Artines Godman, 1901

Medeiros, Adalberto Dantas De, Dolibaina, Diego Rodrigo, Carneiro, Eduardo, Mielke, Olaf Hermann Hendrik & Casagrande, Mirna Martins, 2019, Taxonomic revision of Artines Godman, 1901 (Hesperiidae: Hesperiinae: Moncini) with the description of nine new species, Zootaxa 4614 (1), pp. 1-49 : 4-5

Artines Godman, 1901

Figs 1–80

Artines Godman, 1901 , in Godman & Salvin. Biol. Centr.-Amer, Lep. Rhop. 2, p. 608; type species: Thracides aepitus .— Mabille, 1904, in Wytsman. Gen. Ins. 17, p. 118, 164.— Draudt, 1923, in Seitz. Gross-Schmett. Erde 5, p. 986.—Lindsey, 1925. Ann. Ent. Soc. Amer. 18: 79.— Hayward, 1934. Rev. Soc. ent. arg. 6: 101.— Hayward, 1939. An. Soc. cient. arg. 126: 457.— Hayward, 1941. Rev. Mus. La Plata, n. s., Zool. 2: 320.— Bell, 1946. Bol. Ent. venezol. 5: 177.— Hayward, 1947. Acta zool. Lill. 4: 363.— Hayward, 1950. Gen. Sp. Anim. Arg. 2, p. 306, pl. 2, fig. 19 (brand).— Evans, 1955. Cat. Amer. Hesp. 4, p. 85, 106.— Hemming, 1964. Annot. Lep. 5, p. 165.— Hemming, 1965. Bull. zool. Nomencl. 22: 74; type species.—ICNZ, 1967. Bull. zool. Nomencl. 24: 224; type species: Hesperia aquilina Plötz, 1883 . — Hemming, 1967. Bull. Brit. Mus. (Nat. Hist.), Ent., Suppl. 9: 61.— Cowan, 1968. Annot. Rhop., p. 7, 11.— Lamas, 1969. Biota 7: 338.— Beattie, 1976. Rhop. Direct., p. 7.— Bridges, 1983. Lep. Hesp. 2, p. 3.—Bridges, 1988. Cat. Fam.-Group & Gen.-Group Nam. 4, p. 13; 5, p. 1.—Bridges, 1988. Cat. Hesp. 2, p. 5; App. 2, p. 1.—Llorente-Bousquets; Luis-Martínez & Vargas-Fernández, 1990. Publ. esp. Mus. Zool., Mexico, 1: 20.—J. de la Maza E.; White & R. G. de la Maza E., 1991. Rev. Soc. Mex. Lep. 14: 29.—R. G. de la Maza E. & J. de la Maza E., 1993. Marip. Chiapas, p. 206.— Bridges, 1994. Cat. Fam.-Group, Gen.-Group, Sp.-Group Nam., Hesp. (Lep.) World 4, p. 3; 5, p. 1; 6, p. 7; 9, p. 6.— Warren, 2000, in Llorente; González & Papavero (eds). Biodiv., Tax., Biogeogr. Art. Mex. 2, p. 557.—O. Mielke, 2004. Hesperioidea, p. 62, in Lamas (ed.). Checklist: Part 4A, Hesperioidea-Papilionoidea, in Heppner (ed.). Atlas Neotrop. Lep. 5A; syn.: Itines.—O. Mielke, 2005. Cat. Amer. Hesperioidea 4, p. 832; syn.: Itines.—DeVries; Austin & Martin, 2008. Biol. Jour. Linn. Soc. 94: 727.—Warren; Ogawa & A. Brower, 2008. Cladistics 24: 27.—Warren; Ogawa & A. Brower, 2009. Syst. Ent. 34: 499, 517.— Cock, 2011. Living World 2011: 19.—Carneiro; Mielke & Casagrande, 2015. Zootaxa 3931(2), 199, 210-213.

Itines [sic]; Weeks, 1911. Ill. Diurn. Lep. 2, p. XV; Itines itizies [sic] [= atizies ].

