Dynastor darius (Fabricius, 1775)

Piovesan, Mônica, Casagrande, Mirna Martins, Lamas, Gerardo & Mielke, Olaf Hermann Hendrik, 2024, Systematics of Dynastor Doubleday, [1849] (Lepidoptera: Nymphalidae, Satyrinae, Brassolini), Zootaxa 5492 (4), pp. 451-504 : 464-476

Dynastor darius (Fabricius, 1775)

( Figs 8–18 )

Papilio darius Fabricius, 1775 . Syst. Ent., p. 482, Mus. Banks; Brasilia [ Brazil].—Fabricius, 1807. S yst. Glossat., p. 289.— Hemming, 1943. Proc. r. ent . Soc. London (B) 12 (2): 26. —Zimsen, 1964. Type mat. I. C. Fabricius, p. 514.

Papilio anaxarete Cramer, 1776 . Pap. Exot. 1, p. 148, 151, pl. 95, figs A, B (male d, v); Suriname.—Stoll, 1780, in Cramer. Pap. Exot. 4, p. 167, pl. 374, figs A, B (female d, v).—Schneider, 1785. Nomencl. Ent., p. 36. Syn. nov.

Papilio anaxareta [sic]; Stoll, 1780, in Cramer. Pap. Exot. 4, p. 167.

Papilio anaxarethus [sic]; Herbst, 1793, in Jablonsky. Natursyst. Schmett. 6, p. 65, n. 34, pl. 131, figs 1, 2 (male d, v), 3, 4 (female d, v).

Potamis anaxarete ; Hübner, [1808]. Samml. exot. Schmett. 1, pls. 72, figs 3, 4 [male d, v].

Brassolis anaxarete ; Hübner, [1819]. Verz. bek. Schmett. (2–8), p. 50.

Pavonia anaxarete ; Godart, [1824], in Latreille & Godart. Enc. Méth. 9 (2), p. 438, 452, 807.

Pavonia darius ; Godart, [1824], in Latreille & Godart. Enc. Méth. 9 (2), p. 438, 452, 807.—Doubleday, 1844. List Lep. Brit. Mus. 1, p. 118; syn.: anaxarete .

Dynastor darius ; Westwood, 1851, in Doubleday. Gen. Diurn. Lep. 2, p. 347.—Lucas, 1851, in Chenu (ed.). Enc. d’Hist. nat., Pap. diurn., p. 171.—Herrich-Schäffer, 1865. Corr.-Blatt zool.-min. Ver. Regensburg 19 (5): 64; syn.: anaxarete .— Prittwitz, 1865. Stett. ent. Zeit. 26 (10/12): 310; syn.: anaxarete .—Butler, [1870]. Cat. Diurn. Lep. Fabricius, p. 40; syn.: anaxarete .—Kirby, 1871. Syn. Cat. Diurn. Lep., p. 127; syn.: anaxarete .— Burmeister, 1873. Rev. & Mag. Zool. 3 (1): 44, pl. VI, fig. 3 (larva); syn.: anaxarete .— Burmeister, 1879. Descr. phys. Rép. Arg. 5, Atlas, p. 23, pl. VII fig. 10 (larva); syn.: anaxarete .— Kirby, 1879. Cat. Coll. Diurn. Lep. Hewitson, p. 108.— Godman & Salvin, 1881. Biol. Centr. -Amer., Lep. Rhop. 1 (12), p. 123; syn.: anaxarete , stygianus .—W. Müller, 1886. Zool. Jahrb., Syst. 1 (3/4), p. 172; biol.—Staudinger, 1886, in Staudinger & Schatz. Exot. Schmett. 1 (15), pl. 72 (female d, v).— Staudinger, 1887, in Staudinger & Schatz. Exot. Schmett. 1 (18), p. 214; syn.: stygianus .—Schatz, 1888, in Staudinger & Schatz. Exot. Schmett. 2 (4), pl. 32 (morph.).— Röber, 1889, in Staudinger & Schatz. Exot. Schmett. 2 (5), p. 192.—Godman, 1901, in Godman & Salvin. Biol. Centr.-Amer., Lep. Rhop. 2 (164), p. 664.—Fassl, 1920. Ent. Rdsch. 37 (12): 45.— Campos, 1927. Rev. Col. Nac. V. Rocafuerte, Guayaquil, 9 (27–28): 46.—Costa Lima, 1928. Arch. Esc. Sup. Agr. Med. Vet. 8 (1/2): 138; hostplant.—Köhler, 1928. Publ. Breyer, Buenos Aires, p. 2.—Zikán, 1928. Ent. Rdsch. 45 (3): 10.—Bates, 1932. Bull. Brooklyn Ent. Soc. 27 (3): 155; biol.— Monte, 1934. Bol. Agric. Zoo. Vet. Sec. Agric., Minas Gerais, 21: 72, fig. 69 (female d, larva); biol.—F. Hoffmann, 1937. Int. ent. Ztschr. 50 (6): 57; biol.—Biezanko & Freitas, 1938. Cat. Ins. Pelotas, Fasc. 1, Lepidopteros, Pelotas, p. 13; biol.—Hall, 1939. Agric. Jour. Brit. Guiana 10 (1): 35.— Hayward, 1951. Acta zool. lill. 9: 220; syn.: anaxarete , stygianus , ictericus, faenius.—J. Zikán & W. Zikán, 1968. Pesq. agropec. bras. 3: 49; hostplant.—Lewis, 1973. Butt. World, p. 29, fig. 7.—Blandin & Descimon, 1975. Ann. Soc. ent. France 11 (1): 7.— Blandin, 1977. Publ. Lab. Zool. l’École norm. sup., Paris, 9: 163, 169, 185, 196, fig. 15 (geogr. distr.); biogeogr.— Kesselring & Ebert, [1982]. Rev. Nordestina Biol. 2 (1/2): 106.—Raymond, 1982. Marip. Venezuela, pl. 13, fig. 2 (female d, v).—Álvarez-Sierra & Álvarez-Corral, 1984. Marip. Venezuela, p. 129 (fig. female d); biol., hostplant. —Ackery, 1988. Biol. Jour. Linn. Soc. 33 (2): 100; hostplant.— Penz et al., 1999. Jour. Lep. Soc. 53 (4): 148; hostplant.—Uehara-Prado et al., 2004. Biota Neotrop. 4 (1): 4.—D’Abrera, 2005. World Butt., p. 94, 244, pl. 124, fig. 17 (male d).—Brown et al., 2007. Biol. Jour. Linn. Soc. 92 (3): 471; genet.— Penz, 2007. Syst. Ent. 32 (4): 669, 674, figs 1 (d, v), 8a (male gen.); phylog.— Garzón-Orduña & Penz, 2009. Zootaxa 2134: 1, figs 1e (male d, v), f (female d), 4, 6 (phylogenetic tree), 7 (wing), 8c, e, k (male gen.), 9a, b (female gen.). — Penz et al., 2013. Syst. Ent. 38 (2): 330; phylog.—Orlandin, 2016, in Orlandin et al., Borb. Marip. S. Catarina, p. 110.—Vélez-Estrada & Ríos-Málaver, 2018. Colombia País Marip., p. 462 (fig. v).— Vaughan et al., 2019. Jour. Nat. Hist. 53 (29–30): 1854.

