Opsiphanes Doubleday, [1849]

Piovesan, Mônica, Casagrande, Mirna Martins & Mielke, Olaf Hermann Hendrik, 2022, Systematics of Opsiphanes Doubleday, [1849] (Lepidoptera: Nymphalidae, Satyrinae, Brassolini): an integrative approach, Zootaxa 5216 (1), pp. 1-278 : 14-18

publication ID	https://doi.org/ 10.11646/zootaxa.5216.1.1
publication LSID	lsid:zoobank.org:pub:F9C92B91-42CF-4D2D-B315-14FF801815D4
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scientific name	Opsiphanes Doubleday, [1849]
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Opsiphanes Doubleday, [1849] View in CoL

( Figs 1–248 View FIGURE 1 View FIGURE 2 View FIGURE 3 View FIGURE 4 View FIGURE5 View FIGURE 6 View FIGURE 7 View FIGURE 8 View FIGURE 9 View FIGURE 10 View FIGURE 11 View FIGURE 12 View FIGURE 13 View FIGURE 14 View FIGURE 15 View FIGURE 16 View FIGURE 17 View FIGURE 18 View FIGURE 19 View FIGURE 20 View FIGURE 21 View FIGURE 22 View FIGURE 23 View FIGURE 24 View FIGURE 25 View FIGURE 26 View FIGURE 27 View FIGURE 28 View FIGURE 29 View FIGURE 30 View FIGURE 31 View FIGURE 32 View FIGURE 33 View FIGURE 34 View FIGURE 35 View FIGURE 36 View FIGURE 37 View FIGURE 38 View FIGURE 39 View FIGURE 40 View FIGURE 41 View FIGURE 42 View FIGURE 43 View FIGURE 44 View FIGURE 45 View FIGURE 46 View FIGURE 47 View FIGURE 48 View FIGURE 49 View FIGURE 50 View FIGURE 51 View FIGURE 52 View FIGURE 53 View FIGURE 54 View FIGURE 55 View FIGURE 56 View FIGURE 57 View FIGURE 58 View FIGURE 59 View FIGURE 60 View FIGURE 61 View FIGURE 62 View FIGURE 63 View FIGURE 64 View FIGURE 65 View FIGURE 66 View FIGURE67 View FIGURE 68 View FIGURE 69 View FIGURE 70 View FIGURE 71 View FIGURE 72 View FIGURE 73 View FIGURE 74 View FIGURE 75 View FIGURE 76 View FIGURE 77 View FIGURE 78 View FIGURE 79 View FIGURE 80 View FIGURE 81 View FIGURE 82 View FIGURE 83 View FIGURE 84 View FIGURE 85 View FIGURE 86 View FIGURE 87 View FIGURE 88 View FIGURE 89 View FIGURE 90 View FIGURE 91 View FIGURE 92 View FIGURE 93 View FIGURE 94 View FIGURE 95 View FIGURE 96 View FIGURE 97 View FIGURE 98 View FIGURE 99 View FIGURE 100 View FIGURE 101 View FIGURE 102 View FIGURE 103 View FIGURE 104 View FIGURE 105 View FIGURE 106 View FIGURE 107 View FIGURE 108 View FIGURE 109 View FIGURE 110 View FIGURE 111 View FIGURE 112 View FIGURE 113 View FIGURE 114 View FIGURE 115 View FIGURE 116 View FIGURE 117 View FIGURE 118 View FIGURE 119 View FIGURE 120 View FIGURE 121 View FIGURE 122 View FIGURE 123 View FIGURE 124 View FIGURE 125 View FIGURE 126 View FIGURE 127 View FIGURE 128 View FIGURE 129 View FIGURE 130 View FIGURE 131 View FIGURE 132 View FIGURE 133 View FIGURE 134 View FIGURE 135 View FIGURE 136 View FIGURE 137 View FIGURE 138 View FIGURE 139 View FIGURE 140 View FIGURE 141 View FIGURE 142 View FIGURE 143 View FIGURE 144 View FIGURE 145 View FIGURE 146 View FIGURE 147 View FIGURE 148 View FIGURE 149 View FIGURE 150 View FIGURE 151 View FIGURE 152 View FIGURE 153 View FIGURE 154 View FIGURE 155 View FIGURE 156 View FIGURE 157 View FIGURE 158 View FIGURE 159 View FIGURE 160 View FIGURE 161 View FIGURE 162 View FIGURE 163 View FIGURE 164 View FIGURE 165 View FIGURE 166 View FIGURE 167 View FIGURE 168 View FIGURE 169 View FIGURE 170 View FIGURE171 View FIGURE 172 View FIGURE 173 View FIGURE 174 View FIGURE 175 View FIGURE 176 View FIGURE 177 View FIGURE 178 View FIGURE 179 View FIGURE 180 View FIGURE 181 View FIGURE 182 View FIGURE 183 View FIGURE 184 View FIGURE 185 View FIGURE 186 View FIGURE 187 View FIGURE 188 View FIGURE 189 View FIGURE 190 View FIGURE 191 View FIGURE 192 View FIGURE 193 View FIGURE 194 View FIGURE 195 View FIGURE 196 View FIGURE 197 View FIGURE 198 View FIGURE 199 View FIGURE 200 View FIGURE 201 View FIGURE 202 View FIGURE 203 View FIGURE 204 View FIGURE 205 View FIGURE 206 View FIGURE 207 View FIGURE 208 View FIGURE 209 View FIGURE 210 View FIGURE 211 View FIGURE 212 View FIGURE 213 View FIGURE 214 View FIGURE 215 View FIGURE 216 View FIGURE 217 View FIGURE 218 View FIGURE 219 View FIGURE 220 View FIGURE 221 View FIGURE 222 View FIGURE 223 View FIGURE 224 View FIGURE 225 View FIGURE 226 View FIGURE 227 View FIGURE 228 View FIGURE 229 View FIGURE 230 View FIGURE 231 View FIGURE 232 View FIGURE 233 View FIGURE 234 View FIGURE 235 View FIGURE 236 View FIGURE 237 View FIGURE 238 View FIGURE 239 View FIGURE 240 View FIGURE 241 View FIGURE 242 View FIGURE 243 View FIGURE 244 View FIGURE 245 View FIGURE 246 View FIGURE 247 View FIGURE 248 )

