Ophrynopus Konow, 1897

Ophrynopus Konow, 1897

Type species: Ophrynopus andrei Konow, 1897 = Ophrynopus fulvostigma (Westwood, 1874)

Diagnosis. Head with frons densely sculptured. Female antenna usually with antennomere 9 swollen distally, less swollen in some species (e.g., O. rufocephalus Vilhelmsen, 2020 and O. seagi ( Vilhelmsen 2016)) . Head with postocular carina present, usually extending to a considerable distance above ventral margin of eyes. Labiomaxillary complex well developed, maxillary palps elongate, with five palpomeres. Mesoscutellum with lateral margin indistinct, posterior margin parallel with anterior margin of metanotum for some distance, mesoscutellum not raised relative to adjacent sclerites. Fore wing vein 2r arising from pterostigma at more than 0.6 of pterostigma length. Hind femur with distinct triangular projection on laterodistal corner. Hind margin of tergum 1 with scattered setae.

Key to extant Ophrynopus species, including key to ophrynopine genera (modified from Vilhelmsen 2020)

1. Fore wing vein 2r arising from pterostigma more than 0.6 of pterostigma length from base of latter (Vilhelmsen 2020, Figs 5B View FIGURES 1–6 , 7D View FIGURES 7–10 ; width of pterostigma measured from distal end of costal cell to apex of pterostigma, insertion of vein 2r measured from distal end of costal cell to distal margin of 2r). Hind femur with distinct triangular projection on laterodistal corner (Vilhelmsen 2020, Figs 5C, 6A View FIGURES 1–6 , 7A View FIGURES 7–10 ) or hind margin of tergum 1 with distinct dense transverse fringe of setae. Sternum 9 of male with projections anteromedially and posterolaterally ( Vilhelmsen et al. 2013, fig. 8E).................. 2 (ophrynopine clade)

- Fore wing vein 2r arising from pterostigma less than or equal to 0.6 of pterostigma length from base of latter ( Vilhelmsen 2003, figs 63–71). Hind femur with laterodistal corner rounded and hind margin of tergum 1 at most with scattered setae. Sternum 9 of male without projections..................................................................other Orussidae View in CoL

2. Head with postocular carina distinct but present only ventrally, extending at most slightly above lower margin of eyes ( Vilhelmsen 2003, fig. 20). Labiomaxillary complex reduced, maxillary palp very short, with three palpomeres. Hind femur with laterodistal corner rounded. Tergum 1 with distinct dense transverse fringe of setae on hind margin ( Vilhelmsen 2003, fig. 80). ( Costa Rica, Mexico, southern USA)............................................... Kulcania Benson, 1935

- Head with postocular carina usually extending for a considerable distance above lower margin of eyes (Vilhelmsen 2020, Fig. 6A, C View FIGURES 1–6 ; rarely weakly developed Vilhelmsen 2020, Figs 4A, 5A View FIGURES 1–6 ). Labiomaxillary complex well developed, maxillary palps elongate, with five palpomeres (Vilhelmsen 2020, Figs 4A View FIGURES 1–6 , 7A View FIGURES 7–10 ). Hind femur with laterodistal corner having distinct, triangular projection (Vilhelmsen 2020, Figs 5C, 6A View FIGURES 1–6 , 7A View FIGURES 7–10 ). Hind margin of tergum 1 at most with scattered setae (Vilhelmsen 2020, Figs 5C, 6E View FIGURES 1–6 , 7C View FIGURES 7–10 ).......................................................................................... 3

3. Head with frons having only scattered punctuation, separated by large smooth spaces ( Blank et al. 2010, figs 5, 11, 16, 22, 28, 34, 41). Occipital carina absent ( Blank et al. 2010, fig. 38). Fore wing Rs hyaline or weakly pigmented distally ( Blank et al. 2010, figs 6, 12, 14, 20, 26, 32, 42). (USA: California)................................ Ophrynon Middlekauff, 1983

