron usually with at least ventral margin yellow; if pleuron entirely brown, apex of hind (and sometimes mid) femur brown or mid and hind tibiae entirely brown. Female usually with dorsal half (or less) of gena silvery­tomentose (occasionally only ventral margin shiny)............................................................................................................... 26

26. Male: first flagellomere entirely yellow to white or with dark anteroventral spot; pleuron brown on dorsal half or less; if only dorsal and posterior sutures of anepisternum brown or pleuron entirely yellow, then vibrissa yellow, posterior 2/3 of frons (often) brown, and frons with wide pilose patch on anterior half. Female (only known for C. annulipes ): faded brown spot often present on anterior 2/3 of first flagellomere; pleuron brown on at least dorsal half; if pleuron entirely brown, either mid tibia brown or first flagellomere yellow ............................................................................................. 27

­ Male: first flagellomere white or with dark dorsal spot; pleuron entirely brown or only on dorsal margin of anepisternum; vibrissa yellow to brown; frons variable. Female unknown...................................................................................................................... 29

27. Frons yellow to orange, at least along the anterior margin (usually on anterior 1/3). Surstylus evenly rounded with all margins smooth. Ejaculatory apodeme with “fringe” ( Fig. 88 ). Common from Mexico to Bolivia (Map 13); uncommon in eastern United States.................................................................... C. annulipes (Johnson) comb. nov.

­ Frons entirely dark brown. Surstylus with margins variably sculptured. Ejaculatory apodeme without “fringe” ( Fig. 81 ). Uncommon ...................................................... 28

28. Pedicel yellow. Gena brown. Face light yellow. Genal bristles well developed. Posterior corners of scutum not faded. Pleuron brown on dorsal half of anepisternum and on anterior margin of anepimeron. Surstylus yellow. Surstylus less than half the length of the epandrium with one wide posteroventral lobe and one small thin anteroventral lobe ( Fig. 95 ). Cerci relatively wide, not thin and elongate ( Fig. 96 ). Epandrium height less than half length. Brazil (Map 10) .................................................. C. biloba spec. nov.

­ Pedicel light brown. Gena light yellow. Face with orange tint. Genal bristles hair­like. Posterior corners of scutum with yellowish circles. Pleuron brown on dorsal margin of proepisternum and on posterior margin of anepisternum. Surstylus brownish basally. Surstylus subtriangular in outline with sculpturing on posterior margin ( Fig. 98 ). Cerci long and thin ( Fig. 99 ). Height of epandrium slightly less than length. Ecuador (Map 10) ........................................................................................ C. chauliodon spec. nov.

29. First flagellomere white. Pleuron yellow with dorsal margin of anepisternum brown. Vibrissa brown. Frons with central brown triangular spot on posterior 2/3 and with thin triangular pilose patch on anterior half. Occiput and lower 1/5 of back of head white. Jamaica (Map 10) ................................................ C. brunneivibrissa spec. nov.

­ First flagellomere brown in part. Pleuron entirely or predominantly brown. Vibrissa yellow. Frons brown with only thin yellowish­orange margins, and with wide pilose patch on anterior half. Occiput and back of head dark brown. Bolivia (Map 10) ...... 30

30. First flagellomere with small, well defined spot around base of first flagellomere. Tip of hind tibia and base of hind femur brown. Frons with anterior margin thinly yellow ................................................................................................ C. pullipleura spec. nov.

­ First flagellomere dark brown on dorsal half and white ventrally. Legs entirely yellow to white. Frons with lateral margins yellow on anterior half ........ C. zongo spec. nov.

Figures

FIGURES 7–9: heads; outline drawing, left lateral (arista and bristles removed, excluding vibrissa and fronto­orbital bristles); 7: Craspedochaeta concinna (Williston, 1896) (C. concinna species group); 8: C. pleuralis (Williston, 1896) (C. transversa species group); 9: C. argoniae spec. nov. (C. biseta species group).

FIGURES 67–70: Internal female genitalia; Figs. 67: Craspedochaeta pleuralis (Williston, 1896); Figs. 68: C. pollostos spec. nov.; Figs. 69: C. weemsi spec. nov.; Figs. 70: C. synneura (Hennig, 1938).