Opsiphanes Doubleday, [1849] View in CoL . Gen. diurn. Lep. 1, pl. 57; included species: Opsiphanes sallei Doubleday, [1849] View in CoL , Opsiphanes boisduvallii Doubleday, [1849] , Opsiphanes reevesii Doubleday, [1849] View in CoL .—Westwood, 1851, in Doubleday. Gen. diurn. Lep. 2, p. 344.—Lucas, 1851, in Chenu (ed.). Enc. d‘Hist. nat., Pap. diurn., p. 170.—Herrich-Schäffer, 1865. Corr.-Blatt zool.-min. Ver. Regensburg 19 (5): 64.—Prittwitz, 1865. Stett. ent. Zeit. 26 (10/12): 309.— Kirby, 1871. Syn. Cat. Diurn. Lep., p. 125, 646, 681.— Scudder, 1875. Proc. Amer. Ac. Arts & Sc. 10 (2): 233; type species: sallei View in CoL .—Wallace, 1876. Geogr. Distr. Anim. 2, p. 472.—Kirby, 1877. Syn. Cat. Diurn. Lep., Suppl., p. 716.—F. Müller, 1878. Trans. ent. Soc. London, 1878 (3): 214.— Kirby, 1879. Cat. Coll. Diurn. Lep. Hewitson, p. 107.— Godman & Salvin, 1881. Biol. Centr.- Amer., Lep. Rhop. 1 (12), p. 125.— Aurivillius, 1882. Kongl. Svenska Akad. Handl. 19 (5): 76.—Gerhard, 1883. Berl. ent. Ztschr. 27 (2): 179.—W. Müller, 1886. Zool. Jahrb., Syst. 1 (3/4), p. 170; biol.—Staudinger, 1886, in Staudinger & Schatz. Exot. Schmett. 1 (15), pls. 71, 72 (male, female d, v).— Staudinger, 1887, in Staudinger & Schatz. Exot. Schmett. 1 (18), p. 213.—Glaser, 1887. Cat. etymol. Col. Lep., p. 304; etymol.—Schatz, 1888, in Staudinger & Schatz. Exot. Schmett. 2 (4), pl. 32.— Röber, 1889, in Staudinger & Schatz. Exot. Schmett. 2 (5), p. 190.—Reuter, 1896. Acta Soc. Sc. Fenn. 22 (1): 112, 553; morph.— Stichel, 1902. Berl. ent. Ztschr. 46 (4): 503.— Stichel, 1904, in Wytsman. Gen. Ins. 20, p. 1.— Röber, 1906. Soc. ent. 21 (3): 20; (4): 27.— Stichel, 1907. Soc. ent. 22 (13): 98.— Stichel, 1909. Das Tierreich 25: 51.—Pagenstecher, 1909. Geogr. Verbr. Schmett., p. 413.— Fruhstorfer, 1912, in Seitz. Gross-Schmett. Erde 5, p. 291.— Strand, 1916. Lep. Niep. 2, p. 13— Campos, 1921. Rev. Col. Nac. V. Rocafuerte, Guayaquil, 4: 31.—Jurriaanse, 1923. Tijd. Ent. 66: 147; scent organs.—Schwanwitsch, 1924. Proc. zool. Soc. London 1924 (2): 521; scent organs.— Bates, 1932. Bull. Brooklyn Ent. Soc. 27 (3): 155; biol.—Eltringham, 1935. Proc. r. Soc. London B 117 (806): 478; scent organs.—Hoffmann, 1937. Int. ent. Ztschr. 