Artines sp. J. Zikán, 1928. Ent. Rdsch., Stuttgart, 45: 35.— Lamas, 1994, in Foster et al. RAP Working Papers 6: 175.—C. Mielke, 1995. Revta bras. Zool. 11 (4): 7 63.— Robbins et al., 1996, in Wilson & Sandoval. Manu , p. 248.— Warren, 2000, in Llorente; González & Papavero (eds). Biodiv., Tax., Biogeogr. Art. Mex. 2, p. 557.— Lamas, 2003. Marip. Machu Picchu, p. 190.—Schmidt-Mumm; Schmidt & Salazar, 2003. Bol. Cient. Mus. Hist. Nat. U. de Caldas 7: 87.—O. Mielke; Emery & Pinheiro, 2008. Revta bras. Ent. 52 (2): 286.—Luis-M., Salinas-G. & Llorente-B., 2011, in Álvarez (coord.). Chiapas: estudios diversidad biol., Inst. Biol., UNAM, Mexico, p. 389.—O. Mielke et al., 2012. Revta bras. Ent. 56 (1): 63.—Vargas-Ch. & Salazar, 2014. Bol. Cient. Mus. Hist. Nat. U. de Caldas 18 (1): 277.— Paluch et al., 2016. Biota Neotrop. 16 (1): 5.

Type species. Hesperia aquilina Plötz, 1882 , by designation of ICZN (1967).

Taxonomic history. The genus Artines was described by Godman (1901), who designated Thracides aepitus Geyer, 1832 as the type-species, and provided figures of the venation of the forewing and the male genitalia of a specimen that he thought belonged to this species. However, based on the illustrations provided, this specimen corresponds to Hesperia aquilina Plötz, 1882 . Godman (1901) further provided the description of what he considered a new species, naming it Artines atizies , and illustrating the genitalia and the ventral surface of the wings. However, this species is a synonym of Thracides aepitus , as interpreted by Evans (1955). Thus, Godman (1901) had two species, H. aquilina , mistakenly identified by him as T. aepitus and T. aepitus , mistakenly interpreted by him as a new species. The type species of Artines was selected on the basis of an identification error, which led Hemming (1964, 1965) to suggest the alteration of its type species to Hesperia aquilina , an alteration later formalized by the ICZN (1967).

In addition to Godman (1901), the authors who provided descriptions of species in Artines were Mabille (1904), Draudt (1923), Evans (1955), Mielke (1968, 1992), Steinhauser & Austin (1993) and Dolibaina et al. (2015). The others provided citations in catalogs and taxonomic comments.

Diagnosis. Artines differs from the other genera of Moncini by the third segment of the labial palpus long, erect and pointed; antenna longer than 1/2 length of costa; nudum with 11 to 12 segments, only on apiculus; male with two androconial organs: a sagitated brand on base of CuA 1 –CuA 2 of forewing (absent in the “ aepitus group”) ( Fig. 3 ) and a tuft of elongated scales along 2A on dorsal hind wing ( Fig. 4 ); ventral forewing with a violet sheen at apical and costal areas; ventral hind wing with a central black spot at distal end of the discal cell (absent only in A. bipunctata O. Mielke, 1968 ), distally surrounded by six blue to violet discal spots between Rs–CuA 2, which are externally limited by an arched postdiscal band, from lower 1/2 of Sc+R 1 –Rs to upper 1/2 of CuA 2 –2A; aedeagus with a ventral projection; opening of ejaculatory bulb at the near-dorsal proximal margin; opening of the aedeagus distal; cornuti present; lamella postvaginalis bifid; ostium at the central portion of the sterigma, ventrally projected by folds of the lamellae ante and postvaginalis; ductus bursae long and narrow, first folded in spiral and distally arched to the left.

