Nephochaetopteryx Townsend, 1934

Nephochaetopteryx Townsend, 1934: 203 .

Type species: Nephochaetopteryx pallidiventris Townsend, 1934, by original designation.

Sarcohelicobia Blanchard, 1939: 795 .

Type species: Sarcohelicobia elegans Blanchard, 1939, by original designation.

Generic diagnosis. In order to avoid repetition in the species descriptions, we here present those features found in all the known Nephochaetopteryx species (males and females).

Head. Gena and postgena with black setulae. Parafacial plate with very short, fine setae along lower anterior eye margin. One or two proclinate fronto-orbital setae (also in males). Two orbital setae. Inner vertical setae parallel and outer vertical setae divergent, differentiated from post-oculars. Occiput with black setulae. Scape and pedicel brown, first flagellomere gray, arista plumose in basal half.

Thorax. Brown with gold-gray microtomentum; scutum with three black longitudinal stripes; scutellum with a median black spot on anterior margin. Chaetotaxy: acrostichals = 0+1 (fine); postpronotals = 3; notopleurals = 2 with 1 to 2 subprimary notopleurals; postalars = 2; proepisternals = 2; proepimerals = 1; katepisternals = 3. Postallar wall and metasternum setulose. Scutellum with 1 pair of basal setae, 1 pair of lateral setae, 1 pair of apical setae and no discal setae. Legs. Brown. Fore femur with rows of anterodorsal and anteroventral setae. Fore tibia with one anteroventral seta in distal half and two posterodorsal setae in proximal half. Mid femur with two or three setae on anterior surface, rows of anteroventral and posteroventral setae on proximal half, and two apical setae on posterior surface. Mid femur of male with a ctenidium of rounded spines (circular cross section) on posteroventral surface. Mid tibia with two posterodorsal setae on distal half. Hind coxa with stout setae on posterior surface. Hind femur with two apical setae on posterior surface and rows of anterodorsal and anteroventral setae, first anteroventral seta about two times the length of the other setae. Hind tibia with two anterodorsal and two posterodorsal setae. Apical setae present on all tibiae. Wing. Vein R 1 entirely setulose dorsally. Third costal section bare on ventral surface. Wing hyaline or fumose between apical part of vein R 2+3 and costal vein. Cell r 4+5 open. Tegula blackish. Basicosta white.

Male terminalia. Sternite 5 usually with an arm, a lobe, and a cleft. Syntergosternite 7+8 and epandrium red or black. Basiphallus and distiphallus connected by a desclerotized strip. Acrophallus composed of capitis, hillae, lateral styli and median stylus. Lateral and median styli tubular.

Recognition. Nephochaetopteryx is usually characterized by wing with an apical spot and vein R 1 entirely setulose dorsally (Pape 1996). However, some specimens belonging to this genus have a hyaline wing, and some species of Panava Dodge and Sarcofahrtiopsis Hall also show R 1 entirely setulose (Lopes 1978; Carvalho-Filho et al. 2014). According to the phylogenetic hypothesis of Buenaventura & Pape (2018), Nephochaetopteryx is a natural group of species with three possible autapomorphies: “hind coxa with thick setae posteriorly”, “mid tibia with neither antero-dorsal nor antero-ventral setae” and “wing fumose between apical part of veins R 2+3 and C”.

