Thraulodes, Ulmer, 1920

Kluge, Nikita J., 2020, Systematic position of Thraulodes Ulmer 1920 (Ephemeroptera: Leptophlebiidae) and descriptions of new and little-known species, Zootaxa 4756 (1), pp. 1-142 : 5-21

Species composition of Thraulodes

All species of Thraulodes are distributed in new New World only, being limited by the southern part of North America, Central America (except Antilles) and South America (except Patagonia). Traver & Edmunds (1967) reported a larva from southern Chile, which they determined as belonging to Thraulodes .

The species named Thraulodes marhieus Dubey 1970 was described as female imago from Himalayas; judging by the words «The new species differs from the other Palaearctic species ...» the author confused the New World genus Thraulodes with something other; judging by confusion of vestigial paracercus with ovipositor, he was not familiar with insect morphology. Hubbard and Peters (1978: 25) wrote that «this species is in all probability misplaced at the generic level»; Kluge (2004: 208–209) placed it to Radulapalpata, i.e. Heptageniidae s. str.

The species vitripennis Blanchard in Gay 1851 [ Ephemera ], colombiae Walker 1853 [ Ephemera ], nervosa Eaton 1892 [ Choroterpes ], subfasciata Navás 1934 [ Thraulodes ], humeralis Navás 1935 [ Thraulodes ], and limbatus Navás 1936 [ Thraulodes ] have unclear systematic position within Leptophlebiidae and should be treated as Atalophlebolinguata INCERTAE SEDIS.

The species originally described as Ephemera vitripennis Blanchard in Gay 1851 from Chile was attributed to Thraulodes by Ulmer (1920). This species is know as poorly preserved male subimago, which cannot be reliably determined, and its placement to Thraulodes is probably wrong ( Traver & Edmunds 1967, Dominguez et al. 2006). Most probably, vitripennis [ Ephemera ] belongs to one of Chilean leptophlebiid genera, while Thraulodes is not ordinary for Chile (if occurs there at all).

The species originally described as Ephemera colombiae Walker 1853 was attributed to Thraulodes by Ulmer (1920) and is recently placed to this genus ( Traver & Edmunds 1967, Dominguez et al. 2006). This species is known as female subimago only and has no characters of Thraulode s: in contrast to all known Thraulodes , costal angulation of hind wing is located «about 2/3 of distance from base to apex»; in contrast to nearly all species of Thraulodes , femora are not banded.

The species originally described as Choroterpes nervosa Eaton 1892 is known as female only. Its MP of hind wing is not forked (Eaton 1982: fig. 6a), so it is unclear, why Ulmer (1920) placed it to Thraulodes . Traver and Edmunds (1967) assumed that «it comes closer to Traverella Edmunds », i.e. belongs to Hermanellognatha.

Two species originally named Thraulodes subfasciata Navás 1934 and Th. limbatus Navás 1936 were described as males, but without illustrations of genitalia. Traver & Edmunds (1967) assumed that they «are more likely to belong to the genus Ulmeritus Traver than to Thraulodes »; however, some authors place them to Thraulodes without argumentation (e.g., Dominguez et al. 2006). Genitalia of subfasciata [ Thraulodes ] were described as «forcipe longa, pallida, primo articulo interne basi et apice dilatato» ( Navás 1934: 165), that is not characteristic for Thraulodes .

The species originally named Thraulodes humeralis Navás 1935 also was described as male imago without illustration of genitalia. Recently it is placed to Thraulodes (e.g. Traver & Edmunds 1967). However, according to the original description, its hind wing has no triad of RS ( Navás 1935: fig. 119), in contrast to all known Thraulodes ( Fig. 41 ). Figures made by Navás are often inaccurate, but in his other figures (e.g. fig. 120, in the same paper) triad RA of the hind wing is adequately drawn.

The species originally named Thraulodes irretitus Navás 1924 is recently regarded to belong to Thraulodes , but with uncertain position within this genus, because its genital structure is unknown ( Traver & Edmunds 1967).

All known species of Thraulodes can be distinguished by characters of male imagines, which often have species-specific coloration of body, legs, wings and caudalii, as well as species-specific structure of genitalia. In contrast to this, larvae of Thraulodes have fewer species-specific characters. For most species of Thraulodes larvae remain to be undescribed; in those cases when larvae are described, the descriptions do not allow us to distinguish them from other species. In this situation formal species described as larvae only, appear to be non-comparable with others, and can be qualified as NOMINA DUBIA. These are Thraulodes salinus Kilgore & Allen 1973 , Th. eccentricus Lugo-Ortiz & McCafferty 1996 , Th. grandis Lugo-Ortiz & McCafferty 1996 , Th. tenulineus Lugo-Ortiz & McCafferty 1996 and Th. pacaya McCafferty, Baumbgardner & Guenter 2004 (see below).

The single complete key to male imagines of Thraulodes was published by Traver and Edmunds (1967). It begins with characters of coloration of abdomen, which are peculiar only for males, being different in females. Below, a new key is given, which includes also species described after 1967 and new species described in this paper. In or- der to make this key available for determination not only males, but also some females, it begins with the characters common for males and females; such characters are coloration of legs and wings. Species, whose leg coloration or genital structure is unknown, are not included in this key; these are Thraulodes mexicanus ( Eaton 1884 [Thaulus]) (legs coloration unknown), Th. bomplandi ( Esben-Petersen 1912 [ Thraulus ]) (fore and middle legs missing), Th. irretitus Navás 1924 (genitalia unknown), Th. pedregoso Traver 1946 (legs presumably associated), Th. hilaroides Traver 1946 (legs missing), Th. brunneus Koss 1966 (fore and hind legs missing), Th. lunatus Traver & Edmunds 1967 (fore and hind legs missing), Th. osiris Traver & Edmunds 1967 (legs missing).

This is a nested key, in which the number of antithesis is given in parentheses after the number of thesis, and the next thesis is given just after the correct thesis. Species described in this paper are numbered and marked by bold face.

Key to male imagines of Thraulodes

1 (8) All tibiae either entirely dark ( Figs 467–469 , 491–492 , 526–527 ), or dark in proximal half ( Fig. 437 ).

2 (7) All femora uniformly dark brown ( Figs 436–437 , 467–469 , 491–492 ).

3 (4) Fore wing with costal and subcostal fields contrastingly yellowish ( Figs 427, 435 )..................... 13. Th. niger sp. n.

4 (3) Fore wing with costal and subcostal fields not colored ( Figs 465 , 493 ).

5 (6) Fore wing with dark macula on costal brace ( Fig. 493 ). ♂: Abdominal terga II–VI light ocher, terga VII–X reddish ( Fig. 495 ).................................................................................... 15. Th. nigripes sp. n.

6 (5) Fore wing without dark macula on costal brace ( Fig. 465 ). ♂: Abdominal terga unicolor ( Fig. 466 )................................................................................................... 14. Th. nigrabdominalis sp. n.

7 (2) All femora twice banded, i.e. with small light area at middle of femur between bands ( Figs 526–527 )...................................................................................................... 6. Th. nigrotibialis sp. n.

8 (1) At least middle and hind tibiae either entirely light ( Fig. 29–30 ), or light with dark apex ( Figs 306–307 , 659–660 )

9 (14) Fore tibia entirely dark brown ( Fig. 597 ).

10 (11) At least in male, proximal half of fore wing brown, distal half colorless (female unknown)......................................................................................... Th. basimaculatus Giordano & Dominguez 2005

11 (10) No such difference between proximal and distal halves of fore wing.

12 (13) All femora uniformly dark brown ( Figs 597–598 )..................................... 18. Th. lepidus ( Eaton 1884)

13 (12) Middle and hind femora yellowish with black bands in distal half......................... Th. venezuelana Ulmer 1943

14 (9) Fore tibia at least partly light ( Figs 28 , 305 , 339 , 365 , 574 ) (as well as middle and hind ones).

15 (16) Fore tibia with brown band at middle, apex whitish; hind femur whitish with two narrow black bands close to apex ( Figs 574–575 ).......................................................................... 17. Th. alboniger sp. n.

16 (15) Fore tibia with dark band adjacent to apex ( Figs 28 , 305 , 339 , 365 ). Hind femur not as above.

17 (18) Femora of middle and hind legs yellowish, without dark maculae; only femur of fore leg with dark maculae apically............................................................................ Th. papilionis Traver & Edmunds 1967

1 (17) Femora of middle and hind legs with dark maculae at least in distal half ( Figs 29–30 , 97 , 306–307 , 367 ).

19 (22) Fore femur evenly blackish brown nearly all over its length (paler at extreme base and/or apex).

20 (21) ♂: Penis without ears, medio-ventral ridge forms lapel projected laterad of lateral margin ( Traver & Edmunds 1967: fig. 67)................................................................................... Th. hilaris Eaton 1892

21 (20) ♂: Penis with ears, no lapel projected laterad of lateral margin ( Traver & Edmunds 1967: fig. 51)............................................................................................... Th. arizonicus McDunnough 1942

22 (19) Fore femur with darker maculae at least in distal half ( Figs 28 , 305 , 339 , 365 , 398 ).

23 (66) Middle and/or hind femora, besides dark band and/or maculae in distal half, with dark band or spot in proximal half ( Figs 307 , 341 , 365–367 , 398–401 , 660 )

24 (57) Proximal half of middle and hind femora with band or wide spot ( Figs 307 , 341 , 365–367 , 398–401 , 660 ).

25 (26) Subcostal field of fore wing yellow, apically brown ( Figs 365–367 )................................. Th. flavus sp. n.

26 (25) Subcostal field of fore wing, if colored, with apex not darker than remainder part.

27 (28) Fore wing with dark macula on furcation of MA larger than other maculae. Th. alapictus Lima, Mariano & Pinheiro 2013

28 (27) If dark maculae on fore wing present, macula on furcation of MA not larger than others ( Fig. 398–401 ).

29 (30) Contrasting dark brown spot on whitish background on median furcasternal concavity of mesothorax ( Fig. 400 )........................................................................... 12. Th. spangleri Traver & Edmunds 1967

30 (29) No such unpaired spot on median furcasternal concavity.

31 (32, 33, 34) ♂: Penis with pair of apico-lateral projections directed proximally ( Kimmins 1934: fig. 8).............................................................................................. Th. valens ( Eaton 1892 [ Thraulus ])

32 (31, 33, 34) ♂: Penis with pair of lateral projections directed proximally ( Campos & Mariano 2019: figs 10–12)............................................................................... Thraulodes catoles Campos & Mariano 2019

33 (31, 32, 34) ♂: Penis with pair of lateral projections directed distally ( Traver 1946: fig. 30; Traver & Edmunds 1967: fig. 65)................................................................................... Th. furficulus Traver 1946

34 (31, 32, 33) ♂: Penis without additional lateral projections.

35 (40) ♂: Penis in distal half parallel-sided, not widened apically ( Fig. 310 or similar to Fig. 440 ).

36 (37) ♂: Penis without ears ( Traver & Edmunds 1967: fig. 37)...................... Th. ephippiatus Traver & Edmunds 1967

37 (36) ♂: Penis with ears directed caudally ( Fig. 310 ; Traver & Edmunds 1967: fig. 41).

38 (39) ♂: Abdominal terga II–IX equally dark ( Fig. 343 ).......................... 10. Th. zonalis Traver & Edmunds 1967

39 (38) ♂: Abdominal terga II–V contrastingly lighter than terga VII–X ( Figs 298–301 )................ 9. Th. fascipennis sp. n.

40 (35) ♂: Penis widened apically.

41 (42) ♂: Penis hidden under very wide dorsal projection of styliger (Neito & Dominguez 2001: figs 6–8)....................................................................................... Th. cryptodrilus Neito & Dominguez 2001

42 (41) ♂: Penis not hidden, dorsal projection of styliger smaller.

43 (50) ♂: Penis without lateral pouches.

44 (45) ♂: Dorsal margin of styliger rounded, without median projection ( Traver & Edmunds 1967: fig. 63)............................................................................................ Th. laetus ( Eaton 1884 [ Thraulus ])

45 (44) ♂: Dorsal margin of styliger either pointed, or with median projection ( Traver & Edmunds 1967: fig. 44; Dominguez 1987: figs 6, 8).

46 (47) ♂: Abdominal terga IV–VI with composite dark markings ( Dominguez 1987: figs 19–20). h. liminaris Dominguez 1987

47 (46) ♂: Abdominal terga IV–VI with sublateral dark spots ( Dominguez 1987: figs 15–16).

48 (49) Distributed in South America; genitalia as figured by Dominguez (1987: fig. 6)......... 21. Th. consortis Dominguez 1987

49 (48) Distributed in North America; genitalia as figured by Traver & Edmunds (1967: fig. 44)......... Th. speciosus Traver 1934

50 (43) ♂: Penis with lateral pouches.

51 (52, 53, 54, 55, 56) ♂: Abdominal coloration and genitalia as described by Traver & Edmunds (1967: figs 8–9, 31)................................................................................... Th. packeri Traver & Edmunds 1967

52 (51, 53, 54, 55, 56) ♂: Abdominal coloration and genitalia as described by Traver & Edmunds 1967: fig. 26)................................................................................................. Th. daidaleus Thew 1960

53 (51, 52, 54, 55, 56) ♂: Abdominal coloration and genitalia as described by Traver & Edmunds (1967: figs 4–5, 46)........................................................................................ Th. paysandensis Traver 1964

54 (51, 52, 53, 55, 56) ♂: Abdominal coloration and genitalia as described by Traver & Edmunds (1967: figs 2–3, 66)............................................................................................. Th. traverae Thew 1960

55 (51, 52, 53, 54, 56) ♂: Abdominal coloration and genitalia as described by Lima et al. (2013: figs 10–11, 18–20).......................................................................... Th. luizgonzagai Lima, Mariano & Pinheiro 2013

56 (51, 52, 53, 54, 55) ♂: Abdominal coloration and genitalia as described by Lima et al. (2013: figs 16–17, 31–32)................................................................................ Th. pinhoi Lima, Mariano & Pinheiro 2013

57 (24) Proximal half of middle and hind femora with small spot (besides band in distal half).

