Vanima Zacca, Casagrande & Mielke, 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 : 13-15

Vanima Zacca, Casagrande & Mielke , gen. nov.

Type species. Euptychia labe Butler, 1870 , by present designation.

Etymology. The generic name ‘vanima’ means beautiful in the fictional elvish language Quenya from J.R.R. Tolkien’s Middle-Earth novels. In this case, we explicitly note that we use the name Vanima as a feminine noun in the nominative singular.

Diagnosis. Among the Euptychiina genera, Vanima gen. nov. can be distinguished by the following set of characters: DHW with a large bipupilled ocellus occupying the entire width of cell CuA 1 -CuA 2 (also present in some other euptychines, e.g., ‘ Cissia’ confusa (Staudinger, 1887)) , VHW having a reddish-brown quadrate patch at the tornus formed by the expansion of the marginal line ( Fig. 31 ) (also present in other euptychines, such as ‘Magneuptychia’ andrei Zacca, Casagrande & Mielke, 2017 and related species), and a tiny ocellus in 2A and inner margin in VHW ( Fig. 31 ) (absent in a few individuals of V. labe comb. nov., and present rarely in other euptychines, e.g. Euptychia sophiae Zacca, Nakahara, Dolibaina & Dias, 2015 , and E. attenboroughi Neild, Nakahara, Fratello & LeCrom, 2015 ). This combination of characters is unique within the larger ‘ Megisto ’-clade of which Vanima gen. nov. is a member ( Espeland et al. 2019).

Among Euptychiina , the genitalia of the species of Vanima gen. nov. resembles the otherwise distantly related Omacha pax ( Huertas, Lamas, Fagua & Willmott, 2016) ( Huertas et al. 2016; Andrade-C. et al. 2019) but can be distinguished by the ovoid uncus in dorsal view (elliptical in O. pax ), apex of valva pointed or rounded (double prongs in O. pax ), aedeagus bipartite at the distal region (truncated in O. pax ), papillae anales without posterior apophysis (present in O. pax ) and lateral plate of the 8 th segment with a spiracle dorsally located (spiracle absent in O. pax ), lamella antevaginalis absent (present in O. pax ). Compared to the phylogenetically closest genera, Vanima gen. nov. is easily distinguished from Carminda Ebert & Dias, 1998 by the ocelli on the DW (absent in Carminda ), elongated valva (rhomboid in Carminda ) and absence of lamella antevaginalis (present in Carminda ), and from Yphthimoides by the reddish-brown quadrate patch at the VHW tornus (absent in Yphthimoides ) and the absence of spine-like projections in the apex of the valva (present in Yphthimoides ).

Description. Antennae half length of costal margin of FW, with white scales around base of each flagellomere; chaetosema brown; postgena area with small white or cream scales; labial palp about two times length of head with mixed small white and black scales at distal segment and elongated scales at mid and basal segments. Legs covered with mixed small white, brown and black scales.

Wings. FW sub-triangular, costa slightly convex, apex rounded, external margin slightly convex, without indentations, and inner margin straight. DHW ground color brown with elements of VFW visible through translucence, large bipupilled ocellus between Radius and M 1 with black ocellar spot surrounded by a narrow yellow ocellar ring (except in V. lesbia comb. nov. in which this ocellus is only present on VFW). VFW with three broad straight lines (basal, submedian and median) of variable color (dark brown or rufous), two thin crenulated dark brown lines (submarginal and marginal), space between median and submarginal lines with a dark brown umbra extending from costa to 2A or costa to M 3 -CuA 1 (in latter, with an additional yellow spot between M 3 -CuA 2), two to four ocelli between R-M 1, M 1 -M 2, M 2 -M 3, M 3 -CuA 1; one at M 1 -M 2 is always present and larger with a developed black ocellar spot and two pupils while others are smaller with faded ocellar spot and 1–2 large pupils. HW sub-rectangular, costa convex, apex rounded, external margin convex with slight indentations, inner margin slightly convex and curved at tornus. DHW ground color brown with elements of VHW visible through slightly translucent wings, large ocellus in CuA 1 -CuA 2 with black ocellar spot surrounded by a narrow yellow ocellar ring and 1–2 tiny pupils. VHW similar to VFW except for five ocelli between Rs-CuA 2 and an additional tiny ocellus in 2A and inner margin; bipupilled ocelli in M 1 -M 2 and CuA 1 -CuA 2 are larger than others with developed black ocellar spot surrounded by yellow ocellar ring, ocelli between M 2 -CuA 1 with faded ocellar spot and two large pupils; marginal line enlarged at tornus forming a rufous sub-square.