50 (6): 57; biol.— Hemming, 1943. Proc. r. ent . Soc. London (B) 12 (2): 25.—Barth, 1958. An. Ac. Bras. Ciências 30 (3): 353, 357; scent organs.—Ehrlich, 1958. Univ. Kansas Sci. Bull. 39 (8): 351; morph.—L. Miller, 1968. Mem. Amer. Ent. Soc. 24: 24, 26.—Hemming, 1967. Bull. Brit. Mus. (Nat. Hist.), Ent., Suppl. 9: 88, 321.—de Lesse & K. Brown, 1972. Bull. Soc. ent. France 76 (5/6): 136; genet.—Hayward, 1973. Op. Lill. 23: 242.—Biezanko et al., 1974. Rev. Centro Ciências Rur., Santa Maria , 4: 112.— Young & Muyshondt, 1975. Stud. Neotrop. Fauna 10: 19; biol, host plant.—Bristow, 1981. Zool. Jour. Linn. Soc. 72 (2): 117.—Boos, 1986. Living World 1985-1986: 31.—R. F. de la Maza R., 1987. Marip. Mexicanas, p. 95.— Ackery, 1988. Biol. Jour. Linn. Soc. 33 (2): 99; host plant.— Bristow, 1991. Zool. Jour. Linn. Soc. 101 (3): 203.—R. G. de la Maza E. & Soberón, 1998. Biodiv. Conserv 7 (7): 929; ecol.— Casagrande, 2002. Rev. bras. Zool. 19 (2): 468, 496; biol.— Casagrande, 2004. Brassolini View in CoL , in Lamas (ed.) Checklist: Part 4A, Hesperioidea-Papilionoidea, in Heppner (ed.). Atlas Neotrop. Lep. 5A, p. 203; syn.: Pavonia Godart, [1824] View in CoL preocc. (Lamarck, 1816 [Coelenterata]), Anthomantes Gistel, 1848 repl. name, Caligo Boisduval, 1870 View in CoL preocc. (Hübner, [1819] [ Nymphalidae View in CoL ]).—Freitas & K. Brown, 2004. Syst. Biol. 53 (3): 378, 383, figs A1 (eggs), A14 (fem. gen.); phylog.— Penz, 2007. Syst. Ent. 32 (4): 682; phylog.—Smith et al., 2008. Proc. Nat. Ac. Sc. USA 105 (34): 12362.—Peña & Wahlberg, 2008. Biol. Lett. 4 (3): 275; phylog.—Wahlberg et al., 2009. Proc. r. Soc. B 276 (1677): 4298; phylog.— Penz et al., 2011. Zootaxa 2897: 1, 7, 9.—Cassildé et al., 2012. Ann. Soc. ent. France 48 (1/2): 175.— Chacón et al., 2012. Bull. Allyn Mus. 166: 1, 9.—Penz & Mohammadi, 2013. Biota Neotrop. 13 (3): 163, 169, 180.—Blandin et al., 2014. Eur. Jour. Tax. 71: 2; cit.— Neves & Paluch, 2016. Neotrop. Ent. 45 (1): 13; biol., host plant.—Nobre & Schlindwein, 2016. Borboletas Vale Catimbau, p. 160.—Shirai et al., 2017. Ann. Ent. Soc. Amer. 110 (2): 145, 154, fig. 5 (phylogenetic tree); biol., phylog.—Pinheiro & Cintra, 2017. Jour. Lep. Soc. 71 (2): 110; biol.

