Alesa prema (Godart, [1824])

Santos, Wildio Ikaro Da Graça, Dolibaina, Diego Rodrigo, Dias, Fernando Maia Silva, Mielke, Olaf Hermann Hendrik & Casagrande, Mirna Martins, 2023, A review of the South American metalmark genus Alesa Doubleday, 1847 (Lepidoptera: Riodinidae: Eurybiini) with the description of two new species, Zootaxa 5284 (1), pp. 77-120 : 81-84

Alesa prema (Godart, [1824])

Figs 1–2 , 7 , 12–13 , 61 , 71 , 77 , 82–83 , 92

Erycina prema Godart, [1824] , in Latreille & Godart. Enc. Méth. 9, p. 555, 569; one specimen [holotype], without locality.

Alesa prema ; Doubleday, 1847. List. Lep. Brit. Mus. 2, p. 1.—Westwood, 1851, in Doubleday, 1851. Gen. diurn. Lep. 2, p. 418, pl. 70, fig. 8 (male d).—Bates, 1868. Journ. Linn. Soc. London, Zool., 9 (39): 414; syn.: smaragdifera .—Herrich-Schäffer, 1868. Corr.-Blatt zool.-min. Ver. Regensburg 22: 121.—Kirby, 1871. Syn. Cat. diurn. Lep., p. 287.—Kirby, 1879. Cat. Coll. diurn. Lep. Hewiston, p. 186.—Glaser, 1887. Cat. etymol. Coleopt. Lepid., p. 274.—Staudinger, 1888, in Staudinger & Schatz. Exot. Schmett. 1, p. 242; 2, pl. 88 (male, female d, v).—Mengel, 1905. Cat. Erycinidae , p. 54.— Stichel, 1910, in Wytsman. Gen. Ins. 112A, p. 80, pl. 4, figs 12a, c (palpus), d (leg), g (male gen.).— Seitz, 1916. Gross-Schmett. Erde 5, p. 652, pl. 127 (male, female d).—Stichel, 1925. Ztschr. wiss. Insektenbiol. 20: 21.—Michael, 1928. Erinn. S.- Amer, p. 50, 88.— Stichel, 1930. Lep. Cat. 40, p. 293; syn.: smaragdifera , f. nigra.— Biedermann, 1936. Bull. Soc. ent. France 41 (15): 252, pl. 3, figs 3, 4, 7, 8 (male, female d, v).—Cardoso, 1949. Rev. Entom. 20(1/3): 430.—K. Brown & Mielke, 1967. Jour. Lep. Soc. 21 (3): 146.—Barcant, 1970 Butt. Trinidad and Tobago, p. 226, pl. 9, fig. 16 (male d).—Callaghan, 1983. Jour. Res. Lep. 21 (3): 164, fig. 2.—Brown, 1993. Occas. Paper IUCN Sp. Surv. Comm. 8: 48.—Brown, 1993. Occas. Paper IUCN Sp. Surv. Comm. 8: 151.— d’Abrera, 1994. Butt. Neotrop. Region 6, Riodinidae , p. 933, figs (male, female d).—Lamas, 1994. RAP Work. Pap. 6: 84, 181.— Robbins et al., 1996, in Wilson & Sandoval. Manu, p. 235.— Brévignon, 1997. Lambillionea 97 (3)(1): 331, 334.—O. Mielke & Casagrande, 1998. Revta bras. Zool. 14(4): 998.—Brown & Freitas, 2000. Bol. Mus. Biol. Melo Leitão 11/12: 86.—Andrade, 2002. Biodiversity de las mariposas ( Lepidoptera : Rhopalocera) de Colombia, in Costa et al. (eds.) Proyecto de Red Iberoamericana de Biogeografía y Entomología Sistemática 2: 170.— Hall, 2003. Syst. Ent. 28: 24, 26, fig. 1b (male d).— Callaghan & Lamas, 2004, Riodinidae , p. 148, in Lamas (ed.) Checklist: Part 4A. Hesperioidea— Papilionoidea, in Heppner (ed.) Atlas of Neotropical Lepidoptera 5A; syn.: smaragdifera , ab. nigra.—O. Mielke & Casagrande, 2004. Borboletas, p. 727, 738 figs (male, female d), in Mikich & Bérnils (eds). Livro verm. Fauna ameaç. Est. Paraná; syn.: smaragdifera , ab. nigra, f. sapphirina.—Emery et al., 2006. Rev. Bras. Ent. 50 (11): 88.—Pinheiro & Emery, 2006. Biota Neotrop. 6 (3): 10.— Salazar & Constantino, 2007. Bol. Cient. Mus. Hist. Nat. U. Caldas 11: 168, 169, figs 35, 36 (male, female d).— Nuñes, 2008. Trop. Lep. Res. 18 (2): 81, 84.—Nuñes, 2009. Trop. Lep. Res. 19 (2): 74.— Hall & Ahrenholz, 2010. Trop. Lep. Res. 20 (1): 21.— Dolibaina et al., 2010. Biota Neotrop. 10 (3): 75, 76, 78, 79, figs 16, 17 (female d, v).—Salazar et al., 2010. Bol. Cient. Mus. Hist. Nat. U. Caldas 14(1): 171.—Núñez-Bustos et al. 2011. Trop. Lep. Res. 21 (1): 40.—K. Brown, Schoultz, A. O. Saura & A. Saura, 2012. Hereditas 149: 130.—Nielsen & Salazar, 2014. Bol. Cient. Mus. Hist. Nat. U. Caldas 18 (2), p. 218.—Salik et al., 2014. SHILAP Revta. lepid. 42 (166): 268.—Avigliano & Schenone, 2015. Entomotropica 30(9): 89.— Gallard & Fernandez, 2015. Bull. Soc. ent. France 120 (2): 135, 136, 137, 138, figs 1, 2 (male d, v).—Gonzalo-Andrade et al., 2015. Revta. Colomb. Amaz. 8: 107.— Gallard, 2017. Riodinidae Guyane, p. 56, pl 10, fig. 2K (male d).—Salazar et al., 2017. Bol. Cient. Mus. Hist. Nat. U. Caldas 21 (2), p. 159.—Klimaitis et al., 2018. Mariposas Argentinas, p.106, fig. 280 [1–4] (male and female, d,v).—Dickens et al., 2019. J. Insect Cons. 23: 715, 720, 725.

