Vareuptychia themis (Butler, 1867) Zacca & Casagrande & Mielke & Huertas & Espeland & Freitas & Willmott & Nakahara & Lamas, 2020

Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi & Lamas, Gerardo, 2020, Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae), Zootaxa 4858 (1), pp. 1-34 : 11-13

Vareuptychia themis (Butler, 1867) comb. nov.

( Figs 5–13 , 19–23 , 26–27 , 29 )

Euptychia themis Butler, 1867a: 463 [nom. nud].

Euptychia themis Butler, 1867b: 104 , pl. 12, fig. 13; syntypes: no locality. Lectotype male here designated: no locality; NHMUK (examined).— Butler, 1868: 16.— Kirby, 1871: 48.— Butler, 1877: 118.— Kirby, 1879: 132.—Godman, 1901: 653.— Kaye, 1921: 57.— Barcant, 1970: 159, 161, pl. 13, fig. 11.— Santin, 2004: 1056.

Euptychia similis ; Butler, 1867b: 109, pl. 12, fig. 10 [misidentification].

Euptychia undina Butler, 1870: 252 ; holotype: Nicaragua; NHMUK (examined).— Kirby, 1871: 643.— Butler, 1877: 118.

Euptychia cleophes Godman & Salvin, 1889: 352 (partim) [misidentification].

Cissia themis ; DeVries, 1987: pl. 48, figs. 20–21.—Singer & Ehrlich, 1993: 248.— Lamas, 2004: 218.— Nakahara et al., 2012: 359–362, figs. 1, 2A–2B, 3–4 (male, female, genitalia, distributional map).— Luis-Martínez et al., 2004: 349.—Hernández-Mejía et al., 2008: 125.— Luis-Martínez et al., 2011: 23.— Miller et al., 2012: 58.— Luis-Martínez et al., 2016: 216.— Cock, 2014: 11.— Marín et al., 2017: 774, fig. 4C.—Henao-Bañol & Gantiva, 2020: 187.

Vareuptychia themis ; Llorente-Bousquets et al., 1996: 45.

Vareuptychia undina ; Llorente-Bousquets et al., 1996: 45.

Cissia undina ; Luis-Martínez et al., 2004: 349.— Luis-Martínez et al., 2011: 23.— Luis-Martínez et al., 2016: 216.

Yphthimoides maepius [misidentification]; Montero-Abril et al., 2009: 165.

‘Cissia’ similis ; Zacca et al., 2018b: 351 [misidentification].

Diagnosis. Vareuptychia themis can be distinguished from its congener by the bipupilled ocellus in CuA 1 -CuA 2 and the shape of the lamella antevaginalis ( Fig. 27 ).

Male genitalia ( Figs. 19–23 ). See above under ‘Redescription’ section.

Female genitalia ( Figs. 26–27 ). In addition to the characters mentioned in the generic description, the lateral plate does not reach the 8 th tergite. Lamella antevaginalis is large, wrinkled and half-orbicular in ventral view. The paired signa are dorsally located.

Variation. The VW ground color varies significantly from pale to dark brown, as also observed by Nakahara et al. (2012), and it is not associated with seasonal polyphenism or geographical distribution. Some individuals have orangey yellow patches on the umbra similar to those found in V. similis . Some individuals have an additional ocellus without a pupil in 2A and the inner margin of the VHW ( Fig. 8 ). Examination of the male genitalia of ten individuals from different localities (i.e. Mexico, El Salvador, and Venezuela) showed some variation in the size, shape and width of the tegumen, uncus, valva and aedeagus, in contrast to the observations of Nakahara et al. (2012). Conversely, female genitalia structures were found to be more conservative (mainly the shape of the lamella antevaginalis) but the presence/absence of the posterior apophysis and the size and width of the corpus bursae were variable among some individuals.