Caligo Boisduval, 1870 View in CoL (not Hübner, [1819]). Consid. Lép. env. Guatemala, p. 54; included species: C. cassiae Linnaeus, C. fabricii Boisduval, 1870 , C. berecynthia Cramer , C. tamarindi Boisduval, 1870 , C. sallei Doubleday View in CoL , C. boisduvalii Doubleday ; preocc. (Hübner, [1819] [ Nymphalidae View in CoL ])— Hemming, 1943. Proc. r. ent . Soc. London (B) 12 (2): 25.—Hemming, 1967. Bull. Brit. Mus. (Nat. Hist.), Ent., Suppl. 9: 348.

Type species: Opsiphanes sallei Doubleday, [1849] View in CoL , designated by Scudder (1875).

Pavonia Godart [1824] View in CoL syn. nov., considered synonymous with Opsiphanes Doubleday [1849] ( Casagrande 2004) View in CoL , for which the type species is Papilio actorion Linnaeus 1758 , designated by Froriep (1836, p. 379), is a synonym of Bia Hübner, [1819] View in CoL . Pavonia Godart [1824] View in CoL is also preoccupied by Lamarck (1816) [Coelenterata] and Hübner [1819] [ Nymphalidae View in CoL ]. Gistel (1848) published Anthomantes as a new name for Pavonia Godart [1824] View in CoL and therefore also synonymous with Bia Hübner, [1819] View in CoL . For the same reason, Hemming’s (1943) designation of Papilio idomeneus Linnaeus, 1758 as the type species of Pavonia Godart [1824] View in CoL is invalid, since at the time it was considered synonymous with Opsiphanes View in CoL due to the lack of knowledge of Froriep’s designation (1836).

Caligo Boisduval, 1870 View in CoL , for which the type species is Opsiphanes sallei Doubleday [1849] View in CoL , is an invalid name, as according to Hemming (1943), Boisduval (1870) republished the name previously published by Hübner ([1819]) as his own. Thus, Caligo Boisduval, 1870 View in CoL is a junior homonym of Caligo Hübner, [1819] View in CoL (ICZN 1999, Art. 53).

Diagnosis. Opsiphanes differs from Blepolenis in having the labial palpus with the distal segment measuring half or at most two-thirds the length of the basal segment, while in Blepolenis the corresponding segments are equivalent. DFW, M 2 -M 3 without a whitish spot near the outer margin. FW, the common branch of R 3 +R 4 +R 5 is shorter than the common branch of R 4 +R 5, while in Blepolenis the common branch length of R 3 +R 4 +R 5 is longer than the common branch of R 4 +R 5.

Redescription. Head. Eyes naked; labial palpus with the distal joint about half or at most two-thirds the length of the basal joint.

Thorax. Dark-brown dorsally and ventrally, except orange in Opsiphanes blythekitzmillerae Austin & A. Warren, 2007 and Opsiphanes boisduvallii Doubleday, [1849] ; legs dark-brown except golden yellow in Opsiphanes blythekitzmillerae and Opsiphanes boisduvallii .

Forewing. Length: male 32–53 mm (n=199), female 35–61 mm (n=105). Costal margin with apex projected; outer margin usually concave between M 1 and CuA 2 and then convex to tornus; inner margin straight. DFW between R 3 and M 1 with whitish or yellowish apical spots and a band extending from the middle of the costal margin to the anal angle, varying from white to orange; ground color dark-brown; in Opsiphanes blythekitzmillerae and Opsiphanes boisduvallii , M 1 –M 2 with a dark-brown rounded subapical spot, without a band and with orange ground color. VFW with “c” element forming a band that crosses the discal cell, “d” and “e” elements with variable patterns; males have an androconial brand in the basal half of 2A; postdiscal region with a disruptive pattern, formed by narrow brown lines; subapical eyespot present between M 1 and M 3; with submarginal lines EII and EIII.

