Taxonomic revision and phylogeny of Lycas (Lepidoptera: Hesperiidae: Hesperiinae), with a redescription of Lycas gabrielABSTRACT
Lycas Godman, 1901 (Pericharini: Pericharina) currently comprises five species. Although molecular data have driven numerous taxonomic changes in recent years, this genus still lacks a comprehensive revision based on a robust morphological diagnosis. Here, it is provided a morphological analysis based on 410 specimens alongside a phylogenetic analysis. Lycas is redescribed, including a detailed redescription of Lycas gabriel Grishin, 2025, and diagnostic features are provided for all included species. Finally, lectotypes are designated for Hesperia argentea Hewitson, 1866 and Hesperia ceraca Hewitson, 1866.
KEYWORDS:
Butterflies; lectotype designation; morphology; Neotropical region; skipper
INTRODUCTION
Lycas Godman, 1901 was originally described based on the morphology of the antennae, palpi, wing venation, and legs, and initially included Hesperia argenteaHewitson, 1866 and Hesperia ceraca Hewitson, 1866, without the designation of a type species. Subsequently, Lindsey (1925) designated H. argentea as the type species of Lycas. Later, Eudamus boisduvaliiEhrmann, 1909 was transferred to Lycas, although it was treated as a subspecies of Lycas godart (Latreille, [1824]) by Moss (1949). Finally, Evans (1955) characterized Lycas in greater detail, placing the genus in the “Carystus Group”, synonymizing Goniloba devanesHerrich-Schäffer, 1869 with L. argentea, and maintaining E. boisduvalii as a subspecies of L. godart.
In the genomic era, Lycas was recovered as one of the seven genera within Pericharini Grishin, 2019 (Li et al. 2019, Zhang et al. 2022a) and as one of the five genera of Pericharina Grishin, 2019 (Zhang et al. 2022a). Recently, taxonomic changes were proposed following the examination of the lectotype of Goniloba devanes and specimens of Lycas argentea, resulting in the recognition of both taxa as distinct species; the former occurs exclusively in South America, whereas the latter ranges from Mexico to Colombia (Zhang et al. 2022b). Most recently, Lycas gabriel Grishin, 2025 was described based on a single female specimen from Peru, and Lycas boisduvalii was elevated from subspecies to species status (Zhang et al. 2025).
Despite recent taxonomic advances based on molecular data, species of Lycas still lack robust morphological diagnoses and comprehensive illustrations for accurate identification, a limitation that has historically led to misidentifications within Pericharini and other morphologically similar Hesperiinae genera. Therefore, this study aims to: (1) revise the genus Lycas; (2) present a molecular phylogenetic analysis; (3) provide comprehensive species descriptions, diagnostic characters, and an illustrated identification key; (4) designate necessary lectotypes; and (5) redescribe L. gabriel based on 23 additional specimens, including the previously unknown male.
MATERIAL AND METHODS
Institutional collections
Specimens examined in this study were deposited in the following institutional collections: Carnegie Museum (CMP), Pittsburgh, Pennsylvania, USA; Coleção Entomológica Padre Jesus Santiago Moure, Departamento de Zoologia, Universidade Federal do Paraná (DZUP), Curitiba, Paraná, Brazil; Coleção Olaf Hermann Hendrik Mielke (OM), Curitiba, Paraná, Brazil; Museo de La Plata (MLP), La Plata, Buenos Aires, Argentina; Museum für Naturkunde (MfN), Berlin, Germany; The Natural History Museum (NHMUK), London, United Kingdom; National Museum of Natural History, Smithsonian Institution (USNM), Washington, D.C., USA; and Zoologiska Museum (ZMH), Helsinki, Finland.
Morphological analysis
A total of 410 specimens were examined, and 65 genitalia dissections were performed (42♂ and 23♀). Morphological terminology for genital structures and forewing venation follows Carneiro et al. (2013). Genitalia were removed from the abdomen, cleared in a 10% potassium hydroxide (KOH) solution, and dissected. Dissected genitalia and wing structures were photographed using a Leica M205 C stereomicroscope equipped with a Leica DMC4500 digital camera operated via Leica Application Suite software (version 4.12.0, Build 86). Forewing length, measured from base to apex, was quantified using ImageJ software (Schneider et al. 2012).
Taxon sampling and molecular data
Cytochrome c Oxidase I (COI) was utilized as the sole molecular marker. For phylogenetic reconstruction, mitochondrial DNA barcodes were extracted, amplified, and sequenced at the Canadian Centre for DNA Barcoding (CCDB, Centre for Biodiversity Genomics, University of Guelph, Ontario, Canada) following the protocols described by Hebert et al. (2003). In addition to 12 newly sequenced Lycas specimens, sequences available in GenBank and BOLD Systems from previous studies (Lavinia et al. 2017, Kawahara et al. 2023) were included (Table 1). One sequence not deposited in GenBank was obtained directly from the literature (Zhang et al. 2025). Outgroup selection was based on Zhang et al. (2022a) to preserve tribal relationships, including Perichares adela (Hewitson, 1867) and Orses cynisca (Swainson, 1821) as the sister group to Lycas. All valid species of Lycas, including newly discovered taxa, were sampled. The final dataset comprised 21 terminals (19 ingroup and two outgroups). Whenever possible, full-length sequences of 658 base pairs (bp) were used; however, two shorter sequences consisting of 307 and 656 bp were retained (Table 1).
Specimens, collection codes, GenBank and BOLD Systems accession numbers, localities, and sources of Cytochrome c Oxidase I (COI) sequences used in this study. New sequences generated from specimens deposited in the DZUP and OM collections are indicated as “this study”. All sequences comprised 658 base pairs, except those marked with an asterisk (*), which comprised 656 and 307 base pairs, respectively. Superscript (1) indicates the distinct phenotype recorded in Lycas.
Alignment and phylogenetic inference
Sequences were aligned using the MAFFT online platform, version 7, and genetic distances were calculated using MEGA software (Kumar et al. 2024). Phylogenetic inference was performed under the Maximum Likelihood (ML) criterion using IQ-TREE (Nguyen et al. 2015) integrated within the PhyloSuite platform (Zhang et al. 2020, Xiang et al. 2023). Node support values were calculated using 1,000 replicates of the ultrafast bootstrap (UFboot) approximation (Minh et al. 2013) and 1,000 replicates of the Shimodaira-Hasegawa-like approximate likelihood ratio test (SH-aLRT) (Guindon et al. 2010). The resulting topology was visualized in FigTree (Rambaut 2010) and rooted with Perichares adela and Orses cynisca. Following the criteria of Guindon et al. (2010) and Minh et al. (2013), nodes with SH-aLRT values greater than or equal to 80% and UFboot support greater than or equal to 95% were considered strongly supported.
Geographic distribution and host records
Locality data from DZUP and OM collections were used to generate the distribution map. To minimize taxonomic misidentifications, literature records were included in the geographical database only if accompanied by verifiable photographs. The distribution map was constructed using QGIS Desktop software (version 3.32.1). Published host plant records for two species were excluded from the final analysis due to the lack of photographic evidence to corroborate species identification.
The synonymic bibliography under each species follows the format established by Mielke (2026).
Abbreviations
The following abbreviations are used throughout the text: (DFW) dorsal forewing; (DHW) dorsal hindwing; (FW) forewing; (HW) hindwing; (ICZN) International Commission on Zoological Nomenclature; (VFW) ventral forewing; (VHW) ventral hindwing.
TAXONOMY
Lycas Godman, 1901
Lycas Godman, 1901, in Godman & Salvin. Biol. Centr.-Amer., Lep.-Rhop. 2, p. 618; included species: Hesperia argentea Hewitson, Hesperia ceraca Hewitson.- Mabille, 1904, in Wytsman. Gen. Ins. 17, p. 121, 167.- Draudt, 1923, in Seitz. Gross-Schmett. Erde 5, p. 991.- Lindsey, 1925. Ann. Ent. Soc. Amer. 18: 91.- Hayward, 1934. Rev. Soc. ent. arg. 6: 101, 175, 198, pl. 15, fig. 9 (venation).- Hayward, 1941. Rev. Mus. La Plata, n. s., Zool. 2: 320.- Williams & Hayward, 1944. Acta zool. Lill. 2: 222.- Bell, 1946. Bol. Ent. venezol. 5: 180.- Hayward, 1950. Gen. Sp. Anim. Arg. 2, p. 320.- Evans, 1955. Cat. Amer. Hesp. 4, p. 208, 261.- Hemming, 1967. Bull. Brit. Mus. (Nat. Hist.), Ent., Suppl. 9: 268.- Lamas, 1969. Biota 7: 338.- Beattie, 1976. Rhop. Direct., p. 36.- C. Hoffmann, 1976. Publ. Esp. Soc. Mex. Lep. 1: 156.- Hayward, 1973. Op. Lill. 23: 86.- Okano, 1981. Tokurana 1: 31.- Bridges, 1983. Lep. Hesp. 2, p. 19.- Bridges, 1988. Cat. Hesp. 2, p. 31; App. 2, p. 2.- Bridges, 1988. Cat. Fam.-Group & Gen.-Group Nam., 2nd ed., 4, p. 74; 5, p. 3.- Llorente-Bousquets, Luis-Martínez & Vargas-Fernández, 1990. Publ. esp. Mus. Zool., Mexico, 1: 21.- J. de la Maza E., White & R. G. de la Maza E., 1991. Rev. Soc. Mex. Lep. 14: 35.- Lamas, O. Mielke & Robbins, 1993. Jour. Lep. Soc. 47: 81; ethol.- R. G. de la Maza E. & J. de la Maza E., 1993. Marip. Chiapas, p. 168, 208.- R. G. de la Maza E. & Gutiérrez, [1994]. Rev. Soc. Mex. Lep. 15(1), Add., p. 14.- Bridges, 1994. Cat. Fam.-Group, Gen.-Group, Sp.-Group Nam., Hesp. (Lep.) World 4, p. 17; 5, p. 3; 6, p. 7; 9, p. 35.- Warren, 2000, in Llorente, González & Papavero (eds). Biodiv., Tax., Biogeogr. Art. Mex. 2, p. 560.- O. Mielke, 2004. Hesperioidea, p. 9, 71, in Lamas (ed.). Checklist: Part 4A, Hesperioidea-Papilionoidea, in Heppner (ed.). Atlas Neotrop. Lep. 5A.- O. Mielke, 2005. Cat. Amer. Hesperioidea 1, p. 12; 4, p. 1050.- DeVries, Austin & Martin, 2008. Biol. Jour. Linn. Soc. 94: 728.- Warren, Ogawa & A. Brower, 2008. Cladistics 24: 669.- Warren, Ogawa & A. Brower, 2009. Syst. Ent. 34: 495. 516.- Li et al., 2019. Proc. Nat. Acad. Sc. 116(13): 6232-6237, S1 App., p. 3, 8.- Toussaint & Warren 2019. Jour. Nat. Hist. 53: 35-36, p. 2175; eye, ethol.- Suênia-Bastos et al. 2025. Zootaxa 5604(2): 102 (table 1). Suênia-Bastos et al. 2025. Zootaxa 5604(2): 131.
Lycas sp. Beccaloni et al., 2008. Monografías Tercer Milenio 8: 39; hostplant.
Species included. Lycas argentea (Hewitson, 1866), Lycas boisduvalii (Ehrmann, 1909), Lycas devanes (Herrich-Schäffer, 1869), Lycas godart (Latreille, [1824]), and Lycas gabriel Grishin, 2025.
Diagnosis and remarks. The phylogenetic analysis recovered Lycas as a strongly supported monophyletic clade (Fig. 1). Morphologically, Lycas superficially resembles two genera of Hesperiinae: Aides Billberg, 1820 and Calpodes Hübner, [1819]. However, it can be distinguished from Aides by wing shape and the pattern of the silver bands on the VHW. In Lycas, these bands are arranged longitudinally between the costal and outer margins, whereas in Aides longitudinal bands are absent, being replaced by rounded or irregular silver spots. Compared to Calpodes, Lycas differs by possessing a stigma on the male DFW (Figs 7-8), reddish-brown scales and a yellowish spot between C and Sc on the VFW, and longitudinal silver bands on the VHW. Distinctive characters separating Lycas from other Pericharini genera include: a yellow spot between M1 and M3 on the DHW (Figs 2-6); longitudinal silver bands between the costal and outer margins on the VHW (Figs 2-6). Male genitalia (Fig 9-11) with the gnathos and the uncus almost fused; non-bifid uncus; valva simple, lacking conspicuous subdivisions or dorso-distal spines. Female genitalia (Figs 12, 13) with lamella antevaginalis weakly sclerotized laterally or ventro-laterally, or membranous.
Phylogenetic tree of five Lycas species under Maximum Likelihood (ML) inferred in IQ-TREE, using Cytochrome c Oxidase I (COI) as molecular marker. The node supports are shown below each branch (SH-aLRT ≥ 80% and UFboot ≥ 95%). In pink: Lycas devanes, blue: L. godart, purple: L. boisduvalii, green: Lycas gabriel, and yellow: L. argentea. (*) different phenotype found in Lycas gabriel.
Lycas argentea, dorsal and ventral views: (A-B) male DZ 58.373, from Rio Natal, São Bento do Sul, Santa Catarina, Brazil; (C-D) female DZ 58.366, from Rio Vermelho, São Bento do Sul, Santa Catarina, Brazil. Scale bar: 10 mm.
Etymology. The original etymology was not provided by the author.
Redescription. Head: Antenna dorsally brownish, usually yellowish ventrally; nudum with 14-18 segments. Eyelash absent. Eye red in living individuals (Glassberg [2018]).
Thorax: Mesotibia and metatibia spined. Male DFW (Figs 2-6) brownish with six yellow hyaline spots: one in discal cell, usually three apical between R3 and M1, two discal in M3-CuA1 and CuA1-CuA2, and one opaque yellowish spot in lower half of CuA1-2A; stigma absent in L. argentea and L. devanes, but present, tripartite, and discontinuous between CuA1 and 2A in L. godart, L. boisduvalii, and L. gabriel (Figs 7-8). Female DFW (Figs 2-6) with an additional hyaline spot below CuA2. FW with recurring vein; dcs sometimes present; distance between origins of CuA1-CuA2 wider than origins of M3-CuA1. DHW (Figs 2-6) brownish with opaque yellow spot between M1 and M3. VHW (Figs 2-6) with two longitudinal silver bands between costal margin and tornus (discal and submarginal bands), and a continuous band along the anal margin.