Alesa smaragdifera Westwood, 1851 , in Doubleday. Gen. diurn. Lep. 2, p. 418; Columbia.

Alesa prema ab. nigra Kaye, 1925. Trans. ent. Soc. London 1924(3/4): 415; Trinidad, St. Ann’s , 1000 ft., R. Dick [leg.].— Kaye, 1940. Trans. ent. Soc. London 90: 558 .

Diagnosis. Forewing length, males 18–24 mm, n=52; females 19–23mm, n=12. Forewing and hindwing compact, with pointed forewing apex and hindwing tornus; body and bands of the wings upper side dark blue, wings underside ground color light brown; uncus with posterior margin not lobed; anterior and posterior projections of saccus narrow and long, respectively; valva oval; outer dorsal process of valva short, never exceeding transtilla distally and parallel to inner ventral process of valva; transtilla central process less sclerotized and densely setose; aedeagus longer and narrower than in other species of the group. Females of A. prema are distinguished from those of other members of “ prema group” by the wings upper side ground color brown and bands dark brown; hindwing underside with a row of submarginal ocelli often with metallic blue scales at center; papilla analis triangular; bursa copulatrix twice longer than in A. juliae sp. nov.; corpus bursae with a pair of signa.

Alesa prema is most similar to A. esmeralda and A. beneluzi . Males of A. prema differ from A. esmeralda by the postdiscal and submarginal bands not merged, from A. beneluzi by the blue color of the upper side of the wings, the brown lines on the wings underside, and by the more pointed wings; from A. rothschildi and A. juliae sp. nov. by the base of wings underside without yellow scales, and the absence of gray color.

Type material. Erycina prema Godart, [1824] was described based on a holotype male from an unknown locality. Based on other species described by Godart, the type specimen of E. prema is probably deposited at the MNHN and the type location probably is Rio de Janeiro, Brazil. The type specimen is probably lost, as Callaghan (1995), who was specifically looking for type specimens of Riodinidae at the MNHN, did not find any specimen which could correspond to the holotype at the referred collection. To avoid future uncertainty, a male specimen of Erycina prema Godart [1824] is here designated neotype, with the following labels: / NEOTYPUS / Petrópolis, Independência 900 m [Rio de Janeiro state], 12-IV-1940 / Ex. Col. Gagarin / DZ 34.956 / Neotypus Erycina prema Godart, [1824] W. Santos, Dolibaina, Dias, O. Mielke & Casagrande des. 2021 /. DZUP.

