Systematics of Opsiphanes Doubleday, [1849] (Lepidoptera: Nymphalidae, Satyrinae, Brassolini): an integrative approach Author Piovesan, Mônica 0000-0003-3367-9358 monica367piovesan@gmail.com Author Casagrande, Mirna Martins 0000-0002-6076-8463 mibras@ufpr.br Author Mielke, Olaf Hermann Hendrik 0000-0003-3655-4606 omhesp@ufpr.br text Zootaxa 2022 2022-12-06 5216 1 1 278 http://dx.doi.org/10.1094/PDIS-04-22-0755-PDN journal article 10.11646/zootaxa.5216.1.1 1175-5326 7403532 F9C92B91-42CF-4D2D-B315-14FF801815D4 Opsiphanes invirae pernambucoensis Bristow, 1991 stat. rev. ( Figs 228–230 , 233 , 248 ) Opsiphanes invirae subsp.; D’Abrera, 1987 (Bristow Ms). Butt. Neotrop. Reg. 3 , p. 417 (fig. male, female d); Brazil . Opsiphanes invirae pernambucoensis Bristow, 1991 . Zool. Jour. Linn. Soc. 101 (3): 255, 258, 260, figs 21b (female d, v), 22h (male d, v); holotype male, Brazil , Pernambuco , M. Moss [ leg .]; NHMUK . Taxonomic history. Casagrande (2004) considered Opsiphanes invirae pernambucoensis Bristow, 1991 as synonymous with Opsiphanes invirae remoliatus Fruhstorfer, 1907 . Type material. Opsiphanes invirae pernambucoensis Bristow, 1991 was described based on the holotype male, illustrated by D’Abrera (1987) and Bristow (1991) , and is deposited at the NHMUK, with the following labels: / Holotype / Pernambuco [ Brazil ] 88-10/ pernambucoensis Bristow/ ( Fig. 248 ). Diagnosis. Opsiphanes invirae pernambucoensis stat. rev. differs from Opsiphanes invirae invirae in the DFW oblique band that does not border the discal cell on M 3 -CuA 1 ; and the washed-out appearance of the ground color and band color of DFW and DHW, lighter than in Opsiphanes invirae invirae ( Figs 228–230 ). FIGURE 228. Opsiphanes invirae pernambucoensis Bristow, 1991 stat. rev. : Male, dorsal and ventral views: Brazil: a. (DZ 42.227-DZUP) Rio Grande do Norte, b. (DZ 42.233-DZUP) Paraíba, c. (DZ 42.270-DZUP), d. (DZ 42.267-DZUP) Pernambuco, e. (DZ 42.242-DZUP), f. (DZ 42.240-DZUP) Alagoas. FIGURE 229. Opsiphanes invirae pernambucoensis Bristow, 1991 stat. rev. : Female, dorsal and ventral views: Brazil: a. (DZ 42.251-DZUP) Rio Grande do Norte, b. (DZ 42.228-DZUP) Paraíba. FIGURE 230. Opsiphanes invirae pernambucoensis Bristow, 1991 stat. rev. : Female, dorsal and ventral views: Brazil: a. (DZ 42.272-DZUP) Pernambuco, b. (DZ 42.258-DZUP) Bahia. Variation. FW, length: male: 33–38 mm (n=6), female: 41–43 mm (n=4). DFW, the width of the oblique band. DHW, the submarginal band is generally as wide as DFW oblique band; the submarginal band usually extends from the costal margin to M 2 , with shades of yellow, and between M 2 and CuA 2 , it becomes rusty red; in males, the submarginal band is usually evident up to the anal angle, in females it may extend toward the basal region, where it is limited by the distal portion of the discal cell. VFW, the width of the oblique band, from M 3 -CuA 1 to the anal angle; may have small eyespots between M 2 and CuA 1 ( Fig. 230b ). VHW, the shape of the eyespot between Sc+R 1 and M 1 ( Figs 228–230 ). FIGURE 231. Opsiphanes invirae invirae (Hübner, [1808]) : Male: a. abdomen, lateral view, b. detail of the scent organ, lateral view; genitalia: c. dorsal view, d. ventral view, e. lateral view; aedeagus: f. dorsal view, g. ventral view, h. lateral view (DZ 41.714-DZUP); valva: i. (DZ 45.861-DZUP), j. (DZ 41.991-DZUP), k. (DZ 41.995-DZUP) variations of the dorso-distal portion. FIGURE 232. Opsiphanes invirae invirae (Hübner, [1808]) : Female genitalia: a. lateral view; b. detail of the signa; c. ventral view (DZ 3948-DZUP). FIGURE 233. Geographical distribution of Opsiphanes invirae invirae (Hübner, [1808]) and Opsiphanes invirae pernambucoensis Bristow, 1991 stat. rev. FIGURE 234. Opsiphanes invirae invirae (Hübner, [1808]) . Neotype of Opsiphanes invirae invirae (Hübner, [1808]) (CM- BUT1579-20); a. dorsal view, b. ventral view. FIGURE 235. Opsiphanes invirae invirae (Hübner, [1808]) . Lectotype of Opsiphanes cassina C. Felder & R. Felder, 1862 syn. nov. ; a. dorsal view, b. ventral view. FIGURE 236. Opsiphanes invirae invirae (Hübner, [1808]) . [Holo]type of Opsiphanes invirae intermedius Stichel, 1902 syn. nov. ; a. dorsal view, b. ventral view. FIGURE 237. Opsiphanes invirae invirae (Hübner, [1808]) . Lectotype of Opsiphanes cassina aequatorialis Stichel, 1902 syn. rest. ; a. dorsal view, b. ventral view. FIGURE 238. Opsiphanes invirae invirae (Hübner, [1808]) . Lectotype of Opsiphanes invirae amplificatus Stichel, 1904 syn. nov. ; a. dorsal view, b. ventral view. FIGURE 239. Opsiphanes invirae invirae (Hübner, [1808]) . Neotype of Opsiphanes sticheli Röber, 1906 syn. nov. ; a. dorsal view, b. ventral view. FIGURE 240. Opsiphanes invirae invirae (Hübner, [1808]) . Lectotype of Opsiphanes invirae pseudophilon Fruhstorfer, 1907 syn. rest. ; a. dorsal view, b. ventral view. FIGURE 241. Opsiphanes invirae invirae (Hübner, [1808]) . Lectotype of Opsiphanes invirae remoliatus Fruhstorfer, 1907 syn. rest. ; a. dorsal view, b. ventral view. FIGURE 242. Opsiphanes invirae invirae (Hübner, [1808]) . Lectotype of Opsiphanes invirae agasthenes Fruhstorfer, 1907 syn. rest. ; a. dorsal view, b. ventral view. FIGURE 243. Opsiphanes invirae invirae (Hübner, [1808]) . “Type” of Opsiphanes invirae agasthenes f. rectifasciata Fruhstorfer, 1907 nom. nud. ; a. dorsal view, b. ventral view. FIGURE 244. Opsiphanes invirae invirae (Hübner, [1808]) . Lectotype of Opsiphanes invirae isagoras Fruhstorfer, 1907 ; a. dorsal view, b. ventral view. FIGURE 245. Opsiphanes invirae invirae (Hübner, [1808]) . Neotype of Opsiphanes invirae ledon Fruhstorfer, 1912 ; a. dorsal view, b. ventral view. FIGURE 246. Opsiphanes invirae invirae (Hübner, [1808]) . Holotype of Opsiphanes invirae roraimaensis Bristow, 1991 syn. nov. ; a. dorsal view, b. ventral view. FIGURE 247. Opsiphanes invirae invirae (Hübner, [1808]) . Holotype of Opsiphanes invirae sieberti Bristow, 1991 syn. nov. ; a. dorsal view, b. ventral view. FIGURE 248. Opsiphanes invirae pernambucoensis Bristow, 1991 stat. rev. Holotype of Opsiphanes invirae pernambucoensis Bristow, 1991 stat. rev. ; a. dorsal view, b. ventral view. Comments. In the VHW, the eyespot between Sc+R 1 and M 1 of the holotype of Opsiphanes invirae pernambucoensis stat. rev. was used by Bristow (1991) as a character to separate the subspecies. However, upon examination of a series of specimens, we found that this character is variable. Natural history. Unknown. Geographic distribution. Brazil ( Ceará , Rio Grande do Norte , Paraíba , Pernambuco , Alagoas, Bahia ) ( Fig. 233 ), at altitudes between