Abdomen: Male genitalia (Figs 9-11) with tegumen broader than uncus; uncus simple, long, reaching or nearly reaching apex of valva; uncus and gnathos almost fused; gnathos shorter than uncus; fultura inferior wider posteriorly than anteriorly, anterior portion straight or slightly rounded; valva simple, lacking conspicuous subdivisions or dorso-distal spines; sacculus bearing setae; aedeagus concave ventro-distally, median and distal portions wider than anterior portion; cornuti absent. Female genitalia (Figs 12-13) with lamella antevaginalis weakly sclerotized laterally or ventro-laterally, or membranous, medially connected to lamella postvaginalis; lamella postvaginalis wider than lamella antevaginalis; papilla analis squared with rounded edges; corpus bursae without clear distinction between corpus and ductus.
Key to the species of Lycas
1. VFW with greyish markings in the subapical area; male DFW without stigma (Figs 2, 3) .................... 2
1’. VFW without greyish markings in the subapical area (Figs 5B, D, 6D), or with lilac markings in the subapical area (Figs 4B, D, 6B, F); male DFW with stigma (Figs 7-8) .................... 3
2. Male VHW with the discal silver band connected to the submarginal band in central area, and the discal band continuous from costal margin to the anal margin (Fig. 2B); female VHW with irregular bands, the submarginal band exhibits a small inner projection between M1 and M3 (Fig. 2D) .................... Lycas argentea
2’. Male VHW with the discal silver band not connected to the submarginal band in central area, and the discal band not continuous from costal margin to the anal margin, interrupted by a brownish band in 2A-3A (Fig. 3B-C); female VHW with straight bands (Fig. 3E) .................... Lycas devanes
3. VFW without lilac markings in the subapical area; DFW with large hyaline spots (Fig. 5) .................... Lycas godart
3’. VFW with lilac markings in the subapical area; DFW with small hyaline spots (Figs 4, 6) .................... 4
4. Male DFW stigma with the first part in CuA1-CuA2 broad, almost reaching the origins of CuA1-CuA2 (Fig. 7A-B); female genitalia with lamella postvaginalis with a deep concavity (Fig. 12E) .................... Lycas boisduvalii
4’. Male DFW stigma with the first part in CuA1-CuA2 narrow and not reaching the origins of CuA1-CuA2 (Fig. 8); female genitalia with lamella postvaginalis formed by two plates with median portion folded ventrally (Fig. 13C) .................... L. Gabriel
Lycas argentea (Hewitson, 1866)
Hesperia argentea Hewitson, 1866. Trans. ent. Soc. London (3)2(6): 487; Guatemala; collection Hewitson.- Hewitson, 1872. Illustr. Exot. Butt. 5, p. [102], pl. [53] (Hesperia 5), figs 46, 47 (d, v).- Kirby, 1879. Cat. Coll. Diurn. Lep. Hewitson, p. 216.- Plötz, 1882. Stett. ent. Ztg. 43: 336; syn.: devanes.
Proteides argentea; Kirby, 1871. Syn. Cat. Diurn. Lep., p. 594.- Kirby, 1877. Syn. Cat. Diurn. Lep., Suppl., p. 824.- Staudinger, 1888, in Staudinger & Schatz. Exot. Schmett. 1, p. 297.- Mabilde, 1986. Borb. Est. Rio Grande do Sul, Porto Alegre, p. 127.
Carystus argentea; Mabille, 1878. Ann. Soc. ent. Belg. 21: 36.
Lycas argenteus [sic]; Godman, 1901, in Godman & Salvin. Biol. Centr.-Amer., Lep.-Rhop. 2, p. 618; 3, pl. 104, figs 29 (venation), 30 (male gen.).- Mabille, 1904, in Wytsman. Gen. Ins. 17, p. 167.- Draudt, 1923, in Seitz. Gross-Schmett. Erde 5, p. 991, pl. 191a (d, v).- Lewis, 1973. Butt. World, p. 83, fig. 28 (v), p. 246.- Lewis, 1975. Marip. Mundo, p. 83, fig. 28 (v), p. 246.
Lycas argentea [confirmed records]; Garwood & Lehman, 2005. Butt. Northeastern Mexico, p. 153, fig. (female, v), 188.- Garwood et al., 2007. Butt. South. Amaz., p. 232, figs (female v).- Garwood et al., 2009. Butt. South. Amaz., 2nd ed., p. 320, figs (v).- Garwood & Lehman, 2013. Butt. C. Amer. 3, Hesp., p. 165, figs (female, v).- Glassberg, [2018]. Swift Guide Butt. Mexico and C. America, 2nd ed., p. 259, figs (male, female v).- Palo Jr., 2017. Borb. Brasil 3, p. 2050, 2051, figs (male, female v).- Suênia-Bastos et al. 2025. Zootaxa 5604(2): 102 (table 1), figs 13-20 (male, female, d, v), fig. 63 (palpus), fig. 78 (forewing venation), fig. 85, 90 (male genitalia, female genitalia).
Lycas argenteus [sic] [unconfirmed records]; Costa Lima, 1923. Arch. Esc. Sup. Agric. Vet., Niteroi, 6: 153; hostplant.- Costa Lima, 1928. Arch. Esc. Sup. Agric. Vet., Rio de Janeiro, 8: 139; hostplant.- Ronna, 1928. Ins. Brasil, São Paulo, p. 35; hostplant.- Spitz, 1932. Rev. Mus. Paulista 17: 878.- Ronna, 1933. Egatea, Porto Alegre, 18: 99; hostplant.- Monte, 1934. O Campo, Rio de Janeiro, 5: 72; hostplant.- F. Hoffmann, 1934. Ent. Rdsch. 51: 73.- Hayward, 1935. Rev. Soc. ent. arg. 7: 193.- Kivirikko, 1936. An. Ent. Fenn. 2: 63.- Costa Lima, 1936. Terc. Cat. Ins. viv. Plantas Brasil, p. 231; hostplant.- Hayward, 1937. Rev. Soc. ent. arg. 8: 59, fig. 7 (male gen.).- Hayward, 1941. Rev. Soc. ent. arg. 11: 36.- Forster, 1949. Bol. Ent. venezol. 7: 120.- Biezanko, 1963. Arq. Ent., sér. A, Pelotas, p. 20.- J. Zikán & W. Zikán, 1968. Pesq. agropec. bras. 3: 60.- Silva et al., 1968. Quarto Cat. Ins. viv. Plantas Brasil 2(1), p. 305; syn.: argentea; hostplant.
Lycas argentea [unconfirmed records]; Lindsey, 1925. Ann. Ent. Soc. Amer. 18: 91.- C. Hoffmann, 1941. An. Inst. Biol., Mexico, 12: 282; syn.: argenteus.- Hayward, 1940. An. Soc. cient. arg. 130: 93.- Hayward, 1941. Rev. Mus. La Plata, n. s., Zool. 2: 320; syn.: argenteus, godarti; Hayward, 1934.- Hayward, 1947. Acta zool. Lill. 4: 29, 42; hostplant.- O. Mielke, 1968. Atas Soc. Biol. Rio de Janeiro 12: 77.- Ebert, 1968, in Silva et al. Quarto Cat. Ins. viv. Plantas Brasil 2(2), p. 234.- Ebert, 1969. Jour. Lep. Soc. 23, Suppl. 3: 37.- Hayward, 1969. Inst. Miguel Lillo, Misc. 31: 87.- Hayward, 1973. Op. Lill. 23: 87; syn.: bipunctata, albomarginata.- Biezanko & O. Mielke, 1973. Acta biol. paranaense 2: 91.- K. Brown, 1987. An. Prim. Simp. Pantanal, Brasília, p. 168.- Steinhauser, 1975. Bull. Allyn Mus. 29: 22.- C. Hoffmann, 1976. Publ. Esp. Soc. Mex. Lep. 1: 156; syn.: argenteus.- Bridges, 1983. Lep. Hesp. 1, p. 4, 9, 16, 34; 2, p. 19; syn.: ab. albomarginata, bipunctata, devanes.- J. de la Maza E. & R. G. de la Maza E., 1985. Rev. Soc. Mex. Lep. 9: 44.- Bridges, 1988. Cat. Hesp. 1, p. 6, 15, 26, 53; 2, p. 31; syn.: ab. albomarginata, bipunctata, devanes.- Llorente-Bousquets, Luis-Martínez & Vargas- Fernández, 1990. Publ. esp. Mus. Zool., Mexico, 1: 38.- Lamas, Robbins & Harvey, 1991. Publ. Mus. Hist. Nat., Lima, 40: 18.- Lamas, 1994, in Foster et al. RAP Working Papers 6: 177.- J. de la Maza E., White & R. G. de la Maza E., 1991. Rev. Soc. Mex. Lep. 14: 35.- Stanford & Opler, 1993. Atlas West. USA Butt., p. 63.- R. G. de la Maza E. & Gutiérrez, [1994]. Rev. Soc. Mex. Lep. 15(1): 25, 34; Add., p. 14.- Bridges, 1994. Cat. Fam.-Group, Gen.-Group, Sp.-Group Nam. Hesp. (Lep.) World 7, p. 1; 8, p. 8, 17, 31, 64; 9, p. 35; syn.: devanes bipunctata, ab. albomarginata.- Llorente-Bousquets et al., 1995. Rev. Soc. Mex. Hist. Nat. 46: 41.- C. Mielke, 1995. Revta bras. Zool. 11(4): 765.- Robbins et al., 1996, in Wilson & Sandoval. Manu, p. 251.- Kochalka et al., 1996, in Romero M. Col. Flora Fauna Mus. Nac. Hist. Nat. Paraguay, p. 180.- Vargas-Fernández et al., 1996. Jour. Lep. Soc. 50: 112.- Warren et al., 1996. Dugesiana 3(1): 7.- Llorente-B. et al., 1996. Dugesiana 3(2): 7.- Raguso & Llorente, 1997, in González, Dirzo & Vogt (eds). Hist. Nat. Tuxtlas. Inst. Biol., UNAM, p. 288.- Moreno, Silva & Estévez, 1998. Marip. Ecuador, p. 160.- Warren et al., 1998. Jour. Lep. Soc. 52: 54.- Austin et al., 1999. Trop. Lep. 9, Suppl. 2: 9.- Warren & Llorente-Bousquets, 1999. Dugesiana 6(1): 23.- Maes et al., 1999, in Maes. Cat. Ins. Art. Nicaragua 3, p. 1279; syn.: devanes, bipunctata, albomarginata.- K. Brown & Freitas, 2000. Bol. Mus. Biol. Mello Leitão, n. s., Sta. Teresa, 11/12: 113.- Warren, 2000, in Llorente, González & Papavero (eds). Biodiv., Tax., Biogeogr. Art. Mex. 2, p. 560; syn.: bipunctata, devanes.- Sigrist & M. Sazima, 2003. Revta bras. Bot. 25: 39; ecol.- Llorente et al., 2004. Jour. Lep. Soc. 58: 210.- Luis Martínez et al., 2004, in García-Mendoza, Ordóñez & Briones-Salas. Biodiv. Oaxaca, p. 355.- O. Mielke, 2004. Hesperioidea, p. 71, in Lamas (ed.). Checklist: Part 4A, Hesperioidea-Papilionoidea, in Heppner (ed.). Atlas Neotrop. Lep. 5A; syn.: devanes, form bipunctata, ab. albomarginata.- O. Mielke, 2005. Cat. Amer. Hesperioidea 1, p. 12; 4, p. 1050; syn.: devanes, argenteus, form bipunctata, ab. albomarginata.- Pastrana, 2004. Lep. arg., Plantas hosp., p. 207; hostplant.- Salinas-G., Warren & Luis-M., 2005. Folia Entomol. Mex. 44(3): 318.- Sandoval, Fernandez-Badillo & González, 2007. Alcance, Maracay, 70: 56.- Barrios, Méndez & Austin, 2006, in Cano (ed.). Biodiv. Guatemala 1, p. 436.- Glassberg, 2007. Swift Guide Butt. Mexico and C. America, p. 225, fig. (v); hostplant.- Beccaloni et al., 2008. Monografías Tercer Milenio 8: 39; hostplant.- O. Mielke, Emery & Pinheiro, 2008. Revta bras. Ent. 52(2): 286.- Giovenardi et al., 2008. Revta bras. Ent. 52(4): 602.- Núñez B., 2008. Trop. Lep. Res. 18(2): 80.- Sackis & Morais, 2008. Biota Neotrop. 8(1): 154.- Núñez B., 2009. Trop. Lep. Res. 19(2): 77.- Francini et al., 2011. Revta bras. Ent. 55(1): 67.- Luis Martínez et al., 2011, in Cruz A. (ed.). Biodiv. Veracruz. Apénd. VIII.25, p. 9.- Luis-M., Salinas-G. & Llorente-B., 2011, in Álvarez (coord.). Chiapas: estudios diversidad biol., Inst. Biol., UNAM, Mexico, p. 390.- Núñez B. et al., 2011. Trop. Lep. Res. 21(1): 41.- Núñez B. & Volkmann, 2011. Shilap Revta. lepid. 39(155): 249, 255.- Dolibaina, O. Mielke & Casagrande, 2011. Biota Neotrop. 11(1): 345.- Ritter et al., 2011. Biota Neotrop. 11(1): 363.- Soares et al., 2011. Trop Lep. Res. 21(2): 71.- Giovenardi et al., [2014]. Rev. colomb. Ent. 39(2), Supl., p. 15.- Favretto, Santos & Geuster, 2013. EntomoBrasilis 6(1): 55.- Carneiro et al., 2014. Neotrop. Ent. 43(1), Elect. Suppl. Mat., p. 5.- Carneiro et al., 2014. PLoS ONE 9(10) (e108207), fig. S2.- Thiele et al., 2014. Biota Neotrop. 14(2): 4.- Piovesan et al., 2014. Scientia Plena 10(9): 13.- Núñez B., 2015. Patrim. Nat. Cult. Área Infl. Emb. Yacyretá, Argentina, p. 111.- J. de la Maza & R. G. de la Maza, [2016], in Carabias, J. de la Maza & Cadena (eds.). Conserv. desarr. sust. Selva Lacandona, p. 648.-Luis-Martínez et al., 2016. Southw. Ent. 41(1): 194.- Piovesan & Orlandin, 2016, in Orlandin et al. Borb. Marip. S. Catarina, p. 78.- Leviski et al., 2016. Trop. Lep. Res. 26(2): 66.- Vargas-Fernández et al., 2016, in Conabio. La biodiversidad en Colima: Estudio de Estado. Apéndice 1: 188; hostplant.- Lavinia et al., 2017. PloS ONE 12(10) (e0186845), p. 10.- Uehara, 2018. Butterflies 78: 45.- Dickens, McMahon & Binnie, 2019. Jour. Ins. Cons. 23(4): 712.- Li et al., 2019. Proc. Nat. Acad. Sc. 116(13): 6232-6237, fig. 1 (genomic tree); S1 App., p. 74, 91 (Fig. S1, phylogenetic tree).- Cong et al., 2019. Insecta Mundi 731: 49, fig. 13 (genomic tree).- Orlandin et al., 2020. Pap. Avuls. Zool. 59 (e20195949), p. 14.- Bizarro & Martin, 2020. Guide Butt. Serra dos Orgãos. South-eastern Brazil, p. 271. Zamoner (org.), 2021. Borb. de Curitiba e do Paraná, p. 52, 79.- Orlandin et al., 2022, in Greca de Macedo & Dias. Portaria Nº 1082, Pref. Curitiba, p. 62.- Zhang et al., 2022. Insecta Mundi 921: 95, 124, fig. 17 (genomic tree).- Zhang et al., 2022. Tax. Rept. Intern. Lep. Surv. 10(7): 56, fig. 89 (genomic tree).- Vieira-Silva et al., 2023. Pap. Avuls. Zool. 63: e202363032, p. 6.- Greve et al., 2023. Biota Neotrop. 23(3): e20231487, p. 12. Kawahara et al., 2023. Nat. Ecol. & Evol. 2023. https://doi.org./10.1038/s41559-023-02041-9, fig. S1 (time calibrated tree), table S1 (specimen inf.).
