Dynastor Doubleday, [1849]

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 : 455-458

Dynastor Doubleday, [1849]

( Figs 1–42 )

Dynastor Doubleday, [1849] . Gen. diurn. Lep. 1, pl. 58, fig. 2 (d); Brazil; only included species: Dynastor napoleon .— Westwood, 1851, in Doubleday. Gen. diurn. Lep. 2, p. 346; syn.: Megastes Boisduval, MS. —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): 310.—Butler, [1870]. Cat. Diurn. Lep. Fabricius, p. 40.—Kirby, 1871. Syn. Cat. Diurn. Lep., p. 127.—Burmeister, l873. Rev. & Mag. Zool. 3 (1): 43; biol.— Scudder, 1875. Proc. Amer. Ac. Arts & Sc. 10 (2): 160, 213; syn.: Megastes Boisduval, 1870 .—Wallace, 1876. Geogr. Distr. Anim. 2, p. 472.—Kirby, 1877. Syn. Cat. Diurn. Lep., Suppl., p. 716.— Kirby, 1879. Cat. Coll. Diurn. Lep. Hewitson, p. 108.— Burmeister, 1879. Descr. phys. Rép. Arg. 5, Atlas, p. 23; biol.— Godman & Salvin, 1881. Biol. Centr.-Amer., Lep. Rhop. 1 (12), p. 122.—Gerhard, 1883. Berl. ent. Ztschr. 27 (2): 179.—W. Müller, 1886. Zool. Jahrb., Syst. 1 (3/4), p. 172; biol.— Staudinger, 1887, in Staudinger & Schatz. Exot. Schmett. 1 (18), p. 214.— Glaser, 1887. Cat. etymol. Col. Lep., p. 287; etymol.— Röber, 1889, in Staudinger & Schatz. Exot. Schmett. 2 (5), p. 191.—Reuter, 1896. Acta Soc. Sc. Fenn. 22 (1): 410; morph.— Stichel, 1902. Berl. ent. Ztschr. 46 (4): 523.— Stichel, 1904, in Wytsman. Gen. Ins. 20, p. 8, pl. 1, fig. 2 (venation).—Pagenstecher, 1909. Geogr. Verbr. Schmett., p. 413.— Stichel, 1909. Das Tierreich 25: 20.— Fruhstorfer, 1912, in Seitz. Gross-Schmett. Erde 5, p. 288.— Rothschild, 1916. Novit. Zool. 23 (3): 309.— Stichel, 1932. Lep. Cat. 51, p. 17.—Bates, 1932. Bull. Brooklyn Ent. Soc. 27 (3): 155; biol.—Hemming, 1941. Jour. Soc. Bibl. Nat. Hist. 1 (11): 421.— Hayward, 1951. Acta zool. lill. 9: 219.—Ehrlich, 1958. Univ. Kansas Sci. Bull. 39 (8): 351; morph.—L. Miller, 1968. Mem. Amer. Ent. Soc. 24: 25.—Hayward, 1973. Op. Lill. 23: 241.— DeVries, 1987. Butt. Costa Rica, p. 249; biol., ecol.—Ackery, 1988. Biol. Jour. Linn. Soc. 33 (2): 99; hostplant.—Bristow, 1991. Zool. Jour. Linn. Soc. 101 (3): 214.— Casagrande, 2002. Rev. bras. Zool. 19 (2): 468; biol.— Casagrande, 2004. Brassolini , in Lamas (ed.) Checklist: Part 4A, Hesperioidea-Papilionoidea, in Heppner (ed.). Atlas Neotrop. Lep. 5A, p. 203; syn.: Megastes Westwood, 1851 , unavail. (ICZN, Art. 11.6), Megastes Boisduval, 1870 .— Penz, 2007. Syst. Ent. 32 (4): 670; phylog.—Peña & Wahlberg, 2008. Biol. Lett. 4 (3): 275; phylog.— Wahlberg et al., 2009. Proc. r. Soc. B 276 (1677): 4298; phylog.— Garzón-Orduña & Penz, 2009. Zootaxa 2134: 1, figs 1, 4, 6, 7d, 8c, e, g, h, j–l. — Penz et al., 2013. Syst. Ent. 38 (2): 318; phylog. — Penz & Mohammadi, 2013. Biota Neotrop. 13 (3): 159.— Matos-Maraví et al. 2021. Biol. Jour. Linn. Soc. 133 (3): 711, fig. 2 (phylogenetic tree); phylog.