Another feature found only in species of Nephochaetopteryx is the “inner process of vesica” covered with minute spine-like cuticular projections. However, the “inner process of vesica” is reduced or absent in N. angustifrons and N. pallidifacies . The term “inner process of vesica” is utilized to denominate the structure that Lopes called “base of ventralia” in the description of the male terminalia of his new species Nephochaetopteryx pacatubensis Lopes, 1975a . This term was also utilized by Lopes (1975a) to designate the structure covered with spine-like projections and located inside the distiphallus, attached to the basal portion of the vesica. In the same work, Lopes named a similar structure located inside the distiphallus of N. pallidiventris as “protection of glans”. However, as these structures are located inside the distiphallus and are covered with spine-like projections, we assume that they are homologous. This structure was labeled as “vesical arm-shaped lever” by Buenaventura & Pape (2018, fig. 18B), and according to them it is composed of an “apex” and a “base”. The “apex” is the visible portion of the “vesical arm-shaped lever” body that is joined with the “distal section of vesica”. The base is the portion of the “vesical arm-shaped lever” body that is usually hidden within the paraphallic tube. Based on the figure provided by Bue- naventura & Pape (2018, fig. 18B), it is possible to assesses that the basal portion of the “vesical arm-shaped lever” is the “inner process of vesica”. These authors also mentioned that in Nephochaetopteryx, the “vesical arm-shaped lever” apex is rounded or bilobed. Nephochaetopteryx is a speciose genus with more than 35 described species, but Buenaventura & Pape (2018) analyzed few species (about three). The vesica in Nephochaetopteryx is highly variable in shape and none of them seems to have a rounded or bilobed “vesical arm-shaped lever”. In addition, it is very difficult to assesse the limits between the “vesical arm-shaped lever” and the “distal section of vesica” in most of the species of this genus. For these reasons, we did not utilize the term “vesical arm-shaped lever” and we are utilizing the term “inner process of vesica” to denominate the basal portion of the “vesical arm-shaped lever” sensu Buenaventura & Pape (2018).

Buenaventura & Pape (2018) provided a list of diagnostic features for the genus Nephochaetopteryx, but some of them are not present in all species, such as “male abdominal ST4 [= sternite 4] with a dense patch of erect black setae near posterior margin”, a feature present only in few species. They also mentioned “vesica with distal section ornamented”, but the term “ornamented” was not explained. Based on the SEM images of the male terminalia of Nephochaetopteryx species showed by Buenaventura & Pape (2018, figs 18A–C, 25E), it is not possible to assesses what exactly the ornamentation is.

Species of Nephochaetopteryx are easily recognized by the following combination of features: small to medium-sized flies (4.0– 7.3 mm in length); wing vein R 1 entirely setulose dorsally; wing fumose between apical part of vein R 2+3 and costal vein or more rarely hyaline; male with one or two proclinate orbital setae; notopleuron with subprimary setae; postalar wall setulose; metasternum setulose; male mid femur with a ctenidium; hind coxa with stout setae on posterior surface; male sternite 5 usually with an arm, a lobe and a cleft; basiphallus and distiphallus connected by a desclerotized strip.

Biology. Little is known about the biology of Nephochaetopteryx species, but according to Lopes (1973), many species, in nature, deposit their larvae in feces of mammals and birds. Lopes (1973) obtained many species of this genus in traps baited with human feces in Brazil and one of us (FSCF) collected an adult male on dog feces in a secondary forest in the Brazilian Amazon. Curran & Walley (1934) also collected specimens of Nephochaetopteryx on human feces in Guyana. Lopes (1936, 1973) reared larvae of various species in an artificial medium composed of “gelose plus horse blood serum” or “gelose plus egg albumine”. Pape & Dahlem (2010) reported on larvae bred from vertebrate carcasses, and N. cyaneiventris, N. orbitalis, and N. pallidiventris have been collected from pig carcasses in Brazil (Vairo et al. 2011; Mello-Patiu et al. 2014). Females of Nephochaetopteryx have been collected in traps baited with rotting bovine lung in the Brazilian Amazon (Sousa et al. 2011). However, the natural feeding substrate of the larvae has not yet been reported.

Lopes (1975a) collected many adult Nephochaetopteryx flies in Van Someren-Rydon traps (butterfly trap) bait- ed with rotting banana in Ceará (northern Brazil). Van Someren-Rydon traps baited with pineapple were useful for collecting male specimens in a secondary forest in the Brazilian Amazon (FSCF, personal observation). Pape & Dahlem (2010) mentioned that fermented fruits are useful for attracting specimens of Nephochaetopteryx .

Nephochaetopteryx distincta Dodge, 1968b was collected flying close to trails of army ants of the species Eciton burchelli Westwood ( Hymenoptera: Formicidae) (information from specimen label). Adults of N. limpidipennis Lopes, 1976 were collected on mango flowers ( Mangifera indica; Anacardiaceae) in Mexico (information from specimen label).