Megastes darius ; Boisduval, 1870. Consid. Lép. env. Guatemala, p. 53; syn.: anaxarete .—Salinas-Gutiérrez, 2018. Rev. chil. Ent. 44 (3): 325.

Dynastor darius ictericus Stichel, 1904 . Ins.-Börse 21 (3): 21; 1 male [holotype], Sapucay n. Villa Rica, [Paraguarí, Guairá, Paraguay]; Forster leg, Mus. Tring.—Canals, 2003. Marip. Misiones, p. 361 (figs male d, v); biol.— Casagrande, 2004. Brassolini, in Lamas (ed.) Checklist: Part 4A, Hesperioidea-Papilionoidea, in Heppner (ed.). Atlas Neotrop. Lep. 5A, p. 203; syn.: mardonius .—Bustos, 2009. Trop. Lep. Res. 19 (2): 75. — Garzón-Orduña & Penz, 2009. Zootaxa 2134: 4. — Palo Jr., 2017. Borb. Brasil 2, p. 1453 (figs d, v).— Klimaitis et al., 2018. Marip. Argentina, p. 177 (figs male, female d, v); biol. Syn. nov.

Dynastor darius darius ; Stichel, 1904, in Wytsman. Gen. Ins. 20, p. 9; syn.: anaxarete .— Stichel, 1909. Das Tierreich 25: 21, 23, fig. 6; syn.: anaxarete .— Fruhstorfer, 1912, in Seitz. Gross-Schmett. Erde 5, p. 288.— Rothschild, 1916. Novit. Zool. 23 (3): 310, pl. 6, figs 13 (larva), 11 (pupa); biol.— Stichel, 1932. Lep. Cat. 51, p. 18.—Breyer, 1939. Physis 17 (49): 498.—Silva et al., 1968. Quarto Cat. Ins. viv. Plantas Brasil 2 (1), p. 328; biol., hostplant.—D’Abrera, 1987. Butt. Neotrop. Reg. 3, p. 394.—Urich & Emmel, 1991. Trop. Lep. 2 (2): 145, figs 1 (female d), 2 (egg), 3 (larva), 4, 5 (pupae); biol., hostplant.—Casagrande, 1992. Rev. bras. Ent. 36 (4): 723, fig. 6 (female d, v); biol., hostplant.— Casagrande, 2002. Rev. bras. Zool. 19 (2): 471; biol.— Casagrande, 2004. Brassolini, in Lamas (ed.) Checklist: Part 4A, Hesperioidea-Papilionoidea, in Heppner (ed.). Atlas Neotrop. Lep. 5A, p. 203.—Santin, 2004. Répert. plantes-hôtes subst. chen. lépid. monde (2 nd ed.), 2, p. 858; hostplant.— Romero et al., 2005. Jour. Lep. Soc. 59 (1): 37; biol., hostplant.—Garwood et al., 2007. Butt. South. Amaz., p. 110 (figs female d, v).—Carneiro et al., 2008. Shilap Revta. lepid. 36 (142): 264.— Garwood et al., 2009. Butt. South. Amaz. (2 nd ed.), p. 232 (figs male, female d, v).—Santos et al., 2011. Biota Neotrop. 11 (3): 272.—Penz & Mohammadi, 2013. Biota Neotrop. 13 (3): 175.—Palo Jr., 2017. Borb. Brasil 2, p. 1451 (figs d, v).—Melo et al., 2019. Biota Neotrop. 19 (1): 5.

Dynastor darius f. ictericus; Stichel, 1904, in Wytsman. Gen. Ins. 20, p. 9.

Dynastor darius f. icterica [sic]; Stichel, 1909. Das Tierreich 25: 24.— Stichel, 1932. Lep. Cat. 51, p. 19.

Dynastor mardonius Fruhstorfer, 1911 , in Seitz. Gross-Schmett. Erde 5, pl. 62b, fig. [2] (female d).

Dynastor darius anaxarete ; Fruhstorfer, 1912, in Seitz. Gross-Schmett. Erde 5, p. 289.— Stichel, 1932. Lep. Cat. 51, p. 19.—Blandin, 2003, in Lacomme & Manil. Lép. Guyane 1 (2), p. 18, 77.— Casagrande, 2004. Brassolini, in Lamas (ed.) Checklist: Part 4A, Hesperioidea-Papilionoidea, in Heppner (ed.). Atlas Neotrop. Lep. 5A, p. 203; syn.: populus .— Garzón-Orduña & Penz, 2009. Zootaxa 2134: 4.