sea level and 900 m . Temporal distribution. Throughout the year. Etymology. Reference to the state of Pernambuco , Brazil , the location of the holotype . Specimens examined. Photos of the holotypus of Opsiphanes invirae pernambucoensis Bristow, 1991 stat. rev. , and the following specimens: BRAZIL : Ceará —Jericoacoara, 1-XII-2005 , 1 male , Mielke & Casagrande leg., DZ 42.222 (DZUP); Ubajara ( Parque Nacional Ubajara ), 22–24-IV-2012 , 1 male , Mielke & Casagrande leg., DZ 42.223 (DZUP); Aracoiaba, 120 m , 9–10-I-2013 , 1 female , Dolibaina & Leite leg., DD 486 (DD); Guaramiranga, 900 m , 7–11-I-2012 , 1 male , Dolibaina & Leite leg., DD 485 (DD); Fortaleza, 9-IX-1973 , 1 female , DZ 42.218 (DZUP), 28-XI-2009 , 1 male , Mielke & Casagrande leg., DZ 42.219 (DZUP), 1 male , J. Bustos leg., ex-coll. H. Ebert, DZ 42.220 (DZUP), 26-IV-1976 , 1 male , Oscanua leg., DZ 42.221 (DZUP). Rio Grande do Norte —Natal, 7-IX-2009 , 1 male , L. Merlin leg., DZ 42.226 (DZUP), 16-VII-1978 , 2 males , O. Mielke leg., DZ 42.224 , 42.225 (DZUP); Praia de Ponta Negra, 17-VIII-1987 , 1 male , 7-VIII-1987 , 1 female , Casagrande leg., DZ 42.227 , 42.251 (DZUP). Paraíba —João Pessoa, 19-X-1952 , 1 male , ex-coll. H. Ebert, DZ 42.235 (DZUP), 15-V-1954 , 1 female , 16-V-1953 , 1 male , R. Diniz leg., OM 25, 1259 (DZUP—OM), 1953, 1 female , R. Diniz leg., ex-coll. D’Almeida, DZ 42.231 (DZUP), VIII-1957 , 1 female , 1 male , IX-1956 , 1 female , 10-IX-1973 , 2 males , 1-VII- 1980 , 1 female , I-1978 , 1 male , IX-1975 , 1 male , Kesselring leg., ex-coll. D’Almeida, DZ 42.228–DZ 42.230, 42.232–42.236 (DZUP). Pernambuco —no locality, 250 m , 5-III-1969 , 1 male , H. Ebert leg., ex-coll. H. Ebert, DZ 42.264 (DZUP); Garanhuns, 800–900 m , 26-II-1961 , 1 male , H. Ebert leg., ex-coll. H. Ebert, DZ 42.262 (DZUP); São Lourenço da Mata , 20-VI-1981 , 2 females , Kesselring leg., DZ 42.271 , 42.272 (DZUP); Ipojuca ( Porto de Galinhas ), 4-XII-2012 , 1 female , Mielke & Casagrande leg., DZ 42.274 (DZUP); Recife, 21-VI-1957 , 1 female , 1-VI-1957 , 1 male , ex-coll. H. Ebert, DZ 3947 (DZUP), 9-VIII-1951 , 1 male , 4-IX-1961 , 1 female , 13-VII-1959 , 1 male , 30-VII-1951 , 1 male , 25-VIII-1951 , 1 male , 16-V-1961 , 1 female , 15-VI-1961 , 1 male , 21-V-1961 , 1 female , H. Ebert leg., ex-coll. H. Ebert, DZ 3943, 42.260, 42.261, 42.263, 42.266, 42.267, 42.269, 42.270 (DZUP), 25 m , 15-I-1960 , 1 male , H. Ebert leg., ex-coll. H. Ebert, DZ 42.268 (DZUP); Camaragibe, 20–80 m , 1-IX-1961 , 1 female , H. Ebert leg., ex-coll. H. Ebert, DZ 42.259 (DZUP), 50 m , 2-III-1968 , 1 female , Kesselring leg., ex-coll. H. Ebert, DZ 42.273 (DZUP). Alagoas —Maceió, 1 female , 7-V-1979 , 1 female , 12-III-1977 , 1 male , A. Cardoso leg., DZ 42.237, 42.238, 42.242 (DZUP), VI-1992 , 1 male , Cerqueira leg., DZ 42.246 (DZUP), 22-IV-2004 , 1 female , Santos F. leg., DZ 42.247 (DZUP); Barra de São Miguel , 30-III-2004 , 1 male , 31-III-2004 , 1 male , Santos F. leg., DZ 42.248, DZ 42.249 (DZUP); Pajuçara, 26-III-2001 , 1 male , Mielke & Casagrande leg., OM 53.390 (DZUP—OM), 21-VI-1973 , 1 female , 2 males , 15-I-1974 , 1 male , 3-VI-1972 , 1 male , 20-V-1949 , 1 male , A. Cardoso leg., ex-coll. A. Cardoso, DZ 42.239 - 42.241, 42.243 - 42.245 (DZUP). Bahia —Itaparica, 27-VII-1981 , 1 female , A. Raw