Key to the males of Nephochaetopteryx

This key is based only on males, since females of most species remain unknown. Nephochaetopteryx coxalis Dodge, 1968a and N. maxima Dodge, 1968b were not included because their males are unknown.

1. Wing vein R 2+3 with setulae ventrally...................................................................... 2

- Wing vein R 2+3 without setulae ventrally................................................................... 6

2. Mid femur with small setae, shorter than height of mid femur (Fig. 38B). Vesica in lateral view with an elongate and narrowed tip (Figs 14F, 21F). Ventral margin of distiphallus serrated (Figs 14F, 21F)........................................ 3

- Mid femur with a long seta, longer than or as long as height of mid femur (Fig. 38A). Vesica dome-shaped in lateral view, with an enlarged tip (Figs 10E, 30F, 35F). Ventral margin of distiphallus not serrated (Figs 10E, 30F, 35F)................... 4

3. Tip of vesica in lateral view not curved (Fig. 21F). Apical margin of distiphallus smooth, not corrugated (Fig. 21F)............................................................................................... N. molinai Lopes

- Tip of vesica in lateral view strongly curved (Fig. 14F). Apical margin of distiphallus corrugated (Fig. 14F)..................................................................................................... N. inca sp. nov.

4. Apical margin of distiphallus in lateral view with a preapical concavity close to ventral margin (Fig. 30F). Surstylus with a broad rounded apex (Fig. 30A)............................................................. N. sofiae sp. nov.

- Apical margin of distiphallus entirely rounded in lateral view, without a preapical concavity close to ventral margin (Figs 10E, 35F). Surstylus with a narrowed, rounded apex (Figs 10E, 35F)................................................. 5

5. Cerci in dorsal view with cercal prongs slightly convergent (Fig. 35B). Ventral margin of distiphallus with a glossiform lobe projected anteriorly (Fig. 35F)............................................................ N. travassosi Lopes

- Cerci in dorsal view with cercal prongs parallel (Fig. 10B). Lateral margin of distiphallus with a glossiform lobe curved toward base of distiphallus (Fig. 10E)............................................................. N. distincta Dodge

6. Sternite 4 with a median patch of thick setae near posterior margin (Figs 37A, 37B, 38E)............................ 7

- Sternite 4 without a median patch of thick setae near posterior margin........................................... 11

7. Palpus yellow. Sternite 5 with lobes joined (Dodge 1968a, fig. S)................................ N. juquiana Dodge

- Palpus brown. Sternite 5 with lobes separated (Figs 5E, 16E, 25E).............................................. 8

8. Cercus with pointed tip, strongly curved ventrally and with a small, preapical protuberance (Figs 5 A–B).. N. boruca sp. nov.

- Cercus with tip rounded or quadrate, without a preapical protuberance (Figs 16 A–B, 25A–B, 31A–B).................. 9

9. Apical margin of distiphallus with a prominent concavity (Fig. 16F). Vesica short, with distal portion not reaching ventral margin of distiphallus (Fig. 16F).......................................................... N. limpidipennis Lopes

- Apical margin of distiphallus without a concavity (Figs 25F, 31F). Vesica elongate and angled, with distal portion reaching ventral margin of distiphallus (Figs 25F, 31F).............................................................. 10

10. Postgonite curved anteriorly (Fig. 31D). Vesica with a rectangular median projection (Fig. 31F)......... N. spinosa Dodge

- Postgonite curved posteriorly near middle (Fig. 25D). Vesica with a rounded median projection (Fig. 25F).............................................................................................. N. pallidiventris Townsend

11. Apical margin of distiphallus with two concavities (Fig. 13A). Surstylus about the same length as cercus (Fig. 13A)........................................................................................... N. fuscipennis Lopes

- Apical margin of distiphallus without a concavity (Fig. 9F). Surstylus shorter than cercus (Figs 1A, 2B, 7A)............ 12

12. Cercus with spines on dorsal surface (Figs 1 A–B, 7A–B, 9A–B, 22A–B)........................................ 13

- Cercus without spines on dorsal surface (Figs 2 A–B, 24A–B)................................................. 18

13. Cercal base with a prominent, rounded dorsal projection (Figs 9 A–B).......................... N. cyaneiventris Lopes

- Cercal base without a prominent, rounded dorsal projection (Figs 1 A–B, 7A–B, 22A–B)............................ 14

14. Surstylus with spines (Fig. 7A). Sternite 5 with two strong flattened setae at base of lobe (Fig. 7E)....... N. coendu sp. nov.