Hindwing. Costal and outer margins rounded; the outer margin may be smooth or crenulated; inner margin straight. DHW with a submarginal and/or marginal band and dark-brown ground color, except Opsiphanes blythekitzmillerae and Opsiphanes boisduvallii without a submarginal and/or marginal band and with an orange ground color. Males with a tuft of setae on the distal margin of the humeral cell and with a pair of brush organs; one brush originates in the posterior half of the discal cell and its distal portion is close to CuA 2, in a groove formed by a fold of the alar membrane that contains glandular structures, while the other, located in the basal third of the wing, on 2A–3A acts in conjunction with the abdominal scent organs ( Eltringham 1929; Barth 1952) ( Figs 4a View FIGURE 4 , 11a, b View FIGURE 11 ). VHW, in the basal region has spots and lines that form variable patterns; males and females with an eyespot between Sc+R 1 and M 1, larger than the eyespot between M 3 and CuA 2, both usually rounded; paler disruptive pattern mixed with ground color; with submarginal lines EII and EIII.

Abdomen. Males have scent organs in the pleura of A4 and A5, except Opsiphanes blythekitzmillerae , Opsiphanes boisduvallii and Opsiphanes camena Staudinger, [1886] , which have scent organs only on A4 ( Figs 11a, b View FIGURE 11 , 62a, b View FIGURE 62 ) (for more details see Eltringham 1929 and Barth 1952).

Male genitalia. Tegumen sub-quadrate in lateral view; ventral arms of tegumen and dorsal arms of saccus equal in length; anterior projection of saccus with variable length and width; uncus in dorsal view triangular, with basal half enlarged, tapering distally, and in lateral view curving ventrally; gnathos with expanded proximal and narrowly pointed distal regions, forming varied angles, apex distally projected; valva wider at the base, tapering distally, dorso-distal region asymmetrical, with varied angles on the ventro-distal margin; aedeagus straight, with an anterodorsal or dorsal opening to the ejaculatory bulb, distally pointed, with two sclerotized lateral projections, dorsal and subapical opening for the vesica, and median region with (“ quiteria group”) or without (“ cassiae group”) spines varying in number and arrangement; cordiform fultura inferior, variable in width.

Female genitalia. Tergum VIII rectangular; anterior apophysis atrophied; lamella antevaginalis membranous or sclerotized; lamella postvaginalis sclerotized, with the median portion of different shapes; ostium membranous or sclerotized; ductus bursae with the initial portion membranous or sclerotized; corpus bursae membranous, on the ventral side with two parallel or C-shaped signum; apophysis posterioris almost as long as tergum VIII; papilla analis with sclerotized anterior portion, taller than long, covered with long hair-like scales in the distal region.

Variation. Examination of series of specimens showed that both species and subspecies vary in the following structures: DFW, the size, shape, and color of the apical spots between R 3 and M 1. VFW, the width of the “c” element and the patterns of the “d” and “e” elements; the disruptive pattern in the postdiscal region; and the size and shape of the subapical eyespot between M 1 and M 3. VHW, the patterns of spots and lines in the basal region.

Comments. Characters such as the color, width, and arrangement of the spots that form the bands, mainly the FW oblique band, and the shape of the eyespots were the basis for the description of many Opsiphanes species and subspecies. These characters can be informative in some cases but should not be considered definitive by themselves, because of the variation that can be observed even in small series of specimens ( Stichel 1908). Additionally, the lack of genetic distance between specimens of the same species and from the same location, but with different phenotypes, confirms that these are variations. Molecular analyses, combining one mitochondrial gene and five nuclear genes, recovered Opsiphanes as monophyletic (SH-aLRT 91.4 / UFBoot 96) ( Fig. 2 View FIGURE 2 ), supporting previous hypotheses based on analyses of morphological and molecular data ( Penz 2007; Penz et al. 2013; Matos-Maraví et al. 2021). Considering the molecular analyses, morphological data for adults and immature stages, and larval feeding habits, Opsiphanes contains 23 species and 23 subspecies, in two groups: “ cassiae ” and “ quiteria ”. The “ quiteria ” group is subdivided into seven subgroups: “ boisduvallii ”, “ camena ”, “ zelotes ”, “ sallei ”, “ quiteria ”, “ fabricii ”, and “ invirae ”.