- Head with frons densely sculptured (Vilhelmsen 2020, Figs 4B, 6B View FIGURES 1–6 , 7B View FIGURES 7–10 ). Occipital carina present (Vilhelmsen 2020, Figs 5A, 6C View FIGURES 1–6 ). Fore wing Rs dark (Vilhelmsen 2020, Figs 5B, 6D View FIGURES 1–6 , 7D View FIGURES 7–10 )................................................... 4

4. Mesoscutellum with distinct lateral margin, posterior margin curved, mesoscutellum raised relative to adjacent sclerites ( Vilhelmsen & Smith 2002, fig. 13). Female antenna with antennomere 9 not swollen distally ( Vilhelmsen & Smith 2002, fig. 9) and without carina laterally. ( Australia, Chile, Fiji, New Zealand)............................ Guiglia Benson, 1938

- Mesoscutellum with lateral margin indistinct, posterior margin parallel with anterior margin of metanotum for some distance, mesoscutellum not raised relative to adjacent sclerites ( Vilhelmsen et al. 2013, figs. 11D, 13A). Female antenna usually with antennomere 9 swollen distally (Vilhelmsen 2020, Fig. 6C View FIGURES 1–6 ) and with carina laterally; if antennomere 9 not swollen (Vilhelmsen 2020, Fig. 4C View FIGURES 1–6 ), then groove between dorsalmost coronal teeth present (Vilhelmsen 2020, Figs 4B, 5A View FIGURES 1–6 ) (Central and South America, Far East including Japan, New Guinea)....................................5 ( Ophrynopus Konow, 1897a )

5. Head with posterior coronal teeth separated by narrow longitudinal furrow (Vilhelmsen 2020, Figs 4B, 5A View FIGURES 1–6 ; Vilhelmsen et al. 2013, figs 4B, 5B, 6B, C, 7B, C; Vilhelmsen 2016, figs 4A, 5A). Fore wing vein cu-a inserts on Cu1 distally of vein M, in middle of discal cell 1M (Vilhelmsen 2020, Fig. 5B View FIGURES 1–6 ; Vilhelmsen et al. 2013, figs 5 E, 6 E, 8B; Vilhelmsen 2016, fig. 4A); female antennomere 10 at least 3x as long as broad (Vilhelmsen 2020, Fig. 4C View FIGURES 1–6 ; Vilhelmsen et al. 2013, fig. 5D; Vilhelmsen 2016, fig. 4C).................................................................................................6

- Posterior coronal teeth not separated by narrow longitudinal furrow medially (Vilhelmsen 2020, Figs 6B, C View FIGURES 1–6 , 7B, C View FIGURES 7–10 ). Fore wing vein cu-a usually inserts on Cu1 opposite or at most slightly distal to vein M (Vilhelmsen 2020, Fig. 7D View FIGURES 7–10 ; Vilhelmsen et al. 2013, figs 10D, 13D). Female antennomere 10 less than 3x as long as broad (Vilhelmsen 2020, Fig. 6B, C View FIGURES 1–6 ; Vilhelmsen et al. 2013, figs 10C, 12 E).........................................................................................9

6. Postocular carina present, with dense pilosity between posterior margin of eye and carina ( Vilhelmsen et al. 2013, figs 4A, 5C, 6A, C, 7A, C, 8A). Female antennomere 9 swollen distally ( Vilhelmsen et al. 2013, fig. 5D). Transverse pronotal carina present ( Vilhelmsen et al. 2013, fig. 7D) or absent ( Vilhelmsen et al. 2013, figs 4C, 5C, 6A), if present without median incurvation ( Vilhelmsen et al. 2013, fig. 7C). Hind tibia with only one short apical spur ( Vilhelmsen et al. 2013, figs 6D, 7 E)..........7