58 (61) ♂: Abdominal terga II–IX equally dark.

59 (60) ♂: Penis with lateral pouches wider than apex ( Mariano et al. 2011: figs 9–10)............................................................................. Th. amanda Mariano & Froehlich (in Mariano, Flowers & Froehlich) 2011

60 (59) ♂: Penis with lateral pouches narrower than apex ( Mariano et al. 2011: figs 4–3)........................................................................ Th. xavantinensis Mariano & Froehlich (in Mariano, Flowers & Froehlich) 2011

61 (58) ♂: Abdominal terga II–V contrastingly lighter than terga VII–X.

62 (63) ♂: Dorsal margin of styliger rounded, without median projection ( Lima et al. 2013: fig. 12)..................................................................................... Th. sternimaculatus Lima, Mariano & Pinheiro 2013

63 (62) ♂: Dorsal margin of styliger with median projection.

64 (65) ♂: Abdominal terga II–VI with a pair of dark submedian spots (besides pair of midway spots) ( Souto et al. 2014: figs 1–4)......................... Th. luisae Souto, Da-Silva & Nessimian 2014 and Th. pinga Souto, Da-Silva & Nessimian 2014

65 (64) ♂: Abdominal terga without submedian spots................................................................ Th. insular Domínguez, Molineri & Zúñiga (in Zúñiga, Molineri, Domínguez & Cardona) 2015 and Th. eduardorum Medina & Perez 2010

66 (23) Middle and hind femora light, with dark macula(e) only in distal part ( Figs 30 , 97 ).

67 (68) Hind femur without distinct bands, darker toward apex. ♂: penis unusually sharply widened, with lateral denticles ( Traver 1946: fig. 27; Traver & Edmunds 1967: fig. 49)....................................... Th. prolongatus Traver 1946

68 (67) Hind femur with brown or reddish apical band usually bordered by darker brown on proximal edge ( Figs 30 , 97 ). ♂: Penis not as above.

69 (70) ♂: All abdominal terga I–X dark brown. Dorsal margin of styliger rounded, without median projection; spear-like rolls very long and thick, directed caudally ( Traver & Edmunds 1967: fig. 42)................. Th. regulus Traver & Edmunds 1967

70 (69) ♂: At least abdominal terga II–V light.

71 (72) ♂: Abdominal terga II–VI with grayish brown dome-shaped patches ( Traver & Edmunds 1967: figs 10–11)................................................................................... Th. gonzalesi Traver & Edmunds 1967

72 (71) ♂: Abdominal coloration not as above.

73 (80) ♂: Abdominal terga II–V mostly light, VI–VII mostly colored ( Figs 198 , 232 , 264 ).

74 (75) ♂: Penis long and parallel-sided ( Dominguez 1987: fig. 10)........................ Th. cochunaensis Dominguez 1987

75 (74) ♂: Penis widened apically ( Figs 204 , 242 , 268–269 ).

76 (77) ♂: Spear-like rolls directed dorsally ( Fig. 267 )................................... 8. Th. quevedoensis Flowers 2009

77 (76) ♂: Spear-like rolls directed ventrally ( Figs 203 , 240 ).

78 (79) ♂: Apices of penis lobes as wide as gonostylar cavity of styliger ( Fig. 242 ). Distribution: from Panama to Venezuela............................................................... 7. Th. marreroi Chacon, Segnini & Dominguez 1999

79 (78) ♂: Apices of penis lobes wider than gonostylar cavity of styliger ( Fig. 204 ). Distribution: Peru and Brazil................................................................................. 6. Th. schlingeri Traver & Edmunds 1967

80 (73) ♂: Abdominal terga II–VI mostly light, VII mostly colored ( Figs 25 , 52 , 99 , 136 , 166 ).

81 (84) ♂: Spear-like rolls directed ventrally-proximally, with groove widely opened ( Figs 38 , 60 ).

82 (83) ♂: Spear-like rolls S-shaped; dorsal extension of styliger wide and not separated from styliger ( Fig. 60 )....................................................... 2. Th. sinuosus Mariano & Flowers (in Mariano, Flowers & Froehlich 2011)

83 (82) ♂: Spear-like rolls very short, lanceolate; dorsal extension of styliger finger-like and separated from styliger ( Figs 38, 40 )...................................................................................... 1. Th. ludmilae sp. n.

84 (81) ♂: Spear-like rolls directed medially (medially, medially-ventrally, or medially-dorsally) ( Figs 112–120 , 145 , 169 ).

85 (86) ♂: Penis lobes narrowed apically (Gonçalves et al. 2010: fig. 5)......... Th. jones Gonçalves, Da-Silva & Nessimian 2010

86 (85) ♂: Penis lobes not narrowed apically ( Figs 112–120 , 145 , 169 ).

87 (90) ♂: Abdominal sterna II–VII with 2 or 3 pairs of dark dots.

88 (89) ♂: Penis lobes long, equally narrow all over length ( Traver & Edmunds 1967: fig. 43)........................................................................................ Th. itatiajanus Traver & Edmunds 1967 [ Thraulodes ]

89 (88) ♂: Penis lobes short and greatly expanded laterally at apex ( Dominguez 1987: fig. 7)........... Th. flinti Dominguez 1987

90 (87) ♂: Abdominal sterna II–VII with no more than one pair of dots.

91 (94) ♂: Penis without lateral pouches ( Figs 112–120 ).

92 (93) ♂: Penis lobes short and sharply widened apically ( Boldrini et al. 2018: figs 6–7). Abdominal terga II–VI with reddish brown band adjacent to posterior margin (ibid: figs 2–3).......................... Th. rodrigoi Boldrini, Dantas & Lima 2018

93 (92) ♂: Penis lobes longer and slightly widened apically ( Figs 112–120 ).Abdominal terga II–VI without contrasting dark band ( Figs 97, 99 )........................................................ 3. Th. telegraphicus Needham & Murphy 1924

94 (91) ♂: Penis with lateral pouches ( Figs 144 , 171 ).

95 (101) ♂: Lateral pouches large, so that at midlength penis as wide or wider than at apex.

96 (97) ♂: Dorsal margin of styliger rounded, without median projection ( Chacon et al. 1999: fig. 5)........................................................................................ Th. mucuy Chacon, Segnini & Dominguez 1999

97 (96) ♂: Dorsal margin of styliger with median projection.

98 (99, 100) ♂: Abdominal coloration and genitalia as described by Traver & Edmunds (1967: fig. 33).... Th. ulmeri Edmunds 1950

99 (98, 100) ♂: Abdominal coloration and genitalia as described by Mariano et al. (2011: figs 14–15)................................................................ Th. pelicanus Mariano & Froehlich (in Mariano, Flowers & Froehlich) 2011

100 (98, 99) ♂:Abdominal coloration and genitalia as described by Gonçalves et al. (2013: figs 4–6)..................................................................................... Th. bonito Gonçalves, Da-Silva & Nessimian 2013

101 (95) ♂: Lateral pouches small, so that at midlength penis much narrower than at apex ( Figs 144 , 171 ).

102 (103) ♂: Abdominal terga VIII–X dark reddish brown, tergum VII lighter........................ Th. centralis Traver 1946

103 (102) ♂: Abdominal tergum VII not lighter than tergum VIII ( Figs 136 , 166 ).

104 (105, 110) Fore femur, besides dark band in distal half, with dark band in proximal half................................................................................... Th. trijuncta Banks 1918 and Th. bolivianus Dominguez 1986

105 (104, 110) Fore femur, besides dark band in distal half, with small dark spot in proximal half ( Figs 134 , 160 ).

106 (107) ♂: Pronotum grayish-brown with white spots...................... Th. guanare Chacon, Segnini & Dominguez 1999

107 (106) ♂: Pronotum white ( Figs 134, 136 , 157 ).

108 (109) Macula on costal brace not continuous posteriorly ( Figs 134–136 ). Scutellum white ( Fig. 136 )... 4. Th. panamensis sp. n.

109 (108) Macula on costal brace continuous posteriorly ( Figs 157–159 ). Scutellum brown ( Fig. 159 )....... 5. Th. viviparus sp. n.

110 (104, 105) Fore femur without dark marking in proximal half......................... Th. calori Campos & Mariano 2019

Key to examined larvae of Thraulodes

1 (8) Labrum with sharp median emargination (in anterior-dorsal view), anterior transverse row of setae in median part irregular, with many setae in section ( Figs 190–197 , 209 , 389–395 , 650–653 )

2 (3) Cuticular coloration: tibiae banded ( Figs 642–644 )........................................ 20. Thraulodes sp. «Itaya»

3 (2) Cuticular coloration: tibiae unicolor.

4 (7) Cuticular coloration: pronotum with blanks of characteristic composite form ( Figs 175, 185 , 214, 218, 227 ); abdominal terga IV–V, or at least tergum V contrastingly light medially ( Figs 177, 185 , 215, 219, 227 ).

5 (6) Tergalii with tracheal branches by sides of main trachea ( Figs 178–184 )................................ 6. Th. schlingeri

6 (5) Tergalii without tracheal branches by sides of main trachea ( Figs 220–226 )....................... 7. Thraulodes marreroi

7 (4) Cuticular coloration: pronotum without contrasting blanks ( Fig. 382 ); abdominal terga without large blanks mediad of tergalii bases ( Fig. 383 )......................................................................... 12. Th. spangleri

8 (1) Labrum without median emargination, anterior transverse row of setae regular, with one seta in section ( Figs 86–88 , 563 ).

9 (10) Cuticular coloration: tibiae contrastingly banded ( Figs 548–550 ). Tergalius with main trachea arched anteriorly (i.e. repeating outline of anal margin), with branches directed toward costal margin ( Figs. 551–557 , 585–588 )..... 17. Th. alboniger sp. n.

10 (9) Cuticular coloration: tibiae unicolor ( Fig. 9 ). Tergalius with main trachea either straight, or arched posteriorly, with branches either directed toward both margins, or absent.

11 (14) Cuticular coloration: pronotum entirely light, anterior part of mesonotum contrastingly dark ( Figs 8 , 66 , 70 ).

12 (13) Tergalii narrow, without tracheal branches by sides of main trachea ( Fig. 11 ). On hind tibia stout setae of inner-anterior row variable, long and short ( Fig. 24 )......................................................... 1. Th. ludmilae sp. n.

13 (12) Tergalii wide, with tracheal branches by sides of main trachea ( Fig. 90–93 ). On hind tibia all stout setae of inner-anterior row short ( Fig. 85 ).................................................................... 3. Th. telegraphicus (part)

14 (11) Cuticular coloration: pronotum not contrastingly lighter than mesonotum.

15 (16, 17, 18, 19) Cuticular coloration: abdominal terga mostly light, with pair of large brown maculae on terga VI–VII and previous ones ( Figs 595 , 603 )........................................................................ 18. Th. lepidus

16 (15, 17, 18, 19) Cuticular coloration: on each abdominal tergum area adjacent to anterior margin together with antero-median and sublateral areas brown, remainder part contrastingly light ( Figs 414, 419, 421 )...................... 13. Th. niger sp. n.

17 (15, 16, 18, 19) Cuticular coloration: abdominal terga I–VII contrastingly lighter than terga VIII–IX ( Figs 453, 458, 459 )........................................................................ presumably 14. Th. nigrabdominalis sp. n.

18 (15, 16, 17, 19) Cuticular coloration: abdominal terga IV–V contrastingly lighter than terga VI–VII ( Figs 66 , 81 )............................................................................................. 3. Th. telegraphicus (part)

19 (15, 16, 17, 18) Cuticular coloration: all abdominal terga either uniformly dark with light lateral margins ( Fig. 297 ), or some terga also with blanks medially ( Figs 154–155 , 350 ).

20 (21) Proximal part of fore tibia with inner-anterior row of recurved hairs ( Figs 621–622 )....... 19. Thraulodes sp. «Palo Seco»

21 (20) Fore tibia without inner-anterior row of recurved hairs.

22 (23) Tergalii darkened near costal margin ( Fig. 153 )................................... presumably 5. Th. viviparus sp. n.

23 (22) Tergalii either evenly colored ( Fig. 129 ), or darkened near main trachea ( Figs 291 , 361 , 385 ).

24 (35) Tergalii without visible side tracheae ( Figs 129 , 361 ) or with one proximal-costal trachea on ventral lamella ( Fig. 291 ).

25 (28) On hind tibia stout setae of inner-anterior row variable, long and short ( Figs 51 , 259 ).

26 (27) On fore and middle legs claw with small denticles anteriad of main row of denticles ( Fig. 273 )......... 8. Th. quevedoensis

27 (26) On all legs claw without small denticles anteriad of main row of denticles (as in Fig. 69 )................. 2. Th. sinuosus

28 (25) On hind tibia all stout setae of inner-anterior row short ( Figs 132 , 296 , 336 , 358 ).

29 (30) On femora and hind tibia short stout setae narrowed distally ( Figs 358, 359 )....................... 11. Th. flavus sp. n.

30 (29) On femora and hind tibia short stout setae parallel-sided or widened distally ( Figs 130, 132 , 293, 296 , 303 , 336 ).

31 (32) Tergalii with both lamellae narrow, without side tracheae ( Fig. 129 ).......................... 4. Th. panamensis sp. n.

32 (31) Tergalii with ventral lamella widened proximally ( Figs 284–290 , 321–327 ), sometimes with one side trachea ( Fig. 291 ).