Venation. FW with vein Sc strongly swollen at base, CuA swollen and 2A slightly swollen at base, dcs reduced, dcm concave, dci straight or slightly sinuous ( Fig. 45 ). HW with developed humeral vein ( Figs. 45–46 ).

Male genitalia. Tegumen sub-triangular in lateral view, in dorsal view with a median concavity at posterior region. Uncus about twice length of tegumen, robust at posterior region turning slender from median region to apex in lateral view and dorsally elliptical. Gnathos narrow and curved upwards, almost same length as uncus. Appendices angulares of variable size. Anterior projection of saccus developed and cylindrical. Fultura superior absent. Fultura inferior a thin stripe. Valva sub-triangular covered by long hair-like setae latero-ventrally, and short setae at inner side, costae developed, and apex pointed. Aedeagus straight, shorter than valva, cylindrical, anterior region bottleshaped, posterior region about two times longer than anterior region, with bifid apex in dorsal view, distal opening ventral and almost same length as proximal opening. Vesica without cornuti.

Female genitalia. Eighth tergite rectangular. Papilla anales somewhat triangular, sclerotized at posterior region and covered by long hair-like setae at distal region; posterior apophysis absent. Lateral plate (probably derived from 8 th sternite) can reach or not 8 th tergite, spiracle present. Lamellae antevaginalis and postvaginalis absent. Ductus bursae membranous and of variable size; corpus bursae membranous with paired signa dorsally or dorso-laterally.

Systematic position and discussion. Vanima gen. nov. comprises three species previously treated in Cissia by Lamas (2004): V. labe comb. nov., V. lesbia comb. nov. and V. palladia comb. nov. Previous molecular ( Murray & Prowell 2005; Peña et al. 2006, 2010; Wahlberg et al. 2009) and morphological ( Marín et al. 2017) phylogenies did not sample any species of Vanima gen. nov. Recently, Espeland et al. (2019) included V. labe comb. nov. and V. palladia comb. nov. in a backbone phylogeny of Euptychiina based on hybrid enrichment data. According to that study, V. labe comb. nov. and V. palladia comb. nov. are distantly related to Cissia but sister to a clade that comprises some species of Yphthimoides (e.g. Y. pacta (Weymer, 1911) , Y. borasta (Schaus, 1902)) and Carminda . The results of our present study also corroborate that Vanima gen. nov. is monophyletic (aLRT = 89.8/BS = 94) ( Fig. 87 ). With the inclusion of V. lesbia comb. nov. in the analysis we found that this species is sister to V. labe comb. nov. + V. palladia comb. nov. Some studies have recovered Yphthimoides as polyphyletic ( Marín et al. 2017; Espeland et al. 2019; Barbosa et al., in prep.) and an ongoing study will describe a new genus to contain these species (Barbosa et al., in prep.). Conversely, Carminda is monophyletic based on molecular data and the morphology of the wing pattern, venation, male and female genitalia ( Dias 2011; Aguiar et al., in prep.) are quite distinct from Vanima gen. nov. Although the male genitalia morphology of the species of Vanima gen. nov. shares some similarities with the monotypic Omacha, analysis of sequence data from> 300 species of Euptychiina shows that both genera are not closely related and Omacha appears as sister of the clade that includes Vareuptychia (Espeland et al., in prep.).

Figures

FIGURE 87. Phylogenetic tree showing the position of Vareuptychia stat. rest. (green) and Vanima gen. nov. (purple) in Euptychiina. Numbers above the branches indicate the support values (SH-aLRT support/ultrafast bootstrap support); numbers below 70 are not shown.

FIGURES 30–41. Species of Vanima gen. nov., dorsal and ventral: 30–33. Vanima labe comb. nov.: 30–31. male (Chiriquí, Panama), 32–33. female (Verapaz, Mexico); 34–37. Vanima lesbia comb. nov.: 34–35. male (Amazonas, Brazil), 36–37. female (Amazonas, Brazil); 38–41. Vanima palladia comb. nov.: 38–39. Male (Acre, Brazil), 40–41. female (Acre, Brazil). Scale bar = 1 cm.

FIGURES 45–46. Wing venation of Vanima palladia comb. nov.: 45. male, 46. female. Black arrows indicate generic diagnostic features. Scale bar = 2 mm.