Dynastor stygianus Butler, 1872

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 : 477-482

Dynastor stygianus Butler, 1872 stat. rest.

( Figs 12 , 19–23 )

Dynastor stygianus Butler, 1872 . Cist. Ent. 1 (4): 73; 1 female [holotype]; Coll. Druce; Costa Rica, Cartago.—Butler, 1873, Lepidoptera Exotica, p. 125, pl. 47, fig. 1 (ventral).—Kirby, 1877. Syn. Cat. Diurn. Lep., Suppl., p. 716.

Dynastor darius stygianus ; Stichel, 1904, in Wytsman. Gen. Ins. 20, p. 9, pl. 3, fig. 2 (male d, v).— Stichel, 1909. Das Tierreich 25: 24.— Fruhstorfer, 1912, in Seitz. Gross-Schmett. Erde 5, p. 288.— Stichel, 1932. Lep. Cat. 51, p. 20.— Young & Muyshondt, 1975. Stud. Neotrop. Fauna 10: 27; biol.— DeVries, 1986. Jour. Res. Lep. 24: 326; hostplant.— DeVries, 1987. Butt. Costa Rica, p. 247, 250, figs 31c (pupa), d (larva), 32 e3, pl. 41, fig. 5 (male d); biol., ecol.—D’Abrera, 1987. Butt. Neotrop. Reg. 3, p. 394 (fig. male d, v).—R. F. de la Maza R., 1987. Marip. Mexicanas, p. 96, pl. 23, fig. 7 (male d).—R. G. de la Maza E. et al., 1989. Rev. Soc. Mex. Lep. 12 (2): 61.—R. G. de la Maza E. & J. de la Maza E., 1993. Marip. Chiapas, p. 183.—Moreno et al., 1998. Marip. Ecuador, p. 155.—Maes, 1999. Cat. Ins. Art. Terr. Nicaragua 3, p. 1386.— 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.— Salinas-Gutiérrez et al., 2004. Jour. Lep. Soc. 58 (3): 136. — Santin, 2004. Répert. plantes-hôtes subst. chen. lépid. monde (2 nd ed.), 2, p. 858; hostplant.—J. Miller et al., 2006. 100 Caterpillars, p. 183, fig. 54 (pupa, female d); biol., hostplant.—Chacón & Montero, 2007. Marip. Costa Rica, pl. 169 (figs female d, male v), pl. 172 (fig. larva head); biol.—Glassberg, 2007. Swift Guide Butt. Mexico and C. America (1 st ed.), p. 132 (figs male d, v).—Vargas-Fernández et al., 2008. Nymph. Mexico 2, p. 61, 180, pl. 17, figs 6, 7 (male d, v).— Garzón-Orduña & Penz, 2009. Zootaxa 2134: 4. — J. Miller et al., 2012. Insecta Mundi 205: 57.—J. de la Maza E. & R. G. de la Maza E., 2015, in Carabias, J. de la Maza E. & Cadena (eds.). Conserv. Desarr. sust. Selva Lacandona, p. 639.—Morales-Morales et al., 2018. Ent. Mexicana 5: 565.

Dynastor darius [misidentification]; Aiello & Silberglied, 1978. Psyche 85 (4): 331, figs 1 (male d, v), 2 (eggs), 3–6 (larvae), 9 (larvae, head capsule), 11 (pupae); biol., hostplant.— Ricardo-Molina et al., 2019. Shilap Revta. lepid. 47 (185): 12, fig. 26 (larva); biol.—Henao-B. & Gantiva-Q., 2020. Bol. Cient. Mus. Hist. Nat. U. de Caldas 24 (1): 187.— Matos-Maraví et al. 2021. Biol. Jour. Linn. Soc. 133 (3): 711, fig. 2 (phylogenetic tree); phylog.—R. G. de la Maza E. & Soberón, 1998. Biodiv. Conserv 7 (7): 943; ecol. — Aiello, 2015, in Dyer & Forister (eds). The lives of Lepdopterists, p. 98.—Glassberg, 2018. Swift Guide Butt. Mexico and C. America (2 nd ed.), p. 152 (figs d, v).

Taxonomic history. Dynastor stygianus was treated as a subspecies of Dynastor darius by Stichel (1904) and the subsequent authors in reviews, catalogs, checklists, and natural history studies.