33 (34) Last instar male larva: protogonostyli very short, separated by shallow concavity ( Fig. 313 )....... 9. Th. fascipennis sp. n.

34 (33) Last instar male larva: protogonostyli long, separated by deep emargination ( Fig. 346 )................... 10. Th. zonalis

35 (24) Tergalii with well-visible side tracheae ( Figs 66–68 , 90–93 ).

36 (37) Tergalii very wide; dorsal lamella with costal and anal margins equally convex in proximal and distal parts, abruptly narrowed toward slender apical filament ( Figs 666–672 )................................................ 21. Th.? consortis

37 (36) Tergalii less wide; dorsal lamella widest in proximal or middle part, gradually narrowed toward slender apical filament ( Figs 66–68 , 90–93 , 503–506 , 537–540 ).

38 (39) In last instar larva hypoderm of tibiae more or less darkened with brown or blackish.......................................................................................... 1 5. Th. nigripes sp. n. and 16. Th. nigrotibialis sp. n.

39 (38) Tibia never darker than femur (in mature larva only apex of tibia with dark hypodermal coloration).................................................................................................... 3. Th. telegraphicus (part)

Morphological notes

Traver and Edmunds (1967) gave a detailed review of imaginal, larval and egg structure of Thraulodes , noted significance of certain morphological features for diagnoses of species and discussed individual variability. The new data confirm most statements given in this publication, but some additions and clarifications should be made.

Larval habitus

In all species of Thraulodes larvae have a similar body shape, which is widened and flattened dorsoventrally, an adaptation for inhabiting stone surfaces in rapidly running water. The head is wide, with the clypeus parallel-sided or slightly widened anteriorly, with labrum wider than clypeus. The pronotum is nearly rectangular, with rounded antero-lateral angles and a free anterior margin (i.e. transverse margin between antero-lateral angles and the neck membrane); a transverse ridge runs parallel to the free anterior margin; humeral setae (which are characteristic for Leptophlebiidae ) are stout and located between the free anterior margin and the ridge, being absent in other places; these setae are narrowed distally, either pointed or blunt, either forming a nearly regular row ( Fig. 12 ), or situated rather irregularly (the same in many other Atalophlebolinguata). Legs are large, with widened femora and claws bent perpendicular to the leg flatness.

Structure of labrum

Peters (1980) divided Leptophlebiidae into two subfamilies, Leptophlebiinae and Atalophlebiinae , each of which he regarded to be holophyletic.Among autapomorphies of Atalophlebiinae , he reported the following character: «Anteromedian emargination of labrum with denticles or with acute median cleft ...», contrary to the plesiomorphy of Leptophlebiinae «Anteromedian emargination of labrum smooth, without denticles ...» ( Peters 1980: Table 1 ). According to my opinion ( Kluge 2009), his subfamily Leptophlebiinae is paraphyletic, and his subfamily Atalophlebiinae is really holophyletic and corresponds to the rank-free taxon Atalophleboculata Kluge 2009. However, the character of Atalophlebiinae cited above, seems to be not a good autapomorphy of this taxon. Denticles reported here, are convexities separated by sensilla; these sensilla are found in various taxa of Ephemeroptera , usually are 6 in number and form a regular transverse row on the distal margin of labrum ( Figs 1–3 ); if surface between each two sensilla forms a convexity, labrum bears a row of 5 convexities, or denticles ( Fig. 4 ). Because of this, 5 denticles in the median emargination of labrum are found not only among representatives of Atalophleboculata, but also among representatives of Leptophlebiinae sensu Peters 1980 .

Initially for Atalophlebopectinata (the taxon comprising Atalophleboculata and Habrophlebia /fg1), labrum bears two transverse rows of setae—anterior and posterior ones.

The larval head of Thraulodes is prognathous, and labrum is normally directed anteriorly. In the normal condition, the initial anterior margin of labrum (which bears denticles) is turned ventrally, so that in dorsal view the anterior transverse setal row (or even the posterior setal row) appears located on anterior margin, and the initial anterior margin with denticles is hidden beneath the labrum ( Fig. 86 ). In this case shape and denticulation of the initial anterior margin can be visible in frontal view. In another condition of the same labrum, when it is swollen and straightened, its denticles can be visible in dorsal view ( Figs 87, 88 ).

In all examined species of Thraulodes , the number of denticles is normally 5, but can be less in selected individuals. The posterior transverse setal row is always regular, nearly straight, located on the anterior part of labrum and occupies most of its width ( Fig. 88 ). The anterior transverse setal row is often much shorter, either also regular ( Fig. 88 ), or irregular.

In Th. schlingeri , Th. marreroi , Th. spangleri and Th. sp. «Itaya» the labrum has narrow and deep median emargination ( Fig. 194 ), and 5 denticles are located in this emargination often in such a way that 3 of them are relatively narrow and form the bottom of the emargination, and 2 lateral denticles are wider and form lateral sides of the emargination ( Figs 197 ). Since the emargination with its denticles is directed ventrally, in dorsal view the outline of the labrum appears as a shallow median concavity without denticles ( Fig. 191 , 208 ). In all these four species anterior transverse row of setae in its median part (i.e. just behind the median emargination) represents a field of irregularly situated setae ( Figs 193, 195 ). The same median emargination is recently reported for Thraulodes sternimaculatus Lima et al. 2013 ( Nascimento et al. 2019)

In all other examined species of Thraulodes the labrum has no median emargination, all its 5 denticles are very wide and low, and anterior transverse row of setae is regular in all its parts ( Fig. 88 ); the initial anterior margin, which bears denticles, is either straight, or slightly convex ( Fig. 88 ), or slightly concave ( Fig. 563 ). Possibly, width of the anterior setal row is species-specific: in the examined individuals of Th. ludmilae sp. n., Th. sinuosus , Th. panamensis sp. n. and Th. quevedoensis this row is about as wide as 3 denticles (among 5 existent ones), while in examined individuals of Th. fascipennis sp. n., Th. zonalis , Th. flavus sp. n., Th. niger sp. n., Th. nigrabdominalis sp. n., Th. nigripes sp. n., Th. nigrotibialis sp. n., Th. lepidus, Th. sp. «Palo Seco» and Th. consortis (?) this row is wider, being about as wide as all 5 denticles, or wider. Among examined specimens of Th. telegraphicus , width of the anterior setal row varies from width of 3 denticles ( Fig. 88 ) to width of 5 denticles. In Th. alboniger s. n., the anterior transverse setal row is wider than in all other examined species, being approximately as wide as the posterior transverse setal row ( Fig. 563 ).

Structure of mandibles

Mandibles have the outer margin either entirely convex ( Figs 17–18 ), or with a shallow concavity ( Figs 250– 251 ). In Thraulodes sp. «Palo Seco» the outer margin of each mandible is widened and angulate ( Figs 614–615 ); the same in Thraulodes sp. from Brazil ( Traver & Edmunds 1967: fig. 76).

In all Thraulodes , outer margin of mandible bears long hairs, which are denser and longer nearer the base of incisor, i.e. form a distal tuft ( Figs 17–18 ). In contrast to this, in many other Leptophlebiidae the mandibles either have no long setae on outer margin, or have most long and dense setae at the middle of outer margin. Traver and Edmunds (1967) stated, that «This series of hairs ... is so distinctive of this genus that we believe it to be of major importance in distinguishing Thraulodes nymphs from those of all other genera ...»; as genera whose mandibular setation has some similarity with that of Thraulodes , they reported only Ulmeritus Traver 1956 with allied genera, Massartellopsis Demoulin 1955a and Meridialaris Peters & Edmunds 1972 (termed «Chilean Deleatidium »). Among species examined by me, many Leptophlebiidae have more or less long and dense setae on middle part of outer side of mandible, while the area near base of incisor completely lacks setae; in many cases setae form a distinct tuft at the middle of outer side of mandible, which can be called middle tuft. In fewer cases, mandible also has setae near the base of the incisor, which either form a weak row stretched from the middle tuft toward the base of incisor (e.g., in Massartellopsis and Meridialaris ), or form a second, distal tuft, e.g. in Ulmeritus , Atopophlebia Flowers 1980 , Nathanella Demoulin 1955b , Nonnullidens Grant & Peters 1993 ( Kluge 2013: fig. 3), Kimminsula Peters & Edmunds 1970 , undescribed taxa related to Kimminsula and selected species of Thraulus Eaton 1881 . Besides Thraulodes , in a few other taxa the middle tuft is poorly expressed or absent, while the distal tuft is present. These are Ulmeritoides Traver 1959 , Neochoroterpes Allen 1974 and Petersula courtallensis Sivaramakrishnan 1984 (while an undescribed species of Petersula has both tufts). Thus, presence of the distal tuft combined with absence of the middle tuft is a rare, but not a unique character of Thraulodes ; most probably, it is an autapomorphy of Thraulodes .

Incisors, kinetodontia, prosthecae and molae of mandibles have uniform structure in all examined species of Thraulodes , being similar to that of many other Atalophlebomaxillata; both incisors and left kinetodontium are always terminated by 3 denticles each, right kinetodontium is always terminated by 2 denticles ( Figs 17–18 ).

Structure of maxilla

In all species of Thraulodes the maxillae have a rather uniform structure, which is initial for the taxon Atalophlebomaxillata Kluge 2009 ( Kluge 2012) ( Fig. 19 ). According to Traver and Edmunds (1967), «The slightly undulating row of large pectinate spines below the level of the crown on the ventral surface varies in number of spines from about 14 to 26 or more, a feature that may be of specific value». Among the specimens examined, number of stout pectinate setae (erroneously termed «spines») in this apical-ventral row varies from 10 to 24; in most species it varies individually as 13–18; in two examined specimens of Th. quevedoensis it is 11; in three examined specimens of Th. alboniger sp. n. it is 10; in Th. niger sp. n. and Th. nigrabdominalis sp. n. it varies as 19–21; in Th. lepidus varies as 23–24.

Traver and Edmunds (1967) paid attention to structure of the last (3rd) segment of maxillary palp, which has «a narrow clear area at base of the third segment on the outer side». Before this, some authors regarded this area to be an outgrowth of the 2nd segment: Traver (1935: 552) wrote that «The outer apical margin of the second joint prolonged into a slender process which lies appressed against the side of the short third joint»; Burks (1953: 93) wrote that «maxillary palp forceps-like at the apex». Actually, cuticle of the 3rd palpomere is pigmented on inner side and colorless on outer side of the palpomere; on dorsal side of the palpomere boundary between the pigmented and the colorless areas is sharp ( Figs 19 , 363 ), that makes illusion of a longitudinal line separating two lobes; on ventral side of the palpomere its pigmentation changes gradually ( Fig 362 ). This coloration is present in most of examined species, except for Th. alboniger sp. n., whose colorless area of the 3rd maxillary palpomere is greatly diminished.

Structure of labium

The glossae lie in one plane with the paraglossae; long setae of their ventral side are situated densely and irregularly ( Fig. 19 ); long setae on the dorsal side of glossa form irregular row or stripe of a characteristic horseshoe shape ( Fig. 20 ); the same was observed in Atopophlebia sp. (in contrast to other taxa, whose dorsal setae of glossa are situated irregularly or form a transverse row). According to Traver and Edmunds (1967), «Glossae of labium ... straight on inner margins, well rounded on outer; rather long spines are present along apical margin; a curved row of short spines occurs on dorsal surface and close-set hairs on ventral surface». Actually, in all examined species setae (erroneously termed «spines») on the apical margin are either absent, or short and stout as on inner and outer margins ( Fig. 20 ); the illusion of long apical setae is sometimes caused by the ventral setae (close-set hairs), which can be projected apically. Setae forming an irregular horseshoe row on dorsal side are as long and thin as setae (hairs) on the ventral side; the illusion of «short spines» appears if these setae are directed perpendicular to the surface.

In all species examined, apical (3rd) segment of labial palp is short and conic, much narrower at base than apex of 2nd segment ( Traver & Edmunds 1967: fig. 82).

Setation of larval legs

In larvae of all Leptophlebiidae , fore, middle and hind legs have significant differences in shape and setation ( Table 1 ). Most often and probably initially for Leptophlebiidae are the following differences: the fore femur is widened near the base (where the tibial adductor is attached), with outer margin convex basally ( Fig. 9 ); fore tibia (which always lacks patella-tibial suture) has dense stout pointed setae on inner margin ( Fig. 21 ), while other tibiae have sparser and shorter pointed setae on inner margin ( Figs 22, 24 ); hind tibia has stout setae on outer margin ( Fig. 24 ), while other tibiae have no stout setae on outer margin ( Figs 21, 22 ). Besides stout setae, outer margins of femur, tibia and tarsus of all legs usually bear more or less dense long, thin, projected hairs. Thraulodes has the following modification of these features.

Hairs of outer margin. Long hairs form dense irregular rows or stripes not only on outer margins of femur, tibia and tarsus, but also on corresponding (hind) margin of coxa ( Fig. 77 ).

