Megisto cleophes ( Godman & Salvin, 1889 ), 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 : 23-26

Megisto cleophes ( Godman & Salvin, 1889) comb. nov.

( Figs 74–86 )

Euptychia cleophes Godman & Salvin, 1889: 352 ; syntypes: [ Mexico, Guerrero], Tierra Colorada and Dos Caminos. Lectotype here designated: [ Mexico, Guerrero], Dos Caminos; NHMUK (examined); Godman, 1901 vin: 653, pl. 107, figs. 3–4 (female).—Weymer, 1911: 201.— Gaede, 1931: 443.— D’Abrera, 1988: 761 (male, female).

Cissia cleophes ; Llorente-Bousquets et al., 1996: 45.— Luis-Martínez et al., 1996: 119.— Luis-Martínez et al., 2003: 221.— Lamas, 2004: 218.— Michán et al., 2005: 130.—Llorente-Bousquets et al., 2006: 973.— Luis-Martínez et al., 2011: 23.

Diagnosis. Megisto cleophes comb. nov. resembles M. cymela (Cramer, 1777) but can be easily distinguished from that species by the absence of an ocellus in CuA 1 -CuA 2 on both the DFW and VFW, and by having crenulated submarginal and marginal lines on the VW (straight in M. cymela ). The wing pattern of M. cleophes comb. nov. also resembles that of Cissia rubricata (Edwards, 1871) , but differs by the absence of the orange spot in the median region of the DFW and VFW, and by having the median and submarginal lines running parallel from the costal to the inner margin (the lines converge between 2A and the inner margin in C. rubricata ).

Male genitalia ( Figs. 79–83 ). Tegumen dorsally convex, laterally subtriangular. Uncus robust at posterior region, tapering at apex, dorsally ovoid with apex truncated. Gnathos curved upwards, about 2/3 length of uncus, slightly sinuous, larger at base and tapering at apex. Combination of ventral arm of tegumen and dorsal arm of saccus sinuous. Appendices angulares developed, wider at base with apex curved downwards. Anterior projection of saccus short, laterally lanceolated, and smaller than gnathos. Fultura superior absent. Fultura inferior as a broad sclerotized plate in U-shape. Valva elongated, covered by long hair-like setae latero-ventrally, and short setae at inner side; costae well-developed; dorsal margin sinuous with a dorsal projection at median region, ventral margin concave at median region; apex pointed, dorsally with wide spiny projection. Aedeagus straight, cylindrical, almost same length as valva; anterior region bottle-shape; posterior region about twice longer than anterior region with bifid apex; distal opening ventral and longer than proximal opening. Vesica without cornuti.

Female genitalia ( Figs. 84–85 ). Eighth tergite rectangular. Papillae anales somewhat oblong covered by long hair-like setae at distal region; posterior apophysis absent. Lamella antevaginalis subtriangular in ventral view. Ductus bursae membranous; corpus bursae membranous, longer than ductus bursae, with paired signa ventrally located.

Variation. Females have a rounded FW (triangular in males), and the submedian and median lines are broader than in males. Males have a dark androconial patch in the median region of the DFW and no pupils in the ocelli on both the VFW and VHW ( Fig. 75 ).

Ecology and distribution. This species is endemic to Mexico in the states of Guerrero, Morelos, Oaxaca, Puebla and Veracruz, at altitudes up to 1400 m ( Luis-Martínez et al. 1996, 2003, 2011; Michán et al. 2005; plus examined material) ( Fig. 86 ). Megisto cleophes comb. n. flies from March to October. The immature stages and host plants are unknown, although the related M. cymela has been documented feeding on Xyris L. ( Poales , Xyridaceae ) ( Tietz 1972; Ferris & Brown 1981; Ackery 1988).