FIGURES 71–77: Female internal genitalia (double line indicates the removal of an elongate median section to facilitate illustration); Fig. 71: Craspedochaeta feminea spec. nov.; Fig. 72: C. claripennis (Melander & Argo, 1924); Fig. 73: C. ornata Hennig, 1938; Fig. 74: C. transversa Czerny, 1903; Fig. 75: C. protomis spec. nov.; Fig. 76: C. nigrithorax (Hennig, 1938); Fig. 77: C. apsilutea spec. nov.

FIGURES 106–109: Female internal genitalia; 106: Craspedochaeta concinna (Williston, 1896); 107: C. annulipes (Johnson, 1913); 108: C. fascipennis (Melander & Argo, 1924); 109: C. basalis Czerny, 1903.

FIGURES 10–23: Head and thorax, dorsal colouration of the New World Craspedochaeta without an entirely dark notum; 10: C. xanthonotum spec. nov.; 11: C. festiva (Sóos, 1962); 12: C. loreto spec. nov.; 13: C. trivittata (Sóos, 1962); 14: C. weemsi spec. nov.; 15: C. pacaraima spec. nov.; 16: C. candida spec. nov.; 17: C. apsilutea spec. nov.; 18: C. ornata (Hennig, 1938); 19: C. transversa Czerny, 1903; 20: C. synneura (Hennig, 1938); 21: C. pleuralis (Williston, 1896); 22: C. pollostos spec. nov; 23: C. chela spec. nov, male.

FIGURES 27­29: Craspedochaeta apsilutea spec. nov.; 27: external male terminalia, left lateral; 28: external male terminalia, posterior; 29: hypandrial complex, left lateral.

FIGURES 39–41: Craspedochaeta amazonensis spec. nov.; 39: hypandrial complex, left lateral; 40: external male terminalia, left lateral; 41: external male terminalia, posterior.

FIGURES 42–44: Craspedochaeta melanosoma spec. nov.; 42: hypandrial complex, left lateral; 43: external male terminalia, left lateral; 44: external male terminalia, posterior.

FIGURES 47–52: male terminalia, external: Figs. 47–48: Craspedochaeta pollostos spec. nov.; Fig. 47: left lateral; Fig. 48: posterior; Figs. 49–50: C. chela spec. nov.; Fig. 49: left lateral; Fig. 50: posterior; Figs. 51–52: C. weemsi spec. nov.; Fig. 51: left lateral; Fig. 52: posterior.

FIGURES 30–32: Craspedochaeta nigrithorax (Henng, 1938); 30: external male terminalia, left lateral; 31: external male terminalia, posterior; 32: hypandrial complex, left lateral.

FIGURES 24–26: Left wing, ventral (arrows indicate diagnostic characters mentioned in the key); 24: Craspedochaeta pleuralis (Williston, 1896); 25: C. synneura (Hennig, 1938); 26: C. transversa Czerny, 1903.

FIGURES 53–58: Male terminalia, external: Figs. 53–54: Craspedochaeta transversa Czerny, 1903; Fig. 53: left lateral; Fig. 54: posterior; Figs. 55–56: C. synneura (Hennig, 1938); Fig. 55: left lateral; Fig. 56: posterior; Figs. 57–58: C. pleuralis (Williston, 1896); Fig. 57: left lateral; Fig. 58: posterior.

FIGURES 45–46: Craspedochaeta candida spec. nov.; Fig. 45: external male terminalia, left lateral; Fig. 46: external male terminalia, posterior.

FIGURES 88–90: Craspedochaeta annulipes (Johnson, 1913); 88: hypandrial complex, left lateral; 89: external male terminalia, left lateral; 90: external male terminalia, posterior.

FIGURES 81–83: Craspedochaeta concinna (Williston, 1896); 81: hypandrial complex, left lateral; 82: external male terminalia, left lateral; 83: external male terminalia, posterior.

FIGURES 94–96: Craspedochaeta biloba spec. nov.; 94: hypandrial complex, left lateral; 95: external male terminalia, left lateral; 96: external male terminalia, posterior.

FIGURES 97–99: Craspedochaeta chauliodon spec. nov.; 97: hypandrial complex, left lateral; 98: external male terminalia, left lateral; 99: external male terminalia, posterior.