Dynastor darius mardonius ; Fruhstorfer, 1912, in Seitz. Gross-Schmett. Erde 5, p. 289, two females; Paraguay.— Stichel, 1932. Lep. Cat. 51, p. 20.—Breyer, 1939. Physis 17 (49): 498.— Hayward, 1951. Acta zool. lill. 9: 220.—Biezanko, 1960. Arq. Ent., Esc. Agron. “Eliseu Maciel” (A), Pelotas, 4: 8; biol.—Biezanko, 1960. Arq. Ent., Esc. “Agron. Eliseu Maciel” (B), Pelotas, 4: 8; biol.—Biezanko & Ruffinelli, 1962. Rev. Fac. Agron., Montevideo, 50: 145.—Silva et al., 1968. Quarto Cat. Ins. viv. Plantas Brasil 2 (1), p. 328; biol., hostplant.—Hayward, 1969. Inst. Miguel Lillo, Misc. 31: 74; hostplant.—Hayward, 1973. Op. Lill. 23: 242.— Penz et al., 1999. Jour. Lep. Soc. 53 (4): 148; hostplant.— Casagrande, 2004. Brassolini, in Lamas (ed.) Checklist: Part 4A, Hesperioidea-Papilionoidea, in Heppner (ed.). Atlas Neotrop. Lep. 5A, p. 203.— Pastrana, 2004. Lep. arg., Plantas hosp., p. 180; hostplant.

Dynastor darius icterica [sic]; Fruhstorfer, 1912, in Seitz. Gross-Schmett. Erde 5, p. 289.

Dynastor darius faenius Fruhstorfer, 1912 , in Seitz. Gross-Schmett. Erde 5, p. 289; three males and two females; Candelária, Rio Grande do Sul, Brazil.— Stichel, 1932. Lep. Cat. 51, p. 20.—Silva et al., 1968. Quarto Cat. Ins. viv. Plantas Brasil 2 (1), p. 328; hostplant.— Casagrande, 2004. Brassolini, in Lamas (ed.) Checklist: Part 4A, Hesperioidea-Papilionoidea, in Heppner (ed.). Atlas Neotrop. Lep. 5A, p. 203.—Bonfantti et al., 2009. Checklist 5 (4): 823.— Garzón-Orduña & Penz, 2009. Zootaxa 2134: 4. — Leviski et al., 2016. Trop. Lep. Res. 26 (2): 66. Syn. nov.

Dynastor darius f. mardonius ; Köhler, 1923. Ztschr. wiss. Insektenb. 18 (12), Sonderb., p. 21.

Dynastor populus Röber, 1927 . Int. ent. Ztschr. 2 1 (18): 140, fig. 1; male [holo] type, French Guiana — Casagrande, 2004. Brassolini , in Lamas (ed.) Checklist: Part 4A, Hesperioidea-Papilionoidea, in Heppner (ed.). Atlas Neotrop. Lep. 5A, p. 203.

Dynastor darius f. populus ; Stichel, 1932. Lep. Cat. 51, p. 20.

Dynastor darius ssp.; Lamas, 1985. Rev. peruana Ent. 27: 69.

Taxonomic history. Dynastor darius , described as Papilio darius , was transferred to Pavonia by Godart ([1824]) and to Dynastor by Westwood (1851). Several authors interpreted Papilio anaxarete , and Dynastor stygianus as synonyms of Papilio darius . In addition to Papilio anaxarete and Dynastor stygianus, Hayward (1951) considered Dynastor darius ictericus and Dynastor darius faenius as synonyms of Papilio darius . Casagrande (2004) considered Dynastor populus as a synonym of Papilio anaxarete and Dynastor mardonius as a synonym of Dynastor darius ictericus . Due to the sampling location, the citation by Campos (1927) probably refers to Dynastor stygianus stat. rest.

Type material. Papilio darius Fabricius, 1775 was described based on an unspecified number of specimens from Brasilia [= Brazil, probably from the vicinity of Rio de Janeiro (Ilha Rasa in Guanabara Bay)] in the Banks collection, now in NHMUK. One female syntype was found at the NHMUK, with the following labels: / Type/ B.M. TYPE No. Rh. 6150 Papilio darius ♀ Fab. / Papilio darius Fab. Entom. p. 482, n. 173. To stabilize the nomenclature of this taxon, this specimen is hereby designated as the lectotype of Papilio darius Fabricius, 1775 , and the following labels will be attached: / LECTOTYPUS / Lectotypus Papilio darius Fabricius, 1775 , Piovesan, Casagrande & O. Mielke det. 2024/.

Papilio anaxarete Cramer, 1776 was described based on an unspecified number of specimens from Suriname. One female syntype was found at the NHMUK, with the following labels: / Nº 23. ANAXARETE . Cr. I. 95. A.B. & IV. 374. A.B./ FELDER COLLN./. To stabilize the nomenclature of this taxon, this specimen is hereby designated as the lectotype of Papilio anaxarete Cramer, 1776 , and the following labels will be attached: / LECTOTYPUS / Lectotypus Papilio anaxarete Cramer, 1776 , Piovesan, Casagrande & O. Mielke det. 2024/.

Dynastor darius ictericus Stichel, 1904 was described based on one male [holotype] from Sapucay [= Sapucaí or Sapucái] n[ear] Villa Rica, [Paraguarí, Guairá, Paraguay] leg. Forster, Mus. Tring. The male [holotype] is deposited at the NHMUK, with the following labels: / Sapucay, n. Villa Rica, ( Foster )./ Dynast. darius forma ictericus Stichel Type/.

Dynastor mardonius Fruhstorfer, 1911 was named based on the illustration of at least one female from Paraguay. One female syntype was found at the NHMUK, with the following labels: / Type/ Type H T/ Fruhstorfer Coll. B.M. 1937-285./ Paraguay ex coll. Fruhstorfer/ darius mardonius Fruhst. /. To stabilize the nomenclature of this taxon, this specimen is hereby designated as the lectotype of Dynastor mardonius Fruhstorfer, 1911 , and the following labels will be attached: / LECTOTYPUS / Lectotypus Dynastor mardonius Fruhstorfer, 1911 , Piovesan, Casagrande & O. Mielke det. 2024/.