(no genus) argentea; Beattie, 1976. Rhop. Direct., p. 83.
Taxonomic history. Hesperia argentea Hewitson, 1866 was originally described based on an unknown number of specimens from Guatemala. This species was subsequently transferred to Proteides Hübner, [1819] by Kirby (1871), to Carystus Hübner, [1819] by Mabille (1878), and finally to Lycas by Godman (1901, in Godman and Salvin).
Type material. One male syntype of Hesperia argentea found in NHMUK, with following labels: / Type / Type / orange label without information / 160 [?] / Guatema [Guatemala] / argentea / Hewitson Coll. 79-69 2. Hesperia argentea Hew. /. This syntype is herein designated as lectotype to allow the corrected identification of the species, with the following labels: /LECTOTYPUS/ Lectotype Hesperia argentea Hewitson, 1866 Suênia-Bastos, O. Mielke & Casagrande det. 2025/.
Diagnosis. FW length 21-23 mm. Similar to L. devanes; distinguished from all other congeners by absence of male FW stigma and presence of broad opaque whitish spot above 2A on male VFW, and presence of greyish markings in subapical area on VFW in both sexes (Fig. 2). Male differs from L. devanes by discal silver band connected to submarginal band in central area on VHW, and discal band continuous from costal margin to anal margin (Fig. 2B). Female wing pattern differing from male; silver bands irregular, never connected; submarginal band with small inner projection between M1 and M3 (Fig. 2D). Male genitalia (Fig. 9A-F) similar to L. devanes, with minor differences: gnathos wavy in lateral view; coecum of aedeagus straight; valva more rectangular, posterior margin blunt. Female genitalia (Fig. 12A-B) with informative diagnostic characters: lamella antevaginalis mostly membranous, weakly sclerotized both dorsally and ventrally; lamella postvaginalis with ventro-distal projection in median portion.
Etymology. The original etymology was not explicitly provided by the author. However, the name is a feminine adjective derived from Latin, referring to the silver coloration of the hindwing bands.
Remarks. Hewitson (1866) described L. argentea from Guatemala, explicitly highlighting the diagnostic VHW character in his description as having “two broad silver bands connected in the middle”. Subsequently, in a later publication, Hewitson (1872) illustrated a specimen that may, in fact, represent L. devanes. Additionally, Godman (1901) noted that he examined numerous specimens from both Central and South America, a series that likely included specimens of L. devanes.
Geographical distribution and phenology. Lycas argentea is widely distributed from Central to South America (Fig. 14), occurring at elevations from 10 to 1,300 m, and recorded throughout the entire year. Records are based on examined collection data and literature sources: Mexico - Oaxaca (DZUP), Tamaulipas (OM; Zhang et al. 2022b), Veracruz (OM); Guatemala - without precise locality (Hewitson 1866); Costa Rica - San José (DZUP); Bolivia - Cochabamba (OM); Brazil - Alagoas (DZUP), Minas Gerais (DZUP), Espírito Santo (DZUP), Rio de Janeiro (DZUP; Palo Jr. 2017), Paraná (DZUP), Santa Catarina (DZUP); Paraguay - Guairá (OM; DZUP), Alto Paraná (OM); Argentina - Misiones (OM; Palo Jr. 2017).
Material examined. Besides the figures of the lectotype, 52 more specimens were studied. MEXICO, Oaxaca: Candelaria Loxicha (500 m), 1♂, 15-VIII-1972, Welling leg., DZ 58.260 (DZUP), 1♂, 4-IX-1973, Welling leg., DZ 58.261 (DZUP), 1♂, 5-VIII-1974, Welling leg., DZ 58.262 (DZUP), 1♂, 15-VIII-1975, Welling leg., DZ 58.263 (DZUP); Veracruz: Catemaco (Temolapan, 450 m), 1♂, VII-VIII-2013, G. Noguera leg. OM 77.052 (DZUP-OM); Tamaulipas: (El Encino, 250 m), 1♂, 4-13-VIII-1988, V. O. Becker leg., OM 50.892 (DZUP-OM). COSTA RICA, San José: (San José), 1♂, 12-X-1936, Bollon leg., DZ 58.264 (DZUP). BRAZIL, Alagoas: Junqueiro (Riachão), 1♂, 28-X-1973, A. Cardoso leg., ex-coll. A. Cardoso, DZ 58.277 (DZUP); Minas Gerais: Carangola (Fervedouro), 1♂, 18-II-1972, DZ 58.293 (DZUP), (650 m), 1♂, 1♀, 29-VII-1992, H. H. Ebert leg., ex-coll. H. Ebert, DZ 58.291, DZ 58.292 (DZUP); Espírito Santo: Santa Tereza, 1♂, 25-VIII-1967, 1♀, 18-X-1967, 1♂, 22-IX-1967, C. & C. T. Elias leg., (Parada Santa Lúcia), 1♂, 25-I-2012, N. Tangerini leg., ex-coll. Nirton Tangerini, DZ 58.295-DZ58.298 (DZUP); Rio de Janeiro: Nova Friburgo, 2♂, II-2009, 2♂, VIII-2007, 1♀, VI-2007, 1♂, V-2007, 1♂, XI-2012, E. Grossi leg., (1,100 m), 1♂, 29-3-IV-V-2001, A. Moser leg., (Sítio Caturano, 1,100 m), 1♂, I-2007, E. Grossi leg., DZ 58.299, DZ 58.300, DZ 58.301, DZ 58.304, DZ 58.302, DZ 58.306, DZ 58.308, DZ 58.310, DZ 58.303 (DZUP); Cachoeiras de Macacu (Boca do Mato), 1♂, 29-IV-2008, 1♂, 28-III-2008, N. Tangerini leg., 24-VI-2003,1♂, 30-IV-2002, 1♂, 1♀, 3-IV-2006, Jonas Overney leg., ex-coll. Nirton Tangerini, DZ 58.318, DZ 58.320, DZ 58.319, DZ 58.321, DZ 58.322 (DZUP); Petrópolis, 1♀, 3-X-1959, Gagarin leg., ex-coll. Gagarin, DZ 58.323 (DZUP); Itatiaia (1,200 m), 1♀, 13-V-1996, V. O. Becker leg., OM 50.885 (DZUP-OM); Paraná: Chopinzinho, 1♂, 23-II-1971, Furtado leg., DZ 58.342 (DZUP); Morretes (Morro Alto), 1♂, 11-IV-2015, O. Mielke & Dolibaina leg., DZ 58.347 (DZUP); Paranaguá (Floresta Estadual do Palmito, 25°34’8”S, 48°32’8”W, 10-20 m), 1♀, 25-26-VIII-2014, Leviski, Queiroz-Santos & Santos leg., DZ 58.348 (DZUP); Três Barras do Paraná (Parque Estadual Guarani, 25°26’20”S, 53°09’49”W, 500 m), 1♀, 1-4-XII-2018, O. Mielke & M. Casagrande leg., DZ 58.350 (DZUP); Santa Catarina: São Bento do Sul (Rio Vermelho), 1♂, 29-III-2009, 1♂, 11-IV-1974, 1♂, 16-VI-1973, 1♀, 23-X-1974, Rank leg., DZ 58.369, DZ 58.366, DZ 58.360, DZ 58.364 (DZUP), (Rio Natal), 1♀, 1♂, 20-VIII-2014, (Rio Natal, 500 m), 1♂, 5-IV-1989, 1♂, 23-VI-2006, Rank leg., DZ 58.370, DZ 58.371, DZ 58.375, DZ 58.373 (DZUP); Corupá, 1♂, IV-1966, Maller leg., DZ 58.380 (DZUP); Brusque, 1♂, II-1971, Becker leg., DZ 58.381 (DZUP); Joinville, 1♂, 22-XII-1981, O. Mielke leg., (200 m), 1♂, 21-IV-1967, 1♂, 25-XI-1974, 1♂, 8-VIII-1971, 1♀,14-X-1973, Miers leg., DZ 58.383, DZ 58.384-DZ 58.387 (DZUP). BOLIVIA, Cochabamba: La Paz (Puente Villa, 1,300 m), 1♀, 6-III-1998, Callaghan leg., OM 49.300 (DZUP-OM). PARAGUAY, Guairá: Villa Rica, 1♀, 15-VIII-1968, Anders leg., DZ 58.403 (DZUP). ARGENTINA, Misiones: Puerto Iguazú, 1♀, 5-X-1986, Foerster leg., OM 19.120 (OM).
Lycas devanes (Herrich-Schäffer, 1869)
Goniloba devanes Herrich-Schäffer, 1869. Corr.-Blatt zool.-min. Ver. Regensburg 23: 193; no data; syn.: ? ceraca Hewitson.
Proteides argentea [misidentification]; Mabilde, 1986. Borb. Est. Rio Grande do Sul, Porto Alegre, p. 127.
Lycas argenteus [sic] [misidentifications]; Hayward, 1934. Rev. Soc. ent. arg. 6: 104, 175, 198, pls 12, figs 9, l0 (d, v), 19, fig. 3 (male gen.).
Lycas argenteus [sic] form bipunctata Hayward, 1934. Rev. Soc. ent. arg. 6: 104, 176, 198, pl. 12, fig. 8 (male d); holotype male, Misiones, Argentina; collection Breyer.- Hayward, 1934. Rev. Soc. ent. arg. 6: 218.
Lycas argenteus [sic] ab. albomarginata Kivirikko, 1936. An. Ent. Fenn. 2(2): 63, fig. 3 (d, v); 1 specimen [holotype], 5-V-20-VI- l928, Misiones, Argentina, Kivirikko leg.- Hayward, 1939. Physis 17: 299.
Lycas argentea ab. bipunctata Hayward, 1941. Rev. Mus. La Plata, n. s., Zool. 2: 320.
Lycas argentea ab. albomarginata Hayward, 1941. Rev. Mus. La Plata, n. s., Zool. 2: 320.
Lycas argentea [misidentifications]; Hayward, 1950. Gen. Sp. Anim. Arg. 2, p. 321, pls 14, fig. 2 (male gen.), 25, figs 18, 19, 20 (male d, v, female v); syn.: bipunctata, albomarginata.- Evans, 1955. Cat. Amer. Hesp. 4, p. 262, pl. 71 (male gen.); syn.: devanes, bipunctata, albomarginata.- K. Brown, 1992, in Morellato. Hist. nat. Japi, p. 180, fig. 17 (female v).- Canals, 2003. Marip. Misiones, p. 124, 459, figs (male, female v); hostplant.- Garwood et al., 2007. Butt. South. Amaz., p. 232, figs (male, v).- Garwood et al., 2009. Butt. South. Amaz., 2nd ed., p. 320, figs (v).-O. Mielke, Carneiro & Casagrande, 2012. Revta bras. Ent. 56(1): 64.- Garwood & Lehman, 2013. Butt. C. Amer. 3, Hesp., p. 165, figs (male, v).- Volkmann & Núñez B., 2013. Marip. Serran. Arg. Central. 2, Nymph. Hesp., p. 136, figs ([male, female] v); syn.: devanes, f. bipunctata, ab. albomarginata; biol., hostplant.- Lavinia et al., 2017. PloS ONE 12(10) (e0186845), p. 10.- Palo Jr., 2017. Borb. Brasil 3, p. 2050, 2051, figs (male, female v); p. 2317.- Pérez et al., 2017. Shilap Revta. lepid. 45(179): 441.- Uehara, 2017. Butterflies 76: 47, fig. 61 (male d, v).- Klimaitis et al., 2018. Marip. Argentinas, p. 246, figs (male, female d, v), p. 282 (208).- Toussaint & Warren 2019. Jour. Nat. Hist. 53: 35-36, p. 2168, fig. 2 ([male] v).- Ahlf, Martello & Klein, 2020. Luminária, União da Vitória, 21(1): 35, fig. 3D (d, v).- Orlandin, Piovesan & Carneiro, 2020. Borboletas Meio-Oeste Santa Catarina, p. 63, fig. 26a-d (female d, v, male v); biol., hostplant.- Orlandin et al., 2023, in Straube. Invent. Fauna Curitiba, p. 171.
(no genus) bipunctata; Beattie, 1976. Rhop. Direct., p. 96.
(no genus) albomarginata; Beattie, 1976. Rhop. Direct., p. 72.
(no genus) devanes; Beattie, 1976. Rhop. Direct., p. 126.
Lycas devanes; Zhang et al., 2022. Tax. Rept. Intern. Lep. Surv. 10(7): 56, fig. 89 (genomic tree); LECTOTYPE, type locality South America, MfN.- Vila-Verde et al., 2024. EntomoBrasilis 17: e1097, p. 10.- Suênia-Bastos et al. 2025. Zootaxa 5604(2): 102 (table 1), figs 15-26 (male, female, d, v).
Taxonomic history. Goniloba devanes Herrich-Schäffer, 1869 was described based on an unknown number of specimens without locality. Evans (1955) treated it as a junior synonym of Lycas argentea. Recently, Zhang et al. (2022b), based on genomic data, revalidated Goniloba devanes, confirming that its distribution is restricted to South America, east of the Andes.