Setation of femur. Outer margin of each femur bears irregular outer stripe of thin hairs, bordered by two irregular stripes of stout setae— outer-posterior and outer-anterior ones ( Fig. 83 ). On all legs of all examined species of Thraulodes , stout setae of these stripes irregularly vary from short to very long, with all intermediate forms; their structure and position did not reveal species-specific characters. Thin hairs of the outer stripe are denser and longer on fore and middle legs, but sparser and/or shorter on hind leg. In most species this difference between femora is well expressed ( Fig. 9 ); in some species hairs of all femora are dense and long, but in these cases hairs of hind femur are at least somewhat shorter, than hairs on fore and middle femora ( Fig. 126 ). Besides the hairs on outer margin, femur can bear a row of recurved hairs on anterior surface near inner margin ( Fig. 364 ); this row of recurved hairs is often present on middle and hind legs, being absent on the fore leg; in Th. flavus sp. n., Th. sp. «Palo Seco» and Th. consortis (?) the row of recurved hairs is well developed on all legs; in Th. quevedoensis and Th. alboniger the row of recurved hairs is absent on all legs. Inner margin of femur always bears an irregular inner row of relatively short, stout pointed setae ( Fig. 364 ). Anterior surface of each femur bears numerous irregularly situated stout setae, which are arranged either irregularly, or form more or less distinct longitudinal stripes. The shape of these setae is species-specific: they can be either widened distally with rounded apices ( Figs 130 , 293 ,), or parallel-sided ( Fig. 13 ), or narrowed distally with apices either truncate ( Fig. 359 ) or pointed ( Fig. 560 ). Shape of these stout setae is the same on femora of all leg pairs. Posterior surface of hind femur bears a few small, pointed, curved, pectinate setae, which either form an irregular row parallel to inner margin of femur, or occupy up to half of posterior surface; femora of fore and middle legs usually have no setae on posterior surface.

Setation of tibia. Tibiae of fore, middle and hind legs have significantly different structures and bear the following rows and fields of stout setae and thin hairs. Outer-anterior row of stout setae ( Figs 85 , 89 : o-a.r): absent on fore and middle legs, well-developed on hind leg; consists of species-specific setae, often contains long spoonlike setae ( Fig. 520 ). Outer-posterior row of stout setae ( Figs 85 , 89 : o-p.r): absent on fore and middle legs, welldeveloped on hind leg; consists of species-specific setae, can contain long spoon-like setae ( Fig. 426 ). Inner-anterior row of stout setae ( Figs 85 , 89 : i-a.r): on fore leg either absent (in Th. panamensis — Fig. 131 ), or represented by few setae on proximal part of tibia ( Fig. 21 ); on middle leg present all along tibia, but sparse and often consists of short setae ( Fig. 22 ); on hind leg well-developed ( Fig. 24 ); consists of setae of species-specific shape, never contains spoon-like setae. Inner field of stout pointed setae ( Fig. 89 : i.f): stretched all along inner side of tibia, always consists of pointed setae, which can be either smooth, or pectinate, or heavily dentate; on middle and hind legs these setae are always very small and sparse ( Figs 22, 24 ), on fore leg larger and dense, at least near apex of tibia ( Fig. 21 ). Near the apex of fore tibia, the most posterior row of this field (i.e., inner-posterior row) always consists of 3–4 closely situated arched pointed setae, among which the most proximal seta is significantly larger than others ( Fig. 559 ). Setae of the inner field of fore tibia are either situated irregularly, or form more or less regular longitudinal rows; if setae of these rows differ in structure, setae of the most anterior row are pectinate or dentate, while others are smooth or less heavily pectinate. In Th. spangleri setae of this field are arranged in 2 regular rows with shortened curved pectinate setae in anterior row and long thin straight setae in posterior row ( Fig. 386 ). In Th. flavus sp. n. setae of this field are diminished ( Fig. 630 ); in Th. alboniger sp. n., Th. lepidus and Th. sp. «Palo Seco», this field is vestigial, i.e. its setae are as small and sparse, as on middle and hind legs ( Figs 558 , 599 , 621 ); in all cases dense setae are retained near apex of fore tibia, and the subapical inner-posterior row of arched pointed setae is retained. Posterior pointed pectinate setae ( Fig. 89 : p.r): present on hind tibia only; situated either irregularly, or forming a longitudinal row; on apex of hind tibia such setae are more dense and form a transverse row ( Fig. 23 ). Outer field of hairs ( Fig. 89 : o.h): these hairs (i.e. long and slender setae) are directed nearly perpendicular to tibia surface; present on tibiae of all legs; on hind tibia these hairs are irregularly situated between the outer-anterior and the outerposterior rows of stout setae and form a row posteriad of the outer-posterior row of stout setae. Inner-anterior row of recurved hairs ( Fig. 89 : i-a.h): located on anterior side of tibia, more distally from inner margin than the inneranterior row of stout setae; on fore leg either absent ( Fig. 21 ), or present on proximal part of tibia (in Th. sp. «Palo Seco»— Fig. 622 ); on middle and hind legs always present all along tibia ( Figs 22, 24 ).

Larval claws

In connection with adaptation for inhabiting stone surfaces in rapidly running water, claws are bent perpendicular to the leg flatness, so that when legs are spread, and body and legs are pressed to substrate, claws are directed toward the substrate and have good adhesion with it (a similar form of body and legs evolved independently in various other rheophilous mayfly taxa). In accordance with this, in all species of Thraulodes the claw is distinctly divided into the rigid portion (main part of the claw), and the articulatory portion, located between the rigid portion and the unguitractor ( Fig. 69 ). On its morphologically posterior side (functionally ventral side) the articulatory portion is membranous, and serves rotation of the claw perpendicular to the leg flatness. In various mayfly larvae, which have such or similar rotation mechanism of claw and a row of denticles on inner side of the claw, these denticles are either present on the rigid portion only, or are equally developed on both portions forming an integral row, or have significantly different shapes and sizes on these two portions. In all species of Thraulodes , one longitudinal row of stout denticles is present on inner side of the rigid portion of claw; the articulatory portion of claw often (but not always) bears a longitudinal row of minute, pointed denticles, which are significantly smaller than denticles of the rigid portion ( Figs 69 , 411 ). Traver and Edmunds (1967: 357) reported the denticles of rigid portion as «variable number of large tooth» and the denticles of articulatory portion as «several smaller ones nearer the base». Denticles of articulatory portion are often very small, and the most proximal of them are often fused together, so that their number cannot be adequately counted.

In majority of Thraulodes species, the number of denticles of the rigid portion varies between 5 and 10; usually (but not always) the proximal denticles are smaller and the distal ones are larger; the most distal denticle may or may not be larger than all others; it is located either on the same line as others, or offset from this line; all these characters are not species-specific, but vary individually and on different legs of the same individual.

Some species have significantly diminished number of denticles of the rigid portion: 3–5 in Th. niger sp. n. ( Fig. 411 ), 2–4 in Th. nigrabdominalis sp. n. and constantly 4 in Th. alboniger sp. n.

Allen and Brusca (1978) reported the character «median denticle largest» as a species-specific character for the Mexican species Thraulodes sp. G and figured a claw which has several small denticles between the largest denticle and the claw apex ( Allen & Brusca 1978: fig. 16). Among species examined, the same is found on fore and middle legs of Th. quevedoensis ( Fig. 273 ), while hind legs of the same individuals have claws of a usual structure, without small denticles between the largest denticle and the claw apex.

Larval coloration

The authors, who described larval coloration, did not differentiate between cuticular and hypodermal color patterns. Such descriptions of coloration have no taxonomic significance, because during individual development cuticular coloration becomes more intensive from beginning of each instar to the end of instar, while hypodermal coloration passes from instar to instar and is gradually changed from larva to imago.

Cuticular coloration of larval thorax. In those cases when the cuticle of pronotum is intensively pigmented, its lateral margin is more or less contrastingly colorless ( Figs 75 ). The cuticle of the rest surface of the pronotum and mesonotum can have darker and lighter areas, some of which are connected with muscle attachment; in various species these areas are either contrasting or not.

Among features of the cuticular coloration, Traver and Edmunds (1967) paid special attention to «the U-shaped mark on the mesonotal scutum, 1 arm of the U on each side of the mid-dorsal line, the 2 arms uniting near the posterior portion of the scutum» and noted that «in Thraulodes this U-figure ... is usually more prominent than in other related genera, but is not distinct in all specimens». This is cuticular coloration; the pair of light stripes ( Figs 66–67 ) separate the area of attachment of the median longitudinal muscles from the areas of attachment of other muscles and correspond to the pair of medioparapsidal sutures of the winged stages.

In various species, cuticle of fore protoptera is either nearly uniformly pigmented, or has veins darker than background ( Fig. 175 ), or has veins lighter than background ( Fig. 8 ); in contrast to hypodermal coloration, this cuticular coloration does not depend upon stage of wing development.

Hypodermal coloration of larval thorax. The larval pronotum either has no hypodermal markings ( Figs 66, 68 ), or has blackish stripes bordering colorless lateral margins (see above) ( Figs 332 , 518 , 603 ), or has peculiar blackish markings corresponding to the imaginal ones (compare Figs 543 and 576 ). The mesonotum and thoracic pleura sometimes have irregular blackish markings. Fore and hind protoptera often have blackish hypodermal markings corresponding to certain maculae of the imaginal wing, particularly to the dark brown macula near costal brace ( Figs 620 , 626 ) or others ( Figs 591 , 655 ).

Cuticular coloration of larval legs. In all Leptophlebiidae the fore femur is more or less widened proximally; in some leptophlebiid taxa, its cuticle has a discolored area (blank) on anterior surfaces of the proximal widening; this blank is expressed if surrounding cuticle is pigmented. Presence and shape of this blank is not connected with muscle attachment: all muscle fibers which pass under anterior surface at this area belong to the femoro-tibial flexor and are attached to the proximal, outer and inners margins of the femur, but not to its dorsal surface. In all Thraulodes this proximal forefemoral blank has characteristic shape and lacks stout setae, which are irregularly spread on the anterior surface of the femur (see above). Shape of the proximal forefemoral blank somewhat resembles a triangle, which is stretched obliquely in such a way that its short side is adjacent to the convex proximal-outer margin, and its acute angle touches the inner margin more distally ( Fig. 281 ). Other elements of cuticular coloration of fore, middle and hind femora vary among species often have significant individual variability. The cuticle of tibiae in most species is uniformly pigmented; only in Th. alboniger sp. n. and Th. sp. “Itaya” it has darker and lighter bands ( Figs 548–550 , 642–644 ).

Hypodermal coloration of larval legs. The imaginal femur often has a hypodermal multicolored distal band formed by various combinations of yellowish, reddish and blackish markings, including a blackish stripe along distal part of inner margin (e.g. Figs 28–30 , 97 ); in some species femur has additional middle band, or is entirely darkened (e.g., Figs 468–469 , 597–598 ); in these cases hypodermal coloration of larval femur is limited by blackish pigment, which often (but not always) forms a stripe along distal part of inner margin of the femur ( Figs 452 , 591 ). In Th. alboniger sp. n. imaginal hypodermal coloration of femur is represented by a narrow apical blackish band ( Fig. 574 ), and larval hypodermal coloration of femur is similar to it ( Fig. 543 ).

Hypodermal coloration of larval abdomen. Abdominal terga of imagines have species-specific color patterns. Traver and Edmunds (1967) recognized 9 elements of this coloration. Among them, a blackish or gray posterior margin and a pair of blackish or gray midway spots are present in many species. If these markings are present in imago, they are more or less expressed in larval stage ( Figs 68 , 128 , 155 , 185 , 227 , 292 , 332 , 421 ). Blackish or gray stigmatic dots, if present in imago, are also present in larva; but in contrast to the dark posterior margin and the midway spots, which are located just under cuticle and are well visible, stigmatic dots are situated on tracheae, which in larval stage are located deeply in the body and are poorly visible or invisible. Imagines of some species have paired oblique lateral streaks or darkened posterolateral triangles ( Fig. 611 ); in these cases larvae can have similar blackish or gray markings ( Fig. 603 ). Among species examined, only some individuals of Th. alboniger sp. n. have black median marks ( Fig. 572 ), and these marks are expressed in the larval stage as well ( Fig. 543 ). Some other elements of imaginal coloration, e.g. brown or reddish dorsolateral spots are absent in larval stage and appear only when larva is going to molt to subimago.

Based solely on larval characters, McCafferty, Baumgardner and Guenther (2004) established a formal species name Thraulodes pacaya for the larvae which were described by Allen and Brusca (1978) under arbitrary name Thraulodes sp. C. They report only two characters for this species, «relative narrow elongate gills with indistinct lateral tracheation» and «narrow, dark, transverse band on the posterior border of middle abdominal terga and the pair of antero-sublateral, approximately right-triangle shaped dark marks on middle abdominal terga» McCafferty et al. 2004: 214). Judging by the drawing to which they refer ( Allen & Brusca 1978: fig. 24), these dark markings represent the blackish-brown hypodermal pigmentation, which corresponds to the dark posterior margin and the midway spots on imaginal abdomen ( Traver & Edmunds 1967: fig. 29: dpm, ms). These elements of coloration are present in imagines of many species of Thraulodes and should be expressed in their larvae; most probably, « Th. pacaya » represents a mixture of larvae belonging to various species.

New terms for describing coloration of subimaginal mesonotum

The mesonotum of the mayfly subimago and imago bears the mesonotal suture, which initially represents a transverse furrow crossing the scutum behind the anteronotal transverse impression; in the stage of imago, cuticle anteriad and posteriad of the mesonotal suture usually has the same coloration (brown or ocher), but in the stage of subimago cuticle is usually colored anteriad of the mesonotal suture and colorless posteriad of it. Initially, I called this suture «transverse suture» ( Kluge 1988: figs 90–91: TS), but later suggested the term «mesonotal suture», because in some cases its direction is changed, being more longitudinal than transverse ( Kluge 1994a, 2004). In the cases, when the mesonotal suture is shifted backward, the colored zone anteriad of the mesonotal suture occupies a part of medioscutum and a part of submedioscutum ( Kluge 1994a: fig. 15; 2004: fig. 106D). In some cases, coloration of these zones represents a species-specific character, so these zones should be clearly named in species descriptions. In one of the previous papers, I used the letter «a» for two zones located anteriad of the mesonotal suture and the letter «p» for two zones located posteriad of the mesonotal suture ( Kluge 2013: figs 51–53). However, these two abbreviations are not enough to describe all variants of coloration of subimaginal mesonotum.