Megastes Westwood, 1851 (Boisduval MS), in Doubleday. Gen. diurn. Lep. 2, p. 346; unavail. (ICZN, Art. 11.6 as synonym of Dynastor ).— Scudder, 1875. Proc. Amer. Ac. Arts & Sc. 10 (2): 213.

Megastes Boisduval, 1870 . Consid. Lép. env. Guatemala, p. 53; Brazil, Colombia and Guatemala; included species: Megastes macrosiris and Megastes darius ; preocc. Guenée, 1854 ( Lepidoptera ).— Scudder, 1875. Proc. Amer. Ac. Arts & Sc. 10 (2): 213.— Hemming, 1943. Proc. r. ent. Soc. London (B) 12 (2): 26; type species: Papilio darius Fabricius, 1775 .— Hemming, 1967. Bull. Brit. Mus. (Nat. Hist.), Ent., Suppl. 9: 281.

Type species of Dynastor : Dynastor napoleon Doubleday, [1849] , by monotypy ( Scudder 1875; Hemming 1967).

Type species of Megastes Boisduval, 1870 : Papilio darius Fabricius, 1775 ( Hemming 1943).

Taxonomic history. Doubleday ([1849]) named Dynastor in an illustration of Dynastor napoleon . Westwood (1851), continuing the study, included Papilio darius Fabricius, 1775 in Dynastor and described Brassolis macrosiris from French Guiana. Bates (1864) described Brassolis strix from Guatemala, and Butler (1872) described Dynastor stygianus from Costa Rica. Kirby (1879) retained Papilio darius in Dynastor but transferred Brassolis macrosiris to Opsiphanes Doubleday, [1849] . Godman & Salvin (1881) maintained Dynastor napoleon and Dynastor darius and transferred Brassolis macrosiris and Brassolis strix to Dynastor , based on the rounded wings and the gradually clubbed antennae; Röber (1889) and Stichel (1902) followed this decision. Stichel (1904) was the first to consider species already described for the genus at the subspecific level, with Dynastor darius darius and Dynastor darius stygianus Butler, 1872 and Dynastor macrosiris macrosiris and Dynastor macrosiris strix . Dynastor was revised by Stichel (1909) and Fruhstorfer (1912).

Megastes Westwood, 1851 is unavailable as it was published as a synonym of Dynastor (ICZN 1999, Art. 11.6).

Megastes Boisduval, 1870 , for which the type species is Papilio darius , designated by Hemming (1943), was considered by the same author (1967) as a valid name, not being invalidated under the Law of Homonymy, because Megastes Westwood, 1851 , was unavailable ( Hemming 1967). However, Boisduval’s name is invalid as a junior homonym of Megastes Guenée, 1854 ( Lepidoptera ) ( Scudder [1884]). Strangely, this fact was ignored by Hemming (1967).

Diagnosis. Dynastor species are unique among the Brassolini and are differentiated by their rounded wings, disruptive wing pattern, especially on the hind wings, and by the large size of the body. The larvae are unique in Brassolini in feeding on bromeliads ( Bromeliaceae ).

Redescription. Head. Eyes naked, except in Dynastor hannibal stat. rest. and Dynastor pharnaces stat. nov. which has small setae; labial palpus brown with two lines of cream scales, one lateral and the other ventral, less evident in Dynastor napoleon , with the distal joint about one-third the length of the basal joint.

Thorax. Brown dorsally and ventrally, except Dynastor napoleon , which has dorsally orange prothorax and brown meso and metathorax, ventrally all thoracic segments orange; legs brown except the prothoracic legs of Dynastor napoleon , which are orange.