Type material. Grishin in Zhang et al. (2022b) designated the male syntype of Goniloba devanes, deposited in the MfN, as the lectotype. This specimen bears the following labels: / Typus / Gonil. devanes HS [Goniloba devanes Herrich-Schäffer] / Coll. Staudinger / QR code: http://coll.mfn-berlin.de/u, 449f7e / DNA sample ID: NVG-15036C11, c/o Nick V. Grishin /.
Lycas argenteus fa. bipunctata Hayward, 1934 was described based on the [male] “holotype” deposited in the MLP, with the following labels: / La Plata / Coll. Bryer / Rep. Argentina, Misiones / Tipus / Lycas argenteus fa. bipunctatus [sic] Holótipo, K. J. Hayward det. / 2560 /. According to Article 45.6.2 of the ICZN, this name is regarded as infrasubspecific and, thus, unavailable.
Lycas argenteus ab. albomarginata Kivirikko, 1936 was described based on a single male [“holotype”] deposited in the ZMH; labels are unavailable. According to Article 45.6.2 of the ICZN, this name is regarded as infrasubspecific and, thus, unavailable.
Diagnosis. FW length 19-22 mm. Similar to L. argentea, including the absence of a stigma in the male FW, but it can be distinguished from other species of the genus by the VFW usually presenting a broad, opaque, whitish spot in CuA2-2A in males, and greyish markings in the subapical area (Fig. 3). The male of Lycas devanes differs from L. argentea on the VHW by the discal silver band not connected to the submarginal band in central area, and the discal band interrupted by a brownish band in 2A-3A, thus not continuous from costal margin to the anal margin (Fig. 3B-C). In the female, VHW bands straight and never connected (Fig. 3E). Male genitalia (Fig. 9G-L) similar to those of L. argentea, exhibiting subtle differences: gnathos straight in lateral view; coecum of aedeagus curved dorsally; valva usually trapezoidal with a pointed posterior margin. Female genitalia (Fig. 12C-D) also provide reliable characters and more informative for distinguishing it from L. argentea: lamella antevaginalis with a narrow ventral sclerotization and a broad dorsal sclerotization, and lamella postvaginalis lacking a ventro-distal projection in the median portion.
Variation. Five male specimens (DZ 58.279, DZ 58.327, DZ 58.329, DZ 58.368, DZ 58.408) exhibited a slight connection between the two longitudinal bands on the VHW, but the proximal band remained interrupted by a brownish band (Fig. 3C).
Lycas devanes, dorsal and ventral views: (A-B) male DZ 58.338, from Água Mineral, Tibagi, Paraná, Brazil; (C) red arrow pointing to a variation of the silver band on the VHW; (D-E) female DZ 58.396, from Panambi, Rio Grande do Sul, Brazil. Scale bar: 10 mm.
Etymology. Not stated by the original author.
Geographical distribution and phenology. Lycas devanes is distributed throughout South America (Fig. 15), at elevations from 25 to 1,100 m, and has been recorded throughout the entire year. Records are based on collections and the literature: Peru - Madre de Dios (DZUP; OM; Garwood et al. 2007); Brazil - Acre (DZUP; OM), Maranhão (DZUP), Alagoas (DZUP), Mato Grosso (DZUP), Goiás (DZUP), Distrito Federal (DZUP), Minas Gerais (DZUP, OM), São Paulo (Brown 1992; Palo Jr. 2017), Rio de Janeiro (DZUP), Paraná (DZUP, OM; Ahlf et al. 2020), Santa Catarina (DZUP, OM; Orlandin et al. 2020), Rio Grande do Sul (DZUP); Paraguay - Guaíra (DZUP), Alto Paraná (DZUP), Iguazu (Uehara 2017); Argentina - Misiones (DZUP, OM; Hayward 1934; Kivirikko 1936; Canals 2003; Garwood et al. 2007), Córdoba (Hayward 1934), Corrientes (Klimaitis et al. 2018).
Material examined. In addition to the figures of the lectotype, 143 specimens were examined. PERU, Cuzco: (Cosñipata Valley, Quebrada Quitacalzón, 13°01’S, 71°30’W, 1,100 m), 1♂, 30-X-2016, O. Mielke leg., OM 82.422 (OM); Madre de Dios: (Parque Nacional del Manu, Pakitza, 340 m), 2♂, 1-X-1991, O. Mielke leg., G. Lamas leg., DZ 58.265, DZ 58.266 (DZUP); Alto Rio Madre de Dios (Amazonia Lodge, 2.5 km NO of Atalaya, Cuzco, 12°52’S, 71°22’W, 500 m), 1♂, 23-28-X-2013, O. Mielke leg., OM 75.495 (OM), (Albergue Pantiacolla, 7 km NE of Shintuya, 12°39’21”S, 71°13’53”W, 400 m), 1♂, 26-29-X-2016, Mielke leg., OM 81.869 (OM). BRAZIL, Acre: Bujari (50 km NO, 200 m), 3♂, 10-12-IX-2004, O. Mielke & Casagrande leg., DZ 58.267-DZ58.269 (DZUP); Porto Acre (Reserva Humaitá, 200 m), 4♂, 8-10-IX-2004, 28-31-VII-2008, O. Mielke & Casagrande leg., DZ 58.270-DZ58.273, DZ 74.856 (DZUP), Porto Acre (Reserva Humaitá, 9°45’02”S, 67°40’19”W, 180 m), 17-VII-2019, O. Mielke & Casagrande leg., DZ 74.857 (DZUP); Senador Guiomard (Reserva Catuaba), 3♂, 23-27-VII-2008, 1♂, 29-30-IX-2006, O. Mielke & Casagrande leg., DZ 74.853-DZ 74.855, DZ 74.858 (DZUP), Senador Guiomard (Reserva Catuaba, 10°4’27”S, 67°37’17”W, 200 m), 2♂, 27-5-VIII-IX-2011, D. Dolibaina & D. Moura leg., DZ 74.859, DZ 74.860; Rio Branco, 1♂, 30-IX-1985, Miers leg., OM 13.038 (OM); Maranhão: Imperatriz, 1♂, 4-VII-1974, 1♂, 24-VII-1974, 1♂, 6-VII-1974, Excursão Departamento Zoologia leg., DZ 58.274-58.276 (DZUP); Mato Grosso: Rosário do Oeste, 1♂, V-1969, DZ 58.278 (DZUP); Chapada dos Guimarães (Buriti, 600-800 m), 1♂, 17-I-1970, ex-coll. Nirton Tangerini, DZ 58.279 (DZUP); Diamantino (Fazenda São João, Alto Rio Arinos), 1♂, 24-II-1982, Furtado leg., 1♀, 2-IX-1978, O. Mielke & Furtado leg., DZ 58.280, DZ 58.281 (DZUP); Cáceres, 1♂, 12-XI-1984, Buzzi, O. Mielke & Casagrande leg., DZ 58.282 (DZUP); Nova Xavantina, 1♂, 10-I-1977, ex-coll. Gifford, DZ 58.284 (DZUP); Goiás: Nova Planaltina (Rio Maranhão), 2♂, 23-IX-2003, 2♂, 25-IX-2004, Eduardo Emery leg., ex-coll. Eduardo Emery, DZ 58.286-DZ 58.289 (DZUP); Distrito Federal: 1♀, 4-VII-1951, J. Becker leg., ex-coll. D’Almeida, DZ 58.285 (DZUP); Planaltina (Córrego Grotão), 1♀, 12-IX-2010, Eduardo Emery leg., ex-coll. Eduardo Emery, DZ 58.290 (DZUP); Minas Gerais: Paraisópolis (Cachoeira dos Martins), 1♂, 11-II-2002, N. Tangerini leg., ex-coll. Nirton Tangerini, DZ 58.294 (DZUP); Conceição dos Ouros, 1♂, III-2005, Almeida leg., 1♂, V-2017, Pereira leg., OM 68.147, OM 83.749 (OM); Carmo do Rio Claro, 1♀, 16-II-1959, Mielke leg., OM 2.731 (OM); Rio de Janeiro: 1♀, VIII-IX-1958, ex-coll. Richard Frey, 1♂, 1-VIII-1961, Gagarin leg., ex-coll. Gagarin, DZ 58.314, DZ 58.316 (DZUP); Nova Friburgo, 1♂, VII-2007, 1♂, VI-2005, 1♂, XI-2012, E. Grossi leg., DZ 58.305, DZ 58.307, DZ 58.309 (DZUP); Rio de Janeiro (Gávea), 1♂, 4-IX-1956, ex-coll. Richard Frey, (Represa Rio Grande), 1♀, V-1972, F. M. Oliveira leg., (Botafogo), 1♂, 8-VI-1936, ex-coll. D’Almeida, DZ 58.311, DZ 58.312, DZ 58.313 (DZUP); Duque de Caxias (Imbariê, 25 m), 1♂, 15-I-1970, H. Ebert leg., ex-coll. H. Ebert, DZ 58.315 (DZUP); Cachoeiras de Macacu (Boca do Mato), 1♀, 4-II-1998, N. Tangerini leg., ex-coll. Nirton Tangerini, DZ 58.317 (DZUP); Paraná: Foz do Iguaçu, 1♂, 10-XII-1966, 1♂, 2-XII-1966, D.Z.U.F.P. leg., 1♂, 21-VIII-1971, M. Lauterjung leg., (250 m), 1♀, 17-II-1969, Moure & O. Mielke leg., DZ 58.324, DZ 58.327, DZ 58.326, DZ 58.325 (DZUP), Foz do Iguaçu (Parque Nacional do Iguaçu, 180 m), 1♀, 23-VIII-1998, O. Mielke leg., 1♀, 21-24-IV-1995, O. Mielke & Casagrande leg., 1♀, 20-26-VIII-2000, O. Mielke leg., DZ 58.328, DZ 58.330 (DZUP), OM 51.391 (OM); Céu Azul (Parque Nacional do Iguaçu, Salto Rio Floriano, 210 m), 1♂, 8-IX-1998, O. Mielke leg., DZ 58.329 (DZUP); Campo Mourão (Parque Estadual Lago Azul, 500-600 m), 1♂, 9-11-X-2010, O. Mielke, Dolibaina, Carneiro & M. Maia leg., DZ 58.331 (DZUP); Ponta Grossa, 1♂, II-1959, 1♂, V-1941, ex-coll. Justus Jor, DZ 58.332, DZ 58.333 (DZUP), (20 km N, Piriquitos, 850 m), 1♀, 31-I-1990, O. Mielke leg., OM 25.139 (OM), (Furnas), 1♀, 28-II-1991, Mielke leg., OM 26.870 (OM); Londrina, 1♀, 10-X-1982, O. Mielke leg., 1♀, 25-X-1985, O. Mielke & Casagrande leg., DZ 58.334, 58.335 (DZUP); Guaíra, 1♀, 8-X-1982, O. Mielke leg., DZ 58.336 (DZUP); Tibagi (Água Mineral), 1♀, 2♂, 19-III-1991, O. Mielke & Casagrande leg., DZ 58.337-DZ 58.339 (DZUP); Chopinzinho, 1♂, XI-1969, 1♂, 10-V-1971, Furtado leg., DZ 58.340-DZ 58.341 (DZUP); Terra Boa (Companhia de Melhoramentos do Norte do Paraná), 1♂, 10-XII-1975, Moure, O. Mielke & Wedderhoff leg., DZ 58.343 (DZUP); Ventania (12.5 km N, 1,000 m), 1♂, 22-II-2006, O. Mielke & Casagrande leg., 1♂, 29-30-I-2006, O. Mielke leg., DZ 58.344, DZ 58.345 (DZUP); Campo Bonito (RPPN Hermínio & Maria, 24°49’42”S, 53°01’48”W), 1♀, 9-X-2012, Excursão LABLEP leg., DZ 58.346 (DZUP); Três Barras do Paraná (Parque Estadual Guarani, 25°26’21”S, 53°09’45”W), 2♂, 9-11-X-2012, Excursão LABLEP leg., DZ 58.349, DZ 58.350 (DZUP); São Pedro do Ivaí (RPPN Barbacena, 23°49’45”S, 51° 55’49”W), 1♂, 7-X-2012, Excursão LABLEP leg., DZ 58.351 (DZUP); Curitiba (Jardim das Américas), 1♀, 15-X-1998, Bizarro leg., DZ 58.352 (DZUP), (Passaúna), 1♀, 12-II-1993, Mielke leg., OM 34.558 (OM); Nova Laranjeiras (Rio das Cobras), 1♂, I-1972, L. Widerski leg., ex-coll. Justus Jr, DZ 58.553 (DZUP); Pinhão, 1♂, XII-1979, Schneider leg., OM 48.098 (OM); Lapa, 1♂, 9-IV-1992, Casagrande leg., OM 32.825 (OM); Campo Largo (15 Km NO, Bateias), 1♂, 11-III-2000, Mielke leg., OM 51.771 (OM); Pitanga, 1♂, 15-XII-2000, OM 58.925 (OM); Santa Catarina: Rio Negrinho (850 m), 1♂, 24-27-I-1986, O. Mielke & C. Mielke leg., DZ 58.354 (DZUP); São Bento do Sul (Rio Vermelho, 850 m), 1♀, 21-XII-2014, 2♂, 3-V-1985, 1♀, 12-IX-2014, 2♂, 11-V-1973, 1♂, 22-II-1985, 1♂,17-V-1973, 1♀, 20-V-1973, 1♂,11-II-1973, 1♂, 24-V-1973, 1♀, 26-V-1973, 1♀, 27-IV-1987, 1♀,11-I-2023, 1♀, 15-I-2024, 1♂, 15-VI-2024, Rank leg., DZ 58.372, DZ 58.355, DZ 58.368, DZ 58.356, DZ 58.357, DZ 58.365, DZ 58.358, DZ 58.359, DZ 58.361, DZ 58.362, DZ 58.363, DZ 58.365, DZ 58.367, (DZUP), OM 15.363 (OM), DZ 70.612, DZ 71.633, DZ 73.594 (DZUP), (Rio Natal, 500 m), 1♂, 19-II-2012, 1♂, 12-V-2010, 1♀, 5-IV-1989, 1♀, 8-V-2002, 1♀, 4-IX-2017, Rank leg., OM 21.587 (OM), DZ 58.374 (DZUP), OM 21.588, 58.610, 84.330 (OM); Seara (Nova Teutônia, 450-700 m), 1♂, II-1969, 1♂, VI-1972, (27°11’S, 52°23’W, 300-500 m), 1♀, V-1981, VI-1981, 1♂, F. Plaumann leg., DZ 58.376, DZ 58.377-DZ 58.379 (DZUP); Rio dos Cedros, 1♀, 8-VI-1973, M. Lauterjung leg., DZ 58.382 (DZUP); Taió, 1♀, 29-I-1966, Mielke leg., OM 7.930 (OM); Campo Alegre, 1♀, 14-IV-1990, Mielke leg., OM 25.321 (OM); Rio Grande do Sul: Pelotas, 2♀, 25-III-1967, N. Tangerini leg., 16-III-1967, C. M. Biezanko leg., ex-coll. Nirton Tangerini, DZ 58.388, DZ 58.389 (DZUP), (Monte Bonito), 1♀, 11-VI-1940, ex-coll. D’Almeida, DZ 58.402 (DZUP); Tenente Portela (Parque Florestal Estadual do Turvo), 2♂, 1♀, 10-XI-1985, O. Mielke, Araújo & Casagrande leg., DZ 58.390-DZ 58.392 (DZUP); Ivoti (250 m), 1♀, 7-IV-2000, 1♀,1-II-2002, A. Moser leg., DZ 58.393, DZ 58.394 (DZUP); Iraí, 1♂, 25-XI-1969, Schaal leg., DZ 58.395 (DZUP); Panambi, 1♀, III-1967, 1♂, IV-1969, Schaal leg., DZ 58.396, 58.397 (DZUP); São Luiz Gonzaga (Guarani), 1♂, 24-III-1941, 1♂, 13-V-1941, 1♀, 17-V-1940, 1♀, 19-VIII-1939, Padre Piton leg., ex-col. D’Almeida, DZ 58.398-DZ 58.401. PARAGUAY, Guairá: Villa Rica, 1♂, IV-V-1967, Anders leg., DZ 58.408 (DZUP); Alto Paraná: Itakyri (400 m), 3♀, 3♂, 15-20-I-1980, O. Mielke, C. Mielke & Miers leg., DZ 58.404, DZ 58.406, DZ 58.410, DZ 58.405, DZ 58.407, DZ 58.409 (DZUP), (Estancia Dimas, 25°33’S, 55°13’W), 1♀, 4-6-VI-2007, U. Dreschsel leg., OM 69.090 (OM). ARGENTINA, Misiones: (Parque Provincial Uruguai), 1♂, 28-VIII-1986, Genise leg., OM 40.440 (OM), (Parque Provincial Araucaria), 1♀, 13-I-2002, J. Klimaitis leg., OM 60.928 (OM); Dos de Mayo, 1♂, 10-X-1989, Foerster leg., OM 26.357 (OM); General Belgrano (Almirante Brown, Reserva Yacutinga), 1♂, 2-5-III-2007, O. Mielke & Casagrande leg., DZ 58.411 (DZUP); Puerto Iguazú, 1♂, VIII-2006, Nuñez-Bustos leg., DZ 58.412 (DZUP).