Key to species of Opsiphanes View in CoL

1. DHW with marginal band, without submarginal band ( Fig. 7a View FIGURE 7 ).................................. “ cassiae group” 2

- DHW with marginal and submarginal bands ( Fig. 66a View FIGURE 66 ), with only submarginal band ( Fig. 74b View FIGURE 74 ), or both absent ( Fig. 72 View FIGURE 72 )........................................................................................ “ quiteria group” 4

2(1). VHW with eyespot between Sc+R 1 and M 1 that usually does not reach or exceed M 1; without dark-brown band extending from base of wing to this eyespot ( Fig. 20a View FIGURE 20 )..................................................... Opsiphanes cassiae View in CoL

- VHW with rounded eyespot between Sc+R 1 and M 1, that usually reach or exceed M 1; with dark-brown band extending from base of wing to this eyespot ( Figs 4a View FIGURE 4 , 54a View FIGURE 54 )................................................................. 3

3(2). DFW with continuous oblique band; distal margin of this band is smooth ( Fig. 54a View FIGURE 54 )...... Opsiphanes mexicana stat. nov.

- DFW with discontinuous oblique band; distal margin of this band is uneven ( Fig. 4a View FIGURE 4 )............ Opsiphanes bogotanus View in CoL

4(1). DFW and DHW with orange ground color ( Figs 59 View FIGURE 59 , 61 View FIGURE 61 ) (“ boisduvallii subgroup”)................................. 5

- DFW and DHW with dark-brown ground color ( Fig. 66 View FIGURE 66 )..................................................... 6

5(4). VHW predominantly grayish; eyespot between Sc+R 1 and M 1 with wide dark-brown outline ( Fig. 59a View FIGURE 59 )............................................................................................ Opsiphanes blythekitzmillerae

- VHW predominantly yellow-gold; eyespot between Sc+R 1 and M 1 with narrow dark-brown outline ( Fig. 61a View FIGURE 61 )............................................................................................ Opsiphanes boisduvallii

6(4). DHW with marginal and submarginal bands ( Fig. 66a View FIGURE 66 )..................... Opsiphanes camena View in CoL (“ camena View in CoL subgroup”)

- DHW without marginal band, with ( Fig. 74b View FIGURE 74 ) or without ( Fig. 72 View FIGURE 72 ) submarginal band............................... 7

7(6). VFW and VHW with predominantly white-gray color and violet shades; male, DHW without submarginal band; female DHW with submarginal band ( Figs 72–75 View FIGURE 72 View FIGURE 73 View FIGURE 74 View FIGURE 75 , 80 View FIGURE 80 ) (“ zelotes View in CoL subgroup”).................................................. 8

- VFW and VHW with different color combinations; male and female DHW with submarginal band ( Figs 84 View FIGURE 84 , 87 View FIGURE 87 )......... 9

8(7). Male, DFW with a narrow yellow oblique band not exceeding CuA 2 ( Fig. 72a View FIGURE 72 ).................... Opsiphanes zelotes View in CoL

- Male, DFW with a wide yellow oblique band exceeding CuA 2 ( Fig. 80a View FIGURE 80 ).................. Opsiphanes zelus View in CoL stat. nov.

9(7). HW with strongly crenulated outer margin ( Fig. 84b View FIGURE 84 )....................................................... 10

- HW with smooth outer margin ( Fig. 165 View FIGURE 165 ) or with slight crenulation ( Fig. 187 View FIGURE 187 )................................... 16

10(9). VFW, on CuA 1 -CuA 2, with a yellow spot near the discal cell ( Fig. 84b View FIGURE 84 ); or if without this spot, oblique DFW band interrupted on CuA 1 and this band as wide as that of DHW (“ sallei View in CoL subgroup”)............................................ 11

- VFW, on CuA 1 -CuA 2, without a yellow spot near the discal cell ( Fig. 107a View FIGURE 107 ) (“ quiteria subgroup”).................... 14

11(10). VFW without yellow spot on CuA 1 -CuA 2 near discal cell. DFW with interrupted oblique band on CuA 1 and this band as wide as that of DHW...................................................................... Opsiphanes tiberius

- VFW with yellow spot on CuA 1 -CuA 2 near discal cell...................................................... 12

12(11). DFW with angled oblique band on M 3 -CuA 1 ( Fig. 95a View FIGURE 95 )........................................ Opsiphanes sallei View in CoL

- DFW with non-angled oblique band on M 3 -CuA 1 ( Fig. 84a View FIGURE 84 ).................................................. 13

13(12). DFW oblique band generally as wide as DHW submarginal band. DHW with distinct submarginal band extending from costal margin to anal angle ( Fig. 84a View FIGURE 84 ).................................................. Opsiphanes farrago View in CoL stat. nov.

