Nemopterinae Tillyard, 1925

Subfamily Nemopterinae Tillyard, 1925

Type genus: Nemoptera Latreille, 1802 .

Included genera: The long-abdomen (LA) group: Brevistoma Tjeder, 1967 , Chasmoptera Kirby, 1900 , Derhynchia Tjeder, 1967 , Halter Rambur, 1842 , Lertha Navás, 1915 , Nemeura Navás, 1915 , Nemoptera Latreille, 1802 , Nemopistha Navás, 1910 , Parasicyoptera Tjeder, 1974 , Savignyella Kirby, 1900 , Semirhynchia Tjeder, 1967 , Sinonemoptera gen. nov., Sicyoptera Navás, 1910 , and Stenorrhachus McLachlan, 1886 . The shortabdomen ( SA) group: Afroptera Abdalla & Mansell in Abdalla et al., 2019, Barbibucca Tjeder, 1967 , Halterina Navás, 1912 , Knersvlaktia Picker, 1984 , Nemia Navás, 1915 , Nemopterella Banks, 1910 , Palmipenna , and Siccanda Abdalla & Mansell in Abdalla et al., 2019. Fossil genera: † Marquettia Navás, 1913 and † Paleonemia Claisse, Brisac & Nel, 2019 .

Diagnosis: Forewing basally wide (not narrow); at least five presectoral crossviens present. Hindwing spoon- or ribbonlike, distal part dilated in some species; vein M present. Male gonocoxites 9 posteriorly fused into a single lobe.

Remarks: Based on the phylogeny herein reconstructed, the subfamily Nemopterinae can be divided into two monophyletic groups, i.e. the long-abdomen (LA) group and the shortabdomen ( SA) group. The LA group can be distinguished by the abdomen longer than the head plus thorax, the male gonocoxites 9 anteriorly not protruding ventrad in lateral view, and the male gonostyli 11 laterally protruding as a pair of processes. Notably, the LA group includes two genera that lack a specialized rostrum ( Derhynchia and Stenorrhachus ), and the female of Stenorrhachus walkeri (McLachlan, 1885) has the forewing reduced into a stub. The LA group is widely distributed in arid and warm areas.

The SA group can be distinguished by the abdomen nearly as long as the head plus thorax at most, the male gonocoxites 9 anteriorly protruded ventrad in lateral view, and the male gonostyli 11 fused as a single linguiform structure. Some genera of the SA group ( Afroptera , Knersvlaktia , Nemopterella , Palmipenna , and Siccanda ) are equipped with pleuritocavae on abdominal segment 5, which function for releasing pheromones ( Tjeder 1967). The SA group is endemic to the continent of southern Africa but always in arid areas. The two Oligocene fossil genera, † Marquettia and † Paleonemia , might belong to the LA group because of their long abdomen, although we could not test their phylogenetic positions further owing to lack of sufficient morphological characters preserved in these fossils. Besides, the North American † M. americana seems to be closer to the Australian Chasmoptera rather than Sicyoptera , as mentioned by Sole et al. (2013) based on the dark body, the relatively small eyes, the forewing only with two anal veins, the doubled hindwing dilations, and the male ectoproct possibly with elongate posterior lobe.

Key to extant genera of Nemopterinae

1. Abdomen longer than head plus thorax; male sternum 9 relatively long, ≥1.5 times as long as ectoproct (excluding posterior lobe in some genera) in lateral view; male gonocoxites 9 anteriorly not protruded ventrad in lateral view, and gonostyli 11 laterally with a pair of processes...................................................................................................................................................................2

– Abdomen as long as head plus thorax at most; male sternum 9 relatively short, nearly 1.0–1.5 times as long as ectoproct; male gonocoxites 9 anteriorly protruded ventrad in lateral view, and gonostyli 11 posteriorly fused as a single lingulate structure......................................................................................................................................................................................................... 16

2. Mouthparts not specialized into a rostrum; male tergum 9 fused with ectoprocts ..........................................................................3

– Mouthparts specialized into a rostrum; male tergum 9 distinctly separated from ectoprocts........................................................4