- Postocular carina absent, only scattered hairs present posterior to eye (Vilhelmsen 2020, Figs 4A, 5A View FIGURES 1–6 ; Vilhelmsen 2016, figs 4C, 5A, 6A). Female antennomere 9 not swollen distally (Vilhelmsen 2020, Fig. 4C View FIGURES 1–6 ; Vilhelmsen 2016, fig. 4C). Transverse pronotal carina present, distinct and with distinct incurvation medially (Vilhelmsen 2020, Fig. 5A View FIGURES 1–6 ; Vilhelmsen 2016, figs 5A, 6A). Hind tibia with two apical spurs.......................................................................8

7. Lower frons with broad, distally rounded, leaf-shaped setae ( Vilhelmsen et al. 2013, fig. 4D). Pronotum without distinct transverse carina dorsally ( Vilhelmsen et al. 2013, figs 4C, 5C, 6A). Female predominantly dark brown to black ( Vilhelmsen et al. 2013, figs 4A, 5A, 6A) ( Brazil, French Guiana, Panama)............... .. Ophrynopus amazonicus (Westwood, 1874)

- Lower frons with elongate, distally truncated setae ( Vilhelmsen et al. 2013, fig. 4 E). Pronotum with distinct transverse carina dorsally ( Vilhelmsen et al. 2013, fig. 7D). Female extensively reddish brown on legs and abdomen ( Vilhelmsen et al. 2013, figs 7A, D, E, 8C) ( French Guiana).......................................... Ophrynopus eldorado ( Vilhelmsen, 2013)

8. Body predominantly dark brown to black ( Vilhelmsen 2016, figs 4A. 6A). Dorsal ocellar coronal teeth low, in regular transverse rows ( Vilhelmsen 2016, figs 4B, 5A, 6B). Fore femur without ventral frontal carina. Lateral smooth areas on abdominal tergum 2 distinct ( Vilhelmsen 2016, fig. 5B) ( French Guiana)......................... Ophrynopus seagi ( Vilhelmsen, 2016)

- Head, prothorax and proximal parts of mid and hind legs reddish brown (Vilhelmsen 2020, Fig. 4A View FIGURES 1–6 ). Dorsal ocellar coronal teeth distinct, staggered (Vilhelmsen 2020, Figs 4B, 5A View FIGURES 1–6 ). Distinct longitudinal carina present ventrally on fore femur. Lateral smooth areas on abdominal tergum 2 hardly discernible ( Ecuador)........... Ophrynopus rufocephalus Vilhelmsen, 2020

9. Head with only scattered setae posterior to eyes (Vilhelmsen 2020, Figs 6A View FIGURES 1–6 , 7A, C View FIGURES 7–10 ; Vilhelmsen et al. 2013, fig. 11B)..... 10

- Head with dense pilosity posterior to eyes ( Vilhelmsen et al. 2013, fig. 12C)......................................15

10. Mesepisternal carina entirely absent..................................................................... 11

- Vertical mesepisternal carina developed to various degrees ( Vilhelmsen et al. 2013, figs 9 E, 11C)..................... 12

11. Head with postocular carina developed only ventral to eyes. Scape, all femora and fore and midtibiae reddish brown. Fore wing pterostigma in female black throughout (male unknown) ( Mexico)..... Ophrynopus enigmus ( Vilhelmsen & Smith, 2002)

- Head with well developed postocular carina extending for a considerable distance posterior to eye (Vilhelmsen 2020, Fig. 7A View FIGURES 7–10 ). Scape and all femora and tibiae dark brown (Vilhelmsen 2020, Fig. 7A View FIGURES 7–10 ). Forewing pterostigma in female white basally ( Vilhelmsen & Smith 2002, fig. 26), in male mostly black (Vilhelmsen 2020, Fig. 7D View FIGURES 7–10 ) ( Costa Rica)................................................................................. Ophrynopus gauldi ( Vilhelmsen & Smith, 2002)