Alesa smaragdifera Westwood, 1851 was described based on an unstated number of females from Colombia. A female labeled as holotype, deposited at the NHMUK, is here recognized as a syntype and designated lectotype, to avoid future uncertainty, with the following labels: / Type H. T. / Venezuela [verso of the same label:] 479 / 303 / BMNH (E) 1717344 /, and the following labels will be added: / LECTOTYPUS / Lectotypus Alesa smaragdifera Westwood, 1851 W. Santos, Dolibaina, Dias, O. Mielke & Casagrande des. 2021 /.

Alesa prema ab. nigra was described by Kaye (1925) as a melanic form of the typical phenotype of A. prema from Trinidad and Tobago. The name was originally described as an “aberration”, denoting an infrasubspecific entity and, therefore, invalid and without a formal type ( ICZN 45.6.2) .

Distribution. Alesa prema is widespread in several biomes throughout cis-Andean tropical South America, from Trinidad and Tobago to northern Argentina and southern Brazil ( Fig. 92 ). This species probably occurs in Bolivia, Paraguay, Surinam and Guyana.

Comments. Alesa prema is the type species and the most remarkable species of the genus due to the magnificent blue metallic color of the male and the wide distribution that extends beyond the Amazon where its congeners are confined to. Alesa prema is most similar and probably the most closely related to A. esmeralda . The only unambiguous character that can be used to distinguish these two species is the pattern of bands of the male forewing upper side: not merging together in A. prema , and merged together in A. esmeralda . Based on the current knowledge, these species do not occur in sympatry, even though their ranges overlap.

There is intraspecific variation in bands of the wings upper side of A. prema , especially in males ( Biedermann 1936) ( Fig. 7 ). This variation can be observed even within a single population. A considerable representation of the species variation can be observed in a series of forty males sampled on a same hilltop in western Amazon (Parque Nacional da Serra do Divisor, Mâncio Lima, Acre state) ( Fig. 7 ). The variation includes the color of the bands of the wings upper side, from emerald green to different shades of blue, and the length of the band, mostly the discal and postdiscal bands, which may decrease or increase the area of the ground color between them. Despite the wing pattern variation, no genitalic differences were found.

The wing pattern of the female of A. prema differs greatly from any other known female in the group, having a mostly brown pattern with submarginal ocelli on both sides of the hindwing, resembling some species of satyrs ( Nymphalidae ) and skippers ( Hesperiidae ).

Adults of A. prema are encountered visiting flowers (Brown & Mielke 1968), and, more frequently, in sunny forest gaps on hilltops, where males fly fast, between 2–4 meters from the ground, perching on leaves with open wings ( Gallard & Fernandez 2015; DRD, pers. obs.). Although uncommon, females also can be encountered in hilltops ( Dolibaina et al. 2010). This species is active during early to mid-morning ( Gallard & Fernandez 2015; DRD, pers. obs.).