Lycas boisduvalii (Ehrmann, 1909)
Figs 1, 4, 7A-B, 10A-F, 12E-F, 16
Eudamus boisduvalii Ehrmann, 1909. Can. Ent. 41: 86; [holo]type male, 4-XI-1899, Suapure, Orinoco River, Venezuela, Klagen leg.; collection Ehrmann.- Holland, 1927. Ann. Carnegie Mus. 17: 337.
Epargyreus boisduvalii; Draudt, 1922, in Seitz. Gross-Schmett. Erde 5, p. 861.
Lycas godart boisduvali [sic]; Moss, 1949. Acta zool. Lill. 7: 75; hostplant.- Silva et al., 1968. Quarto Cat. Ins. viv. Plantas Brasil 2(1), p. 306; hostplant.- Ebert, 1968, in Silva et al. Quarto Cat. Ins. viv. Plantas Brasil 2(2), p. 234.
Lycas godarti [sic] boisduvalii; Evans, 1955. Cat. Amer. Hesp. 4, p. 262.
(no genus) boisduvalii; Beattie, 1976. Rhop. Direct., p. 97.
Lycas godart boisduvalii; Lamas, 1981. Rev. Soc. Mex. Lep. 6: 38.- de Jong, 1983. Tijd. Ent. 126: 247.- Bridges, 1983. Lep. Hesp. 1, p. 17; 2, p. 19.- Bridges, 1988. Cat. Hesp. 1, p. 27; 2, p. 31.- Lamas, 1994, in Foster et al. RAP Working Papers 6: 177.- Bridges, 1994. Cat. Fam.-Group, Gen.-Group, Sp.-Group Nam. Hesp. (Lep.) World 8, p. 32; 9, p. 35.- Robbins et al., 1996, in Wilson & Sandoval. Manu, p. 251.- O. Mielke, 2004. Hesperioidea, p. 71, in Lamas (ed.). Checklist: Part 4A, Hesperioidea-Papilionoidea, in Heppner (ed.). Atlas Neotrop. Lep. 5A; syn.: boisduvali.- O. Mielke, 2005. Cat. Amer. Hesperioidea 4, p. 1052; syn.: boisduvali.- Beccaloni et al., 2008. Monografías Tercer Milenio 8: 39; hostplant.- Bauder, Warren & Krenn, 2014. Jour. Res. Lep. 47: 68.- Grados et al., 2015, in Montoya et al. (eds). Parque Nac. Bahuaja Sonene. Invent. biol. rápidos. Anexo 13, p. [5].- Palo Jr., 2017. Borb. Brasil 3, p. 2317.- Khyade, 2019. World Sc. News 126: 110.- Cong et al., 2019. Insecta Mundi 731: 49, fig. 13 (genomic tree).- Zhang et al., 2022. Insecta Mundi 921: 95, 124, fig. 17 (genomic tree).- Suênia-Bastos et al. 2025. Zootaxa 5604(2): 102 (table 1), figs 19-20 (male, female, d, v).
Lycas godart godart [misidentifications]; Vargas, 2008. Bol. Cient. Mus. Hist. Nat. U. de Caldas 12: 209, fig. 1 (male v).
Lycas godart; Garwood & Lehman, 2013. Butt. C. Amer. 3, Hesp., p. 166, figs (male v).
Lycas boisduvalii; Murray, 2000. Jour. Res. Lep. 35: 53.
Lycas boisduvalii; Zhang et al., 2025. Tax. Rept. Intern. Lep. Surv. 12(8): 145, fig. 125.
Taxonomic history. Eudamus boisduvaliiEhrmann, 1909 was described based on the male [holo]type from Venezuela. This species was later transferred to Epargyreus Hübner, [1819] (Draudt 1922), and then to Lycas (Moss 1949). Later, Moss (1949) treated it as a subspecies of L. godart, a status maintained by Evans (1955). Most recently, Grishin in Zhang et al. (2025) treated L. boisduvalii as a distinct species from L. godart.
Type material. The male [holo]type of Eudamus boisduvalii deposited in the CMP bears the following labels: / E. boisduvalii Ehr. ♂. Type E.a. Klages Coll., 11/4/18[99], Suapure, Venez. [Venezuela] / Type / Ehrman Coll. Carn. Mus. Acc. 7815. /.
Diagnosis. FW length 22-25 mm. Lycas boisduvalii is distinct from L. godart by having smaller yellow hyaline spots on the FW, with the discal cell spot excavated at the outer margin and truncated at the inner margin (Fig. 4A, C), and VHW silver bands broader in most specimens. DFW stigma larger in CuA1-CuA2 than in L. godart and L. gabriel; its first part almost touches the origins of CuA1-CuA2, the second thicker, squared, and more robust than in other species, and the third rounded (Fig. 7A-B). VFW exhibits lilac markings in the submarginal area, usually between costal margin and CuA1 (Fig. 4B, D). Male genitalia (Fig. 10 A-F) with rounded tegumen not well delimited in dorsal view, and features a ventral projection; ventral arm of tegumen and dorsal arm of saccus rounded; gnathos straighter and longer than in L. godart and in L. gabriel; aedeagus with a coecum sometimes dorsally curved. Female genitalia (Fig. 12E-F) with the lamella postvaginalis composed of narrower lateral plates, forming a V-shape; lamella antevaginalis wider than the lamella postvaginalis, semi-sclerotized ventrally; the opening of the lamella antevaginalis square-shaped.
Variation. Some specimens exhibit variation in the number of apical hyaline spots on the FW; two specimens from Saül (French Guiana) and Acre (Brazil) possess only two hyaline spots, whereas the others have three. Additionally, the width of the silver bands varies: specimens from Brazil (Acre and Mato Grosso) and Peru (Cuzco and Madre de Dios) display broader silver bands, whereas specimens from Venezuela (Jusepín), French Guiana (Saül), and Brazil (Maranhão) show narrower silver bands.
Lycas boisduvalii, dorsal and ventral views: (A-B) male DZ 58.217, from 50 km NO Bujari, Acre, Brazil; (C-D) female DZ 58.233, from Rio Vermelho, Coronel Rio Branco, Cáceres, Mato Grosso, Brazil. Scale bar: 10 mm.
Etymology. Not stated by the original author. However, the name is a noun in genitive case that honors Jean Baptiste Boisduval, a renowned French lepidopterist.
Biology. Males and females were collected on flowers at 5:00 am in Coronel Rio Branco, Cáceres (Mato Grosso, Brazil).
Geographical distribution and phenology. Lycas boisduvalii is distributed throughout northern South America (Fig. 16), at elevations from 200 to 500 m, and has been recorded in January, February, April, and from June to November. Records are based on collections and original description: French Guiana (DZUP); Venezuela - Bolívar (Ehrmann 1909), Monagas (DZUP); Peru - Madre de Dios (DZUP, OM), Cuzco (OM), San Martín (OM); Brazil - Acre (DZUP), Maranhão (DZUP), Mato Grosso (DZUP).
Material examined. In addition to the figure of the [holo]type, 121 specimens were examined. French Guiana, Saint-Laurent-du-Maroni: Saül, 1♂, 27-IV-2011, S.E.A.G. leg, OM 74.080 (OM), (Belvédère), 1♂, 11-VIII- 2010, 1♂, 4-IX-2010, 2♂, 5-IX-2010, S.E.A.G. leg., DZ 58.188-DZ 58.191 (DZUP).
VENEZUELA, Monagas: Jusepín, 1♂, 10-IX-1965, 1♂, 2♀, 8-X-1965, F. Fernándes-Y. & C. J. Rosales leg., DZ 58.192-DZ 58.195 (DZUP). PERU, Cuzco: Cuzco (Cosñipata Valley, Pilcopata, Villa Carmen, 12°53’41”S, 71°24’12”W, 500 m), 1♂, 2♀, 31-2-I-II-2020, O. Mielke leg., OM 90.587, OM 90.567, OM 90.567 (OM), (Cosñipata Valley, Chontachaca, 12°56’S, 71°24’W, 950 m), 2♂, 12-X-2024, O. Mielke leg., OM 95.044, OM 95.269 (OM); Madre de Dios: (Parque Nacional del Manu, Pakitza, 11°55’48”S, 71°15’18”W, 340 m), 1♀, 1-X-1991, G. Lamas leg., 1♂, 26-IX-1991, DZ 58.196, DZ 58.199 (DZUP); Alto Rio Madre de Dios (Amazonia Lodge, 2.5 km NO of Atalaya, Cuzco, 12°52’S, 71°22’W, 500 m), 2♂, 23-28-X-2013, 1♂, 29-IV-2015, O. Mielke leg., OM 76.452, OM 76.462, OM 77.432 (OM), (1 km NE Atalaya, 12°52’S, 71°22’W, 500 m), 3♂, 7-11-XI-2012, OM 74.033, OM 74.043, OM 73.913 (OM), (Albergue Pantiacolla, 7 km NE of Shintuya, 12°39’21”S, 71°13’53”W, 400 m), 3♂, 26-29-X-2016, 2♂, 26-29-X-2016, 1♂, 4-8-XI-2017, 5♂, 14-18-X-2024, O. Mielke leg., OM 81.819, OM 81.829, OM 81.866, OM 86.024, OM 81.719, OM 86.024, OM 95.381-OM 95.383, OM 95.393, OM 95.394 (OM); San Martín: Moyobamba (6°02’13”S, 76°58’22”W, 800 m), 1♂, X-2016, J. Pintado leg., OM 80.998 (OM), (Calzada, 6°01’51”S, 77°03’54”W, 850 m), 1♂, 6, X-2016, J. Pintado, OM 83.626 (OM). BRAZIL, Acre: Porto Acre (Reserva Humaitá, 200 m), 9♂, 8-10-IX-2004, 2♂, 28-31-VII-2008, O. Mielke & Casagrande leg., 2♂, 16-VII-2004, O. & C. Mielke leg., DZ 58.201-DZ 58.208, DZ 58.210, DZ 74.861, DZ 74.862 (DZUP), OM 64.973, OM 65.022 (OM); Porto Acre (Reserva Humaitá, 9°45’02”S, 67°40’19”W, 180 m), 1♂ 17-VII-2019, O. Mielke & Casagrande leg., DZ 74.872 (DZUP); Bujari (50 km NO, 200 m), 2♂, 18-20-IX-2003, 7♂, 10-12-IX-2004, O. Mielke & Casagrande leg., 15♂, 20-23-VII-2004, O. & C. Mielke leg., DZ 58.211, 58.212, DZ 58.214, DZ 58.216-DZ 58.219 (DZUP), OM 65.565, OM 65.537, OM 65.460, OM 65.691, OM 65.663, OM 65.698, OM 65.642, OM 65.789, OM 65.635, OM 65.726, OM 65.523, OM 65.600, OM 65.670, OM 65.831, OM 65.670 (OM); Senador Guiomard (Reserva Catuaba, 200 m), 8♂, 2-5-IX-2004, 10♂, 23-27-VII-2008, O. Mielke & Casagrande leg., DZ 58.220-DZ 58.223, DZ 58.225, DZ 58.227, DZ 58.228, DZ 58.230, DZ 74.863-DZ 74.871, DZ 74.889 (DZUP), Senador Guiomard (Reserva Catuaba, 10°04’27”S, 67°36’15”W, 200 m), 4♂, 16, 18-21-VIII-IX-2011, O. Mielke & Casagrande leg., DZ 74.873-DZ 74.876 (DZUP), Senador Guiomard (Reserva Catuaba, 10°04’27”S, 67°37’17”W, 200 m), 12♂, 27-5-VIII-IX-2011, D. Dolibaina & D. Moura leg., DZ 74.877-DZ 74.888 (DZUP); Maranhão: Açailândia, 1♂, 23-VIII-1974, O. Mielke leg., DZ 58.232 (DZUP); Mato Grosso: Cáceres (Coronel Rio Branco, 400 m), 1♂, 30-VI-1972, 2♀, 1♂, 3-VII-1972, 10♂, 9♀, 2-VII-1972, 1♂, 4♀, 1-VIII-1972, DZ 58.249, DZ 58.240, DZ 58.244-DZ 58.245, DZ 58.234-DZ 58.236, DZ 58.247-DZ 58.248, DZ 58.250, DZ 58.255-DZ 58.258, DZ 58.233, DZ 58.242, DZ 58.243, DZ 58.246, DZ 58.251-DZ 58.254, DZ 58.259, DZ 58.239, DZ 58.237, DZ 58.238, 58.241 (DZUP).