Here I suggest the following new terms ( Fig. 94a ): the term « chromozone » for the zone anteriad of the mesonotal suture; the term « achromozone » for the zone posteriad of the mesonotal suture; and the term « chromozonal suture » as additional term for the mesonotal suture to indicate that it is a line separating the chromozone and the achromozone of subimaginal cuticle. The terms «chromozone» and «achromozone» should be used for certain zones, independently if they are actually colored or not. In Leptophlebiidae and some other taxa, the chromozonal suture is shifted backward and crosses the medioparapsidal suture, which separates medioscutum (zone of attachment of the median tergal muscle) and submedioscutum (zone of attachment of the scuto-episternal muscle) (the terms «medioscutum» and «submedioscutum» were introduced in Kluge 1994); in this case, 4 zones can be distinguished, which can be termed «chromozone of medioscutum», «chromozone of submedioscutum», «achromozone of medioscutum» and «achromozone of submedioscutum»; two branches of the chromozonal suture can be termed «medioscutal chromozonal suture» and «submedioscutal chromozonal suture» ( Fig. 94a ).

In the stage of subimago, the cuticle of the chromozone(s) is usually more or less intensively colored by brown, while the cuticle of the achromozone(s) is usually colorless. In some taxa the cuticle of the chromozone(s) is also colorless. In some taxa the cuticle of achromozone(s) is colored; in this case its color is different from the color of the chromozone (e.g., Kluge 2013: fig. 53).

In the primitive condition, the chromozonal (i.e., mesonotal) suture represents a deep furrow running transversely across the anterior part of scutum shortly posteriad of the anteronotal transverse impression, being either medially stretched backward into a short point ( Kluge 1994a: fig. 1; Kluge 2004: figs 6), or entirely transverse ( Kluge 2004: fig. 63). In these cases the subimaginal chromozone is located anteriad of the chromozonal suture, being not divided into medioscutal and submedioscutal zones, and the achromozone is located posteriad of the mesonotal suture, including either both medioscutum and submedioscutum ( Kluge 1994a: fig. 12; Kluge 2004: figs 18E, 19H, 21A, D–E, 23F, 25C, 30D, 32B, 33D–E, 34A, F, 39C–E, 54I, 63A, C, 64A, D, 90D; Kluge 1996: fig. 25) or medioscutum only, while submedioscutum is occupies by another pigmented zone ( Kluge 1994a: figs 13–14; Kluge 2004: figs 17D, 21B, 40D).

In certain mayfly taxa the shape of the chromozonal (mesonotal) suture is markedly changed, or this suture is lost. Particularly, in Fimbriatotergaliae and Leptophlebiidae the chromozonal suture is initially shifted backward by the side of the median line, so that on each side of the mesonotum it crosses the medioparapsidal suture ( Kluge 1994a: fig. 15; Kluge 2004: figs 68F, 71E–F, 74A, 83F, 106D). In these cases there can be distinguished chromozones of medioscutum and submedioscutum, and achromozones of medioscutum and submedioscutum. Both chromozones (medioscutal and submedioscutal ones) can be either equally pigmented ( Fig. 337 ; Kluge 2013: fig. 51), or colorless ( Fig. 368 ), or differ by coloration ( Fig. 94 ). Both achromozones (medioscutal and submedioscutal ones) can be either colorless, or pigmented, sometimes differently one from another ( Kluge 2013: fig. 52).

In many representatives of Leptophlebiidae , including the most primitive taxa ( Leptophlebiinae and Habrophlebiinae sensu Kluge 1994c), the chromozonal suture passes at a distance from median longitudinal suture, medioparapsidal suture and lateroparapsidal suture, so that both chromozones and both achromozones (medioscutal and submedioscutal ones) are well expressed ( Kluge 1994a: fig. 12, 1994c: fig. 47; 2004: fig. 40D; Kluge 2012: fig. 37, 92; 2013: figs 51–53; Kluge 2014: fig. 39). In some leptophlebiid taxa the chromozonal suture approaches the medioparapsidal suture, so that the chromozone is narrowed; particularly, this occurs in Terpidinae ( Kluge 2009: fig. 14) and in Hagenulus /fg2 ( Kluge 1994b: figs 20, 50, 87, 113, 201, 222). In Thraulodes the submedioscutal chro- mozonal suture approaches the lateroparapsidal and antelateroparapsidal sutures and repeats their shape, so that the submedioscutal achromozone looks as a narrow light stripe bordering dark lateroparapsidal and antelateroparapsidal sutures ( Fig. 94 ). The taxon Hermanellonota Kluge 2008 is characterized by loss of the chromozonal suture, so that in subimago the whole medioscutum and submedioscutum are either entirely colorless ( Kluge 1994b: fig. 13), or entirely colored ( Kluge 2008: fig. 28).

Fore wing venation

Venation of fore wing is uniform among Thraulodes , and only a few characters are used in species diagnoses.

A generally accepted species-specific character is presence/absence of cross veins in the proximal half of costal field (i.e. proximad of bulla). However, among species examined, none lacks these veins completely; this is true for the species which were described as lacking these veins— Th. telegraphicus , Th. sinuosus , Th. schlingeri , Th. marreroi , Th. quevedoensis and Th. zonalis . In all these cases, costal cross veins proximad of bulla are present, but few in number (one, two or more), colorless and much thinner than other cross veins, and can be visible only at magnification higher than necessary for examination of other veins ( Figs 32 , 142 , 344–345 ); in these species only selected individuals or individual wings lack these veins. Some authors use exact number of cross veins to separate species of Thraulodes (e.g. Souto et al. 2014). This is also wrong, because the number of cross veins in all fields of Leptophlebiidae and most other mayflies varies individually. Species-specific characters connected with cross veins, are only thickness and/or coloration of cross veins and their approximate number or density in certain fields.

One more character in wing venation was reported as species-specific for Thraulodes liminaris Dominguez 1987 : in this species vein ICu 2 is joined at base to vein CuP (Domínguez 1981: fig. 3; Domínguez et al. 2006: fig. 187D, in contrast to other Thraulodes , where ICu 2 is joined to ICu 1, but not to CuP. Among the species examined, the same condition, when the vein ICu 2 is joined at base to vein CuP, was found on one wing of one individual of Th. niger sp. n. ( Fig. 435 ), while on another wing of this individual ICu 2 is joined only to ICu 1, ( Fig. 434 ). Probably, this character is individual.

Genital structure

Styliger. Thraulodes differs from other taxa by the structure of the styliger, which has a common unpaired gonostylar cavity [termed by Traver and Edmunds (1967) «forceps plate»]. The gonostylar cavity is exposed ventrally-caudally ( Fig. 112 ), being well visible in ventral view ( Fig. 114 ), so that its dorsal margin is projected caudally. In various species this dorsal margin of styliger is either smoothly convex ( Fig. 534 ), or medially stretched into an unpaired dorsal projection ( Fig. 110 ), whose shape is species-specific or varies individually; in some species this projection is separated from the dorsal edge of styliger, shifted more dorsally and proximally, so that in ventral view its base is partly hidden by styliger ( Figs 38, 40 ). Since the bases of gonostyli are brought together in the gonostylar cavity, the pair of gonostylar muscles, which run from the styliger to the bases of gonostyli, are convergent ( Figs 102 , 110 ).

Gonostyli. The gonostylus retains all 4 primary segments, but the initial 1st and 2nd segments are not separated and often have equal thickness, and so can be taken for one segment; Traver and Edmunds (1967) called them together «basal segment». The 1st and 2nd segments are often bent in an obtuse angle; both distal segments (3rd and 4th ones) are short and always well separated ( Fig. 102 ). About the development of gonostyli, Traver and Edmunds (1967: 355) wrote «In male nymphs, only tips of forceps seen externally; dissection shows long basal segment to be strongly folded on itself in a U-shaped fashion, opening of U toward midline. In all other Neotropical genera of this family that we have studied, forceps relatively much longer and not folded on themselves as in Thraulodes ». Larval protogonostyli are really short ( Figs 106, 108 ) and in some species reduces to low protuberances ( Fig. 313 ). Subimaginal gonostyli developing inside are located mainly proximad of the protogonostyli, being folded in the junction of initial 1st and initial 2nd segments ( Fig. 108 ).

Musculature of penis. As in most mayflies, in Leptophlebiidae penis has two pairs of extrinsic muscles attached to its base: the pair of large median muscles, which are attached to anterior-lateral part of ninth abdominal sternum ( Figs 101 , 111 : m.med), and the pair of small lateral muscles, which are attached to lateral parts of ninth tergum ( Fig. 101 : m.lat) ( Grandi 1960). In Thraulodes , the median penial muscles are attached to the base of penis medially, close together. Penis lobes are separated nearly up to the base, so contraction of the median penial muscles should provide an active movement, bringing penis lobes together. In connection with this, median sides of penis lobes are often strengthened by a pair of medio-ventral ridges, or recurved folds ( Traver & Edmunds 1967); each of these ridges forms the median margin of the penis lobe and is projected on its ventral side; it stretches from the base of penis (where the median muscle is attached) in distal direction ( Figs 103 , 111 ). In some species the free ventral margin of this medio-ventral ridge extends laterally, so that the ridge represents a flap overlapping some part of the ventral surface of penis ( Figs 530 , 534 ); a certain kind of such recurved flap is termed lapel ( Traver & Edmunds 1967). The medio-ventral ridges are either developed only in basal part of penis ( Fig. 38 ), or are stretched distally, sometimes up to the apex. Length and shape of the medio-ventral ridge is regarded as an important species-specific character ( Traver & Edmunds 1967); in some species it is such, but in Th. telegraphicus it demonstrates a great individual variability ( Figs 114–120 ).

Spermatic pump. Leptophlebiidae have aflagellate spermatozoons ( Soldan 1979, Gaino & Mazzini 1991); in connection with this, their spermatic ducts are equipped by ring muscles, which serve as spermatic pumps ( Grimm 1985; Brito et al. 2012). In many species of Thraulodes , basal part of each penis lobe forms a ventro-lateral swelling, which Traver & Edmunds (1967) named basal lobe. Inside the basal lobe and proximad of it, the gonoduct is widened ( Traver & Edmunds 1967: fig. 30) and surrounded by more or less thick layer of ring-like muscles, which forms the spermatic pump ( Figs 101, 103 , 111 ). Apart of the muscles of spermatic pump located in the basal lobe, the penis of Thraulodes has no any other intrinsic muscles.

Spear-like rolls. The pair of pointed processes, or «the spears, a very characteristic feature of this genus» ( Traver & Edmunds 1967: 353), were characterized as the following: «Each penis lobe bears a slender spearlike process arising dorsally from distal margin, the points of the spears directed inward and toward one another» (ibid, 352) and «The narrow portion of [spermatic] duct can be traced apicad from the basal lobe as far as the base of the spear» (ibid, 353). Some other authors call these processes «spines» ( Dominguez 1987). In all publications these processes are described and figured as integral spines. On some figures such spine is crossed by an oblique line, but without comments (e.g. Zúñiga et al. 2015: fig. 8D).

Actually, these processes are not integral, but each represents a tube with longitudinal cleft stretched up to base. Such an incomplete tube can be interpreted as a rolled plate, whose margins are either separated ( Figs 104 , 144 , 171 , 311 , 439 ,), or contiguous ( Figs 200 , 372 , 405 , 535 ,), or overlap one another ( Figs 270 , 582 ). Apex of this roll is always obliquely truncate, forming a sharp point. The spermatic duct is opined into the canal of this roll, so that sperm escapes from the apical foramen and the cleft of the roll ( Fig. 472 ).

The spear-like processes are attached either on apical, or apical-dorsal side of the penis, and in most species are directed more or less dorsally ( Figs 143 , 267 ); in some species they are inclined ventrally ( Figs 203 , 239 , 240 ); this is an important species-specific character, which is often overlooked or confused on the drawings where genitalia are shown in ventral or dorsal view.

In the overwhelming majority of species, the spear-like rolls are directed medially (i.e., inward and toward one another) and each has a narrow cleft exposed caudally-laterally-dorsally ( Figs 112–114 ). Only in two closely related species, Th. sinuosus and Th. ludmilae sp. n., these processes are directed ventrally and unrolled, so that each process represents a concave plate directed by its concavity ventrally-caudally ( Fig. 60 ) or ventrally-laterally ( Figs 38, 40 ).

Larval protopenis and development of spear-like rolls. Traver and Edmunds (1967: 355) wrote that «In male nymphs, ... Each penis lobe bears a slender elongate projection». This is true for the protopenis (i.e., non-functional larval penis) of the majority of the species examined ( Fig. 107 ), but in some species these projections are not expressed ( Fig. 271 ). The shape of the protopenis is species-specific. In all cases, the apex of each protopenis lobe bears a foramen, which leads to a narrow, long, thin-walled, cuticular tube, corresponding to the imaginal spermatic duct. If the apical projection is present, the foramen is located on its top, being exposed either caudally ( Fig. 313 ), or caudally-laterally ( Fig. 205 ).

When the subimaginal penis lobe is developed under the larval cuticle, its spear-like roll appears as inserted into the apical projection of protopenis, and the cuticular tube of the protopenis is inserted into the canal of the spear-like roll ( Fig. 107 ). During subsequent growth, the spear-like roll is elongated, being inserted by its base into the growing penis, and remains to be directed caudally with the cuticular tube of protopenis inside ( Figs 109 , 347 , 500–501 , 607 ). During the molt from larva to subimago, the spear-like rolls turn medially and get definite position when subimaginal genitalia become free from cuticle of larval protopenis, even when subimaginal abdomen is not yet taken out from the larval cuticle (observed for Th. telegraphicus ).

In subimago, the spear-like rolls have the same size and shape as in imago; their cuticle is rigid and retains its shape on subimaginal exuviae after molt from subimago to imago, while remainding exuviae of the subimaginal penis is very delicate and is crumpled during the molt ( Fig. 475 ).