3. Labrum visible in frontal view; poststigmal area with single row of cells ( Tjeder 1967: figs 1887, 1888; Miller and Stange 2012: figs 1, 6, 10, 14).................................................................................................................................... Stenorrhachus McLachlan

– Labrum enfolded by clypeus and not visible in frontal view; poststigmal area with two rows of cells ( Tjeder 1967: figs 1881, 1883, 2024).................................................................................................................................................................... Derhynchia Tjeder

4. Body mostly black dorsally ( Fig. 5c View Figure 5 ); vertex with many irregular wrinkles ( Fig. 6E View Figure 6 ); apical labrum split ( Fig. 6E View Figure 6 ); forewing with five ra-rp1 crossveins at most ( Fig. 5c View Figure 5 ) and with only two anal veins; ectoproct with an elongate posterior lobe ( Fig. 8F View Figure 8 ) ......................................................................................................................................................................................... Chasmoptera Kirby

– Body dark brown to yellowish brown with markings dorsally; vertex relatively smooth; apical labrum not split, relatively rounded or truncate; forewing with six ra-rp1 crossveins at least, with three anal veins; ectoproct without posterior elongation.........................................................................................................................................................................................................5

5. Basal c-sc and m-av crossveins without markings; male abdominal segments 3–8 nearly 1.2–1.5 times as long as wide .......... ............................................................................................................................................................................................................................6

– Basal c-sc and m-av crossveins each suffuse a marking; male abdominal segments 3–8 ≥1.5 times as long as wide.............. 11

6. Head naked around eyes; male gonocoxites 11 concave in cephalic view, gonostyli 11 extremely swollen, present as a pair of wide lingulate processes ( Fig. 8G, H View Figure 8 ) ........................................................................................................................................................7

– Head setose around eyes; male gonocoxites 11 not concave in cephalic view, gonostyli 11 not swollen ....................................8

7. Hindwing without expanded dilation ( Tjeder 1967: fig. 2072).......................................................................... Brevistoma Tjeder

– Hindwing with an expanded dilation anterior to Sc-R fusion ( Fig. 7K View Figure 7 ).................................................................. Halter Rambur

8. Forewing RP approximating MA, CuA and MP diverging at basal one-eighth ( Tjeder 1967: figs 2074, 2075) ......................... ............................................................................................................................................................................................ Savignyella Kirby

– Forewing RP not approximating MA, CuA and MP diverging at basal one-fifth or more distal ...................................................9

9. Forewing prefork area longer than half of forewing length; lateral part of male gonostyli 11 elongated and upcurved ( Tjeder 1974: figs 14, 19, 20).............................................................................................................................................. Parasicyoptera Tjeder

– Forewing prefork area not longer than half of forewing length; lateral part of male gonostyli 11 relatively short and not upcurved........................................................................................................................................................................................................ 10

10. Forewing mostly hyaline; male gonocoxites 9 without plate-like structure at tip, gonostyli 9 absent ( Monserrat 1988: figs 18–22) ....................................................................................................................................................................................... Lertha Navás

– Forewing pale yellow with many brown markings ( Fig. 5e View Figure 5 ); male gonocoxites 9 with plate-like structure at tip, gonostyli 9 present ( Fig. 8B View Figure 8 ) ......................................................................................................................................................... Nemoptera Latreille

11. Forewing mediocubital crossveins and first cup-a1 crossvein without marking ............................................................................ 12

– Forewing mediocubital crossveins and first cup-a1 crossvein with marking .................................................................................. 14

12. Rostrum nearly half length of head width ( Tjeder 1967: figs 1891, 1897); hindwing without expanded double dilations ( Tjeder 1967: figs 2049–2051) ............................................................................................................................. Semirhynchia Tjeder

– Rostrum longer than half of head width; hindwing with expanded double dilations ................................................................... 13

13. Vertex medially concave; hindwing with double dilations, and each nearly as long as wide ( Tjeder 1967: figs 2222, 2223, 2232)................................................................................................................................................................................... Sicyoptera Navás

– Vertex not concave; hindwing with double dilations, and each longer than wide ( Tjeder 1967: figs 2113, 2116)..................... ................................................................................................................................................................ Nemeura Navás (partial species)

14. Forewing markings suffuse basal four mediocubital crossveins at most ( Fig. 2E, G View Figure 2 ); male sternum 9 bifurcated on distal half ( Fig. 3C, G View Figure 3 )......................................................................................................................................................... Sinonemoptera gen. nov.