Lycas godart (Latreille, [1824])
Figs 1, 5, 7C-D, 10G-L, 13A-B, 16
Hesperia godart Latreille, [1824], in Latreille & Godart. Enc. Méth. 9, p. 722, 762; 1 male [holotype], Brazil, Langsdorff leg.; collection Latreille.- Viette, 1956. Lambilleonea 56: 92; TYPE BM(NH).
Goniloba godartii [sic]; Westwood, 1852, in Doubleday & Westwood. Gen. Diurn. Lep. 2, p. 514.
Hesperia ceraca Hewitson, 1866. Trans. ent. Soc. London (3)2: 488; Rio de Janeiro, [Brazil]; collection Hewitson.- Hewitson, 1872. Illustr. Exot. Butt. 5, p. 101, pl. 53 (Hesperia 5), figs 42, 43 (d, v).- Kirby, 1879. Cat. Coll. Diurn. Lep. Hewitson, p. 217.
Goniloba ceraca; Herrich-Schäffer, 1869. Corr.-Blatt zool.-min. Ver. Regensburg 23: 193.
Carystus godartii [sic]; Kirby, 1871. Syn. Cat. Diurn. Lep., p. 591.
Proteides ceraca; Kirby, 1871. Syn. Cat. Diurn. Lep., p. 593.- Möschler, 1877. Verh. zool.-bot. Ges. Wien 26: 332.- Kirby, 1877. Syn. Cat. Diurn. Lep., Suppl., p. 823.
Hesperia godartii [sic]; Plötz, 1882. Stett. ent. Ztg. 43: 333; syn.: ceraca.
Lycas ceraca; Godman, 1901, in Godman & Salvin. Biol. Centr.-Amer., Lep.-Rhop. 2, p. 619.
Thracides godartii [sic]; Mabille, 1904, in Wytsman. Gen. Ins. 17, p. 179.
Lycas ceraca; Mabille, 1904, in Wytsman. Gen. Ins. 17, p. 167.
Lycas godarti [sic]; Draudt, 1923, in Seitz. Gross-Schmett. Erde 5, p. 991, pl. 191a (d, v); syn.: ceraca.- Riley, 1926. Trans. ent. Soc. London 74: 237; syn.: ceraca.- Holland, 1927. Ann. Carnegie Mus. 17: 338; syn.: boisduvalii, godart, caraca.- J. Zikán, 1928. Ent. Rdsch. 45: 35.- Spitz, 1932. Rev. Mus. Paulista 17: 878.- F. Hoffmann, 1934. Ent. Rdsch., Stuttgart, 51: 73.- J. Zikán & W. Zikán, 1968. Pesq. agropec. bras. 3: 60.- Hayward, 1969. Inst. Miguel Lillo, Misc. 31: 87.
Epargyreus godarti [sic]; Shepard, 1931. Lep. Cat. 47, p. 51; syn.: ceraca, boisduvalii.
Lycas godart; Hayward, 1941. Rev. Mus. La Plata, n. s., Zool. 2: 322; syn.: ceraca, godarti.- Bell, 1946. Bol. Ent. venezol. 5: 180; syn.: ceraca, boisduvalii.- Garwood et al., 2007. Butt. South. Amaz., p. 232, fig. (v).- Beccaloni et al., 2008. Monografías Tercer Milenio 8: 39; hostplant.- Pérez et al., 2017. Shilap Revta. lepid. 45(179): 441.- Sambhu & Nankishore, 2018. Zootaxa 4371(1): 26.
Lycas godarti [sic] godarti [sic]; Evans, 1955. Cat. Amer. Hesp. 4, p. 262, pl. 71 (male gen.); syn.: ceraca.- Hayward, 1973. Op. Lill. 23; 86; syn.: godart, caraca.- K. Brown, 1987. An. Prim. Simp. Pantanal, Brasília, p. 168.- Canals, 2003. Marip. Misiones, p. 459.
Lycas caraca; Hayward, 1973. Op. Lill. 23; 86; as synonym of Lycas godarti [sic] godarti [sic].
(no genus) godarti [sic]; Beattie, 1976. Rhop. Direct., p. 153.
(no genus) ceraca; Beattie, 1976. Rhop. Direct., p. 108.
Lycas godart godart; Bridges, 1983. Lep. Hesp. 1, p. 23, 49; 2, p. 19; syn.: godarti, ceraca.- Bridges, 1988. Cat. Hesp. 1, p. 37, 77; 2, p. 31; syn.: ceraca, godarti.- Bridges, 1994. Cat. Fam.-Group, Gen.-Group, Sp.-Group Nam. Hesp. (Lep.) World 8, p. 32, 44, 92; 9, p. 35; syn.: ceraca, caraca.- K. Brown & Freitas, 2000. Bol. Mus. Biol. Mello Leitão, n. s., Sta. Teresa, 11/12: 113.- O. Mielke, 2004. Hesperioidea, p. 71, in Lamas (ed.). Checklist: Part 4A, Hesperioidea-Papilionoidea, in Heppner (ed.). Atlas Neotrop. Lep. 5A; syn.: godartii, ceraca, godarti.- Pastrana, 2004. Lep. arg., Plantas hosp., p. 207; hostplant.- O. Mielke, 2005. Cat. Amer. Hesperioidea 4, p. 1051; syn.: godartii, ceraca, godarti.- O. Mielke, Carneiro & Casagrande, 2010. Biota Neotrop. 10(4): 297.- Palo Jr., 2017. Borb. Brasil 3, p. 2317.- Orlandin et al., 2022, in Greca de Macedo & Dias. Portaria Nº 1082, Pref. Curitiba, p. 62.- Suênia-Bastos et al. 2025. Zootaxa 5604(2): 102, figs 17-18 (male, female, d, v).
Taxonomic history. Hesperia godartLatreille, [1824] was described based on a single male [holotype] from Brazil. Westwood (1852, in Doubleday and Westwood) described Goniloba and transferred H. godart to it. Hesperia ceraca was described by Hewitson (1866) based on an unknown number of specimens from Rio de Janeiro, Brazil. Subsequently, H. ceraca was transferred to Goniloba (Herrich-Schäffer, 1869) and to Proteides Hübner, [1819] (Kirby 1871), while H. godart was transferred to Carystus Hübner, [1819] (Kirby 1871). Plötz (1882) treated H. ceraca as a junior synonym of H. godart. Godman (1901), when described Lycas, included H. ceraca based on a female from Panama and two other specimens from South America. Mabille (1904) transferred H. godart to Thracides Hübner, [1819]. Draudt (1923) recognized that both were the same species and synonymized H. ceraca with H. godart. Shepard (1931) synonymized Eudamus boisduvalii with H. godart (misspelled as godarti) and transferred it to Epargyreus Hübner, [1819]. Hayward (1941) maintained H. godart in Lycas, as first proposed by Draudt (1923). Evans (1955) accepted the synonymies proposed by Plötz (1882), Draudt (1923), and Hayward (1941), and treated E. boisduvalii as a subspecies of L. godart.
Type material. The male [holotype] of Hesperia godart deposited in the NHMUK bears the following labels: / Type / I-I. Godartii Lat. Bresil / R. Oberthur Coll. Brit. Mus. 1931-136 /.
One female syntype of Hesperia ceraca deposited in the NHMUK bears the following labels: / Type / Type / Rio R. / ceraca / Rio R. Hewitson Coll. 79-69 2. Hesperia ceraca, Hew /. This syntype is herein designated as lectotype to fix the identity of the species, and bears the following labels: / LECTOTYPUS / Lectotype Hesperia ceraca Hewitson, 1866 Suênia-Bastos, Mielke & Casagrande det. 2025 /.
Diagnosis. FW length 23-27 mm. Easily distinguished from all other Lycas species by the large yellow hyaline spots on the FW (Fig. 5A, C), and the discal cell spot not as strongly excavated as in L. boisduvalii (Fig. 4A, C). DFW stigma narrower in CuA1-CuA2 than in L. boisduvalii, but larger than in L. gabriel; second part of stigma in the upper half of CuA2-2A almost touches CuA2 (Fig. 7C-D). VFW lacks lilac markings in the subapical area (Fig. 5B, D). DHW features a rounded, opaque, yellow spot between M1-M3. VHW bands narrower than those of L. boisduvalii. Although this species inhabits a different biome and its wing pattern differs from that of L. gabriel, male genitalia (Fig. 10G-L) similar to those of the latter, usually differentiated by a curved gnathos and a narrow aedeagus coecum with a rounded anterior portion. Female genitalia (Fig. 13A, B) larger than those of other species, and the lamella postvaginalis consists of lateral plates wider than those of L. boisduvalii; lamella antevaginalis oval-shaped, slightly sclerotized ventro-laterally, and narrower than in L. boisduvalii. It is similar to L. gabriel, with lamella postvaginalis formed by two well-sclerotized plates that form a narrow, triangle-shaped space in the median portion.
Lycas godart, in dorsal and ventral views: (A-B) male DZ 58.167; (C-D) female DZ 58.164, both specimens from Joinville, Santa Catarina, Brazil. Scale bar: 10 mm.
Etymology. The name used by Latreille honors Jean-Baptiste Godart, who, according to the original author, assisted with the preparation of the Encyclopédie. The name is a noun in apposition.
Geographical distribution and phenology. Lycas godart is restricted to the southern Atlantic Forest (Fig. 16), at elevations from 10 to 1,200 m, and has been recorded throughout the entire year. Records are based on collections and the literature: Brazil - Minas Gerais (OM), São Paulo (DZUP), Rio de Janeiro (DZUP; OM; Hewitson 1866), Paraná (DZUP, OM), Santa Catarina (DZUP, OM).
Material examined. In addition to the figures of the holotype of Hesperia godart and the lectotype of Hesperia ceraca, herein designated, 51 specimens were examined. BRAZIL, Minas Gerais: Conceição dos Ouros, 1♂, 19-IX-2021, Pereira leg., OM 87.205 (OM); São Paulo: São Paulo (Ipiranga), 1♀, 12-V-1941, Travassos Filho leg., ex-coll. D’Almeida, DZ 58.148 (DZUP); Rio de Janeiro: 1♀, no data, OM 11.304; Petrópolis (900 m), 1♂, 6-IX-1959, Gagarin leg., ex-coll. Gagarin, DZ 58.152 (DZUP), (Independência, 900 m), 1♀, 2-XI-1934, Gagarin leg., ex-coll. Gagarin, DZ 58.149 (DZUP), 1♀, 27-VIII-1939, Gagarin leg., DZ 58.150 (DZUP), 1♂, 20-V-1965, O. Mielke leg., OM 6.997 (OM); Rio de Janeiro (Parque Nacional da Tijuca), 1♂, 3-V-1936, R. D’Almeida & Nelson D’Almeida leg., ex-coll. D’Almeida, DZ 58.151 (DZUP); Nova Friburgo, 1♂, 10-VIII-2005, E. Grossi leg., DZ 58.153 (DZUP); Itatiaia (Parque Nacional do Itatiaia, 1,000-1,200 m), 1♂, 25-VIII-1963, (880 m), 1♂, 13-I-1969, 1♂, 16-IV-1965, H. Ebert leg., ex-coll. H. Ebert, DZ 58.154, DZ 58.155, DZ 58.156 (DZUP); Cachoeiras de Macacu (Boca do Mato), 1♀, 15-V-2008, Jonas Overney leg., ex-coll. Nirton Tangerini, DZ 58.157 (DZUP); Paraná: Antonina (Reserva Natural Guaricica), 1♂, 31-4-I-II-2022, A. P. Pinto leg., DZ 58.158 (DZUP); Curitiba (Centro Politécnico), 1♀, 27-XI-2002, O. Mielke leg., OM 58.860 (OM); Morretes (Marumbi), 1♂, 18-VI-1966, O. Mielke leg., OM 9.609 (OM); Santa Catarina: Joinville, 1♀, 8-IV-1968, Miers leg., DZ 58.159 (DZUP), (10-200 m), 1♀, 23-II-1973, 1♂, 9-III-1973, 1♀, 23-III-1973, 1♂, 10-VII-1973, 1♀,7-X-1973, 1♀, 25-IX-1975, 1♀, 4-II-1976, 1♂, 17-III-1976, 1♀, 2-IV-1977, 1♂, 1-XII-1990, 1♀, 4-XII-1991, 1♂, 26-II-1991, 1♂, 12-I-1992, 1♂, (without day, month and year), 1♂, 7-IX-1972, Miers leg., DZ 58.164, DZ 58.166, DZ 58.169, DZ 58.167, DZ 58.163, DZ 58.161, DZ 58.160, DZ 58.165, DZ 58.168 (DZUP), 1♀,16-X-1971, 1♂, 22-V-1969, O. Mielke & Miers leg., DZ 58.170, DZ 58.173 (DZUP), 1♀, 21- VIII-1972, 1♀, 2- XII-1978, 1♂, O. Mielke leg., DZ 58.171, DZ 58.172, (DZUP), OM 26.946, OM 28.937, OM 27.104, OM 28.893, OM 66.878, OM 66.871 (OM); São Bento do Sul (Rio Vermelho, 850 m), 1♀, 11-II-1973, 1♂, 11-III-1973, Rank leg., DZ 58.175, 58.176 (DZUP), 1♂, 25-IV-1985, Rank leg., DZ 58.174 (DZUP), (Rio Natal, 500 m), 1♂, 21-II-1991, 1♂, 9-V-200, 1♂, 4-II-2009, 2♂, 18-II-2009, 1♀, 3-IX-2014, 1♂, 11-XII-2014, 1♀, 2-II-2015, 1♂, 7-II-2015, 1♂, 20-II-2015, 1♂, 6-VIII-2015, Rank leg., OM 26.400, OM 51.501 (OM), DZ 58.179, DZ 58.177, DZ 58.178, DZ 58.180, DZ 58.181, DZ 58.185, DZ 58.182, DZ 58.184, DZ 58.183 (DZUP).