- DFW oblique band generally twice as wide as DHW submarginal band. DHW with distinct submarginal band extending from the costal margin to M 3 ( Fig. 92a View FIGURE 92 )........................................................ Opsiphanes mutatus View in CoL

14(10). Male, DFW with narrow oblique band, interrupted on CuA 2 -2A ( Fig. 107a View FIGURE 107 ). Female with whitish oblique band, with uneven distal margin ( Fig. 108a View FIGURE 108 )........................................................ Opsiphanes badius View in CoL stat. rest.

- Male, DFW with wide continuous oblique band ( Fig. 123 View FIGURE 123 ). Female with whitish ( Fig. 126 View FIGURE 126 ) or yellow ( Fig. 138 View FIGURE 138 ) oblique band, with smooth distal margin............................................................................. 15

15(14). Male, DFW with oblique band yellow or orange, narrow or imperceptible on CuA 2 -2A ( Fig. 123a View FIGURE 123 ). Female, DFW with oblique band whitish or pale-yellowish; DHW in postdiscal region, between M 2 and CuA 2, rusty red ( Fig. 126a View FIGURE 126 )........................................................................................... Opsiphanes quirinus View in CoL stat. rest.

- Male, DFW with oblique band whitish or yellow, wide and distinct on CuA 2 -2A ( Fig. 134a View FIGURE 134 ). Female, DFW with oblique band yellow or orange ( Fig. 138a View FIGURE 138 ), if whitish, DHW in postdiscal region, between M 2 and CuA 2, without rusty red color ( Fig. 145a View FIGURE 145 ).............................................................................. Opsiphanes quiteria View in CoL

16(9). DFW with fully developed band inside discal cell ( Fig. 165b View FIGURE 165 ) (“ fabricii View in CoL subgroup”)............................... 17

- DFW with isolated spot ( Fig. 223h–k View FIGURE 223 ) or without spot inside discal cell (“ invirae subgroup”)........................ 22

17(16). VFW and VHW with violet shades ( Fig. 165 View FIGURE 165 )............................................................. 18

- VFW and VHW without violet shades ( Fig. 169 View FIGURE 169 ).......................................................... 20

18(17). Oblique band of DFW does not border discal cell on M 3 -CuA 1 ( Fig. 165a View FIGURE 165 ).............. Opsiphanes caliensis stat. nov.

- Oblique band of DFW borders discal cell on M 3 -CuA 1 ( Fig. 169a View FIGURE 169 )............................................. 19

19(18). Male and female, HW with smooth or slightly crenulated outer margin ( Figs 206–207 View FIGURE 206 View FIGURE 207 ). Female with yellow DFW and DHW bands ( Fig. 207 View FIGURE 207 )............................................................ Opsiphanes periphetes View in CoL stat. nov.

- Male and female, HW with crenulated outer margin ( Figs 187–188 View FIGURE 187 View FIGURE 188 ). Female, DFW with oblique whitish band and DHW with submarginal rusty red band ( Fig. 188a View FIGURE 188 )................................................. Opsiphanes jacobsorum

20(17). Female, DFW with band inside discal cell parallel to oblique band, tapering near costal margin; oblique band monochromatic yellow, nearly twice as wide as DHW submarginal band. DHW, submarginal band yellow and continuous, evident from costal margin to M 2 and from there to anal angle practically imperceptible ( Fig. 163 View FIGURE 163 )............ Opsiphanes barkeri stat. nov.

- DFW and DHW with another combination of characters.................................................... 21

21(20). Male, DFW with oblique band continuous on CuA 1 ( Fig. 169a View FIGURE 169 ). Female, corpus bursae with C-shaped signum ( Fig. 180b View FIGURE 180 )..................................................................................... Opsiphanes fabricii View in CoL

- Male: DFW with oblique band discontinuous on CuA 1 ( Fig. 192a View FIGURE 192 ). Female, corpus bursae with two parallel signa ( Fig. 200a View FIGURE 200 )......................................................................... Opsiphanes merianae stat. rest.