Forewing. Length: male 44–72 mm (n=24), female 55–87 mm (n=24). Costal margin convex; outer margin straight, slightly crenulated, or convex; inner margin straight. DFW between R 3 and M 1 with orange ( Dynastor napoleon ), whitish or yellowish (other species) apical spots, species may or may not show a whitish or yellowish oblique band; ground color dark-brown. VFW basal and discal regions mostly with a disruptive pattern; postdiscal region with a conspicuous or not oblique band and colorful spots or patches concentrated mainly in the apical region. Hindwing. Costal and outer margins rounded; the outer margin may be smooth or slightly crenulated; inner margin is less rounded than the costal and outer margins. DHW with a marginal band and dark-brown ground color, Dynastor darius and Dynastor stygianus stat. rest. have also a postdiscal band. VHW with a conspicuous ripple pattern in Dynastor macrosiris , Dynastor pharnaces stat. nov., Dynastor hannibal stat. rest., and Dynastor strix stat. rest.; in Dynastor napoleon resembles a dead leaf; in Dynastor stygianus stat. rest. and Dynastor darius the VFH is similar to Dynastor napoleon , but without CuA well marked; rounded, elongated, or other shaped spots in Sc+R 1 –Rs, M 1 –M 2, and CuA 1 –CuA 2.

Abdomen. Brown, except Dynastor napoleon (orange), Dynastor darius , and Dynastor stygianus Butler, 1872 stat. rest. (ochre) ventrally.

Male genitalia. Tegumen sub-quadrate in lateral view; dorsal arms of saccus slightly longer than ventral arms of tegumen; 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 in lateral view forming varied angles, distally bifurcated, the bifurcated regions may be proportional in length, or the ventral may be longer than the dorsal; valvae wider at the base, tapering distally, elongated or triangular from medial to distal region, dorso-distal region asymmetrical; aedeagus nearly straight, narrower distally, with or without spines along the median region, with an anterodorsal or dorsal opening to the ejaculatory bulb, and dorsal or subapical opening for the vesica; triangular fultura inferior, variable in width.

Female genitalia. Tergum VIII rectangular; anterior apophysis atrophied; lamella antevaginalis membranous, with sparse sclerotization in the wrinkled median region; lamella postvaginalis sclerotized, with the median portion of different shapes; ostium membranous; ductus bursae membranous, basal region of ductus seminalis sclerotized; corpus bursae membranous, rounded or elongated, on the ventral or lateral side with two parallel signa, or without signa; 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 a series of specimens showed that males and females of all species vary in the size, shape, and color of the apical spots between R 3 and M 1 of the DFW.

Comments. Molecular analyses combining one mitochondrial gene and four nuclear genes recovered Dynastor as monophyletic (SH-aLRT 100 / UFBoot 96) ( Fig. 1 ), 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 and adult morphology, we treat Dynastor as containing seven species and eight synonyms (two of them generic synonyms).

Etymology of Dynastor : From Greek, the name means a ruler or sovereign ( Glaser 1887).

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 2. Dynastor napoleon Doubleday, [1849]: Male, a. dorsal, b. ventral views; Brazil, Santa Catarina (DZ 49.739– DZUP). The black arrow indicates the diagnostic character of the species.

FIGURE 3. Dynastor napoleon Doubleday, [1849]: Female, a. dorsal, b. ventral views; Brazil, Rio de Janeiro (UFFLMNH MGCL 1107864–MGCL).

FIGURE 4. Dynastor napoleon Doubleday, [1849]: Male genitalia: a. dorsal view, b. ventral view, c. lateral view; aedeagus: d. dorsal view, e. ventral view, f. lateral view (DZ 49.740–DZUP).

FIGURE 5. Dynastor napoleon Doubleday, [1849]: Female genitalia: a. lateral view; b. detail of the corpus bursae without signa; c. ventral view (DZ 49.351–DZUP).

FIGURE 6. Geographical distribution of Dynastor napoleon Doubleday, [1849]. Asterisk (*) inside the sampled locality indicates that the geographical distribution data was retrieved from the bibliography.

FIGURE 7. Dynastor napoleon Doubleday, [1849]. Lectotype of Dynastor napoleon Doubleday, [1849]; a. dorsal view, b. ventral view.

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.

FIGURE 23. Dynastor stygianus Butler, 1872 stat. rest. [Holotype] of Dynastor stygianus Butler, 1872 stat. rest.; a. dorsal view, b. ventral view.

FIGURE 24. Dynastor macrosiris (Westwood, 1851): Male, a. dorsal, b. ventral views; Brazil, Pará (DZ 54.260–DZUP). The black arrows indicate the diagnostic characters of the species.