Description. Male. Head: dorsally dark brown, with some white and elongated scales at center of vertex, on transfrontal and fronto-clypeal sutures and on the paraocular area; gena white. Labial palpus brown, densely speckled with white scales, darker at apex; first segment short; second segment twice as long as first; third segment about 2/3 of second, erect and pointed. Antenna 1/2 length of costa, brown dorsally, white ventrally at base of antennomeres and club, cream at apiculus; club 2/3 of the shaft lengh; nudum with 11 to 12 segments, only on apiculus.

Thorax: dorsally dark brown; ventrally grayish. Legs externally brown, internally white, somewhat cream on tibiae and tarsi; foretibia with epiphysis; midtibia with spines and a pair of apical spurs; hind tibia with a pair of median and apical spurs.

Forewing length: 12–15.5 mm. Forewing triangular; anterior margin slightly convex; apex obtuse and rounded; outer margin convex, more projected at the end of M 2 and M 3; tornus rounded; anal margin straight. Hindwing rounded; anterior margin convex; apex rounded; outer margin convex, slightly concave from CuA 2 to near tornus; tornus rounded; anal margin straight.

Dorsal forewing: ground color dark brown with yellow scales scattered on costal area and proximal 1/3 of CuA 2 –2A; two groups of white spots: one apical, consisting of three opaque and minute spots in R 3 –R 4, R 4 –R 5 and R 5 –M 1, another discal, consisting of two larger, opaque to semi-hyaline spots, one smaller at the end of the proximal 1/3 of M 3 –CuA 1, the other below dci in CuA 1 –CuA 2; there may be a small, diffuse yellowish spot in CuA 2 –2A; brand, when present, sagitated, located at the base of CuA 1 –CuA 2 and with two distal projections, the upper one longer than the lower one ( Figs 3 , 21–26 ); fringe light tan, darker at end of veins.

Dorsal hind wing: ground color as on the dorsal forewing; tufts of elongated scales along 2A ( Fig. 4 ); fringes white to light brown.

Ventral forewing: ground color as on dorsal surface, paler on anal area, sometimes with large, diffuse white spot on distal 1/2 of CuA 2 –2A; costal area violet with gray to yellow scattered scales; three to four bluish white apical spots in R 2 –R 3, R 3 –R 4, R 4 –R 5 and R 5 –M 1, first one displaced to base; discal spots as on dorsal surface; apical area with two violet bands separated by a ground color stripe, one submarginal between R 4 –CuA 2 (less evident in CuA 1 –CuA 2) and another apical between R 3 –CuA 2; marginal line dark brown: fringes as on dorsal surface.

Ventral hind wing: ground color as on ventral forewing, appearing violet and with grayish scales scattered on basal 1/2, between costal margin and upper 1/2 of CuA 2 –2A, and between 3A and anal margin; paler area between lower 1/2 of CuA 2 –2A and 3A; apical portion of discal cell dark brown with yellow to orange scales; discal area with central black spot separated from discal cell by bluish band on dcm and dci (absent in A. bipunctata ); six blue to violet discal spots between Sc+R 1 –CuA 2 surrounded by a ground color border, the spot in Sc+R 1 –Rs shifted to base; postdiscal band yellow or gray from lower 1/2 of Sc+R 1 –Rs to the upper 1/2 of CuA 2 –2A, limited distally by a stripe with the same color as ground, and externally to this, a violet submarginal band between Rs and anterior 1/3 of CuA 2 –2A; marginal line as on forewing; fringes as on dorsal surface.

Abdomen: dorsally dark brown, white to grayish ventrally, with median dark brown stripe.

Genitalia: tegumen more or less quadrate to rectangular; ventral projection of tegumen fused with dorsal projection of saccus, assuming that the boundaries between these structures are at the angle between them. Fenestra absent. Saccus ranging from shorter to longer than tegumen+uncus. Uncus more or less quadrate, rectangular or tapered, simple or bifid. Gnathos divided, not exceeding uncus and with a central membranous area in lateral view. Valva of variable shape; costa narrow, continuous with or separated from ampulla; harpe simple or bifid, dorsally fused or separated from ampulla. Aedeagus with a ventral projection; opening of ejaculatory bulb at dorso-proximal end, sometimes separated from the anterior margin by a short distance; opening of aedeagus distal; cornuti as numerous spines, either over sclerotized plate or directly over membrane (sclerotized plate absent in A. bipunctata ). Fultura inferior in dorsal view formed by a pair of stems, one on each side of aedeagus, connected by membranous area ( Figs 27 K–35K), in lateral view with anterior end simple or bifid.