Dynastor darius faenius Fruhstorfer, 1912 was described based on three males and two females from Candelária, Rio Grande do Sul, [ Brazil]. Two male syntypes were found at the NHMUK, with the following labels (1): / Type/ Rio Grande Brasil. Fruhstorfer/ faenius. 5-129 Fruh./ Fruhstorfer Coll. B.M. 1937-285./; and (2): / Type/ Rio Grande Brasil. Fruhstorfer/. D. darius faenius Fruhst. / Fruhstorfer Coll. B.M. 1937-285./ To stabilize the nomenclature of this taxon, the first syntype is hereby designated as the lectotype of Dynastor darius faenius Fruhstorfer, 1912 , and the following labels will be attached: / LECTOTYPUS / Lectotypus Dynastor darius faenius Fruhstorfer, 1912 , Piovesan, Casagrande & O. Mielke det. 2024/. The second syntype becomes automatically a paralectotype, and the following labels will be attached: / PARALECTOTYPUS / Paralectotypus Dynastor darius faenius Fruhstorfer, 1912 , Piovesan, Casagrande & O. Mielke det. 2024/.

Dynastor populus Röber, 1927 was described based on one male [holo]type from Französ. Guayana [= French Guiana]. M. M. Casagrande, O. H. H. Mielke, and G. Lamas did not find the male [holo]type. It is possible that, as for other species described by Röber, the [holo]type of Dynastor populus became lost as a direct influence of war ( Müller & Tennent 2018). Therefore, a female neotype of Dynastor populus Röber, 1927 is hereby designated to stabilize the nomenclature of this taxon. The specimen selected as neotype has the same morphological characters as illustrated in the original description and is deposited at the MfN, with the following labels: / coll. v. Schenck Surinam / populus Int. en. Z. Guben v. 21 Röber 1927/; the following labels will be attached: / NEOTYPUS / Neotypus Dynastor populus Röber, 1927 , Piovesan, Casagrande & O. Mielke det. 2024/.

Diagnosis. Dynastor darius differs from Dynastor stygianus stat. rest., a morphologically similar and closely related species ( Fig. 1 ), in having the VFW apical patch generally extending only to the apex, while Dynastor stygianus stat. rest. usually has this patch extending to the outer margin ( Figs 8–9 , 19–20 ). The male genitalia of both species are very similar ( Figs 10 , 21 ), but the female genitalia are not. While the female of Dynastor darius has a larger corpus bursae with two parallel signa and a broader lamella antevaginalis, the female of Dynastor stygianus stat. rest. has a smaller corpus bursae without signa and a smaller lamella antevaginalis ( Figs 11 , 22 ).

Variation. FW, length: male 46–56 mm (n=5), female 55–62 mm (n=5); width and color of the band that extends from the costal margin to M 3; size and shape of the spots between M 3 and 2A. VFW, shape and color of postdiscal patches and spots. DHW, width and color of the band that extends from the costal margin to CuA 1. VHW, number, size, and shape of the spots.

Comments. Dynastor darius shows variation in the wing pattern, which may explain the number of names proposed as subspecies. The examination of a series of specimens from several locations and the low DNA barcode divergence between specimens from different locations showed that many of the characters that supported the proposals of names, such as differences in the general color, the color of spots and bands, the shape of spots, the width of bands, and body size are variable. An example of variation is that some specimens from different locations in Brazil (Pará, Mato Grosso, Rio de Janeiro ) and Peru (Huánuco) have a narrower band and smaller spots on the dorsal surface of the forewings. However, genitalia does not present differences compared to other specimens of Dynastor darius , and even genetic distances do not corroborate the separation of species/subspecies. Therefore, we consider as synonymous with Dynastor darius the following names: Papilio anaxarete Cramer, 1776 , Dynastor darius ictericus Stichel, 1904 , Dynastor mardonius Fruhstorfer, 1911 , Dynastor darius faenius Fruhstorfer, 1912 and Dynastor populus Röber, 1927 . Dynastor darius is sympatric with Dynastor macrosiris and Dynastor napoleon .

Natural history. Information on the immature stages of Dynastor darius can be found in Burmeister (1873, 1879), Monte (1934), Rothschild (1916), Biezanko (1960a; b), Urich & Emmel (1991b), Romero et al. (2005); hybridization with Dynastor napoleon in Otero & Casagrande (1992). Dynastor darius flies in the morning and evening twilight on the coast of Paraná, south of Brazil, at an elevation of 100 m. According to Biezanko (1960b), it is bivoltine, flying in March, April, May, October, and November; based on the label data, we found that the species flies throughout the year. Urich & Emmel (1991b) mentioned that Dynastor darius and other Dynastor species do not feed in adulthood. However, Romero et al. (2005) observed adults at dusk feeding on decaying mangoes in two locations in Venezuela: Palmichal (Carabobo) and Choroní (Aragua).

Hostplants: Bromeliaceae : Bromelia antiacantha Bertol. , Ananas sativus Schult. & Schult.f. ( Biezanko 1960a; b), Billbergia nutans H.H.Wendl. ex Regel , Billbergia speciosa Thunb. , Aechmea fasciata (Lindl.) Baker , Aechmea gamosepala Wittm. ( Biezanko 1960b) , Ananas comosus (L.) Merril ( Penz et al. 1999; Romero et al. 2005), Aechmea nudicaulis (L.) Griseb ( Urich & Emmel 1991b), Vriesea Lindl. ( Otero & Casagrande 1992) .

Geographic distribution. Ecuador (Napo, Morona-Santiago); Peru (Huánuco, San Martín, Junín, Loreto, Cuzco, Madre de Dios); Bolivia (Santa Cruz, La Paz); Trinidad and Tobago (Tunapuna-Piarco); Venezuela (Caracas, Aragua, Carabobo); Guyana (Mazaruni-Potaro); Suriname (no data); French Guiana (no data); Brazil (Pará, Amazonas, Mato Grosso, Mato Grosso do Sul, Ceará, Pernambuco, Alagoas, Bahia, Tocantins, Distrito Federal, Minas Gerais, Espírito Santo, Rio de Janeiro , São Paulo , Paraná, Santa Catarina, Rio Grande do Sul); Paraguay (Iguazú, Cordillera, Central, Guairá); Argentina (Misiones (Palo Jr. 2017), Corrientes ( Klimaitis et al. 2018), Chaco, Formosa, Santa Fe ( Pastrana 2004), Salta, Tucumán) ( Fig. 12 ), at elevations between 10 and 2,200 m.