FIGURE 25. Dynastor macrosiris (Westwood, 1851): Female, a. dorsal, b. ventral views; Peru, Río Huallaga (AMNH_IZC 00352908–AMNH).

FIGURE 26. Dynastor macrosiris (Westwood, 1851): Male genitalia: a. dorsal view, b. ventral view, c. lateral view; aedeagus: d. dorsal view, e. ventral view, f. lateral view (DZ 54.260–DZUP). The black arrows indicate the diagnostic characters of the species.

FIGURE 27. Dynastor macrosiris (Westwood, 1851): Female genitalia: a. lateral view; b. detail of signa; c. ventral view (OM 78.930–DZUP–OM). The black arrow indicates one of the diagnostic characters of the species.

FIGURE 28. Geographical distribution of Dynastor macrosiris (Westwood, 1851), Dynastor pharnaces Stichel, 1908 stat. nov., Dynastor hannibal Oberthür, 1881 stat. rest., and Dynastor strix (Bates, 1864) 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 29. Dynastor macrosiris (Westwood, 1851). Female holotype of Dynastor macrosiris (Westwood, 1851); a. dorsal view, b. ventral view.

FIGURE 30. Dynastor pharnaces Stichel, 1908 stat. nov.: Female, a. dorsal, b. ventral views; Peru, Chanchamayo (USNMENT01589343–USNM). The black arrows indicate the diagnostic characters of the species.

FIGURE 31. Dynastor pharnaces Stichel, 1908 stat. nov.: female genitalia: a. lateral view; b. detail of signa; c. ventral view (USNMENT01589343–USNM). The black arrow indicates one of the diagnostic characters of the species.

FIGURE 32. Dynastor pharnaces Stichel, 1908 stat. nov. Male lectotype of Dynastor pharnaces stat. nov.; a. dorsal view, b. ventral view.

FIGURE 33. Dynastor hannibal Oberthür, 1881 stat. rest.: Male, a. dorsal, b. ventral views; Colombia, Boyacá (DZ 52.734– DZUP; CMBUT1655-20). The black arrows indicate the diagnostic characters of the species.

FIGURE 34. Dynastor hannibal Oberthür, 1881 stat. rest.: Female, a. dorsal, b. ventral views; Colombia, Boyacá (DZ 3969– DZUP).

FIGURE 35. Dynastor hannibal Oberthür, 1881 stat. rest.: Male genitalia: a. dorsal view, b. ventral view, c. lateral view; aedeagus: d. dorsal view, e. ventral view, f. lateral view (DZ 52.734–DZUP; CMBUT1655-20). The black arrow indicates one of the diagnostic characters of the species.

FIGURE 36. Dynastor hannibal Oberthür, 1881 stat. rest.: Female genitalia: a. lateral view; b. detail of signa; c. ventral view (DZ 3969–DZUP). The black arrow indicates one of the diagnostic characters of the species.

FIGURE 37. Dynastor hannibal Oberthür, 1881 stat. rest. Lectotype of Dynastor hannibal Oberthür, 1881 stat. rest.; a. dorsal view, b. ventral view.

FIGURE 38. Dynastor strix (Bates, 1864) stat. rest.: Male, a. dorsal, b. ventral views; El Salvador, Ahuachapán (FLMNH MGCL 1107875–MGCL). The black arrows indicate the diagnostic characters of the species.

FIGURE 39. Dynastor strix (Bates, 1864) stat. rest.: Female, a. dorsal, b. ventral views; El Salvador, Ahuachapán (FLMNH MGCL 1107876–MGCL).

FIGURE 40. Dynastor strix (Bates, 1864) stat. rest.: Male genitalia: a. dorsal view, b. ventral view, c. lateral view; aedeagus: d. dorsal view, e. ventral view, f. lateral view (DZ 52.735–DZUP; CMBUT1654-20). The black arrow indicates one of the diagnostic characters of the species.

FIGURE 41. Dynastor strix (Bates, 1864) stat. rest.: Female genitalia: a. lateral view; b. detail of signa; c. ventral view (OM 41.943–DZUP–OM). The black arrows indicate the diagnostic characters of the species.

FIGURE 42. Dynastor strix (Bates, 1864) stat. rest. Lectotype of Dynastor strix (Bates, 1864) stat. rest.; a. dorsal view, b. ventral view.