Female. Forewing length: 13.5–14 mm. Similar to male, differing in the absence of androconial organs ( Figs 3–4 ).

Genitalia: tergum VIII with complete spiracular opening (except in A. panama Medeiros & Carneiro sp. nov.). Region between seventh sternite and lamella antevaginalis membranous or sclerotized. Sterigma completely or partially sclerotized; lamella antevaginalis developed, with or without proximo-ventral folds; lamella postvaginalis bifid or with disto-lateral projections, lateral folds represented by a pair of flaps, projected ventral and medially ( Fig. 61 A–B). Bursa copulatrix globular; ostium at central portion of sterigma, ventrally projected by folds of the lamellae ante and postvaginalis, surrounded by membranous or sclerotized area; ductus bursae long, membranous or with basal portion sclerotized, first folded in spiral and distally arched to the left, signa represented by two lateral stripes of small spines along the ductus bursae. Papilla analis rectangular, wider than long.

Figures

FIGURES 1–2. Color pattern of ventral wings of Artines: 1. Artines bamba sp. nov.; 2. Artines aquilina.

FIGURES 3–4. Androconial organs present in the species of Artines. 3. Brand (with scales) on dorsal forewing of Artines aquilina (absent in “aepitus group”). 4. Tuft of elongated scales along 2A of the dorsal hind wing of Artines liege sp. nov. (presente in all species of Artines).

FIGURES 5–8. Species of the “aepitus group” in dorsal and ventral views. 5. Artines aepitus, A–B: male, Cueva de Las Pavas, Tingo Maria, Huánuco, Peru (OM 80.822), C–D: female, Ilha de Maracá, Alto Alegre, Roraima, Brazil (DZ 41.790). 6. Artines panama sp. nov., A–B: holotype male, C–D: allotype female. 7. Artines solange sp. nov., A–B: holotype male, C–D: tentative female, Sumaúma, Maranhão, Brazil (DZ 41.800). 8. Artines delfos sp. nov., A–B: holotype male. Scale bar = 1 cm.

FIGURES 9–13. Species of the “aepitus group” in dorsal and ventral views. 9. Artines maya, A–B: holotype male. 10. Artines litoralis sp. nov., A–B: holotype male, C–D: allotype female. 11. Artines rica, A–B: male, Turrialba, Cartago, Costa Rica (DZ 41.890), C–D: female from same locality of male (DZ 41.840). 12. Artines liege sp. nov., A–B: holotype male, C–D: allotype female. 13. Artines bamba sp. nov., A–B: holotype male, C–D: tentative female, Moyobamba, San Martin, Peru (OM 79.413). Scale bar = 1cm.

FIGURES 14–19. Species of the “aquilina group” (14–18) and “bipunctata group” (19) in dorsal and ventral views. 14. Artines aquilina, A–B: male, Joinville, Santa Catarina, Brazil (DZ 41.321), C–D: female from same locality of male (DZ 41.220). 15. Artines angelica sp. nov., A–B: holotype male, C–D: allotype female. 16. Artines focus, A–B: male, Ilha de Maracá, Alto Alegre, Roraima, Brazil (DZ 41.233), C–D: female, Pacaraima, Roraima, Brazil (DZ 41.333). 17. Artines cofus sp. nov., A–B: holotype male, C–D: allotype female. 18. Artines donia sp. nov., A–B: holotype male. 19. Artines bipunctata, A–B: male, Jardim Zoológico, Brasília, Distrito Federal, Brazil (DZ 30.961), C–D: female, Córrego Bananal, Brasília, Distrito Federal, Brazil (DZ 41.224). Scale bar = 1cm.