Temporal distribution. Throughout the year.

Etymology. The name probably refers to the ancient Persian kings.

Specimens examined. Photos of the lectotype of Dynastor darius ; lectotype of Papilio anaxarete ; holotype of Dynastor darius ictericus ; lectotype of Dynastor mardonius ; lectotype of Dynastor darius faenius ; neotype of Dynastor populus ( Figs 13–18 ), and the following specimens: ECUADOR: Ecuador Oriente —no locality, 2 males, ex-coll. F. Johnson, AMNH_IZC 00352870, 00352871 (AMNH). Napo —no locality, Yanahurco, IX-1978, 1 female, Lefebre leg., DZ 54.264 (DZUP). [Morona-Santiago]—Macas, VII-1921, 1 female, E. W. Rorer leg., USNMENT01589337 (USNM). PERU: no department—no locality, 1980, 1 male, ex-coll. M. Simon, UFFLMNH MGCL 1107799 (MGCL); Río Huallaga , 1 male, ex-coll. F. Johnson, AMNH_IZC 00352861 (AMNH). Loreto — Iquitos, 1 male, ex-coll. F. Johnson, AMNH_IZC 00352864 (AMNH), 100 m, VII-1988, 1 male, Büche leg., OM 35.126 (DZUP-OM) Yurimaguas, 05°54’S, 76°06’W, 150 m, n.d., 1 male, 1 female, O. Razzeto leg. (MUSM). Huánuco —no locality, X-1976, 1 male, USNMENT00941129 (USNM); Río Huallaga (15 km N of Tingo María), 15–22-VIII-1981, 1 female, ex-coll. M. Rojas, UFFLMNH MGCL 1107801 (MGCL); Tingo María, 670 m, IX-1990, 1 female **, Büche leg., OM 35.156 (DZUP-OM), 30-VII-1980, 1 female, D. & J. Jenkins leg., UFFLMNH MGCL 1107835 (MGCL), [09°18’S, 76°00’W, 670 m], n.d., 1 male (MUSM); Cordillera del Sira, ca. 09°25’S, 74°45’W, 800 m, IX-1987 – VIII-1988, 1 male, Exp. Univ. Viena leg. (MUSM). San Martín —Moyobamba (Jepelacio), 1 male, ex-coll. F. Johnson, AMNH_IZC 00352860 (AMNH). Tarapoto, 06°29’S, 76°22’W, 350 m, 9-VI-2005, 1 female, A. Silva leg (MUSM). Junín —Satipo, 1 male, 1 female, ex-coll. F. Johnson, AMNH_IZC 00352866, 00352867 (AMNH), V-1944, 1 male, II-1945, 1 female, P. Paprzycki leg., AMNH_IZC 00352862, 00352868 (AMNH); Chanchamayo, 1 male, ex-coll. F. Johnson, AMNH_IZC 00352863 (AMNH), (La Merced) [11°03’S, 75°19’W], 800 m, 1932, 2 males MHN 1597 (MUSM), H[acien]da Chiriyacu [11°05’S, 75°18’W], 1200 m, 6-II-1942, 1 male, J. M. Schunke leg., MHN 1597 (MUSM), Río Perené [ca. 11°09’S, 74°18’W, 600 m], n.d., 1 female, MHN 1596 (MUSM), n.d., 2001, 1 female, H. Rojas leg. (MUSM). Cuzco —Urubamba Valley, (Quillabamba), 12º51’, 72º41’, 2,200 m, 1 male, R. E. Espinoza leg., OM 69.331 (DZUP-OM) [the altitude given is spurious]. Madre de Dios —Puerto Maldonado, VII-1988, 1 male, Büche leg., OM 35.519 (DZUP-OM). BOLIVIA: La Paz —Yungas de La Paz, 1,000 m, 2 males, H. Rolle leg., UFFLMNH MGCL 1107802, 1107804 (MGCL); Caranavi, (Caranavi Area), 1,100 –2,100 m, XII-2002 – I-2003, 2 females, Tello leg. OM 61.275, OM 61.303 (DZUP-OM). Santa Cruz —Buenavista (Ichilo), 400 m, II-1958, 1 female, UFFLMNH MGCL 1107836 (MGCL). TRINIDAD AND TOBAGO: Tunapuna-Piarco —Saint Augustine, 12-III-1937, 1 female, UFFLMNH MGCL 1107812 (MGCL). VENEZUELA: Caracas —no locality, 1 male, AMNH_IZC 00352858 (AMNH). Aragua —El Limón, 4-I-1971, 1 male, Pantoja leg., DZ 54.261 (DZUP), 26-IX-1975, 1 female, Fernández H. leg., DZ 54.262 (DZUP); Maracay, 450 m, 28-II-1975, 1 female, Fernández B. leg., DZ 54.263 (DZUP). Carabobo —Puerto Cabello, 16-VII-1979, 1 male, J. B. Sullivan, USNMENT01589324 (USNM). GUYANA: no subdivision —Mazaruni-Potaro (Kaieteur Falls), 5º 14’ N, 59º 33’ W, 200–450 m, 18–25-XII-1989, 1 male, S. Fratello leg., USNMENT01589325 (USNM). BRAZIL: Pará —no locality, 1 female, ex-coll. F. Johnson, AMNH_IZC 00352890 (AMNH); Rio Arapiuns , VII-1978, 1 female, DZ 54.265 (DZUP), I-1982, 1 male, III-1980, 1 female, VIII-1982, 1 female, Kesselring leg., OM 91.408, OM 12.315, OM 91.448 (DZUP-OM); Rio Arapiuns-Rio Tapajós , IX-1927, 1 female *, DZ 54.266 (DZUP); Óbidos, 1958, 1 female, DZ 54.267 (DZUP), I-VIII-1982, 1 male *, Kesselring leg., OM 91.418 (DZUP-OM), 21-IX-1983, 1 male, Miers leg., OM 91.398 (DZUP-OM); Oriximiná (Porto Trombetas), 11-VII-1980, 1 female **, Gifford leg., ex-coll. Gifford, DZ 52.732 (DZUP); Santo Antonio de Tauá (Reserva Sonho Azul), 11-IV-2005, 1 male */**, P. Jauffret leg., DZ 52.727 (DZUP); Itaituba ( Rio Tapajós ), 1 male, ex-coll. LeMoult UFFLMNH MGCL 1107772 (MGCL). Amazonas -Manaus, INPA sede, 3º5’44”S, 59º59’15”W, 07-VII-1996, 1 female, C. S. Motta leg., 15- X-1976, 1 female, L. Albuquerque leg. (INPA); Maués, 