Vanima palladia (Butler, 1867) comb. nov.

(Figs. 38–41, 44–46, 52–56, 66–67, 70, 73)

Euptychia palladia Butler, 1867a: 461, pl. 39, fig. 21; syntypes: [Brazil, Pará], Tapajós.— Butler, 1868: 16.— Kirby, 1871: 48.— Butler, 1877: 118.—Weymer, 1911: 200, pl. 47b.— Kaye, 1921: 57.— Riley & Gabriel, 1924: 42.— Aurivillius, 1929: 158.— Gaede, 1931: 459.— Bryk, 1953: 60.— Barcant, 1970 [misidentification]: 143, 161, pl. 13, fig. 9 (male).

Argyreuptychia palladia; Forster, 1964: 123.— Lewis, 1973: pl.58, fig. 17.

Cissia palladia; Lamas, 1981: 29.— Singer et al., 1983: 109, figs. 2E, 3E, 4E; lectotype female: [Brazil, Pará], Tapajós; NHMUK (examined).— DeVries, 1987: 274, pl. 48, fig. 28 (male).—Singer & Ehrlich, 1993: 250.— Lamas, 1994: 165.— Robbins et al., 1996: 231. — Ramos, 2000: 40.— Lamas, 2004: 218.— Beccaloni et al., 2008: 329.— Brévignon & Benmesbah, 2012: 40, pl. 4, fig. 5a (male genitalia).— Montero-Abril & Ortiz-Pérez, 2013: 58.— Cock, 2014: 11.

Euptychia pallidia [sic]; D’Abrera, 1988: 761 (male, female).

Diagnosis. Vanima palladia comb. nov. differs from its congeners by its smaller size (FW length: 16–18 mm), the VFW having two developed but faded ocelli in M 2 -M 3 and M 3 -CuA 1 and by having the rufous spot at the tornus smaller and more elongate rather than quadrate (well developed and quadrate in V. labe comb. nov. and V. lesbia comb. nov.) (Figs. 39, 41).

Male genitalia (Figs. 52–56). In addition to the characters mentioned in the generic description, the apex of valva is broad and strongly serrated.

Female genitalia (Figs. 66–67). In addition to the characters mentioned in the generic description, the lateral plate reaches the 8 th tergite, the ductus bursae is short and the signa are located dorso-laterally.

Variation. Females have a rounded FW (triangular in males), well developed subapical ocellus in M 1 -M 2 on the DFW (absent in males), and the VFW with the median and submarginal lines farther away from each other than they are in males. In both sexes, the ocelli in M 2 -M 3 and M 3 -CuA 1 on the VFW can be faded in some individuals, but always present.

Ecology and distribution. This species is distributed widely from Nicaragua to Colombia, Trinidad and Tobago, French Guiana (St-Laurent du Maroni), Brazil (Acre, Roraima, Rondônia, Amazonas, Maranhão, Pernambuco, Alagoas, Goiás and Mato Grosso), Peru (Cuzco, Loreto and Madre de Dios) and Bolivia (El Beni) (Singer et al. 1983; DeVries, 1987; Singer & Ehrlich 1993; Brévignon & Benmesbah, 2012; plus examined material) at altitudes up to 1400 m (Fig. 70). According to DeVries (1987), V. palladia comb. nov. is rare in Costa Rica, being known only from remnant forests near Atenas. The species flies throughout the year. Similar to V. labe comb. nov., the peak of abundance of V. palladia comb. nov. is during the dry season in Trinidad (Singer et al. 1983). The species occurs very locally in association with deciduous and premontane forests, along riparian edges (Fig. 73) (Singer et al. 1983; DeVries, 1987). Recorded larval host plants include undetermined species of Cyperus L. and Seleria (Cyperaceae), Ichnanthus pallens (Sw.), Lasiacis sloanei (Griseb.), Oplismenus hirtellus (L.), Panicum pilosum Sw., P. polygonatum Schrad., Paspalum conjugatum P. J. Bergius, Pasp. convexum Willd. ex Döll, Pasp. decumbens Sw., Setaria paniculifera (Steud.) E. Fourn. (= S. palmifolia (J. Koenig) Stapf and Tripsacum sp. ( Poaceae) (Singer et al. 1971; Singer et al. 1983; DeVries 1987; Ackery 1988; Singer & Erhlich 1993; Beccaloni et al. 2008). Nevertheless, most of these records, and perhaps all of them, come from larvae reared in captivity, and it is not clear whether the host plant used in nature is known. Descriptions of the egg and all four instars (coloration and head capsule shape) are given by Singer et al. (1983).

Type material, lectotype designation and taxonomic history. Euptychia palladia Butler, 1867 was described based on an unstated number of specimens collected by H. W. Bates in Tapajós, Pará, Brazil. The female lectotype of E. palladia (Fig. 44) was designated by Singer et al. (1983) and is deposited at the NHMUK .

The specimen cited and illustrated as E. palladia in Weymer (1911: 200, pl. 47b) corresponds to ‘ Cissia’ myncea (Cramer, 1780) (see discussion below), as also pointed out by Singer & Ehrlich (1993). Weymer described his specimen as having five ocelli on the VHW, which disagrees with the original description of E. palladia, which noted six ocelli on the VHW, including a reduced ocellus in 2A and the inner margin (Butler, 1867a: 462). Barcant (1970) also misidentified ‘ Cissia’ myncea as E. palladia . Brown et al. (2007) cited Cissia sp. nr. palladia from Quibdó, Chocó, Colombia, but it has not been possible to locate this specimen.

Forster (1964) transferred E. palladia to Argyreuptychia . Subsequently, this species was transferred to Cissia (Singer et al. 1983) and placed in the ‘ labe subgroup’ together with C. labe and C. penelope . This classification was followed by Lamas (2004), but recently Zacca et al. (2018b) suggested that the species would need to be removed from Cissia based on morphological (wing pattern, venation, male and female genitalia) and molecular data (nuclear and mitochondrial markers).

Examined material. 41 males and 56 females (7 specimens dissected)—see Supporting Information (S2).