– Forewing markings suffuse more than five mediocubital crossveins; male sternum 9 not bifurcated on distal half 15

15. Forewing prefork and basal cubital crossveins each suffuse a small dark marking ( Fig. 7A View Figure 7 ); hindwing dilated nearly at distal one-quarter or more distal ( Fig. 5g View Figure 5 ); male sternum 9 with lateral distal margin extended ( Kimmins 1938: fig. 2; Tjeder 1967: figs 2203, 2211).................................................................................................................................................. Nemopishta Navás

– Forewing prefork and basal cubital crossveins without spot ( Tjeder 1967: figs 2136, 2137, 2139, 2140, 2142, 2143); hindwing dilated nearly at distal one-third ( Tjeder 1967: fig. 2109); male sternum 9 with lateral distal margin not extended ..................................................................................................................... Nemeura Navás (partial species, including genus type)

16. Diameter of eye less than head width; many setae present around eyes (Supporting Information, Fig. S1F View Figure 1 )......................... 17

– Diameter of eye as long as or longer than head width; no setae around eyes ( Abdalla et al. 2019: fig. 9) ................................ 20

17. Forewing basal three-quarters brown, not hyaline; hindwing not dilated, pterostigma indistinct (Supporting Information, Fig. S1G View Figure 1 ; Picker 1984: fig. 6) ................................................................................................................................... Knersvlaktia Picker

– Forewing mostly hyaline; hindwing dilated, pterostigma distinct .................................................................................................... 18

18. Hindwing completely not hyaline, brown on basal half, only narrowly dilated near pterostigmal part ( Tjeder 1967: figs 2306, 2307) ............................................................................................................................................................................... Barbibucca Tjeder

– Hindwing partly hyaline on basal half, with an expanded foliate dilation ....................................................................................... 19

19. Male antenna basally with relatively long, stout setae ( Tjeder 1967: figs 2254, 2255); hindwing pterostigma large, costal crossveins and anal crossveins on distal dilation connected by many oblique veinlets ( Tjeder 1967: figs 2263–2266) ......................................................................................................................................................................................... Palmipenna Tjeder

– Male antenna basally short, without stout setae ( Tjeder 1967: fig. 2286); hindwing pterostigma indistinct, costal crossveins and anal crossveins on distal dilation barely connected by oblique veinlets ( Tjeder 1967: figs 2289, 2292)............................... ............................................................................................................................................................................................... Halterina Navás

20. Most forewing costal cells each with a pale marking, a dark brown marking suffused on pterostigma ( Abdalla et al. 2019: fig. 158); male abdominal pleuritocavae absent ...................................................................................................................... Nemia Navás

– Forewing costal cells without a pale marking, no marking suffused on pterostigma; male abdominal pleuritocavae present.. ......................................................................................................................................................................................................................... 21

21. Distal flagellomere with partly membranous structure, more conspicuous in male ( Tjeder 1967: figs 2364–2381; Abdalla et al. 2019: fig. 39); many forewing costal crossveins each with a spot, pterostigma dark brown ( Abdalla et al. 2019: fig. 38) ....................................................................................................................................................................... Afroptera Abdalla & Mansell

– Distal flagellomere completely sclerotized ( Tjeder 1967: figs 2362, 2363; Abdalla et al. 2019: fig. 8 View Figure 8 ); forewing costal crossveins without spot, pterostigma pale ( Abdalla et al. 2019: figs 17–19) .................................................................................. 22

22. Tip of distal flagellomere with acute tooth ( Tjeder 1967: fig. 2363); forewing anal area without marking ( Abdalla et al. 2019: fig. 24) ........................................................................................................................................................... Siccanda Abdalla & Mansell

– Tip of distal flagellomere without tooth ( Tjeder 1967: fig. 2362); forewing basal anal area with a brown marking ( Abdalla et al. 2019: fig. 7 View Figure 7 )............................................................................................................................................................. Nemopterella Banks