3-III-2008, 1 female, OM 70.538 (DZUP-OM). Mato Grosso —Diamantino, 2-V-1990, 1 female, DZ 54.270 (DZUP), ( Fazenda São João ), 300–400 m, 11-II-1978, 1 male, E. Furtado leg., ex-coll. H. Ebert, DZ 54.271 (DZUP), 15-VI-1978, 1 female, Furtado leg., OM 91.428 (DZUP-OM). Ceará —Crateús (RPPN Serra das Almas ), 630 m, 23-V-2014, 1 male */**, Melo & Rosa leg., DZ 52.725 (DZUP). Pernambuco —no locality, 1 female, ex-coll. F. Johnson, AMNH_IZC 00352887 (AMNH); Pombos, 30- IV-2002, 2 females **, N. Souza leg., DZ 52.729, 52.730 (DZUP); Camaragibe, (Aldeia Bonsai), X-2021, 1 male, P. Grossi leg., OM 93.763 (DZUP-OM). Alagoas — Rio Largo (Fazenda Riachão) , 4-VIII-1992, 1 male, Rego leg., DZ 54.272 (DZUP); Maceió, 26-VI-1946, 1 female, V-1967, 1 male, 7-XI-1980, 1 male, A. Cardoso leg., ex-coll. A. Cardoso, DZ 54.273, 54.274, 51.848 (DZUP). Bahia —Camacan (Reserva Serra Bonita ), 200 m, 3–10-XII-2016, 1 male **, Mielke, Carneiro, Dias, Dolibaina & Santos leg., DZ 52.731 (DZUP). Tocantins —Porto Nacional, 27- VI-1974, 1 female, DZ 54.279 (DZUP). Distrito Federal — Rio Maranhão, 27-VII-2001, 1 female, Emery leg., ex-coll. E. Emery, DZ 54.275 (DZUP); Brasília (Reserva Ecológica IBGE), 2-IX-2011, 1 male **, O. Mielke leg., DZ 52.728 (DZUP), (UnB), 16-V-1979, 1 female, Rocha leg., DZ 54.276 (DZUP), 10-X-1977, 1 female, 18-III-1977, 1 female, A. Negrett leg., ex-coll. Gifford, DZ 54.277, 54.278 (DZUP); Planaltina (Córrego Grotão), 912 m, 4-II-2012, 1 male */**, Emery leg., ex-coll. E. Emery, DZ 52.733 (DZUP). Mato Grosso do Sul —Bela Vista, 1931, 2 females, ex-coll. A. Cardoso, DZ 54.268, 54.269 (DZUP). Minas Gerais —Pirapora, 16-XI-1977, 1 male, Moure & Sakakibara leg., DZ 54.281 (DZUP); Barbacena ( Serra da Mantiqueira ), 1,100 m, 5-X-1952, 1 male, H. Ebert leg., ex-coll. H. Ebert, DZ 54.280 (DZUP); Uberaba, 1 male, ex-coll. LeMoult, UFFLMNH MGCL 1107774 (MGCL); Carmo do Rio Claro , 11-II-1959, 1 male, 06-II-1959, 1 female, O. Mielke leg., OM 2617, OM 2616 (DZUP-OM); Conceição da Aparecida, 04-II-1960, 1 male, O. Mielke leg., OM 3173 (DZUP-OM); Conceição dos Ouros, 25-XII-2022, 1 male, Pereira leg., OM 92.358 (DZUP-OM), ( Serra Grande ), 1,365 m, 08-II-2023, 1 male, O. Mielke & Casagrande leg., OM 59.064 (DZUP-OM). Espírito Santo —Santa Teresa, 14-XI-1967, 1 female, C. & C.T. Elias leg., DZ 54.282 (DZUP); Conceição da Barra, 26-VIII-1969, 1 male *, C. & C.T. Elias leg., DZ 2338 (DZUP). Rio de Janeiro —no locality, 1 male, USNMENT01589326 (USNM), 1 male, ex-coll. W. Schaus, USNMENT01589329 (USNM), 1928, 1 male, UFFLMNH MGCL 1107791 (MGCL), 1 male, ex-coll. A. Cardoso, DZ 54.284 (DZUP), 150 m, 17-III-1951, 1 male, 6–9-II-1968, 1 female, H. Ebert leg., ex-coll. H. Ebert, DZ 54.293, 54.294 (DZUP); Saquarema, 23-VI-1941, 1 male, DZ 54.283 (DZUP); Nova Friburgo, II-2007, 1 male **, 1 female **, E. Grossi leg., DZ 52.724, 51.858 (DZUP); Rio de Janeiro (Jacarepaguá—Três Rios), 14-VI-1965, 1 female, ex-coll. D’Almeida, DZ 54.285 (DZUP), ( Leme ), 2-XII-1961, 1 female, ex-coll. R. Frey, DZ 54.286 (DZUP), (Piedade), 10-VI-1924, 1 male, 2-I-1925, 1 female, Ferreira D’Almeida, ex-coll. D’Almeida, DZ 54.287, 54.288 (DZUP), ( Tijuca ), 1970, 1 male, C. Callaghan leg., UFFLMNH MGCL 1107788 (MGCL); Petrópolis, 5-IV-1962, 1 male, 30-X-1959, 1 female, 20-X-1965, 1 male, ex-coll. Gagarin, DZ 54.290 –54.292 (DZUP), (Independência), 900 m, 15-XI-1935, 1 female, Gagarin leg., ex-coll. Gagarin, DZ 54.289 (DZUP); Itatiaia, (Parque Nacional de Itatiaia), 900 m, 28-VI-1962, 1 female, 1 male, O. Mielke leg., OM 2232, OM 4477 (DZUP-OM). São Paulo —no locality, 28-I-1974, 1 female, ex-coll.A. Cardoso, DZ 54.295 (DZUP), 2 males, ex-coll. F. Lemmer, AMNH_IZC 00352877, 00352880 (AMNH), (Alto da Serra ), 4-I-1937, 1 male, 22-XII-1936, 1 male, Travassos F. leg., ex-coll. D’Almeida, DZ 54.298, 54.299 (DZUP); Rio Claro , 600 m, 4-X-1969, 1 female, H. Ebert leg., ex-coll. H. Ebert, DZ 54.297 (DZUP); Campinas, X-1976, 1 male, Ebert leg., ex-coll. H. Ebert, DZ 54.302 (DZUP), 750 m, 6-XII-1976, 1 male, 26-XII-1976, 1 female, Ebert leg., ex-coll. H. Ebert, DZ 54.303, 54.304 (DZUP); Mogi das Cruzes, 600 m, 6-VIII-1927, 1 male, ex-coll. H. Ebert, DZ 54.301 (DZUP); São Sebastião ( Praia de Juqueí ), 22-II-1977, 1 female, A. Cardoso leg., ex-coll. A. Cardoso, DZ 54.296 (DZUP); Brotas (Campo Alegre), 250 m, 1-IV-1966, 1 male, Ebert leg., ex-coll. H. Ebert, DZ 54.300 (DZUP). Paraná —no locality, 1 female, ex-coll. F. Johnson, AMNH_IZC 00352876 (AMNH); Campina Grande do Sul, 2-II-2021, 1 male, 26-I-2021, 1 female, M. Casagrande leg., DZ 52.565, 52.566 (DZUP); Querência do Norte (RPPN Fazenda Santa Fé), 23°05’37”S, 53°35’43”W, 230 m, 11–12-IX-2018, 1 male, Casagrande, Mielke, Carneiro e Orlandin leg., DZ 57.267 (DZUP); Paranaguá ( Ipanema ), 20-XI-1993, 1 male, O. Mielke leg., OM 37.927 (DZUP-OM), (Floresta Estadual do Palmito), 10–20 m, 26-VIII-2014, 1 male, Leviski, Queiroz-Santos & Santos leg., DZ 54.305 (DZUP); Guaíra, 29-V-1983, 1 male *, Lingnau leg., DZ 54.306 (DZUP); Pontal do Paraná (Atami), 10 m, 26-XII-2007, 1 male, Mielke leg., DZ 54.307 (DZUP); Tibagi, 2-X-2003, 1 male, 15-XI-2003, 1 female, O.—C. Mielke leg., OM 63.038, OM 63.361 (DZUP-OM), 3-V-2003, 1 male, O. Mielke leg., OM 61.322 (DZUP-OM), 27- X-2003, 1 male, 1 female, 1-XI-2003, 1 female, 6-XI-2003, 1 male, O. & C. Mielke leg., DZ 54.308 –54.311 (DZUP); Guarapuava, 900 m, 13-II-1980, 1 female, 18-II-1980, 2 males *, 27-II-1980, 1 male, 28-II-1980, 2 females, O.-C. Mielke & Miers leg., DZ 54.312 –54.317 (DZUP); Ponta Grossa, XI-1939, 1 male, ex-coll. D’Almeida, DZ 54.318 (DZUP), IV-1941, 1 female, Hatschbach leg., ex-coll. D’Almeida, DZ 54.326 (DZUP), I-1954, 1 female, II-1944, 1 female, X-1954, 1 female, IX-1954, 1 male, IX-1940, 1 male, IX-1957, 1 male, VII-1939, 1 male, ex-coll. F. Justus Jr, DZ 54.319 –54.325 (DZUP), IX-1952, 1 female, X-1952, 1 female, F. J. Jr. leg., UFFLMNH MGCL 1107792, 1107793 (MGCL);Arapoti, XII-1947, 1 female, ex-coll.F.Johnson,AMNH_IZC 00352886 (AMNH); Castro, 1 male, ex-coll. W. Schaus, USNMENT01589332 (USNM). Santa Catarina —no locality, 1 male, USNMENT01589331 (USNM), 1 male, 1 female, ex-coll. E. T. Owen, USNMENT01589333, 01589334 (USNM), 1 male, H. Gerstner leg., ex-coll. Emem, UFFLMNH MGCL 1107798 (MGCL), 1 female, AMNH_IZC 00352882 (AMNH), 1 female, ex-coll. H. Edwards, AMNH_IZC 00352884 (AMNH), 2 males, ex-coll. F. Johnson, AMNH_IZC 00352873, 00352874 (AMNH); Santa Catarina — Seara (Nova Teutônia), II-1982, 1 male, 1 female, I-1982, 2 males, Fritz Plaumann leg., DZ 54.330, 54.327–54.329 (DZUP); São Bento do Sul, 850 m, XII-1973, 1 male, XII-1972, 1 male, ex-coll. H. Ebert, DZ 54.331, 54.332 (DZUP), 10-IX-1970, 1 female *, Weiss leg., ex-coll. Gagarin, DZ 2531 (DZUP), 26-III-1971, 1 female, ex-coll. Gagarin, DZ 54.333 (DZUP), ( Rio Vermelho ), 850 m, 8-XII-1999, 1 male, 12-II-1972, 1 female, 24-XI-1971, 1 male, 22-II-1971, 1 male, Rank leg., DZ 54.334 –54.337 (DZUP), X-1969, 1 male, USNMENT00941195 (USNM); Florianópolis, Lagoa do Peri, 19-X-2003, 1 male **, Mielke & Carneiro leg., DZ 52.726 (DZUP); Mafra, XI-1969, 2 males, 2 females, Weiss leg., ex-coll. H. Ebert, DZ 54.338 –54.341 (DZUP), 9-III-1997, 1 female, Reynavo leg., DZ 54.342 (DZUP); Joinville, VIII-1970, 1 female *, 19-X-1980, 1 female *, Miers leg., DZ 2337, 54.343 (DZUP), 17-IX-1970, 1 male *, ex-coll. Gagarin, DZ 54.344 (DZUP), II-1984, 2 males, Miers leg., ex-coll. D. & J. Jenkins, UFFLMNH MGCL 1107794, 1107795 (MGCL), II-1984, 2 males, Miers leg., UFFLMNH MGCL 1107829, 1107832 (MGCL), VI-1969, 1 female, ex-coll. Flaspohler, UFFLMNH MGCL 1107797 (MGCL), 21-IX-1973, 1 male, 2-XI-1973, 1 male, 1 female, USNMENT01589328, 00941193, 00941194 (USNM); Corupá, XII-1968, 1 male, ex-coll. Flaspohler, UFFLMNH MGCL 1107796 (MGCL), 1 male, ex-coll. F. Johnson, AMNH_IZC 00352875 (AMNH), 1 male, 1 female, AMNH_IZC 00352879, 00352883 (AMNH). Rio Grande do Sul —Pelotas, 18-III-1939, 1 male, Rosa Figueiredo leg., DZ 54.345 (DZUP), 3-IV-1956, 1 male, 18- I-1956, 1 female, 30-XI-1957, 1 female, 1-XI-1956, 1 male, C. Biezanko leg., AMNH_IZC 00352892–00352895 (AMNH). PARAGUAY: [Alto Paraná]—Iguazú, 24-X-2007, 1 female, UFFLMNH MGCL 1107833 (MGCL). Cordillera —Santísima Trinidad, 25º 15’ S, 57º 38’ W, VI, 1 male, B. Podtiaguin leg., AMNH_IZC 00352897 (AMNH). Central —Nueva Itália, 23-XI-1940, 1 male, P. Willim leg., ex-coll. F. E. Church, AMNH_IZC 00352898 (AMNH). ARGENTINA: Salta —Tartagal, 500 m, 11-III-1975, 1 female, B. N. MacPherson leg., UFFLMNH MGCL 1107837 (MGCL). Tucumán —Yerba Buena, 700 m, 8-II-1991, 1 male, K. Johnson leg., AMNH_IZC 00352899 (AMNH).