Lycas gabriel Grishin, 2025
Lycas godart [misidentification]; Garwood et al., 2009. Butt. South. Amaz., 2nd ed., p. 319, 320, figs (d, v).
Lycas godart boisduvalii [misidentification]; Uehara, 2020. Butterflies 84: 55, fig. 107 (male d, v).
Lycas gabriel Grishin, 2025, in Zhang et al. Tax. Rept. Intern. Lep. Surv. 12(8): 145, figs 125 (nuclear genome and mitochondrial genome trees), 126 (female d, v); holotype female, Peru, Madre de Dios Region, 30 km SW of Puerto Maldonado, 300 m, 27-X-1983, S. S. Nicolay leg.; USNM.
Taxonomic history. Lycas gabriel Grishin, 2025 was described through genomic analysis based on a single female from Peru.
Type material. The female holotype of Lycas gabriel, deposited in the USNM, bears the following labels: / PERU, 300 m, 30 km S.W., Pto. [Puerto] Maldonado, 27 Oct. ’83, S. S. Nicolay / Lycas ♀ argentea, Det. Hew., S. S. Nicolay / DNA sample ID: NVG-23122G07, c/o Nick V. Grishin / HOLOTYPE ♀ Lycas gabriel Grishin /.
Diagnosis. This species is remarkable for its resemblance to two other congeners: the male wing pattern, in both shape and number of hyaline spots (Fig. 6A, E), resembles L. boisduvalii from Amazonia, whereas the male genitalia (Fig. 11A-F) closely resemble those of L. godart from the southern Atlantic Forest. This species can be easily distinguished from L. godart by the smaller hyaline spots on FW (Fig. 6A, E), and from L. boisduvalii by the narrower stigma (Fig. 8). In contrast, the female wing pattern (Fig. 6C-D) similar to both L. argentea and L. devanes, while the genitalia (Fig. 13C-D) similar to those of L. godart. However, females are distinguished from L. argentea and L. devanes by the genitalia, which, although similar to those of L. godart, feature a well-sclerotized lamella postvaginalis formed by two plates with median portion folded ventrally and square-shaped in median portion (Fig. 13C-D), and an entirely membranous lamella antevaginalis (with an inconspicuous lateral sclerotized line) with a square-shaped opening to the ostium, whereas it is rounded in L. godart.
Male description. Head: antenna dorsally and ventrally brownish, club and apiculus yellowish; nudum with 14-18 segments.
Thorax: FW length 21-26 mm. FW brownish, with six small yellow hyaline spots: one in the discal cell (excavated outwardly, with the upper half longer than the lower half), three apical located between R3 and M1, two discal in M3-CuA1 and CuA1-CuA2, and one opaque spot in CuA2-2A (Fig. 6A-B). DFW stigma narrower than in other species between CuA1 and 2A (Fig. 8), composed of three parts: (1) a narrow one not reaching the origin of CuA1-CuA2; (2) a small below CuA2, almost touching the vein; and (3) a nearly inconspicuous line below the second part in the same space. VFW central area brownish; marginal, apical, and costal areas ferruginous, with lilac markings along the costal margin and submarginal area, usually between costal margin and CuA1 (Fig. 6B, F); an opaque yellowish square spot in CuA2-2A, sometimes larger than that shown in the figure (Fig. 6B); fringes brownish; marginal line brownish. DHW brownish, with a yellowish spot between M1 and M3, sometimes subtly divided, and costal margin yellowish; anal margin light brownish; fringes mixed brownish and yellowish (Fig. 6A). VHW brownish; marginal line brownish; fringes mixed with brownish and yellowish; two longitudinal silver bands extending from costal to tornus (Fig. 6B), discal band extending from costal margin to above or below CuA2, and a submarginal band between apex of the costal margin and below CuA2, whether faded or not when reaching below CuA2; a continuous band along the anal margin.
Abdomen: Dorsally brownish, ventrally yellowish, with a continuous central brownish stripe. Male genitalia (Fig. 11) with tegumen clearly separated from uncus dorsally; ventral arm of tegumen obtusely inclined; anterior projection of saccus longer than the uncus, completely straight or tiny curved dorsally; uncus simple, truncate (Fig. 11A shows a concavity, which is actually an artifact of the angle of the photograph), almost reaching the tip of the valva; gnathos shorter than the uncus, straight, thumb-shaped; fultura inferior wider posteriorly than anteriorly, anteriorly straight or concave; valva rectangular and distally pointed, varying in width; aedeagus with coecum slightly or strongly curved dorsally (Fig. 11F, L).
Female redescription. FW length 20 mm, smaller than males. Head: Antenna dorsally brownish, ventrally yellowish; nudum with 16 segments.
Thorax: FW brownish with seven small yellow hyaline spots: one in discal cell (excavated outwardly), three apical between R3 and M1, three discal in M3-CuA1, CuA1-CuA2, and in the upper half of CuA2-2A, and an opaque spot in the lower half of CuA2-2A (Fig. 6C-D). VFW central area brownish, with slight lilac markings on costal margin and submarginal area (see Zhang et al. 2025); this region appears damaged, and it is possible that the scales have rubbed off the wing in the female specimen deposited in the DZUP (Fig. 6D). DHW brownish, with a yellow opaque spot between M1 and M3; costal margin yellowish; fringes yellowish, only apparent in the female deposited in the DZUP. VHW brownish; marginal line indistinct; fringes yellowish; two longitudinal opaque silver bands extending from costal to tornus, the discal band between costal margin and 2A strongly or slightly directed toward outer margin, and a submarginal band between costal margin and below CuA2; a continuous band along the anal margin.
Lycas gabriel, dorsal and ventral views: (A-B) male DZ 58.213, from 50 km NO of Bujari, Acre, Brazil; (C-D) female DZ 58.283, from Tangará da Serra, Mato Grosso, Brazil. (E-F) Different phenotype of Lycas gabriel, dorsal and ventral views of the male OM 95.395, from Albergue Pantiacolla (3.3 km NE of Shintuya, Cusco), Madre de Díos, Peru. Scale bar: 10 mm.
Forewing stigma morphology: (A-C) Lycas gabriel; (D-E) Lycas gabriel, different phenotype. Scale bar: 2 mm.
Male genitalia of Lycas argentea, DZ 58.260: (A) dorsal view; (B) lateral view; (C) ventral view; (D) valva, inner view; (E-F) aedeagus in ventral and left lateral views. Male genitalia of Lycas devanes, DZ 58.305: (G) dorsal view; (H) lateral view; (I) ventral view; (J) valva, inner view; (K-L) aedeagus in ventral and left lateral views. Scale bar: 1 mm.
Male genitalia. (A-F) Lycas boisduvalii, DZ 58.258: (A) dorsal view; (B) lateral view, white arrow indicating some diagnostic characters; (B) ventral view; (D) valva, inner view; (E, F) aedeagus in ventral and left lateral views. (G-L) Lycas godart, DZ 58.155: (G) dorsal view; (H) lateral view; (I) ventral view, white arrow indicating the dorsal arm of saccus extremely curved; (J) valva, inner view; (K-L) aedeagus in ventral and left lateral views. Scale bar: 1 mm.
Male genitalia. (A-F) Lycas gabriel, (DZ 58.198), white arrow indicating the diagnostic character: (A) dorsal view; (B) lateral view; (C) ventral view; (D) valva, inner view; (E, F) aedeagus in ventral and left lateral views. (G-L) Male genitalia of the different phenotype, OM 95.395: (G) dorsal view; (H) lateral view; (I) ventral view; (J) valva, inner view; (K-L) aedeagus in ventral and left lateral views. Scale bar: 1 mm.
Female genitalia, ventral and lateral views: (A-B) Lycas argentea, DZ 58.320, white arrow indicating the projection on the lamella postvaginalis in median portion; (C-D) Lycas devanes, DZ 58.323, white arrow indicating the absence of a projection on the lamella postvaginalis in median portion; (E-F) Lycas boisduvalii, DZ 58.195. Scale bar: 2 mm.
Female genitalia, ventral and lateral views: (A-B) Lycas godart, DZ 58.185, white arrow indicating the lamella postvaginalis with a narrow triangle-shape in the median portion; (C-D) Lycas gabriel, DZ 58.283, white arrow indicating the lamella postvaginalis square-shaped in the median portion. Scale bar: 2 mm.
Distribution of Lycas species: (14) Lycas argentea; (15) Lycas devanes; (16) Lycas godart and Lycas boisduvalii; (17) Lycas gabriel. The points with a black dot represent data from the literature.
Abdomen: Dorsally brownish, ventrally whitish with a central brownish line. Female genitalia (based only on the female deposited in the DZUP, Fig. 13C-D): lamella postvaginalis composed of two plates with median portion folded ventrally, similar to L. godart; lamella antevaginalis entirely membranous (with an inconspicuous lateral sclerotized line), opening square-shaped and broader than in L. godart; ductus bursae showing a clear distinction between the ductus and the corpus bursae.
Etymology. See Zhang et al. (2025).
Variation. The hyaline spots on the FW vary in size, especially the apical ones. One male specimen presents only two apical spots between R3 and M1. Additionally, the first part of the stigma sometimes appears slightly larger (OM 85.974, OM 95.378, DZ 58.200, DZ 58.209, DZ 58.215, DZ 58.226, DZ 58.231, DZ 74.893, DZ 74.894) (Fig. 8C), similar to the phenotype mentioned below. The yellowish spot between M1 and M3 on the DHW also varies in size, from a complete spot to a subtly divided one. As observed in L. boisduvalii, the width of the silver bands on the VHW varies from broad to narrow in specimens from the same locality. Regarding the male genitalia, the curvature of the coecum is variable, as is the anterior projection of the saccus; however, as mentioned in the diagnosis, the inclination of the ventral arm of the tegumen is a stable character. One specimen (OM 95.395) (Fig. 6E-F) displays a distinct phenotype from the remaining L. gabriel, with FW discal cell spot reduced to two points and the male genitalia (Fig. 11G-L) exhibiting an almost horizontal ventral arm of the tegumen. On the other hand, the male FW stigma (Fig. 8D-E) is similar to some L. gabriel specimens, and in the phylogenetic tree (Fig. 1) is grouped with L. gabriel. Since there is only a single specimen with this subtle difference, thus it will be provisionally interpretated as a strong intraspecific variation in L. gabriel.
Geographical distribution and phenology. Lycas gabriel is restricted to the Amazonian region (Fig. 17), at elevations from 200 to 400 m, and has been recorded from June to November. Records are based on collections and the literature: Peru - Madre de Dios (DZUP; OM; Uehara 2020; Zhang et al. 2025); Brazil - Amazonas (DZUP), Acre (DZUP; OM), Mato Grosso (DZUP).
Material examined. In addition to the figure of the holotype of L. gabriel, 23 specimens were examined. PERU, Madre de Dios: (Parque Nacional del Manu, Pakitza, 11°55’48”S, 71°15’18”W, 340 m), 1♂, 8-X-1991, G. Lamas leg., DZ 58.197, (DZUP), (Reserva Tambopata, 12°50’S, 69°17’W, 300 m), 1♂, 25-X-1991, O. Mielke leg., DZ 58.198 (DZUP); Alto Río Madre de Dios (Albergue Pantiacolla, 7 km NE of Shintuya, 12°39’21”S, 71°13’53”W, 400 m), 2♂, 4-8-XI-2017, 1♂, 18-22-VI-2019, 1♂, 14-18-X-2024, O. Mielke leg., OM 86.034, OM 85.974, OM 88.857, OM 95.378, OM 95.395 (OM). BRAZIL, Amazonas: Manaus, 1♂, DZ 58.200 (DZUP); Acre: Porto Acre (Reserva Humaitá, 200 m), 1♂, 8-10-IX-2004, O. Mielke & Casagrande leg., DZ 58.209 (DZUP); Bujari (50 km NO, 200 m), 2♂, 10-12-IX-2004, 1♂, 20-23-VII-2004, O. Mielke & Casagrande leg., DZ 58.215 (DZUP), OM 65.796 (OM), DZ 58.213 (DZUP); Senador Guiomard (Reserva Catuaba, 10°4’27”S, 67°37’17”W, 200 m), 3♂, 2-5-IX-2004, 5♂, 23-27-VII-2008, 1♂, 16,18-21-VII-2019, O. Mielke & Casagrande leg., 1♂, 27-5-VIII-IX-2011, D. Dolibaina & D. Moura leg., DZ 58.224, DZ 58.226, DZ 58.229, DZ 74.891-DZ 74.895, DZ 74.896, DZ 74.890 (DZUP), Santa Rosa do Purus (9.2 km SE), 1♂, 6-8-VIII-2008, O. Mielke & Carneiro leg., DZ 58.231 (DZUP); Mato Grosso: Tangará da Serra, 1♀, 1-VI-2014, B. F. Camera & D. Krinski leg., DZ 58.283 (DZUP).
DISCUSSION
The phylogeny presented in this study recovers Lycas as a strongly monophyletic clade (Fig. 1), encompassing all recognized species of the genus, including the newly described taxon. Previous studies (Cong et al. 2019, Zhang et al. 2022a) focused on relationships at the tribal or generic levels and included fewer representative taxa of Lycas; nevertheless, the genus was consistently recovered with strong support in all analyses. In the present study, L. devanes is recovered as sister to all other Lycas species, rather than clustering with the morphologically similar L. argentea, differing from the results obtained by Zhang et al. (2025), which L. argentea is siter to L. devanes in both nuclear and mitochondrial genomes trees. In contrast, L. argentea is recovered as sister to L. boisduvalii + L. gabriel, although with weak support (Fig. 1). Morphological evidence, however, suggests a closer affinity between L. argentea and L. devanes, especially with respect to the wing pattern in both sexes and the absence of a stigma on the male forewings. Similar to the topology obtained by Zhang et al. (2025) using mitochondrial genomes, L. gabriel is here recovered as sister to L. boisduvalii.
Zhang et al. (2022b) sequenced the type specimen of Goniloba devanes (type locality unknown) and specimens of Lycas argentea, revealing that the South American clade represents a species distinct from the North and Central American clade. They recognized the latter as L. argentea, whereas the South American clade corresponded to L. devanes. Our morphological examination expanded the known distribution of L. argentea from North and Central America to South America, where it occurs sympatrically with L. devanes in southern and southeastern Brazil, as well as in Argentina and Paraguay. As illustrated in Fig. 14, the distribution of L. argentea is discontinuous across South America, occurring only in Central America, Bolivia, and from northeastern to southern Brazil. Due to the lack of photographs in the literature and the limited availability of specimens in collections, determining the full geographic distribution of this species remains a challenge. Despite its similarity to L. devanes, L. argentea is easily distinguished by the ventral hindwing pattern in both sexes. Additionally, the female genitalia appear to be more informative than the male genitalia, confirming clear distinctions among Lycas species.