The slender projection of protopenis can be interpreted as a larval precursor of the spear-like roll. Length of this larval projection does not correlate with length of imaginal spear-like roll: for example, in Th. quevedoensis the projections of larval protopenis are absent, while the imaginal spear-like rolls are well developed.

Unfortunately, development of unusual penes of Th. sinuosus and Th. ludmilae sp. n. was not examined.

Judging by their structure and transformations in ontogenesis, the spear-like rolls of Thraulus are homologous to the tubular telopenes of Hermanellonota Kluge 2008. Their presence is a synapomorphy of Thraulodes and Hermanellonota, which allows uniting them to a new taxon Hermanellandria (see below).

Egg structure

In all species of Thraulodes , knob-terminated coiled threads (KTC) are evenly dispersed all over egg surface, equally on equator and pole regions, independently from shape of the egg, which varies from ellipsoid ( Fig. 589 ) to barrel-shaped, i.e. with flat pole areas ( Fig. 348 ). Each KTC is located in a round cavity and coiled as a regular spiral always directed anticlockwise ( Fig. 150 ). The knob, which terminates this coiled thread, is located in the middle of the spiral. The coiled thread can be either located openly [in Th. zonalis — Fig. 349 ; in Th. paysandensis ( Dominguez 1987: fig. V.4)], or hidden by a ring-like cover, which surrounds the knob (in other examined species). In various species, this ring-like cover can be narrower or wider, smooth or rugose.

Other chorion is either smooth ( Fig. 121 ) or bears round protuberances, which can be large or smaller, located either evenly ( Fig. 249 ), or concentrated around KTCs ( Figs 274–275 ), or, vice verse, located at maximum distance from KTCs forming rows ( Figs 315–316 ); in some species rows of protuberances are fused into integral ridges, which form a net-like relief with KTC in each cell ( Figs 377–378 ).

Among the species examined, one species Thraulodes viviparus sp. n. is probably ovoviviparous; in a single examined female imago of this species, abdomen is filled by completely grown eggs lacking chorion.

Systematic position and diagnosis of Thraulodes

Thraulodes belongs to subsequently subordinated taxa: Leptophlebiidae > Atalophleboadentata Kluge 2009 > Atalophlebopectinata Kluge 2009 > Atalophleboculata Kluge 2009 > Atalophlebomaxillata Kluge 2009 > Atalophlebolinguata Kluge 2009. General system of the taxon Atalophlebolinguata remains poorly elaborated. One of the taxa established within Atalophlebolinguata is Hagenulus /fg1, or the tribe Hagenulini Kluge 1994b ( Kluge 2008) .

Hagenulus /fg1

Tribe Hagenulini Kluge 1994b: 250 ;

Hegenulus/fg1: Kluge 2008: 387.

Autapomorphy. Patella-tibial suture is lost on all legs of all stages—larva ( Figs 21–24 ), subimago ( Fig. 31 ) and imago ( Figs 28–30 ).

Other characters. Hind wing has more or less prominent costal projection; vein Sc terminates on costal margin just distad of costal projection, far from wing apex ( Fig. 41 ) (apomorphy, independently occurred in various taxa). Imaginal and subimaginal claws of all legs are ephemeropteroid (plesiomorphy).

Distribution. Neotropical Region.

Composition. The taxon Hagenulus /fg1 comprises Hagenulus /fg2 (incl. Borinquena , Turquinophlebia , Poecilophlebia , Careospina , Traverina , Hagenulopsis , Neohagenulus ), Ulmeritus /g1 (incl. Ulmeritoides ), Miroculus/ g1, Hermanellandria taxon nov. and some other taxa. The new taxon Hermanellandria comprises Thraulodes and Hermanellonota:

Hermanellandria taxon nov.

Thraulodes group: O’Donnell & Jockusch 2008: 663;

Circumscriptional name: Hermanellandria taxon nov.;

Hierarchical name: Hermanella /f1= Thraulodes /g1;

Possible rank-based name: subtribe Hermanellina .

Autapomorphy of Hermanellandria. Each of two penis apices is produced into a slender sclerotized tubular process— telopenis, inside which the genital duct passes; apex of telopenis is obliquely truncated and pointed. In imago telopenes are sharply bent at their bases and/or shifted from apices of penis lobes, so that are directed either medially (in most Thraulodes ), or laterally (in Farrodes ), or ventrally (in Hermanellognatha), or even ventrallyanteriorly, i.e. toward penis base. Unlike imago, in mature larva telopenes represent apical continuations of penis lobes, being directed caudally, so that gonopores occupy apical position; larval protopenis has developed gonopores with ducts lined by larval cuticle ( Figs 107, 109 , 347 , 500–501 , 607 ; Kluge 2008: Figs 9–11 , 15 , 27–28, 32, 34 ). In subimago telopenes either have the same position as in imago, or position intermediate between larval and imaginal ones ( Kluge 2008: Fig. 12 ). Subimaginal telopenes are sclerotized, while the rest subimaginal cuticle of penis is membranous.

In Hermanellognatha and Simothraulopsis each telopenis represents a tube with apical opening. In all Thraulodes each telopenis is a roll, i.e. a tube cleft up to the base. In Farrodes telopenes lack gonoducts, and genital openings are located at their bases.

Telopenes can be confused with superficially similar processes, which occur in some other taxa and arise from penis apex toward its base; but unlike telopenes, penial processes of other Leptophlebiidae do not bear genital ducts and in larval stage develop in the same position as they are in imago.

Distribution. Neotropical Region.

Composition. The taxon Hermanellandria (or Hermanella /f1= Thraulodes /g1) comprises Thraulodes and Hermanellonota Kluge 2008 (or Hermanella /f2=g1). In its turn, Hermanellonota is divided into Simothraulopsis Demoulin 1966 , Farrodes Peters 1971 and Hermanellognatha Kluge 2008 (or Hermanella /f3=g2).

Comments. O’Donnell and Jockusch (2008) established the « Thraulodes group» based on similarity of histone H3 and 28S ribosomal DNA of three examined species, Thraulodes sp., Farrodes sp. and Traverella albertana Edmunds (which belongs to the « Hermanella complex», i.e. to Hermanellognatha Kluge 2008). These authors stated that their analysis did not support hypothesis about relationship of these species with Meridialaris lineage, which was proposed by Flowers and Dominguez (1991). O’Donnell and Jockusch (2008) also wrote that «this grouping is additionally supported by the similarity of maxillary palps in Thraulodes spp. and Farrodes spp.», but did not report concrete synapomorphies in the maxillary palp structure. The taxon with the new circumscriptional name Hermanellandria corresponds to the « Thraulodes group» established by O’Donnell and Jockusch (2008).

The idea about relationship of Thraulodes with the Notogean genera Meridialaris and Massartellopsis suggested by Flowers and Dominguez (1991) was based on parsimony analysis of a limited number of characters of several New World genera. Generally speaking, the method of parsimony analysis is not based on any scientific theory; a tree built by this method depends on occasional set of characters, their arbitrary formulation and an arbitrarily chosen method of calculation, but not on revealed apomorphies ( Pesenko 1989). I the tree suggested by Flowers and Dominguez (1991: Fig. 3 ), the branch comprising Thraulodes , Meridialaris and Massartellopsis and the subordinated branch comprising Thraulodes and Meridialaris were argued by characters «1» (wide labrum), «3» (hood of labrum), «30» (dorsal setae on fore tibia), «50» (spines on penis) and «25» (absence of setae on submentum). Each of these characters was also reported for some other taxa, so no one autapomorphy of these branches was reported; the character «3» was also reported as lost in Thraulodes ( Flowers & Dominguez 1991: Table 1 ). The character «spines on penis» was reported, besides Thraulodes , Meridialaris and Massartellopsis , for Massartella Lestage 1930 , for all taxa belonging to Hermanellognata, for Traverina Peters 1971 and for Hagenulus Eaton 1882 ; among Hagenulus , it was erroneously reported as absent in H. morrisonae Peters & Alayo (in Peters 1971) , while actually this species has diminished spines ( Kluge 1994b: Fig. 37 ). These «spines» of Thraulodes and «spines» of Hermanellognatha are homologous, being telopenes with sperm canal inside (see above). In contrast to them, the «spines» of Traverina and Hagenulus are a pair of simple denticles on ventral side of penis far from the genital opening. The «spines» of Meridialaris , Massartellopsis and Massartella are various small processes on margins of genital opening; in various species these processes are soft or sclerotized, pointed or blunt, stout or hair-like; no one of these processes contains sperm canal, neither in imaginal penis, not in larval protopenis. So the idea about a relationship of Thraulodes with the Meridialaris lineage is groundless.

Earlier, Flowers (1987) proposed a relationship between Thraulodes and Atopophlebia Flowers 1976 . Unfortunately, I am unable to examine all stages of development of Atopophlebia , being limited by a single undetermined female larva from Peru. Its examination reveals the following characters common for Atopophlebia and Thraulodes : horseshoe row of hairs on dorsal side of glossa (see above); dense setation on distal part of outer side of mandible (the same in some other taxa—see above); absence of patella-tibial suture on at least middle leg of larva (the same in all other Hagenulus /fg1 and some other taxa). The penis of Atopophlebia has a pair of apical processes directed ventrally; their structure and development are not described, so it is unclear, if they are telopenes, or not. At present, the position of Atopophlebia in Hermanellandria is an open question.

Thraulodes Ulmer 1920

Hierarchical name: Thraulodes /g2;

Rank-based name: Thraulodes Ulmer 1920: 33 (imago); Needham & Murphy 1924 (imago, larva); Traver & Edmunds 1967 (imago, larva, eggs; species revision)

Type species: Thraulus laetus Eaton 1884 .

Diagnosis.

Autapomorphies:

(1) Bases of gonostyli are brought together and inserted in a common unpaired gonostylar cavity of styliger; dorsal margin of styliger is projected caudally, and the gonostylar cavity is exposed ventrally-caudally ( Figs 102 , 110 , 112 ).

(2) Telopenis (see autapomorphy of Hermanellandria) is tubular with a longitudinal cleft up to its base, i.e. represents a roll ( Figs 103–105 ) (in contrast to integral tube of Hermanellognatha and Simothraulopsis ).

(3) On hind tibia, stout setae of outer margin (initial for Leptophlebiidae ) form two separated rows, outer-anterior and outer-posterior ones; long hairs (also initial for Leptophlebiidae ) are situated between these two rows of stout setae and form one more row posteriad of them; row of recurved hairs is located on anterior side of tibia, more distally from inner margin than the inner-anterior row of stout setae, which is present on middle and hind legs ( Fig. 89 ).

Characters of unclear phylogenetic status:

(4) Outer side of each mandible with distal setal bunch, but without middle setal bunch: long hairs forming a row or a field on outer margin of mandible, are denser and longer nearer the base of incisor ( Figs 17–18 ). (the same in some other taxa—see above).

(5) Long setae of dorsal side of glossa form irregular row or stripe of a characteristic horseshoe shape ( Fig. 20 ) (the same in Atopophlebia —see above).

(6) Swimming larva makes lateral movements with its abdomen (in contrast to most other Ephemeroptera , which make dorso-ventral swimming movements). Among the taxa whose larval swimming was observed, Kimminsula makes the same movements as Thraulodes .

Plesiomorphy:

(7) Hind wing always retains furcations of RS and MP (while furcation of MA is lost in all Ephemeroptera ) ( Fig. 41 ).

Figures

FIGURE 121. Thraulodes telegraphicus, eggs.

FIGURES 35–41. Thraulodes ludmilae sp. n.: 35, genitalia of male imago in ventral view; 36, subimaginal exuviae of styliger and gonostyli at the same magnification; 37, larval exuviae of protogonostyli and protopenis at the same magnification; 38, imaginal styliger and penis enlarged; 39, subimaginal exuviae of penis at the same magnification; 40, genitalia of male imago in lateral view; 41, hind wing (35–39, 41, holotype). Abbreviations: bl, basal lobe; gr, groove of telopenis; dp, dorsal projection of styliger; m-v.r, medio-ventral ridge (recurved flap); tp, telopenis.

FIGURES 464–473. Thraulodes nigrabdominalis sp. n. (holotype). 464, left half of subimaginal exuviae of mesonotum; 465, 466, male imago; 467–469, fore, middle and hind legs of male imago; 470, subimaginal exuviae of fore leg; 471, genitalia of male imago; 472, apex of penis; 473, subimaginal exuviae of tarsus.

FIGURES 491–497. Thraulodes nigripes sp. n. 491–492, fore and middle legs of female imago; 493–494, female imago in dorsal and lateral view; 495, abdominal terga and sterna of male subimago extracted from larva; 496, genitalia of male subimago extracted from larva; 497, right half of subimaginal exuviae of mesonotum (495–496, holotype).

FIGURES 522–531. Thraulodes nigrotibialis sp. n. 522, right half of subimaginal exuviae of mesonotum; 523, head and thorax of male imago in lateral view; 524–525, female imago in dorsal and lateral view; 526–527, fore and middle leg of male imago; 528, abdominal terga and sterna of male imago; 529–530, genitalia of male imago with focus on styliger and on penis; 531, genitalia of male subimago extracted from larva (522–523, 526–530, holotype).

FIGURES 427–438. Thraulodes niger sp. n. 427–428, head and thorax of male imago in lateral and dorsal view; 429, abdominal terga and sterna of male imago; 430, the same of male subimago; 431, subimaginal mesonotum; 432–433, genitalia of male imago with focus on styliger and on penis; 434, female imago; 435, its left fore wing; 436–437, its fore and middle legs; 438, subimaginal exuviae of fore wing (427–429, 432–433, 438, holotype).