Type material. Dynastor stygianus Butler, 1872 was described based on one female [holotype] in Coll. Druce. The female holotype is deposited at the NHMUK, with the following labels: / Dynastor stygianus Butl. type./ Dynastor stygianus , Type. Butl./ Costa Rica. Van Patten. Druce Coll./ B.M. TYPE No. Rh. 6098 Dynastor stygianus , ♀ Butl./ Type H. T./ ♀ / B.C.A.Lep.Rhop. Dynastor darius, Fabr. Godman-Salvin Coll. 1919 .—6./.

Diagnosis. Dynastor stygianus stat. rest. differs from Dynastor darius , a morphologically similar and closely related species ( Fig. 1 ), in having the VFW apical patch generally extending to the outer margin, while Dynastor darius usually has this patch extending only to the apex ( 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 stygianus stat. rest. has a smaller corpus bursae without signa and a smaller lamella antevaginalis, the female of Dynastor darius has a larger corpus bursae with two parallel signa and a broader lamella antevaginalis ( Figs 11 , 22 ).

Variation. FW, length: male 44–53 mm (n=5), female 56–65 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. Almost since its description, Dynastor stygianus stat. rest. has been treated as a subspecies of Dynastor darius . However, the wing color pattern, the morphology of the female genitalia, and the molecular data do not support this status. Besides all variations in the wing color patterns, a character that is usually present in Dynastor stygianus stat. rest. is the VFW costal patch generally extending to the outer margin. Another important character is the corpus bursae of the female genitalia without signa. DNA barcode divergence (4.0–5.0%) between trans-Andean ( Costa Rica and Panama) and cis-Andean ( Brazil and Peru) specimens agree with the differences found in the morphology. Dynastor stygianus stat. rest. is sympatric with Dynastor strix stat. rest. and Dynastor hannibal stat. rest.

Natural history. Detailed information on the immature stages of Dynastor stygianus stat. rest. can be found in Aiello & Silberglied (1978). This species flies at dusk and dawn, and inhabits forests, second-growth, and urban areas where a suitable hostplant occurs; the larvae are seen at most times of the year ( DeVries 1987), remaining in the middle of the leaf when not feeding ( DeVries 1986). Larvae are parasitized by Winthemia sp. ( Tachinidae ) ( Miller et al. 2006).

Hostplants: Bromeliaceae : Bromelia pinguin L., Aechmea magdalenae (Andre)Andre ex Baker ( DeVries 1986), Aechmea bracteata (Sw.) Griseb. ( Janzen & Hallwachs 2009) , Ananas comosus (L.) Merril ( Aiello & Silberglied 1978; Janzen & Hallwachs 2009), Bromelia karatas L. ( Penz et al. 1999; Miller et al. 2006).

Geographic distribution. Mexico (Chiapas (Maza R. 1987; Maza E. & Maza E. 1993), Oaxaca, Guerrero, Quintana Roo); Guatemala (El Petén, Izabal); El Salvador (San Salvador); Nicaragua (Granada); Costa Rica (Alajuela, Puntarenas, Guanacaste); Panama (Colón, Chiriquí, Panamá); Colombia (Antioquia, Boyacá, Valle del Cauca); Ecuador (Pichincha) ( Fig. 12 ). Dynastor stygianus stat. rest. occurs at elevations between sea level and 1,100 m ( Young & Muyshondt 1975; DeVries 1987).

Temporal distribution. Throughout the year.

Etymology. Probably means a dweller of Styx, one of the rivers of the Greek mythology underworld.