Examined material. TRINIDAD AND TOBAGO — Holdo, XI.1969, 1 male, R. Dick leg., no data, 1 male and 1 female, R. Dick leg. (MGCL). Sangre Grande: Sans Souci, XI.1969, 1 male, C. Urich leg., Laloja South road, mile 2.4, 7.III.1982, 1 male. F. Preston leg. (MGCL). EQUADOR — Napo: Puerto Misahuallí (8Km W), 450 m, VII.1988, 1 female, J. P. DeVries leg., (USNM). PERU — Moyatta, 14.VI.1948, 1 male, Klug leg. (MGCL). BRAZIL — Amazonas: São Paulo de Olivença, 19.III.1951, 1 male, DZ 34.896 (DZUP). Acre: Mâncio Lima (Parque Nacional da Serra do Divisor, Porç„o Norte, Rio Moa), 200–400 m, 10–21.IX.2011, 5 males, D. Dolibaina & D. Moura leg., DZ 34.894, DZ 34.864, DZ 34.886, DZ 34.856, DZ 35.044; 20–27.VI.2013, 14 males, Mielke, Casagrande, Carneiro, Dias, Dolibaina leg., DZ 35.054, DZ 35.014, DZ 34.904, DZ 34.994, DZ 35.024, DZ 34.824, DZ 34.796, DZ 34.814, DZ 34.786, DZ 35.004, DZ 34.844, DZ 34.846, DZ 35.074, DZ 34.806 (DZUP), 23– 30.VIII.2014, 12 males, Mielke, Casagrande, Carneiro, Dias, Dolibaina, Siewert & Salik leg., DZ 34.964, DZ 34.794, DZ 34.934, DZ 34.874, DZ 34.884, DZ 35.034, DZ 34.834, DZ 35.064, DZ 34.804, DZ 34.914, DZ 34.854, DZ 34.784 (DZUP), 7–10,13. X.2018, 8 males, O. Mielke, Casagrande, Dolibaina & Medeiros leg., DZ 48.037, DZ 48.137, DZ 48.147, DZ 48.097, DZ 48.167, DZ 48.217, DZ 48.067, DZ 48.087 (DZUP). Bujari (50 Km NW), 200 m, 18–20.IX.2003, 1 female, Mielke & Casagrande leg., DZ 35.264 (DZUP). Rondônia: Cacaulândia, 21.X.1993, 1 male, 8.X.1995, 2 males, 16.IV.1997, 1 male, M. O. Gomes leg., 1.IX.1991, 1 female, 17.XI.1991, 1 female, 25.XII.1991, 1 female, T. Schmitz leg. (MGCL). Mato Grosso: Barra do Garça, 16.I.1977, 1 female, DZ 35.214, 19.IV.1978, 2 females, no collector, DZ 35.254, DZ 35.234 (DZUP). Bonito, 10.XII.2016, 1 male, no collector, DZ 48.277, ex. coll. E. Emery (DZUP). Goiás: Pirenópolis, 1200 m, 2.V.1991, 2 males, no collector (USNM). Distrito Federal: Brasília (Mata Rio Maranh„o), 4.IX.2002, 1 male, E. Emery leg., DZ 48.127, ex. coll. E. Emery (DZUP). Planaltina, 1000 m, 1.V.1991, 1 male, no collector (USNM). Maranhão: Riachão (Poço Azul), 12.IV.2011, 1 female, Mielke & Casagrande leg., DZ 34.941 (DZUP). Alagoas: Maceió, 6.II.1949, 1 female, II.1964, 1 male, A. Cardoso leg., DZ 35.244, DZ 34.816 (DZUP). Bahia: Rio de Contas (Pico das Almas), 28.I.2005, 1 female, Mielke & Casagrande leg., DZ 35.224 (DZUP). Morro do Chapéu, 25.IV.1991, 1 male, Robbins & Becker leg. (USNM). Minas Gerais: Belo Horizonte, 15–16.IV.1973, 3 males, 28.IV.1975, 1 male, 27.XII.1975, 1 male, Callaghan leg. (USNM). Paracatu, 16.V.1972, 2 males, Mielke & Brown leg., DZ 34.926, DZ 34.866 (DZUP); (BR 40, Km 490), 13.V.1969, 1 male, Nicolay leg., (USNM). Espírito Santo: Conceição da Barra, 15.VIII.1968, 1 female, XII.1971, 1 male, C. & C. T. Elias leg., DZ 34.954, DZ 34.946 (DZUP). Linhares, VIII.1975, 1 female, C. & C. T. Elias leg., DZ 34.944 (DZUP). Rio de Janeiro: Guapimirim, 16.X.1940, 1 male, no collector, DZ 34.826 (DZUP). Petrópolis (Independência), 900 m, 5.IV.1931, 1 female, 12.IV.1940, 2 males, no collector, ex. col. Gagarin, DZ 34.924, DZ 34.956, DZ 34.936 (DZUP). São Paulo: Teodoro Sampaio (Parque Estadual do Morro do Diabo), 17–19.VIII.1989, 1 male, 5–8.XII.1989, 20–23.II.1990, 1 female, 2 males, 15–18.III.1991, 1 male, Mielke & Casagrande leg., DZ 34.836, DZ 34.906, DZ 34.816, DZ 34.876, DZ 34.974 (DZUP), 16.III.1991, 2 males, R. Robbins leg. (USNM). Paraná: Terra Rica (Parque Municipal dos Três Morrinhos) , 17.XII.2009, 1 female, Carneiro & Dolibaina leg., DZ 34.984 (DZUP).