iNaturalist records: BRAZIL: Mato Grosso — Alta Floresta , 9°35’50.4”S, 55°55’49.1”W, 16-XII-2021 , Diego Ferreira (available at: inaturalist.org/observations/104314945. Accessed June 30, 2023) .

Figures

FIGURE 1. Phylogenetic tree (Maximum Likelihood) inferred with IQ–TREE based on five molecular markers: cytochrome c oxidase subunit I (COI) (mitochondrial), elongation factor 1–alpha (EF–1α), ribosomal protein S5 (RpS5), wingless (wg), Glyceraldehyde–3–phosphate dehydrogenase (GAPDH) (nuclear), rooted in Bia actorion (Linnaeus, 1763). Numbers indicate SH-aLRT/UFBoot.The scale bar represents the expected number of nucleotide substitutions per site. Data of the specimens used are described in Table 1.

FIGURE 8. Dynastor darius (Fabricius, 1775): Male, a. dorsal, b. ventral views; Brazil, Paraná (DZ 52.565–DZUP). The black arrows indicate the diagnostic characters of the species.

FIGURE 9. Dynastor darius (Fabricius, 1775): Female, a. dorsal, b. ventral views; Brazil, Paraná (DZ 52.566–DZUP).

FIGURE 10. Dynastor darius (Fabricius, 1775): Male genitalia: a. dorsal view, b. ventral view, c. lateral view; aedeagus: d. dorsal view, e. ventral view, f. lateral view (DZ 54.314–DZUP).

FIGURE 11. Dynastor darius (Fabricius, 1775): Female genitalia: a. lateral view; b. detail of the signa; c. ventral view (DZ 54.266–DZUP). The black arrows indicate the diagnostic characters of the species.

FIGURE 12. Geographical distribution of Dynastor darius (Fabricius, 1775) and Dynastor stygianus stat. rest. Asterisk (*) and at (@) inside the sampled locality indicate that the geographical distribution data was retrieved from the bibliography and iNaturalist (www.inaturalist.org), respectively.

FIGURE 13. Dynastor darius (Fabricius, 1775). Lectotype of Dynastor darius (Fabricius, 1775); a. dorsal view, b. ventral view.

FIGURE 14. Dynastor darius (Fabricius, 1775). Lectotype of Papilio anaxarete Cramer, 1776; a. dorsal view, b. ventral view.

FIGURE 15. Dynastor darius (Fabricius, 1775). [Holotype] of Dynastor darius ictericus Stichel, 1904; a. dorsal view, b. ventral view.

FIGURE 16. Dynastor darius (Fabricius, 1775). Lectotype of Dynastor mardonius Fruhstorfer, 1911; a. dorsal view, b. ventral view.

FIGURE 17. Dynastor darius (Fabricius, 1775). Lectotype of Dynastor darius faenius Fruhstorfer, 1912; a. dorsal view, b. ventral view.

FIGURE 18. Dynastor darius (Fabricius, 1775). Neotype of Dynastor populus Röber, 1927; a. dorsal view, b. ventral view.

FIGURE 19. Dynastor stygianus Butler, 1872 stat. rest.: Male, a. dorsal, b. ventral views; Costa Rica, Guanacaste (USNMENT01589299–USNM). The black arrow indicates the diagnostic character of the species.

FIGURE 20. Dynastor stygianus Butler, 1872 stat. rest.: Female, a. dorsal, b. ventral views; Costa Rica, Guanacaste (USNMENT01589316–USNM).

FIGURE 21. Dynastor stygianus Butler, 1872 stat. rest.: Male genitalia: a. dorsal view, b. ventral view, c. lateral view (the uncus is damaged); aedeagus: d. dorsal view, e. ventral view, f. lateral view (USNMENT01589295–USNM).

FIGURE 22. Dynastor stygianus Butler, 1872 stat. rest.: Female genitalia: a. lateral view; b. detail of corpus bursae without signa; c. ventral view (USNMENT01589316–USNM). The black arrows indicate the diagnostic characters of the species.