FIGURE 20. Variation in the color pattern of the ventral forewing of Artines aquilina. A: Joinville, Santa Catarina, Brazil; B: Lençóis, Bahia, Brazil and C: Caranavi, La Paz, Bolivia. Scale bar = 1 cm.

FIGURES 21–26. Male brands (no scales) on dorsal forewing of the “aquilina group” (21–25) and “bipunctata group” species (26). 21. Artines aquilina, Joinville, Santa catarina, Brazil, (DZ 41.341). 22. Artines angelica sp. nov., paratype, Fazenda Ter- rasse, Santa Luzia, Maranhão, Brazil (DZ 41.273) 23. Artines focus, Ilha de Maracá, Roraima, Brazil (DZ 41.233). 24. Artines cofus sp. nov., paratype, Santa Rosa do Purus, Acre, Brazil (DZ 41.303). 25. Artines donia sp. nov., paratype, Santa Rita do Araguaia, Goiás, Brazil (OM 66.000). 26. Artines bipunctata, Ponte Funda, Vianópilis, Goiás, Brazil (DZ 41.264). Scale bar = 1 mm.

FIGURES 27–29. Male genitalia of the “aepitus group” species. 27. Artines aepitus, Jusepín, Monagas, Venezuela (OM 29.417). 28. Artines panama sp. nov., holotype. 29. Artines solange sp. nov., holotype. A) lateral view of tegumen, saccus, uncus and gnathos. B) ventral view of saccus. C) dorsal view of tegumen, uncus and gnathos. D) ventral view of tegumen, uncus and gnathos. E) interior view of right valva. F) dorsal view of right valva. G) dorsal view of aedeagus and vesica. H) left lateral view of aedeagus. I) ventral view of aedeagus. J) right lateral view of aedeagus. K) dorsal view of fultura inferior. L) lateral view of fultura inferior. Scale bar = 1 mm.

FIGURES 30–32. Male genitalia of the “aepitus group” species. 30. Artines delfos sp. nov., holotype. 31. Artines maya, holotype. 32. Artines litoralis sp. nov., paratype, Linhares, Espírito Santo, Brazil (DZ 41.215). For reference of the letters see legend of the figure 27. Scale bar = 1 mm.

FIGURES 33–35. Male genitalia of the “aepitus group” species. 33. Artines rica, Turrialba, Cartago, Costa Rica (DZ 4.061). 34. Artines liege sp. nov., holotype. 35. Artines bamba sp. nov., paratype, Pakitza, Parque Nacional del Manu, Madre de Díos, Peru (DZ 9.728). For reference of the letters see legend of the figure 27. Scale bar = 1 mm.

FIGURES 36–37. Male genitalia of the “aquilina group” species. 36. Artines aquilina, Caminho da Lagoinha, Cachoeiras de Macacú, Rio de Janeiro, Brazil (DZ 41.196). 37. Artines angelica sp. nov., holotype. M) dorsal view of vesica showing the cornuti. For reference of the letters see legend of the figure 27. Scale bar = 1 mm.

FIGURES 38–39. Male genitalia of the “aquilina group” species. 38. Artines focus, Reserva Duque, Manaus, Amazonas, Brazil (DZ 41.192). 39. Artines cofus sp. nov., paratype, Tambopata, Puerto Maldonado Madre de Díos, Peru (MUSM). For reference of the letters see legend of the figures 27 and 36. Scale bar = 1 mm.

FIGURES 40–41. Male genitalia of the “aquilina group” (40) and “bipunctata group” (41) species. 40. Artines donia sp. nov., paratype, Santa Rita do Araguaia, Goiás, Brazil (DZ 66.000). 41. Artines bipunctata, Brasília, Distrito Federal, Brazil (DZ 41.194). For reference of the letters see legend of the figures 27 and 36. Scale bar = 1 mm.

FIGURES 42–48. Dorsal view of vesica and cornutus of the "aepitus group" species. 42. Artines aepitus. 43. Artines panama sp. nov.. 44. Artines delfos sp. nov.. 45. Artines litoralis sp. nov.. 46. Artines rica. 47. Artines liege sp. nov. 48. Artines bamba sp. nov. Scale bar = 0,2 mm.