As pointed out by Zhang et al. (2025), L. godart and L. boisduvalii are distinct species based on molecular data. Through morphological characters, their distinction is readily noticeable, representing allopatric species, a status further supported by the significant genetic distance (4.5-4.8%) reported in this work. Lycas godart is endemic to the southern Atlantic Forest, while L. boisduvalii is an Amazonian species. Despite the unusual practice of describing sexually dimorphic species based on a single specimen, Lycas gabriel was originally described from a single female. Here, several male specimens are paired to this holotype through mitochondrial DNA, along with an additional female specimen from Mato Grosso (Brazil). Lycas gabriel was previously misidentified as L. boisduvalii due to similarities in wing patterns. Supported by morphological evidence, allied with the genetic distances (3.67-4.88%) and strong support in the phylogenetic tree (Fig. 1), it is confirmed its status as a valid species. Despite their sympatric distribution in Madre de Dios (Peru) and Acre (Brazil), the stigma on the male forewing and the structures of the genitalia provide distinctive characters to reliably differentiate these two species. Moreover, the known distribution of L. gabriel is herein expanded from Peru (Madre de Dios) to three states in Brazil (Amazonas, Acre, and Mato Grosso).
The single specimen displaying a different phenotype, collected in the same locality as L. gabriel, presents a similar male stigma shape and low genetic distance; however, its forewing discal cell spot and male genitalia differ from all other L. gabriel specimens examined. This morphotype could potentially represent a distinct species, but the inclusion of more specimens is desirable before making such a restrictive taxonomic decision. Thus, for the time being, this specimen is interpreted as a strong intraspecific variation within L. gabriel. In conclusion, this study provides a comprehensive taxonomic revision of Lycas by integrating molecular and morphological lines of evidence. While molecular data represent an invaluable tool for elucidating species relationships, they may not always provide definitive taxonomic resolution on their own. This work highlights the critical importance of detailed morphological studies-such as the highly informative female genitalia illustrated here for all species-in providing reliable, recognizable diagnoses. Establishing these clear morphological characteristics is essential for grounding future research in natural history, tritrophic interactions, biogeography, and ecology. Through this integrative approach, it was possible to validate L. gabriel morphologically, resolve its sexual dimorphism, and demonstrate that L. argentea is widely and sympatrically distributed with L. devanes in South America.
ACKNOWLEDGMENTS
We thank Diego R. Dolibaina and Adalberto D. de Medeiros for their assistance with the molecular data, and the meetings/discussions that provided fruitful insights. We are also grateful to Suianne Cajé for her help with the phylogenetic analyses. Specimen genetic data were registered in the Sistema Nacional de Gestão do Patrimônio Genético e do Conhecimento Tradicional Associado (SisGen) under the code RCB424E.
LITERATURE CITED
- Ahlf MA, Martello AR, Klein AL (2020) Lepidopterofauna diurna em remanescente de floresta ombrófila mista no sul do Paraná, Brasil. Luminária 21(1): 28-38.
- Brown KS (1992) Borboletas da Serra do Japi: diversidade, hábitats, recursos alimentares e variação temporal. In: Morellato LPC (Ed.) História natural da Serra do Japi: ecologia e preservação de uma área florestal no Sudeste do Brasil. Editora UNICAMP, Campinas, 142-187.
- Canals GR (2003) Mariposas de Misiones. Literature of Latin America, Buenos Aires, 476 pp.
- Carneiro E, Mielke OHH, Casagrande MM (2013) Thorax and abdomen morphology of some Neotropical Hesperiidae (Lepidoptera). Insecta Mundi 327: 1-47.
- Cong Q, Zhang J, Shen J, Grishin NV (2019) Fifty new genera of Hesperiidae (Lepidoptera). Insecta Mundi 731: 1-56.
- Draudt MWK (1921-1924) Grypocera, breitköpfige Tagfalter. In: Seitz A (Ed.) Die Gross Schmetterlinge der Erde. Alfred Kernen, Stuttgart, vol. 5, 833-1011.
- Ehrmann GA (1909) New species of exotic Lepidoptera. The Canadian Entomologist 41(3): 85-87.
- Evans WH (1955) A catalogue of the American Hesperiidae indicating the classification and nomenclature adopted in the British Museum (Natural History). Part IV. Hesperiinae and Megathyminae. British Museum (Natural History), London, 499 pp.
- Garwood K, Lehman R, Carter W, Carter G (2007) Butterflies of Southern Amazonia: A photographic checklist of common species. RiCalé Publishing, McAllen, 273 pp.
- Glassberg JA (2018) A swift guide to the butterflies of Mexico and Central America. Princeton University Press, Princeton, 2nd ed., 304 pp.
- Godman FD (1883-1901) Hesperiidae. In: Godman FD, Salvin O (Eds) Biologia Centrali-Americana, Insecta, Lepidoptera-Rhopalocera 2. Dulau & Co., Bernard Quaritch, London, 244-742.
-
Guindon S, Dufayard J-F, Lefort V, Anisimova M, Hordijk W, Gascuel O (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology 59(3): 307-321. https://doi.org/10.1093/sysbio/syq010
» https://doi.org/10.1093/sysbio/syq010 - Hayward KJ (1934) Lepidopteros Argentinos. Familia Hesperidae. IV. Subfamilia Pamphilinae. V. resúmen, clave, apéndices e índice. Revista de la Sociedad Entomológica Argentina 6(2-4): 97-233.
- Hayward KJ (1941) Hesperiidarum Argentinae Catalogus. Revista del Museo de La Plata (Zoología) 2(14): 227-340.
-
Hebert PDN, Cywinska A, Ball SL, deWaard JR (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society of London, Series B, 270(1512): 313-321. https://doi.org/10.1098/rspb.2002.2218
» https://doi.org/10.1098/rspb.2002.2218 - Herrich-Schäffer GAW (1869) Prodromus systematis Lepidopterorum. Versuch einer systematischen Anordnung der Schmetterlinge. Correspondenz-Blatt des zoologisch mineralogischen Vereines in Regensburg 23(12): 184-204.
- Hewitson WC (1866) Descriptions of new Hesperiidae. Transactions of the Entomological Society of London (3)2(6): 479-501.
- Hewitson WC (1872-1877) Illustrations of new species of exotic butterflies, a selected chiefly from the collections of W. Wilson Saunders and William C. Hewitson. John Van Voorst, London, vol. 5, 128 pp.
-
Kawahara AY, Storer C, Carvalho APS, Plotkin DM, Condamine FL, et al. (2023) A global phylogeny of butterflies reveals their evolutionary history, ancestral host and biogeographic origins. Nature Ecology and Evolution 7(6): 903-913. https://doi.org/10.1038/s41559-023-02041-9
» https://doi.org/10.1038/s41559-023-02041-9 - Kirby WF (1871) A synonymic catalogue of diurnal Lepidoptera. John Van Voorst, London, 690 pp.
- Kivirikko E (1936) Beobachtungen über die Tagfalterfauna (Lep., Diurna) des Territoriums Misiones (Rep. Argentina) in der Zeit 5.V.-20.VI.1928. Annales Entomologici Fennici 2(2): 49-63.
- Klimaitis JF, Núñez-Bustos EO, Klimaitis CL, Güller RM (2018) Mariposas Argentinas, Guía de identificación. Vázquez Mazzini Editores, Buenos Aires, 328 pp.
-
Kumar S, Stecher G, Suleski M, Sanderford M, Sharma S, Tamura K (2024) Molecular Evolutionary Genetics Analysis Version 12 for adaptive and green computing. Molecular Biology and Evolution 41(12): 1-9. https://doi.org/10.1093/molbev/msae263
» https://doi.org/10.1093/molbev/msae263 - Latreille PA (1824) In: Latreille PA, Godart JB (Eds) Encyclopédie Methodique. Histoire naturelle. Entomologie, ou histoire naturelle des crustacés, des arachnides et des insects 9(2): 329-828.
-
Lavinia PD, Núñez-Bustos EO, Kopuchian C, Lijtmaer DA, García NC, et al. (2017) Barcoding the butterflies of southern South America: species delimitation efficacy, cryptic diversity and geographic patterns of divergence. PLOS One 12(10): e0186845. https://doi.org/10.1371/journal.pone.0186845
» https://doi.org/10.1371/journal.pone.0186845 -
Li W, Cong Q, Shen J, Zhang J, Hallwachs W, et al. (2019) Genomes of skipper butterflies reveal extensive convergence of wing patterns.Proceedings of the National Academy of Sciences 116(13): 6232-6237. https://doi.org/10.1073/pnas.1821304116
» https://doi.org/10.1073/pnas.1821304116 - Lindsey AW (1925) The types of hesperioid genera. Annals of the Entomological Society of America 18(1): 75-106.
- Mabille MP (1878) Catalogue des hespérides du Musée Royal d’Historie Naturelle de Bruxelles. Annales de la Société Entomologique de Belgique 21(1-2): 12-44.
- Mabille MP (1903-1904) Lepidoptera Rhopalocera. Fam. Hesperidae. In: Wytsman P (Ed.) Genera Insectorum. L. Desmet-Verteneuil, Bruxelles, vol. 17, 210 pp.
-
Mielke OHH (2026) Catalogue of the American Hesperiidae. Volume 10. Hesperiinae-Part 4. Megathymini, Pericharini. https://sites.google.com/site/lablepneo/publica%C3%A7%C3%B5es/livros [Accessed: 10/06/2026]
» https://sites.google.com/site/lablepneo/publica%C3%A7%C3%B5es/livros -
Minh BQ, Nguyen MAT, Haeseler A (2013) Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution 30(5): 1188-1195. https://doi.org/10.1093/molbev/mst024
» https://doi.org/10.1093/molbev/mst024 - Moss AM (1949) Biological notes on some Hesperiidae of Para and the Amazon. Acta Zoologica Lilloana 7: 27-80.
-
Nguyen L-T, Schmidt H A, Haeseler A, Minh BQ (2015) IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32(1): 268-274. https://doi.org/10.1093/molbev/msu300
» https://doi.org/10.1093/molbev/msu300 - Orlandin E, Piovesan M, Carneiro E (2020) Borboletas do Meio-Oeste de Santa Catarina: História natural e guia de identificação. Clube dos Autores, Joaçaba, 398 pp.
- Palo Jr H (2017) Borboletas do Brazil: Hesperiidae. Vento Verde Editora, São Carlos, vol. 3, 1729-2400.
- Plötz C (1882) Die Hesperiinem-Gattung Hesperia Aut. und ihre Arten. Stettiner Entomologische Zeitung 43(7/9): 314-344.
-
Rambaut A (2010) FigTree v1.3.1. Institute of Evolutionary Biology, University of Edinburgh, Edinburgh. http://tree.bio.ed.ac.uk/software/figtree/ [Accessed: 01/06/2025]
» http://tree.bio.ed.ac.uk/software/figtree/ -
Schneider CA, Rasband WS, Eliceiri KW (2012) NIH Image to ImageJ: 25 years of image analysis. Nature Methods 9(7): 671-675. https://doi.org/doi:10.1038/nmeth.2089
» https://doi.org/doi:10.1038/nmeth.2089 - Shepard HH (1931) Lepidopterorum Catalogus 47, Hesperidae: Subfamilia Pyrginae I. Junk, Berlin, 144 pp.
- Uehara J (2017) Collecting butterflies in Paraguay. Butterflies (Tokyo) 76: 36-51.
- Uehara J (2020) Collecting butterflies in S. E. Pelu [sic] (2). Butterflies (Tokyo) 84: 48-57.
- Westwood JO (1852) In: Doubleday E, Westwood JO (Eds) (1850-1852) The genera of diurnal Lepidoptera: comprising their generic characters, a notice of their habits and transformations, and a catalogue of the species of each genus. Longman, Brown, Green, and Longmans, London, vol. 2, 511-514.
-
Xiang C-Y, Gao F, Jakovlić I, Lei H-P, Hu Y, et al. (2023) Using PhyloSuite for molecular phylogeny and tree-based analyses. iMeta 2(1): e87. https://doi.org/10.1002/imt2.87
» https://doi.org/10.1002/imt2.87 -
Zhang D, Gao F, Jakovlić I, Zou H, Zhang J, et al. (2020) PhyloSuite: An integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Molecular Ecology Resources 20(1): 348-355. https://doi.org/10.1111/1755-0998.13096
» https://doi.org/10.1111/1755-0998.13096 - Zhang J, Cong Q, Shen J, Grishin NV (2022a) Taxonomic changes suggested by the genomic analysis of Hesperiidae (Lepidoptera). Insecta Mundi 921: 1-135.
- Zhang J, Cong Q, Shen J, Song L, Gott RJ, et al. (2022b) Taxonomic discoveries enabled by genomic analysis of butterflies. The Taxonomic Report of the International Lepidoptera Survey 10(7): 1-59.
- Zhang J, Cong Q, Shen J, Song L, Grishin NV (2025) New butterfly taxa and findings from genomic analyses. The Taxonomic Report of the International Lepidoptera Survey 12(8): 1-154.
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ZooBank register
https://zoobank.org/9D3CE3AB-6080-46FB-9144-90F3E57AD723
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Data Availability Statement
All data generated and/or analyzed are included in this article. All Cytochrome c Oxidase I sequences analyzed in this study, including newly generated and previously published sequences, are listed in Table 1.
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Funding Statement
This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES, Finance Code 001), through a research fellowship to ASB (CAPES, 88887.949958/2024-00). MMC is supported by a research fellowship from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, 310227/2021-3).
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Ethical Statement
Not applicable. This study was based exclusively on specimens deposited in scientific collections and did not involve field collection or handling of live vertebrate animals.
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AI Statement
No artificial intelligence tools were used in the preparation of this article.
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How to cite this article
Suênia-Bastos A, Mielke OHH, Fåhraeus C, Casagrande MM (2026) Taxonomic revision and phylogeny of Lycas (Lepidoptera: Hesperiidae: Hesperiinae), with a redescription of Lycas gabriel. Zoologia 43: e25072. https://doi.org/10.1590/S1984-4689.v43.e25072
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Published by
Sociedade Brasileira de Zoologia at Scientific Electronic Library Online - https://www.scielo.br/zool
All data generated and/or analyzed are included in this article. All Cytochrome c Oxidase I sequences analyzed in this study, including newly generated and previously published sequences, are listed in Table 1.