Type material, lectotype designation and taxonomic history. Euptychia cleophes Godman & Salvin, 1889 was described based on at least one male and one female from Tierra Colorada and Dos Caminos, Mexico. Two female syntypes from Tierra Colorada and Dos Caminos were found at NHMUK, and one of them is herein designated as the lectotype of E. cleophes to fix the identity of the species ( Fig. 74 ); this specimen has the following labels: /Type / Type of species/ E[uptychia]. cleophes type ♀ / B. M. Type No Rh 3144 Euptychia cleophes , ♀ G[odman]. & S[alvin]./ ♀ / Dos Caminos, Guerrero, 2700 ft. [822 m], Sept[ember]., H. H. Smith./ Godman-Salvin Coll. 1904.–1. B[iologia]. C[entrali]. A[mericana]. Lep[idoptera]. Rhop[alocera]. Euptychia cleophes , G[odman]. & S[alvin]./ Sp. figured/ BMNH(E) 1267115/; and two other labels will be added later: / Lectotypus / Lectotypus Euptychia cleophes Godman & Salvin, 1889 . T. Zacca det. 2020/. NHMUK. The female syntype from Tierra Colorada will be labeled as a paralectotype. Another female specimen found at the NHMUK, from Acahuizotla, also from Smith’s collection, is not part of the type series because it was not mentioned in the original description and is, in fact, Vareuptychia themis comb. nov. As clarified by Godman (1901: 653), what Godman & Salvin (1889) supposed was the male of E. cleophes also turned out to be V. themis comb. nov.

Based on the wing pattern, Godman & Salvin (1889) and Godman (1901) suggested that this species was closely allied to ‘Cissia’ myncea and Vanima labe comb. nov. . Later, Euptychia cleophes was transferred to Cissia by Singer et al. (1983) based on the immature stages, and this classification was followed by subsequent authors. Recently, Zacca et al. (2018b) suggested that the species needed to be removed from Cissia , mainly based on morphological data (wing patterns, venation, male and female genitalia), since DNA sequences were not available for this species. In the present study, we place this species in Megisto based on morphological similarities with M. cymela (see illustrations in Miller 1976: 6, figs. 7–12), including the wing pattern, venation, male genitalia with elongated gnathos, short anterior projection of saccus and absence of cornuti and female genitalia with a developed lamella antevaginalis. Although M. cleophes comb. nov. shows some similarities in wing pattern with Cissia rubricata , these might be the result of convergent evolution; the presence of androconial scales on the DFW is also shared with M. cymela , and absent in all species of Cissia . Superficial similarity in wing pattern is also true of M. cleophes comb. nov. and Llorenteana pellonia (Godman, 1901) , but wing venation and morphology of the male genitalia are helpful to distinguish both taxa. Until very recently, L. pellonia was treated as ‘ incertae sedis’ ( Lamas 2004), but the species was recently transferred to the new genus Llorenteana Viloria & Luis-Martínez, 2019, in the subtribe Ypthimina ( Viloria & Luis-Martínez 2019) . Nevertheless, no morphological or molecular phylogenetic analysis has yet confirmed this tentative subtribal placement.

Systematic position and discussion. Megisto is sister to the Oriental Palaeonympha opalina ( Peña et al. 2006, 2010; Espeland et al. 2019), with the split of both genera from the remaining South American Euptychiina genera (except Euptychia ) estimated to have occurred in the early Miocene ( Peña et al. 2010). Unfortunately, we were unable to obtain DNA sequences for M. cleophes comb. nov. and our classification of this taxon is based only on comparison of morphological characters. Clearly, DNA sequence data for this species, as well as the enigmatic Llorenteana pellonia , would be valuable to test these taxonomic proposals and better understand the relationships of these restricted Mexican species.

Examined material. 3 males and 19 females (2 specimens dissected)—see Supporting Information (S2).

Figures

FIGURE 86. Map showing the distribution of Megisto cleophes comb. nov..

FIGURES 74–78. Megisto cleophes comb. nov.: 74. lectotype of Euptychia cleophes Godman & Salvin, 1889—dorsal, ventral and labels; 75–76. male (Guerrero, Mexico), dorsal and ventral; 77–78. female (Guerrero, Mexico). Scale bar = 1 cm.

FIGURES 79–83. Male genitalia of Megisto cleophes comb. nov. (BMNH#1420652): 79. lateral, 80. dorsal, 81. ventral, 82. aedeagus, dorsal, 83. aedeagus, lateral. Black arrows indicate generic diagnostic features. Scale bar = 1 mm.

FIGURES 84–85. Female genitalia of Megisto cleophes comb. nov. (BMNH#1420655): 84. lateral, 85. ventral. Black arrows indicate generic diagnostic features. Scale bar = 1 mm.