FIGURES 25–34. Thraulodes ludmilae sp. n.: 25, 27, male imago; 26, apex of fore wing; 28–30, fore, middle and hind legs of male imago; 31, subimaginal exuviae of hind leg; 32, costal margin of fore wing (arrows show proximal costal cross veins); 33, genitalia of male imago; 34, left half of subimaginal exuviae of mesonotum (25–26, 28–34, holotype).

FIGURES 297–308. Thraulodes fascipennis sp. n. 297, left half of subimaginal exuviae of mesonotum; 298–300, male imago in dorsal, ventral and lateral view; 301, abdominal terga and sterna of male imago; 302–303, male imaginal genitalia in ventral and lateral view; 304, male imaginal penis; 305–307, fore, middle and hind legs of male imago; 308, subimaginal exuviae of fore leg (301, holotype).

FIGURES 654–665. Thraulodes? consortis. 654, larval protogonostyli and protopenis; 655, fore protopteron; 656, hind protopteron; 657, larva with ocher cuticular coloration and blackish hypodermal markings on head and abdomen; 658–660, fore, middle and hind legs of male subimago; 661, abdominal terga and sterna of male subimago; 662, anterior side of hind femur; 663, anterior side of fore tibia; 664, hind tibia with focus on outer-posterior row of stout setae; 665, the same with focus on outer-anterior and inner-anterior rows of stout setae.

FIGURES 591–602. Thraulodes lepidus. 591, larva in dorso-lateral view with brown cuticular markings and blackish hypodermal markings on head, thorax, protoptera, legs and abdomen; 592–598, larval exuviae at the same magnification; 597–598, fore and hind larval legs with subimaginal legs inside; 599, larval fore tibia; 600, anterior side of hind femur; 601, hind tibia with focus on outer-posterior row of stout setae; 602, the same with focus on outer-anterior and inner-anterior rows of stout setae.

FIGURES 337–345. Thraulodes zonalis. 337, left half of subimaginal exuviae of mesonotum; 338, female imago with its subimaginal exuviae; 339–341, fore, middle and hind legs of female imago; 342, hind wing; 343, abdominal terga and sterna of male subimago extracted from larva (sterna VIII–IX with larval cuticle); 344–345, anterior-proximal part of fore wing.

FIGURES 365–370. Thraulodes flavus sp. n. 365–366, male imago in lateral and dorsal view; 367, female imago; 368, left half of subimaginal exuviae of mesonotum; 369, subimaginal exuviae of fore leg of male; 370, genitalia of male imago (368–370, holotype).

FIGURES 569–577. Thraulodes alboniger sp. n. 569, female subimago extracted from larva; 570, apex of fore wing; 571–573, abdominal terga of three different male imagines; 574–575, two male imagines in lateral view; 576, head and thorax of male imago in dorsal view; 577, right half of subimaginal exuviae of mesonotum (570, 572, 577, holotype).

FIGURES 94–100. Thraulodes telegraphicus. 94–95, right half of subimaginal exuviae of mesonotum of two different specimens; 96, apex of fore wing; 97–98, male imago in lateral and ventral view; 99, abdominal terga and sterna of male imago (black dots are algae under cuticle); 100, female imago. Abbreviations: acz.ms, achromozone of medioscutum; acz.sms, achromozone of submedioscutum; cz.ms, chromozone of medioscutum; cz.sms, chromozone of submedioscutum; mp.s, medioparapsidal suture; ms.cz.s, medioscutal chromozonal suture; sms.cz.s, submedioscutal chromozonal suture.

FIGURES 396–402. Thraulodes spangleri. 396–397, left half of subimaginal exuviae of mesonotum of two different specimens; 398–400, male imago in lateral, dorsal and ventral view; 401, female imago; 402, genitalia of male imago.

FIGURES 209–314. Thraulodes fascipennis sp. n. 309, hind wing; 310, genitalia of male imago in ventral view; 311, apex of penis in dorsal view; 312, genitalia in lateral view; 313, larval exuviae of protogonostyli and protopenis; 314, subimaginal exuviae of genitalia (313, holotype).

FIGURES 439–447. Thraulodes niger sp. n. 439, apex of penis in dorsal view; 440, genitalia of male imago in ventral view; 441, their subimaginal exuviae; 442, larval protogonostyli and protopenis; 443, hind wing; 444–447, tergalii I, III, VI and VII.

FIGURES 198–202. Thraulodes schlingeri. 198–199, male imago in lateral and ventral view; 200, penis in apical view; 201, penis in ventral view; 202, left half of subimaginal exuviae of mesonotum.

FIGURES 232–239. Thraulodes marreroi. 232–234, male imago in dorsal, ventral and lateral view; 235, left half of subimaginal exuviae of mesonotum; 236–239, male imaginal genitalia in apical, dorsal ventral and lateral view.

FIGURES 260–266. Thraulodes quevedoensis. 260–261, genitalia of male imago with focus on penis and on styliger; 262, right half of subimaginal exuviae of mesonotum; 263–264, male imago in ventral and lateral view; 265, abdominal terga and sterna of male imago; 266, apex of fore wing.

FIGURES 203–206. Thraulodes schlingeri. 203–204, male imaginal genitalia in lateral and ventral view; 205, larval protogonostyli and protopenis; 206, hind wing.

FIGURES 240–245. Thraulodes marreroi. 240, genitalia of male imago in lateral view; 241, apex of penis in dorsal view; 242, genitalia in ventral view; 243, sibimaginal exuviae of styliger, gonostyli and telopenes; 244, larval protogonostyli and protopenis; 245, hind wing.

FIGURES 267–273. Thraulodes quevedoensis. 267, genitalia of male imago in lateral view; 268, styliger and penis in ventral view; 269, genitalia of another specimen in ventral view; 270, the same, apex of penis in dorsal view; 271, larval protogonostyli and protopenis; 272, hind wing; 273, larval claw of middle leg.

FIGURES 52–59. Thraulodes sinuosus. 52–53, male imago in dorsal and ventral view; 54, genitalia of male imago with focus on gonostyli; 55, the same, with focus on penis; 56, right half of subimaginal exuviae of mesonotum; 57–58, female imago in ventral and lateral view; 59, pterostigma of fore wing.

FIGURES 133–142. Thraulodes panamensis sp. n. 133, pterostigma; 134–136, male imago in lateral in ventral and dorsal view; 137–138, left half of subimaginal exuviae of mesonotum of two different specimens; 139, female imago; 140–141, genitalia in lateral and ventral view; 142, costal margin of fore wing (arrows show proximal costal cross veins) (133, 134, 136, 137, 141, 142, holotype).

FIGURES 156–168. Thraulodes viviparus sp. n. 156, pterostigma; 157–159, head and thorax of male imago in lateral in ventral and dorsal view; 160–161, fore and hind legs of male imago; 162–164, female imago in ventral in dorsal and lateral view; 165, left half of subimaginal exuviae of mesonotum; 166, abdominal terga and sterna; 167–168, penis with focus on dorsal and ventral side (156–161, 165–168, holotype).

FIGURES 60–63. Thraulodes sinuosus. 60, genitalia of male imago in ventral view; 61, their subimaginal exuviae at the same magnification; 62, larval exuviae of protogonostyli and protopenis at the same magnification; 63, hind wing.

FIGURES 112–120. Thraulodes telegraphicus. 112, genitalia of male imago in lateral view; 113, apex of penis with spear-like roll in dorsal view; 114, genitalia of the same specimen in ventral view; 115, their subimaginal exuviae at the same magnification; 116, hind wing; 117–120, penis of four different specimens. Abbreviations: dp, dorsal projection of styliger; tp, spear-like roll (telopenis).

FIGURES 143–148. Thraulodes panamensis sp. n. 143, genitalia of male imago in lateral view; 144, apex of penis with spearlike roll in dorsal view; 145, genitalia of the same specimen in ventral view; 146, subimaginal exuviae of genitalia at the same magnification; 147, larval exuviae of protogonostyli and protopenis at the same magnification; 148, hind wing (146, 147, holotype). Abbreviations: lp, pateral pouch; tp, telopenis.

FIGURES 169–173. Thraulodes viviparus sp. n. 169, genitalia of male imago in ventral view; 170, their subimaginal exuviae at the same magnification; 171, apex of penis in dorsal view; 172, protogonostyli and protopenis of presumably associated larva; 173, hind wing (169–171, 173, holotype). Abbreviations: lp, pateral pouch; tp, telopenis.

FIGURES 190–197. Thraulodes schlingeri, labrum. 190–193, labrum on slide: 190, total view; 191, anterior margin with focus on posterior transverse setal row; 192–193, with focus on ventral side, median emargination and anterior setal row (represented by field of irregularly situated setae); 194–197, labrum of another specimen in anterior view: 194, total view; 195, median part with focus on anterior and posterior setal rows; 196, the same with focus on median emargination; 197, the same enlarged (arrows show six sensilla separating five denticles). Abbreviations: anter.r, anterior transverse row of setae (actually field of setae); poster.r, posterior transverse row of setae.

FIGURES 208–231. Thraulodes marreroi. 208, margin of labrum with focus on posterior transverse setal row; 209, the same with focus on anterior transverse setal row (represented by field of irregularly situated setae) (arrows show four sensilla separating three denticles). 210–215 and 216–219, larval exuviae of two different specimens at the same magnification; 220–226, tergalii I–VII; 227, immature larva (with ocher and whitish cuticular coloration and blackish paired hypodermal markings on abdomen); 228, anterior side of hind femur; 229, anterior side of fore tibia; 230, hind tibia with focus on outer-posterior row of stout setae; 231, the same with focus on outer-anterior and inner-anterior rows of stout setae.

FIGURES 389–395. Thraulodes spangleri, labrum. 389–391, one specimen: 389, general view with focus on dorsal side and posterior transverse setal row; 390, focus on ventral side and denticles; 391, outline of denticles separated by four sensilla (shown as dots). 392–395, another specimen: 392, focus on dorsal side and posterior setal row; 393, focus on anterior transverse setal row (transformed to field of irregularly situated setae); 394, focus on denticles and tips of irregularly situated setae of the anterior row; 395, outlines of vestigial denticles separated by incisions and two sensilla.

FIGURES 650–653. Thraulodes sp. «Itaya», labrum. 650, general view; 651, focus on ventral side and six sensilla (shown by arrows); 652, with focus on anterior transverse setal row (transformed to field of irregularly situated setae); 653, with focus on posterior transverse setal row.

FIGURES 637–649. Thraulodes sp. «Itaya», larva. 637, tergalius III; 638–645, larval exuviae at the same magnification: 638, part of head with labrum; 639–640, left and right mandibles; 641, right half of pronotum and mesonotum; 642–644, fore, middle and hind legs; 645, abdominal terga and sterna; 646, anterior side of hind femur; 647, anterior side of fore tibia; 648, hind tibia with focus on outer-posterior row of stout setae; 649, the same with focus on outer-anterior and inner-anterior rows of stout setae.

FIGURES 174–189. Thraulodes schlingeri. 174–184, larval exuviae at the same magnification: 174, head with labrum and mandibles; 175, right pronotum and mesonotum; 176, part of sternum and left pleura with half of metanotum and hind protopteron, left fore, middle and hind legs; 177, abdominal terga and sterna; 178–184, tergalii I–VII; 185, immature larva (with ocher and whitish cuticular coloration and blackish paired hypodermal markings on abdomen); 186, anterior side of hind femur; 187, anterior side of fore tibia; 188, hind tibia with focus on outer-posterior row of stout setae; 189, the same with focus on outeranterior and inner-anterior rows of stout setae.

FIGURES 379–388. Thraulodes spangleri. 379–383, larval exuviae at the same magnification: 379–381, parts of head with labrum and mandibles; 382, right half of pronotum and mesonotum; 383, abdominal terga and sterna; 384, abdominal terga and sterna of male subimago extracted from larva; 385, tergalius; 386, larval fore tibia with subimaginal tibia inside and hypodermal dark brown markings; 387, anterior side of larval hind femur; 388, anterior side of larval hind tibia.

FIGURES 86–93. Thraulodes telegraphicus. 86, median longitudinal section of labrum in usual condition; 87, the same, in spread condition; 88, labrum in spread condition in dorsal view (setal rows shown as dotted lines); 89, transverse section of hind tibia; 90–93, tergalii I, III, VI and VII. Abbreviations: anter.r, anterior transverse row of setae; dent., five denticles separated by six sensilla; poster.r, posterior transverse row of setae; i.f, inner field of stout pointed setae; i-a.h, inner-anterior recurved hairs; i-a.r, inner-anterior row of stout setae; o-a.r, outer-anterior row of stout setae; o.h, outer hairs; o-p.r, outer-posterior row of stout setae; p.r, row of posterior pointed pectinate setae.

FIGURES 563–568. Thraulodes alboniger sp. n. 563, labrum; 564–565 and 566–567, male imaginal genitalia of two different specimens; 568, fragment of imaginal caudalius (563, 566–568, holotype).Abbreviations: anter.r., anterior transverse setal row; poster.r., posterior transverse setal row; other arrow show six sensilla separating five denticles.

FIGURES 543–562. Thraulodes alboniger sp. n. 543, larva with ocher cuticular coloration and blackish hypodermal markings on thorax, legs and abdomen, 544–557, larval exuviae at the same magnification; 551–557, tergalii I–VII; 558, fore tibia; 559, its inner-apical margin in anterior view; 560, anterior side of fore femur; 561, hind tibia with focus on outer-posterior row of stout setae; 562, the same with focus on outer-anterior and inner-anterior rows of stout setae (544–562, holotype).

FIGURES 578–588. Thraulodes alboniger sp. n. 578, hind wing; 579, genitalia of male imago in ventral view; 580, dorsal margin of styliger and penis of another specimen; 581, subimaginal exuviae of genitalia; 582, penis in lateral view; 583, posterior margin of IX sternum of female larva; 584, the same of male larva with protogonostyli and protopenis; 585–588, tergalii I, III, VI and VII (578, 580–581, 584–588, holotype).