- Surstylus without spines (Figs 1A, 22A). Sternite 5 without strong flattened setae at base of lobe (Figs 1E, 22E)......... 15

15. Ventral margin of distiphallus not serrated (Fig. 1F). Cercus in dorsal view mostly narrowed, in lateral view strongly curved anteriorly (Figs 1 A–B)..................................................................... N. affinis Lopes

- Ventral margin of distiphallus serrated (Figs 22F, 27F). Cercus in dorsal view mostly broadened, in lateral view not so strongly curved anteriorly (Figs 22 A–B, 27A–B).................................................................. 16

16. Cercus in lateral view with a single, rounded preapical projection (Fig. 32A). Pregonite narrowed with rounded apex, slightly curved posteriorly..................................................................... N. subaurata (Engel)

- Cercus in lateral view with two or three rounded projections (Figs 22A, 27B). Pregonite with a different combination of features.............................................................................................. 17

17. Cercus in lateral view with three rounded projections (Fig. 27B). Pregonite subrectangular (Fig. 27C).................................................................................................. N. psittacocercus sp. nov.

- Cercus in lateral view with two rounded projections (Fig. 22A). Pregonite claw-shaped (Fig. 22C)................................................................................................ N. orbitalis (Curran & Walley)

18. Basiphallus as long as or longer than distiphallus. Ventroapical margin of distiphallus with a strong projection (Figs 2F, 24F). Palpus yellow....................................................................................... 19

- Basiphallus shorter than distiphallus. Ventroapical margin of distiphallus without a strong projection (Figs 4F, 23D). Palpus brown or yellow..................................................................................... 20

19. Ventral margin of distiphallus membranous (Fig. 2F). Vesica with a median spiny projection (Fig. 2F). Lateral stylus as long as vesica (Fig. 2F)...................................................................... N. angustifrons Lopes

- Ventral margin of distiphallus sclerotized (Fig. 24F). Vesica without a median spiny projection (Fig. 24F). Lateral stylus shorter than vesica (Fig. 24F)................................................................. N. pallidifacies Lopes

20. Cercus with one or two dorsal excavations (Figs 4 A–B)...................................................... 21

- Cercus without a dorsal excavation (Figs 3B, 15B, 19B)..................................................... 22

21. Cercus with one large dorsal excavation (Figs 4 A–B). Apical margin of distiphallus with a fissure (Fig. 4F).................................................................................................... N. biculcita Dodge

- Cercus with two dorsal excavations (Fig. 23A). Apical margin of distiphallus without a fissure (Fig. 23D)................................................................................................. N. pacatubensis Lopes

22. Cercus (lateral view) with a preapical protuberance on dorsal margin (Figs 15A, 19A).............................. 23

- Cercus (lateral view) without a preapical protuberance on dorsal margin (Fig. 3A)................................. 25

23. Sternite 5 with lobe almost triangular and bare (Fig. 15E). Surstylus with rounded apex, not curved posteriorly (Fig. 15A). 24

- Sternite 5 with lobe rounded and with a tuft of setulae (Fig. 19E). Surstylus with pointed apex, strongly curved posteriorly (Fig. 19A)................................................................................ N. matinta sp. nov.

24. Vesica in lateral view with a strong triangular basal projection and not strongly arched distally (Fig. 15F)..................................................................................................... N. lamasi sp. nov.

- Vesica in lateral view without a triangular basal projection and strongly arched distally (Hime 1985, figs 7–9)................................................................................................ N. panamensis Hime

25. Lateral wall of distiphallus with many tiny cuticular spines (Figs 3F, 8E). Ventral margin of distiphallus with two strong, pointed projections (Figs 3F, 8E)........................................................................ 26

- Lateral wall of distiphallus without cuticular spines (Figs 26F, 28E). Ventral margin of distiphallus different............ 27

26. Proximal posterior margin of distiphallus with many small cuticular spines (Fig. 3F). Cercus without a preapical tuft of hair-like setulae (Figs 3 A–B). Distal portion of vesica without cuticular spines (Fig. 3F).................... N. aurescens Dodge

- Proximal posterior margin of distiphallus without small cuticular spines (Fig. 8E). Cercus with a preapical tuft of hair-like setulae (Figs 8 A–B). Distal portion of vesica with many cuticular spines (Fig. 8E)..................... N. cuzco sp. nov.