Ecology and distribution. This species is distributed widely from Mexico to Colombia and Venezuela, and in Trinidad and Tobago ( Barcant 1970; Cock 2014; DeVries 1987; Llorente-Bousquets 1996; Luis-Martínez et al. 2004; Hernández-Mejía et al. 2008; Luis-Martinez et al. 2011; Nakahara et al. 2012; Singer & Ehrlich 1991; plus examined material), at altitudes up to 2000 m ( Fig. 29 ). Vareuptychia themis comb. nov. is sympatric and syntopic with V. similis in several localities in Mexico, such as Jalisco, Malinalco, Oaxaca, Sonora, Veracruz and Yucatán ( Llorente-Bousquets 1996; Luis-Martinez et al. 2004; Hernández-Mejía et al. 2008; Luis-Martínez et al. 2011; E. Pfeiler pers. comm.; plus examined material), Guatemala (Baja Verapaz), Honduras (San Pedro Sula) and Venezuela (Aragua). Vareuptychia themis comb. nov. is multivoltine, flying all year. It has been recorded in deciduous and semideciduous forests along forest edges, and occasionally in open areas. In Costa Rica, this species is most abundant during the rainy season (Jun–Aug), and becomes less abundant as the dry season progresses ( DeVries 1987). Similar to V. similis comb. rest., V. themis comb. nov. has been recorded feeding on decaying fruits, such as graviola ( Annonaceae ), sapote ( Sapotaceae ) and mango ( Anacardiaceae ) ( Hernández-Mejía et al. 2008). Recorded larval host plants include undetermined species of Poaceae ( DeVries 1987; Santin 2004).

Type material, lectotype designation and taxonomic history. Butler (1867a) first introduced the name Euptychia themis as a nomen nudum, since no description or illustration was provided by the author, which fails to conform to Article 13 of the International Code of Zoological Nomeclature (ICZN 1999). The name became available when Butler (1867b) published an illustration of the species (pl. 12, fig. 13) in the supplementary material to his “Monograph of the genus Euptychia ”, although the locality of the specimen illustrated was not provided. One male specimen that agrees with the illustration of E. themis ( Butler, 1867b, pl. 12, fig. 13) was found at NHMUK. This specimen has no locality label, although ‘Mexico’ is handwritten (but has been erased) on one label ( Fig. 10 ), which also states that the specimen came from Hewitson’s collection, agreeing with the information provided by Butler (1867a). To fix the identity of the name, this male specimen is herein designated as the lectotype of E. themis ; it has the following labels: /Type ♂ / Hewitson Coll. 79–69. Euptychia themis . Butl[er]. 1./ Euptychia themis Butler / B. M. Type No. Rh. 3201 Euptychia themis , ♂ Butl[er]./ BMNH(E) 982889/; and two others will be added later: / Lectotype / Lectotype Euptychia themis Butler, 1867 . T. Zacca det. 2020/. NHMUK.

Butler (1870) named Euptychia undina based on a single specimen (holotype) from Nicaragua ( Fig. 9 ), being part of the type series of E. similis (see discussion under V. similis ) and bearing Butler’s distinctive blue label for types. Examination of the male holotype of E. undina at the NHMUK confirmed that this specimen is, in fact, E. themis since it has two silvery pupils in ocellus CuA 1 -CuA 2 on the VHW.

Examined material. 339 males and 125 females (21 specimens dissected)—see Supporting Information (S2).

Figures

FIGURE 29. Map showing the distribution of Vareuptychia themis comb. nov.

FIGURES 1–8. Species of Vareuptychia stat. rest., dorsal and ventral: 1–4. Vareuptychia similis comb. rest.: 1–2. male (Guerrero, Mexico), 3–4. female (San Jerónimo, Guatemala); 5–8. Vareuptychia themis comb. nov.: 5–6. male (San Mateo, Costa Rica), 7–8. female (Ilopango, El Salvador). Scale bar = 1 cm.

FIGURES 9–11. Lectotypes and original illustration of species of Vareuptychia stat. rest.: 9. Lectotype of Euptychia similis Butler, 1867 and labels; 10. Lectotype of Euptychia themis Butler, 1870 and labels; 11. Original illustration of Euptychia themis.

FIGURES 12–13. Wing venation of Vareuptychia themis comb. nov.: 12. male; 13. female. Black arrows indicate generic diagnostic features. Scale bar = 2 mm.

FIGURES 14–23. Male genitalia of species of Vareuptychia stat. rest.: 14–18. Vareuptychia similis comb. rest. (BMNH#1420973): 14. lateral, 15. dorsal; 16. ventral; 17. aedeagus, dorsal; 18. aedeagus, lateral; 19–23. Vareuptychia themis comb. nov. (BMNH#1420750): 19. lateral; 20. dorsal; 21. ventral; 22. aedeagus, dorsal; 23. aedeagus, lateral. Scale bar = 1 mm.

FIGURES 24–27. Female genitalia of species of Vareuptychia stat. rest.: 24–25. Vareuptychia similis comb. rest. (DZUP 27.891): 24. lateral, 25. ventral; 26–27. Vareuptychia themis comb. nov. (DZUP 27.897): 26. lateral; 27. ventral. Scale bar = 1 mm.