Specimens examined. Photos of the holotype of Dynastor stygianus ( Fig. 23 ), and the following specimens: MEXICO: Oaxaca — Santa María Chimalapa , VIII-1951, 2 males, IX-1963, 1 male, VII-1956, 1 male, UFFLMNH MGCL 1107757–1107760 T. Escalante leg., (MGCL). Guerrero —Acahuizotla, III-1958, 1 female, T. Escalante leg., UFFLMNH MGCL 1107807 (MGCL). Quintana Roo —Nuevo Durango, 27-X-1978, 1 male, OM 41.895 (DZUP-OM); Nuevo X-Can, 18-XI-1979, 1 female, VIII-1983, 1 female Welling, leg., OM 41.481, OM 91.438 (DZUP-OM). GUATEMALA: El Petén —Sayaxché, 20-IX-1963, 1 male, E. C. Welling leg., AMNH_IZC 00352865 (AMNH), 18-VIII-1963, 1 female, OM 41.911 (DZUP-OM). Izabal —Cayuga, VIII, 1 male, Schaus & Barnes leg., USNMENT01589291 (USNM); Quiriguá, VI, 1 female, Schaus & Barnes leg., USNMENT01589292 (USNM). COSTA RICA: Puntarenas —La Guardia de Barranca, 200 m, 20-XI-1982, 1 male, R. Hesterberg leg., ex-coll. R. Hesterberg, UFFLMNH MGCL 1107762 (MGCL). Guanacaste —Área de Conservación Guanacaste (Sendero Bejuquilla), 280 m, 10-IX-2008, 1 male *, 1 female, L. Ríos leg., USNMENT01589293, 01589315 (USNM), (Sendero Mora), 480 m, 21-XI-2006, 2 males *, J. A. Sánchez leg., USNMENT01589294, 01589295 (USNM), (Tajo Ángeles), 540 m, 14-I-2006, 1 male, 1 female, O. Espinoza leg., USNMENT01589297, 01589302 (USNM), 29-I-2006, 1 female *, E. Araya leg., USNMENT01589314 (USNM), (Tangelo), 410 m, 22-IV-2009, 2 males, R. Moraga leg., USNMENT01589313, 01589304 (USNM), (Sendero Tigre), 280 m, 9-XI-2000, 1 female, 17-X-2001, 1 male, 20-XI-2000, 1 male, L. Ríos leg., USNMENT01589305, 01589298, 01589307 (USNM), (Área Administrativa), 295 m, 6-XII-1999, 2 females *, G. Pereira leg., USNMENT01589316, 01589310 (USNM), (Quebrada Costa Rica), 275 m, 29-XI-1999, 1 female, 1 male, M. Pereira leg., USNMENT01589311, 01589300 (USNM), (Cafetal), 280 m, 5-XI-2004, 1 male, R. Franco leg., USNMENT01589301 (USNM), (Quebrada Oro), 290 m, 2-X-2003, 1 male, L. Ríos leg., USNMENT01589299 (USNM), (Estación Maritza), 570 m, 4-III-1995, 1 female, R. Moraga leg., USNMENT01589306 (USNM), (Sendero Natural), 290 m, 29-XI-1999, 1 female, M. Pereira leg., USNMENT01589308 (USNM), (Quebrada García), 495 m, 3-IV-2007, 1 female, O. Espinoza leg., USNMENT01589312 (USNM), (Bosque Transición), 3-IV-2012, 1 female, G. Sihezar leg., USNMENT01589363 (USNM). Alajuela —El Coco, V-1998, 1 male, 1 female, L. Kassarov leg., UFFLMNH MGCL 1107761, 1107809 (MGCL); Área de Conservación Guanacaste (Puente Palma ), 460 m, 06-X-2004, 1 female, E. Araya leg., USNMENT01589296 (USNM), ( Vochysia ), 320 m, 5-IV-2004, 1 female, J. Pérez leg., USNMENT01589303 (USNM), (Sendero Pinyal), 630 m, 2-VII-1998, 1 male, O. Espinoza leg., USNMENT01589309 (USNM). PANAMA: Colón —no locality, 457,2 m, 1 female, USNMENT01589319 (USNM). Chiriquí —no locality, 1 male, USNMENT01589318 (USNM); Santa Cruz, VIII-1975, 1 male, G. B. Small leg., USNMENT01589317 (USNM). Panamá —Gamboa, 1 female, O. Seitz leg., USNMENT01589320 (USNM); Diablo, 1 female, USNMENT01589321 (USNM); Las Cumbres, 8-XII-1958, 1 male, 12-XII-1960, 1 female, UFFLMNH MGCL 1107765, 1107810 (MGCL). COLOMBIA: Antioquia —no locality, 1 female, UFFLMNH MGCL 1107811 (MGCL). Boyacá —Muzo, VI-1924, 1 male, USNMENT01589322 (USNM), 1 male, 1 female, ex-coll. F. Johnson, AMNH_IZC 00352848, 00352857 (AMNH). Valle del Cauca —no locality, 2 females, ex-coll. F. Johnson, AMNH_IZC 00352855, 00352856 (AMNH). ECUADOR: Pichincha —Toachi, 9-IX-1974, 1 male, Vénédictoff leg., ex-coll. Vénédictoff, UFFLMNH MGCL 1107771 (MGCL).

iNaturalist records: EL SALVADOR: San Salvador — Tonacatepeque , 13°43’28.2”N, 89°06’00.7”W, 17-XII-2017 , Ares-afc. NICARAGUA: Granada — Nandaime , 11°45’34.42”N, 86° 3’37.85”W, 22-VIII-2003 , Eric van den Berghe (available at: inaturalist.org/observations/8848362; inaturalist.org/observations/50661799. 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 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.