Figures

FIGURES 7. Variation of wings upper side of males of Alesa prema. A, Brasil, Rio de Janeiro: Guapimirim (DZ 34.826). B, Brazil, Acre: Mâncio Lima, Parque Nacional da Serra do Divisor (DZ 35.034). C, Brazil, Amazonas: S„o Paulo de Olivença (DZ 34.896). D–G, Brazil, Acre: Mâncio Lima, Parque Nacional da Serra do Divisor (DZ 34.804, DZ 35.024, DZ 35.074, DZ 34.814). H, Brazil, Distrito Federal: Brasília, Rio Maranh„o (DZ 48.127).

FIGURES 92. Geographic distribution of Alesa prema and A. esmeralda.

FIGURES 1–6. Males of species of “prema group”, part I, in dorsal and ventral views (left to right). 1–2, A. prema Brazil, Acre: Mâncio Lima, Parque Nacional Serra do Divisor (DZ 34.804). 3–4, A. esmeralda Colômbia, Cauca: river Villalobos (holotype, CJS). 5–6, A. beneluzi French Guiana, Saint-Laurent-du-Maroni: route d’Apatou (holotype, MNHN). Scale bar: 1cm.

FIGURES 12–17. Females of species of “prema group”, in dorsal and ventral views (left to right). 12–13, A. prema Brazil, Maranh„o: Riach„o (DZ 34.941). 14–15, A. rothschildi (SF). 16–17, A. juliae sp. nov. Brazil, Mato Grosso, Diamantino, rio Arinos (USNM). Scale bar: 1cm.

FIGURES 61–63. Male genitalia of species of “prema group”. 61, A. prema Brazil, Acre: Mâncio Lima, Parque Nacional da Serra do Divisor (DZ 34.884). 62, A. rothschildi Brazil, Pará: Óbidos (OM 67.441). 63, A. juliae sp. nov. Brazil, Acre: Mâncio Lima, Parque Nacional da Serra do Divisor (DZ 35.184). Left to right: dorsal view of tegumen and uncus, lateral and ventral views of genitalia, and dorsal view of valvae and transtilla. Scale bar: 1mm.

FIGURES 71–76. Female genitalia of Alesa species. 71, A. prema Brazil, Mato Grosso: Barra do Garça (DZ 35.254). 72, A. juliae sp. nov. Brazil, Mato Grosso: Rio Arinos (DZ 35.144). 73, A. telephae Brazil, Acre: Mâncio Lima (DZ 35.094). 74, A. amesis Brazil, Amazonas: Tefé (DZ 35.116). 75, A. lipara Brazil, Amazonas: Fonte Boa (DZ 48.237). 76, Putative female of A. hemiurga Brazil, Amazonas: Tefé (DZ 35.266). Up to bottom: lateral view of papilla analis and ventral view of sterigma and bursa copulatrix. Scale bar: 1mm.

FIGURES 77–81. Signum of species of Alesa. 77, A. prema Brazil, Mato Grosso: Barra do Garça (DZ 35.254). 78, A. telephae Brazil, Acre: Mâncio Lima (DZ 35.094). 79, A. amesis Brazil, amazonas: Tefé (DZ 35.116). 80, A. lipara Brazil, Amazonas: Fonte Boa (DZ 48.237). 81, Putative female of A. hemiurga Brazil, Amazonas: Tefé (DZ 35.266). Scale bars: 1mm.

FIGURES 82–85. Species of Alesa in situ, part I. 82–83, A. prema, male from Peru, San Martín, Tarapoto (by Ken Kertell, https://www.inaturalist.org/observations/72706630) (82), female from Brazil, Paraná, Foz do Iguaçu (by Roberto Greve) (83). 84, A. juliae sp. nov., male from Ecuador, Napo, Tena (by Robert Carrion, https://www.inaturalist.org/observations/27548806). 85, A. telephae, male from Peru, San Martín, Rumipata-Mishquiyacu area (by David Geale, https://calydna.com/taxon/4372- alesa-telephae).