27. Sternite 5 with a lobe and with a broad cleft (Fig. 26E) or without a cleft (Fig. 33E). Pregonite without a blunt projection basally (Figs 26C, 28C)..................................................................................... 28

- Sternite 5 without a lobe and with a very narrow cleft (Fig. 12E). Pregonite with a blunt projection basally (Fig. 12C)......................................................................................... N. flavipalpis Lopes

28. Pregonite with a prominent glossiform projection on anterior margin (Fig. 6C). Ventral margin of distiphallus serrated (Fig. 6F).................................................................................... N. canga sp. nov.

- Pregonite without a prominent glossiform projection on anterior margin. Ventral margin of distiphallus not serrated (Figs 26F, 33F)............................................................................................... 29

29. Cercal prongs not separated in dorsal view (Fig. 33B). Sternite 5 with lobe very short and rectangular, and without a cleft (Fig. 33E).................................................................................. N. tembe sp. nov.

- Cercal prongs widely separated in dorsal view (Figs 26B, 28B). Sternite 5 with lobe well developed and not rectangular, and with a broad cleft (Figs 26E)........................................................................... 30

30. Sternite 5 with lobe pointed and bare (Fig. 26E). Surstylus with pointed apex, curved posteriorly (Fig. 26A).................................................................................................. N. paraensis Dodge

- Sternite 5 with lobe rounded and with a tuft of setulae (Figs 17E, 34E). Surstylus with rounded apex, not curved posteriorly (Figs 11A, 17A, 28A, 34A)............................................................................ 31

31. Cercus in lateral view with a cluster of stout and pointed preapical setae on ventral margin (Fig. 28A) N. rettenmeyeri Dodge

- Cercus in lateral view without a cluster of stout and pointed preapical setae on ventral margin (Figs 11A, 17A, 34A)..... 32

32. Anterior margin of pregonite without small, pointed projections (Figs 11C, 18C, 29C, 34C, 36C). Vesica with a simple (nonbifid) tip in lateral view (Figs 11E, 18F, 29F, 34F, 36F)...................................................... 33

- Anterior margin of pregonite with small, pointed projections (Fig. 17C). Vesica with a bifid tip in lateral view (Fig. 17F)........................................................................................... N. lopesi Dodge

33. Anterior margin of pregonite with grooves or granulations (Figs 18C, 36C). Tip of vesica strongly curved anteriorly in lateral view (Figs 18F, 36F).................................................................................. 34

- Anterior margin of pregonite without grooves or granulations (Figs 11C, 29C, 34C). Tip of vesica curved posteriorly in lateral view (Figs 11E, 29F, 34F)............................................................................. 35

34. Palpus brown. Vesica with a large, rounded projection in lateral view (Fig. 18F). Ventral margin of distiphallus with a claw-like projection (Fig. 18F).................................................................... N. marianae Dodge

- Palpus yellow. Vesica with a small triangular projection in lateral view (Fig. 36F). Ventral margin of distiphallus with a pointed projection (Fig. 36F).......................................................... N. utinguensis Tibana & Hime

35. Pregonite with a prominent, rounded projection on posterior margin (Fig. 11C).................. N. equatoriana sp. nov.

- Pregonite without a prominent, rounded projection on posterior margin (Figs 29C, 34C)............................ 36

36. Cercal prongs with convergent tips in dorsal view (Fig. 34B). Surstylus with setulae covering all of basal half (Fig. 34A). Distal portion of pregonite widened (Fig. 34C)................................................... N. tinguensis Dodge

- Cercal prongs with parallel tips in dorsal view (Fig. 29B). Surstylus with setulae restricted to posterobasal corner (Fig. 29A). Distal portion of pregonite narrowed (Fig. 29C)............................................... N. similis sp. nov.