12. Mesepisternal carina prominent, present along dorsal half of anterior edge of mesepisternum ( Vilhelmsen et al. 2013, fig. 9 E). Pronotum dorsally almost without setae ( Vilhelmsen et al. 2013, fig. 9A, D)..................................... 13

- Mesepisternal carina inconspicuous, absent along dorsal half of anterior edge of mesepisternum ( Vilhelmsen et al. 2013, fig. 1C). Pronotum dorsally with scattered white setae ( Vilhelmsen et al. 2013, figs 10A, 11D) ( Bolivia)........................................................................... Ophrynopus guarani Blank, Smith & Vilhelmsen, 2013

13. Antennomeres 6–7 mostly creamy white, contrasting with the rest of the antennomeres being dark brown (Vilhelmsen 2020, Fig. 6A–C View FIGURES 1–6 ). Sculpture on frons and thorax less coarsely areolate (Vilhelmsen 2020, Fig. 6B, C View FIGURES 1–6 ) ( French Guiana)............................................................................... Ophrynopus savinai Vilhelmsen 2020

- Antennae brown throughout, no antennomeres significantly differently colored ( Vilhelmsen et al. 2013, fig. 9A). Sculpture on frons and thorax coarsely areolate ( Vilhelmsen et al. 2013, fig. 9B, D, E) ( Brazil)................................. 14

14. Pronotum of approximately equal length in dorsal view ( Vilhelmsen et al. 2013, fig. 9D). Sculpture on lateral of mesepisternum with irregular coarsely areolate pits ( Vilhelmsen et al. 2013, fig. 9 E). Median metanotal carina present.............................................................................. Ophrynopus carinatus Vilhelmsen & Smith, 2002

- Pronotum in dorsal view longer sublaterally than medially ( Fig. 3 View FIGURES 1–6 ). Sculpture on the lateral of mesepisternum with shallower rounded pits and, with some spaces between pits. Median metanotal carina absent ( Fig. 3 View FIGURES 1–6 )..... Ophrynopus aridus sp. nov.

15. Head anteriorly dull black, without metallic sheen ( Vilhelmsen et al. 2013, fig. 11A). Median longitudinal carina dorsally on pronotum present ( Vilhelmsen & Smith 2002, fig. 18)....................................................... 16

- Head anteriorly dark green with metallic luster ( Vilhelmsen et al. 2013, fig. 12B). Median longitudinal carina dorsally on pronotum absent..................................................................................... 19

16. Head with lateral longitudinal carina weakly developed or absent. Fore wing with two broad infuscate bands traversing entire length of wing ( Indonesia, Papua New Guinea)............................ Ophrynopus maculipennis (F. Smith, 1859)

- Head with lateral longitudinal frontal carina well developed ( Vilhelmsen & Smith 2002, fig. 66). Fore wing either with infuscate area proximal to pterostigma reduced ( Vilhelmsen & Smith 2002, fig. 30), not traversing wing, or proximal infuscate area entirely absent....................................................................................... 17

17. Head with considerable area devoid of sculpture dorsal to median ocellus ( Vilhelmsen & Smith 2002, fig. 6). Hind tibia with lateral carina (e.g., Vilhelmsen & Smith 2002, fig. 22; usually less developed in male). Fore wing infuscate area proximal to pterostigma usually developed anteriorly, at least in female ( Indonesia, Laos, Malaysia, Philippines, Vietnam).................................................................................... Ophrynopus kohli (Konow, 1897b)

- Head without distinct smooth area dorsal to median ocellus (e.g., Vilhelmsen & Smith 2002, fig. 5). Hind tibia without lateral carina. Fore wing with only one infuscate area, situated distal to pterostigma..................................... 18

18. Fore wing pterostigma predominantly white ( Japan).................. Ophrynopus tosensis (Tosawa & Sugihara, 1934)

- Fore wing pterostigma entirely black (e.g., Vilhelmsen & Smith 2002 fig. 30) ( Japan).. Ophrynopus apicalis (Togashi, 2000)