FIGURES 5–13. Thraulodes ludmilae sp. n., larval exuviae of holotype (5–10 at the same magnification): 5–7, parts of head with labrum and mandibles; 8, right half of pronotum and mesonotum; 9, part of sternum and left pleura with half of metanotum and hind protopteron, left fore, middle and hind legs; 10, abdominal terga and sterna; 11, tergalii II–III; 12, part of pronotum; 13 anterior side of fore femur.Abbreviation: f.a.m, free anterior margin of pronotum.

FIGURES 66–69. Thraulodes telegraphicus. 66–67, last instar larvae with intensive cuticular coloration and poorly visible hypodermal coloration; 68, younger larva with light cuticle and well visible blackish hypodermal markings on abdomen; 69, claw. Abbreviations: artic.p., articulatory portion; rigid p., rigid portion of claw.

FIGURES 70–85. Thraulodes telegraphicus, larval exuviae (70–82 at the same magnification): 70–75, right half of pronotum and mesonotum; 76, head with labrum and mandibles; 77–78 and 79–80, fore and hind legs of two specimens; 81–82, abdominal terga; 83, anterior side of hind femur; 84, anterior side of fore tibia; 85, anterior side of hind tibia. Abbreviations: i-a.r, inneranterior row of stout setae; o-a.r, outer-anterior row of stout setae; o-p.r, outer-posterior row of stout setae.

FIGURES 21–24. Thraulodes ludmilae sp. n., exuviae of tibiae of holotype (at the same magnification): 21, anterior side of fore leg; 22, anterior side of middle leg; 23, posterior side of hind leg; 24, anterior side of hind leg.

FIGURES 603–612. Thraulodes lepidus. 603, larva with brown cuticular markings and blackish hypodermal markings; 604– 605, cuticle of larval protogonostyli and protopenis; 606–607, subimaginal styliger with gonostyli and penis extracted from larva; 608, tergalius III; 609–610, subimaginal thorax extracted from larva; 611, abdom of male subimago extracted from larva; 612, its enlarged fragment. Abbreviations: cut.subim, brown subimaginal cuticle with small paired blanks; hyp.w, non-pigmented area of hypoderm visible through blanks of cuticle; hyp.b, dark brown hypodermal coloration.

FIGURES 411–426. Thraulodes niger sp. n. 411, larval claw; 412–413, 414–418 and 419–420, larval exuviae of three specimens at the same magnification; 421, larval abdomen with ocher cuticular markings and blackish hypodermal markings; 422, anterior side of hind femur; 423, anterior side of fore tibia; 424, the same with focus on more posterior setae; 425, hind tibia with focus on outer-posterior row of stout setae; 426, the same with focus on outer-anterior and inner-anterior rows of stout setae (412–413, holotype).

FIGURES 450–463. Larvae, presumably attributed to Thraulodes nigrabdominalis sp. n. 450–455, parts of last instar larva at the same magnification: 450, fragment of head with labrum and mandibles; 451, pronotum and mesonotum; 452, hind leg; 453, abdominal terga; 454–455, tergalii IV–V; 456–458, exuviae of young larva at the same magnification: 456–457, fore and hind legs; 458, abdominal terga and sterna; 459, young larva with brown cuticular coloration and blackish hypodermal coloration; 460, larval protogonostyli and protopenis; 461, anterior side of hind femur; 462, hind tibia with focus on outer-posterior row of stout setae; 463, the same with focus on outer-anterior and inner-anterior rows of stout setae.

FIGURES 151–155. Larva presumably attributed to Thraulodes viviparus sp. n. 151, anterior side of hind tibia; 152, anterior side of hind femur; 153, tergalius VII; 154, general view; 155, abdominal terga spread on slide (with brown and whitish cuticular coloration and blackish paired hypodermal markings).

FIGURES 350–360. Thraulodes flavus sp. n. 350–356, larval exuviae at the same magnification: 350, abdominal terga; 351– 353, parts of head with labrum and mandibles; 354, left half of pronotum and mesonotum; 355–356, fore and hind legs; 357, hind tibia with focus on outer-posterior row of stout setae; 358, the same with focus on outer-anterior and inner-anterior rows of stout setae; 359, anterior side of fore femur; 360, anterior side of fore tibia (350–356, 359–360, holotype).

FIGURES 613–625. Thraulodes sp. «Palo Seco», larva. 613, clypeus and labrum; 614–615, left and right mandibles; 616–617, larvae with brown cuticular coloration and blackish hypodermal markings; 618, fragment of fore protopteron with venation of pterostigma; 619, hind protopteron; 620, fore protopteron; 621, fore tibia with dark brown hypodermal marking; 622, anterior side of fore tibia; 623, hind tibia with focus on outer-posterior row of stout setae; 624, the same with focus on outer-anterior and inner-anterior rows of stout setae; 625, anterior side of hind femur.

FIGURES 122–132. Thraulodes panamensis sp. n. 122–127, larval exuviae at the same magnification: 122–124, parts of head with labrum and mandibles; 125, left half of pronotum and mesonotum; 126, part of sternum and left pleura with half of metanotum and hind protopteron, left fore, middle and hind legs; 127, abdominal terga and sterna; 128, immature larva with light cuticle and well visible blackish hypodermal markings on abdomen; 129, tergalius III; 130, anterior side of hind femur; 131, anterior side of fore tibia; 132, anterior side of hind tibia (122–127, 130, holotype).

FIGURES 276–296. Thraulodes fascipennis sp. n. 276–283, larval exuviae at the same magnification; 284–290, tergalii I–VII; 291, ventral lamella of tergalius IV; 292, young larva with light cuticle and well visible blackish hypodermal markings; 293, anterior side of hind femur; 294, anterior side of fore tibia; 295, hind tibia with focus on outer-posterior row of stout setae; 296, the same with focus on outer-anterior and inner-anterior rows of stout setae (276–283, 293–296, holotype).

FIGURES 361–364. Thraulodes flavus sp. n. (holotype). 361, tergalii; 362–363, distal segment of maxillary palp in ventral and dorsal view; 364, hind femur in anterior view.

FIGURES 42–51. Thraulodes sinuosus, larval exuviae (42–47 at the same magnification): 42–44, parts of head with labrum and mandibles; 45, abdominal terga; 46, part of sternum and left pleura with half of metanotum and hind protopteron, left fore, middle and hind legs; 47, right half of pronotum and mesonotum; 48, tergalii V–VI; 49, anterior side of hind femur; 50, anterior side of fore tibia; 51, anterior side of hind tibia.

FIGURES 250–259. Thraulodes quevedoensis. 250–255, larval exuviae at the same magnification: 250–251, parts of head with labrum and mandibles; 252, right half of pronotum and mesonotum; 253–254, fore and hind legs; 255, abdominal terga; 256, anterior side of hind femur; 257, anterior side of fore tibia; 258, hind tibia with focus on outer-posterior row of stout setae; 259, the same with focus on outer-anterior and inner-anterior rows of stout setae.

FIGURES 317–336. Thraulodes zonalis. 317–331, larval exuviae at the same magnification; 332, young larva with light cuticle and well visible blackish hypodermal markings; 333, anterior side of hind femur; 334, anterior side of fore tibia; 335, hind tibia with focus on outer-posterior row of stout setae; 336, the same with focus on outer-anterior and inner-anterior rows of stout setae.

FIGURES 346–347. Thraulodes zonalis. 346, larval protogonostyli and protopenis in dorsal view with developing subimaginal gonostyli and penis; 347, optic section of through protopenis with developing penis. Abbreviations: gd, larval cuticle of gonoduct; tp, subimaginal telopenis.

FIGURES 666–672. Thraulodes? consortis, tergalii I–VII.

FIGURES 498–507. Thraulodes nigripes sp. n. 498, larval protogonostyli and protopenis; 499, subimaginal styliger and gonostyli extracted from larva; 500–501, subimaginal penis inside larval protopenis in ventral and dorsal view; 502, larval protogonostyli with subimaginal gonostyli and styliger at earlier stage of development; 503–506, tergalii I, III, VI and VII (498–501, 503–506, holotype).

FIGURES 532–540. Thraulodes nigrotibialis sp. n. 532, larval protogonostyli and protopenis; 533, subimaginal exuviae of genitalia; 534, genitalia of male imago in ventral view; 535, apex of penis on dorsal view; 536, hind wing; 537–540, tergalii I, III, VI and VII (532–536, holotype).

FIGURES 1–4. Median emargination of labrum: 1, Siphlonurus aestivalis Eaton 1903; 2, Leptophlebia marginata (Linnaeus 1767); 3, Habroleptoides pontica Kluge 1994c; 4, Habrophlebia fusca (Curtis 1834). Arrows show six sensilla, which on Fig. 4 separate five marginal denticles.

FIGURES 14–20. Thraulodes ludmilae sp. n., exuviae of mouthparts of holotype (14, 17–19 at the same magnification): 14, labrum; 15, its anterior margin with focus on dorsal side; 16, the same, with focus on ventral side; 17–18, left and right mandibles; 19, labium and maxilla; 20, dorsal side of glossa.Abbreviations: anter.r, limits of anterior transverse row of setae; poster. r, posterior transverse row of setae; other arrows show six sensilla separating five denticles.

FIGURES 510–521. Thraulodes nigrotibialis sp. n. 510 and 511–517, larval exuviae of two specimens at the same magnification: 510 and 511–512, parts of head with labrum and mandibles; 513, abdominal terga and sterna; 514–516, fore, middle and hind legs; 517, left half of pronotum and mesonotum; 518, young larva with brown cuticle and blackish hypodermal markings on head, thorax and abdomen; 519, anterior side of hind femur; 520, hind tibia with focus on outer-posterior row of stout setae; 521, the same with focus on outer-anterior and inner-anterior rows of stout setae (511–517, holotype).

FIGURES 626–636. Thraulodes sp. «Palo Seco», larva. 626, hind protopteron; 627–633, tergalii I–VII; 634–635, protogono- styli and protopenis at the same magnification; 636, protopenis enlarged.

FIGURES 106–111. Thraulodes telegraphicus. 106, protogonostyli of last instar larva (hypoderm of gonostyli and styliger and hidden cuticle of protopenis shown by interrupted lines); 107, enlarged cuticle of protopenis and hypoderm of developing penis inside; 108–109, the same, at later stage of development; 110–111, genital of male imago (110, styliger ventrally; 111, penis dorsally at left and ventrally at right). Abbreviation: bl, basal lobe; dp, dorsal projection of styliger; gd, gonoduct; m.gs, gonostylar muscle; m.med, median penial muscle; m.s, styligeral muscle; m-v.r, medio-ventral ridge (recurved flap); sp, seminal pump; tp, spear-like roll (telopenis).

FIGURES 101–105. Thraulodes telegraphicus. 101–102, genitalia of the same specimen; 103, penis of another specimen; 104– 105, part of penis with telopenis forming spear-like roll.Abbreviations: bl, basal lobe; dp, dorsal projection of styliger; gd, gonoduct; gs1, gs2, gs3, gs4, segments of gonostylus; m.gs, gonostylar muscle; m.lat, lateral penial muscle; m.med, median penial muscle; m.s, styligeral muscle; m-v.r, medio-ventral ridge (recurved flap); sp, seminal pump; tp, spear-like roll (telopenis).

FIGURES 371–376. Thraulodes flavus sp. n. 371, genitalia of male imago in lateral view; 372, apex of penis in dorsal view; 373, genitalia of male imago in ventral view; 374, their subimaginal exuviae; 375, larval protogonostyli and protopenis; 376, hind wing (372–376, holotype).

FIGURES 403–408. Thraulodes spangleri. 403–404, genitalia of male imago in lateral and ventral view; 405, apex of penis in dorsal view; 406, subimaginal exuviae of styliger, gonostylus and spear-like rolls; 407, larval protogonostyli and protopenis; 408, hind wing.

FIGURES 474–478. Thraulodes nigrabdominalis sp. n. 474, genitalia of male imago in ventral view; 475, their subimaginal exuviae; 476, hind wing; 477–478, protogonostyli and protopenis of presumably associated larva, with subimaginal gonostyli and styliger at early stage of development shown by interrupted line (474–476, holotype).

FIGURES 589–590. Thraulodes alboniger sp. n., eggs.

FIGURES 348–349. Thraulodes zonalis, eggs.

FIGURES 149–150. Thraulodes panamensis sp. n., egg.

FIGURES 248–249. Thraulodes marreroi, eggs.

FIGURES 274–275. Thraulodes quevedoensis, eggs.

FIGURES 315–316. Thraulodes fascipennis sp. n., eggs.

FIGURES 377–378. Thraulodes flavus sp. n., eggs.

Tables

TABLE 1. Inequality of larval legs in Thraulodes.

		Fore leg	Middle leg	Hind leg
	Size			longest
	Form	widest proximally	widest at middle	widest at middle
	Proximal blank	+	–	–
	Outer-anterior stripe of stout setae	+	+	+
Femur	Outer-posterior stripe of stout setae Outer stripe of thin hairs	+ long, dense	+ long, dense	+ shorter, sparser
	Row of recurved hairs	– (+)	+ (–)	+ (–)
	Stout setae on inner margin	+	+	+
	Stout setae on anterior surface	+	+	+
	Setae on posterior surface	– (+)	– (+)	+
	Outer-anterior row of stout setae	–	–	+
	Outer-posterior row of stout setae	–	–	+
Tibia	Inner-anterior row of stout setae Inner field of stout pointed setae	vestigial or absent dense, especially apically	sparse sparse	+ sparse
	Posterior pointed pectinate setae	–	–	+
	Outer hairs	+	+	+
	Inner-anterior row of recurved hairs	– (+)	+	+