22(16). DFW without isolated spot inside discal cell; yellow oblique band may or may not reach 2A ( Fig. 213a View FIGURE 213 ).......................................................................................... Opsiphanes cuspidatus View in CoL stat. nov.

- DFW with or without isolated spot, of variable proportions, inside discal cell; yellow oblique band reaches 2A ( Fig. 219a View FIGURE 219 )...................................................................................... Opsiphanes invirae View in CoL

“ cassiae View in CoL group”

Species included:

1. Opsiphanes bogotanus Distant, 1875

a) Opsiphanes bogotanus bogotanus Distant, 1875

Opsiphanes tamarindi var. alba Stichel, 1902 nom. nud.

Opsiphanes tamarindi tamarindi View in CoL ab. spadix Stichel, 1902

Opsiphanes aurivillii Röber, 1906 View in CoL

Opsiphanes bogotanus phrataphernes Fruhstorfer, 1912 View in CoL syn. nov.

Opsiphanes bogotanus blandini Bristow, 1991 syn. nov.

b) Opsiphanes bogotanus castaneus Stichel, 1904 stat. rest.

Opsiphanes cassiae alajuela Bristow, 1991 syn. nov.

2. Opsiphanes cassiae ( Linnaeus, 1758) View in CoL

a) Opsiphanes cassiae cassiae ( Linnaeus, 1758)

Papilio glycerie Fabricius, 1787

Opsiphanes crameri C. Felder & R. Felder, 1862 View in CoL syn. rest.

Opsiphanes cassiae cassiculus Stichel, 1904 View in CoL

Opsiphanes cassiae rubigatus Stichel, 1904 View in CoL syn. nov.

Opsiphanes cassiae View in CoL f. pudicus Fruhstorfer, 1907

Opsiphanes cassiae f. decentius Fruhstorfer, 1907

Opsiphanes cassiae lucullus Fruhstorfer, 1907

Opsiphanes cassiae strophios Fruhstorfer, 1907 syn. nov.

Opsiphanes tamarindi xiphos Fruhstorfer, 1907 syn. nov.

Opsiphanes tamarindi pseudocassiae Fruhstorfer, 1907

Opsiphanes tamarindi cherocles Fruhstorfer, 1912 View in CoL

Opsiphanes cassiae appias J. Zikán & W. Zikán, 1968 nom. nud.

b) Opsiphanes cassiae incolumis Stichel, 1904 stat. nov.

Opsiphanes tamarindi latifascia Rothschild, 1916 syn. rest.

Opsiphanes bogotanus peruanus Rothschild, 1916 View in CoL

Opsiphanes tamarindi incolumis f. profana Stichel, 1936

Opsiphanes tamarindi douglasi Bryk, 1953

c) Opsiphanes cassiae tamarindi C. Felder & R. Felder, 1861 stat. nov.

Caligo tamarindi Boisduval, 1870 preocc.

Opsiphanes tamarindi corrosus Stichel, 1904 View in CoL syn. nov.

Opsiphanes tamarindi kleisthenes Fruhstorfer, 1912 View in CoL syn. nov.

Opsiphanes tamarindi sikyon Fruhstorfer, 1912 View in CoL

Opsiphanes tamarindi terenzius Fruhstorfer, 1912 View in CoL

Opsiphanes tamarindi tamarindi View in CoL f. mesomerista Stichel, 1925

3. Opsiphanes mexicana Bristow, 1991 stat. nov.

Diagnosis. The species of the “ cassiae group” are distinguished from their congeners by the following characters: known larvae feed on Zingiberales ; in the last instar, on the head, they have four pairs of scoli that are developed and larger than in the species of the “ quiteria group” and lack “whiskers” (set of setae near stemmata 1–4, present from the second instar in the species of the “ quiteria group”) ( Fig. 3 View FIGURE 3 ). DHW with a marginal band ( Fig. 7a View FIGURE 7 ) and aedeagus without spines on the surface ( Fig. 11f View FIGURE 11 ). Adults of all species with dark-brown ground color. Molecular analyses recovered the “ cassiae group” as monophyletic (SH-aLRT 100 / UFBoot 100) ( Fig. 2 View FIGURE 2 ).