19. Mesoscutum with distinct posterior pilosity ( Vilhelmsen et al. 2013, fig. 13A). Hind coxa with dense lateral pilosity ( Vilhelmsen et al. 2013, fig. 10B). Female abdomen with lateral setose patches on terga 3–7 ( Vilhelmsen et al. 2013, fig. 12A), becoming progressively broader posteriorly.........................................................................20

- Mesoscutum with only scattered setae. Hind coxa with at most scattered setae. Female abdomen without lateral setose patches on terga............................................................................................ 24

20. Head in lateral view with depression below ventral coronal tooth ( Vilhelmsen et al. 2013, fig. 12C). Hind femur dark brown to black. Fore wing apex infuscate, with hyaline spot subapically ( Vilhelmsen et al. 2013, fig. 13D) ( Brazil).................................................................. Ophrynopus rupestris Vilhelmsen, Costa & Alvarenga, 2013

- Head in lateral view without depression. Hind femur red or reddish brown. Fore wing apex hyaline, infuscate areas more proximal ( Vilhelmsen et al. 2013, fig. 10D)............................................................... 21

21. Median longitudinal carina on mesoscutum distinct ( Fig. 9 View FIGURES 7–10 ); mesepisternal carina absent ( Fig. 14 View FIGURES 11–16 ); ventral denticles on hind femur absent............................................................. Ophrynopus caatinguensis sp. nov.

- Median longitudinal carina on mesoscutum less developed; mesepisternal carina present; ventral denticles on hind femur present..............................................................................................22

22. Fore wing pterostigma predominantly black, at most with small white area basally ( Brazil, French Guiana)................................................................................ Ophrynopus batesianus (Westwood, 1874)

- Fore wing pterostigma predominantly white............................................................... 23

23. Antenna with some basal antennomeres light brown or reddish brown. Fore and mid tibiae reddish brown, hind tibia dark brown ( Costa Rica, Trinidad).......................................... Ophrynopus hansoni Vilhelmsen & Smith, 2002

- Antenna black to dark brown. Fore and mid tibiae dark brown or black, hind tibia black ( Brazil, Ecuador, French Guiana)................................................................... Ophrynopus fulvostigma (Westwood, 1874)

24. Postocular carina diverging from lower half of eye, space between eye and carina broadest toward ventral half of eye. Mesopleuron with scattered punctures, separated by large shiny interspaces ( Argentina)................................................................................................ Ophrynopus wagneri (du Buysson, 1910)

- Postocular carina not diverging from lower half of eye, space between eye and carina broadest at mid-level of eye or toward upper half of eye, or eye and carina parallel. Mesopleuron with dense punctation, punctures separated by narrow ridges without conspicuous interspaces............................................................................... 25

25. Head with ventral coronal tooth conspicuously larger than remaining teeth, with distinct depression situated immediately below ocellar corona ( Smith 1988, fig. 114). Postocular carina parallel to hind margin of eye or space between eye and carina widest at mid-level of eye. Hind femur in female usually predominantly reddish brown ( Argentina, Brazil)........................................................................................... Ophrynopus depressatus Smith, 1988

- Head with ventral coronal tooth about equal in size to remaining teeth, without depression below ocellar corona ( Smith 1988, fig. 115). Postocular carina diverging from upper half of eye, space between eye and carina widest toward upper half of eye. Hind femur in female always black to dark brown.......................................................... 26

26. Punctures on mesoscutellum much closer together than puncture diameters, separated by sharp ridges. Tarsi usually dark brown ( Costa Rica, Ecuador, Guatemala, Mexico, Panama, USA: Texas)............... Ophrynopus nigricans (Cameron, 1883)

- Punctures of mesoscutellum sparser, mostly as widely separated as puncture diameters, and separated by shiny flat interspaces. Tarsi always reddish brown ( Brazil, Paraguay)................................. Ophrynopus plaumanni Smith, 1988