leg., DZ 42.258 (DZUP). General comments The present study increased the number of species of Opsiphanes , from 14 to 23, and more than halved the number of subspecies, from 60 to 23. Analysis of information on natural history, morphological and molecular, and geographic distribution revealed that many proposed hypotheses, mainly for subspecies, were based on variable phenotypic data. The phenotypic variation, mainly in the color and width of the wing bands, made it difficult to understand the diversity of the genus, and only after analyzing more than 5,500 specimens was it possible to develop a new classification for Opsiphanes . However, characters related to the host plants of immatures and the secondary sexual characters of males may be a consequence of important evolutionary processes in the genus diversification, deserving further investigation. Apparently, the feeding habit of the larvae can be used to separate the Opsiphanes groups; while the “ cassiae group” feeds on members of Zingiberales ( Musaceae , Cannaceae , Heliconiaceae ), the “ quiteria group” feeds on members of Arecales (Arecaceae) , although some contradictory information has been reported (e.g., Beccaloni et al . 2008 ; Pulido & Cárdenas 1979 ). Only one study ( Young & Muyshondt 1975 ) tested the food preference of Opsiphanes , including Opsiphanes cassiae tamarindi stat. nov. and Opsiphanes fabricii fabricii ( Boisduval, 1870 ) , representatives of the “ cassiae ” and “ quiteria ” groups, respectively. The authors found that Opsiphanes cassiae tamarindi stat. nov. did not feed on members of Arecales and Opsiphanes fabricii fabricii did not feed on members of Zingiberales . However, food preference tests should be conducted with other species of Opsiphanes in different geographic locations. Co-evolution between host plants and insects or a change of host plants by a certain group of herbivores may be involved in the diversification of species ( Becerra 2015 ; Winter et al. 2017 ; Allio et al. 2021 ). Studies of the relationship of Opsiphanes with their host plants may provide support for formulating and testing hypotheses about the role of feeding habits in the diversification of Opsiphanes lineages. Other striking features in Opsiphanes males are the systems for producing and dispersing scent, supposedly involved in the communication within species ( Young & Muyshondt 1975 ). Chemical communication takes place in many organisms, and this type of signal can contain information about the identity and quality of potential sexual partners, playing a key role in sexual selection ( Nieberding et al. 2008 ; Bacquet et al. 2015 ). Few studies have explored the scent organs of Opsiphanes in detail, and the largest contributions to the knowledge of these structures have been made through analysis of the internal anatomy ( Eltringham 1929 ; Barth 1952 ). Nothing is known about the nature of chemical compounds in scent organs or their influence on the behavior of species of the genus. Because different species of Opsiphanes fly at the same time and in the same location ( Orlandin et al . 2020 ), it is likely that chemical compounds play a fundamental role in intra- and inter-specific communication and that this strategy is also related to the diversification of the species of the genus.