FIGURES 49–53. Ventral view of vesica and cornutus of the “aquilina group” species. 49. Artines aquilina. 50. Artines angelica sp. nov.. 51. Artines focus. 52. Artines cofus sp. nov.. 53. Artines donia sp. nov.. Scale bar = 0,2 mm.

FIGURES 54–60. Female genitalia of the “aepitus group” species. 54. Artines aepitus, Jusepín, Monagas, Venezuela (OM 30. 125). 55. Artines panama sp. nov., allotype. 56. Artines solange sp. nov. (tentative female), Sumaúma, Maranhão, Brazil (DZ 41.800). 57. Artines litoralis sp. nov., paratype, Linhares, Espírito Santo, Brazil (DZ 41.820). 58. Artines rica, Turrialba, Cartago, Costa Rica (DZ 41.840). 59. Artines liege sp. nov., paratype, Parque do Gama, Brasília, Distrito Federal, Brazil (DZ 41.830). 60. Artines bamba sp. nov. (tentative female), Moyobamba, San Martin, Peru (OM 81.229). A) lateral view of tergum VIII, sterigma and papilla analis; B) ventral view of esterigma. Scale bar = 1 mm.

FIGURES 61–65. Female genitalia of the “aquilina group” and “bipunctata group” species. 61. Artines aquilina, Cananéia, São Paulo, Brazil (OM 49.831). 62. Artines angelica sp. nov., paratype, Tambopata, Puerto Maldonado, Madre de Díos, Peru (DZ 41.222) 63. Artines focus, Pacaraima, Roraima, Brazil (DZ 41.333). 64. Artines cofus sp. nov. paratype, Tambopata, Puerto Maldonado, Madre de Díos, Peru (DZ 41. 293). 65. Artines bipunctata, Reserva IBGE, Brasília, Distrito Federal, Brazil (DZ 41.165). For reference of the letters see legend of the figure 54. Scale bar = 1 mm.

FIGURES 66–72. Female genitalia of the “aepitus group” species (ventral view of sterigma and bursa copulatrix). 66. Artines aepitus, Jusepín, Monagas, Venezuela (OM 30. 125). 67. Artines panama sp. nov., allotype. 68. Artines solange sp. nov. (tentative female), Sumaúma, Maranhão, Brazil (DZ 41.800). 69. Artines litoralis sp. nov., paratype, Linhares, Espírito Santo, Brazil (DZ 41.820). 70. Artines rica, Turrialba, Cartago, Costa Rica (DZ 41.840). 71. Artines liege sp. nov., paratype, Parque do Gama, Brasília, Distrito Federal, Brazil (DZ 41.830). 72. Artines bamba sp. nov. (tentative female), Moyobamba, San Martin, Peru (OM 81.229). Scale bar = 1 mm.

FIGURES 73–77. Female genitalia of the “aquilina group” and “bipunctata group” species (ventral view of sterigma and bursa copulatrix). 73. Artines aquilina, Cananéia, São Paulo, Brazil (OM 49.831). 74. Artines angelica sp. nov., paratype, Tambopata, Puerto Maldonado, Madre de Díos, Peru (DZ 41.222). 75. Artines focus, Pacaraima, Roraima, Brazil (DZ 41.333). 76. Artines cofus sp. nov. paratype, Tambopata, Puerto Maldonado, Madre de Díos, Peru (DZ 41. 293). 77. Artines bipunctata, Reserva IBGE, Brasília, Distrito Federal, Brazil (DZ 41.165). Scale bar = 1 mm.

FIGURE 78. Geographic distribution of the “aepitus group” species. The symbols with white inferior half represent records from literature.

FIGURE 79. Geographic distribution of the “aepitus group” species. The symbols with white inferior half represent records from literature.

FIGURE 80. Geographic distribution of the “aquilina group” and “bipunctata group” species. The symbols with white inferior half represent records from literature.