Revision of the poorly known Neotropical butterfly genus Zischkaia Forster, 1964 (Lepidoptera, Nymphalidae, Satyrinae), with descriptions of nine new species Nakahara, Shinichi Zacca, Thamara Dias, Fernando M. S. Dolibaina, Diego R. Xiao, Lei Espeland, Marianne Casagrande, Mirna M. Mielke, Olaf H. H. Lamas, Gerardo Huertas, Blanca Kleckner, Kaylin Willmott, Keith R. European Journal of Taxonomy 2019 2019-09-13 551 1 67 Dolibaina, Dias & Zacca Dolibaina, Dias & Zacca 2019 [456,1029,1802,1829] Insecta Nymphalidae Zischkaia Animalia Lepidoptera 50 51 Arthropoda species mielkeorum sp. nov.   Figs 1, 3I–L, 5B, 7E–F, 14    Diagnosis See relevant sections of  Z. saundersii,  Z. josti sp. nov., and  Z. warreni sp. nov.    Etymology This species is named to honor three generations of German-Brazilian naturalists: Carl Helmuth Theodor Mielke, Olaf Hermann Hendrik Mielke and Carlos Guilherme Costa Mielke. The specific epithet is a masculine noun in the genitive case.    Type material examined   Holotype   BRAZIL • ♂; “// Brasil, Espírito Santo, [ Sooretama,] Res[erva]. Ecológica Sooretama, 19°03′25”S 40°08′50”W,  19-26-II-2013, Mielke & Casagrande leg. // DZ 36.970 // BC-DZ Willmott 248 //”; DZUP.   Paratypes( 3 ♂♂, 6 ♀♀)   BRAZIL:  Espírito Santo• 1 ♂; Conceição da Barra; [ 18°34′ S, 39°45′ W];  6 Oct. 1969; C.-C. Eliasleg.; DZ-21200; DZUP•  1 ♀; Linhares; [ 19°23′ S, 40°4′ W];  Sep. 1981; Eliasleg.; DZ-36629; DZUP•  3 ♀♀; Sooretama, Reserva Ecológica de Sooretama; [ 19°3′25″ S, 40°8′50″ W];  100 m;  19–26 Feb. 2013; O.H.H. Mielkeand M.M. Casagrandeleg.; DZ-36618, DZ-36688, DZ-36968; DZUP•  1 ♀; Santa Teresa, [ 19°56′ S, 40°36′ W];  25–29 Mar. 1970; Ebertleg.; DZ-36711; DZUP. –   Minas Gerais• 1 ♂;  12 kmS of Teofilo Otoni;  600 m;  11 Feb. 2007; O.H.H. Mielkeand M.M. Casagrandeleg.; DZ-36599; DZUP. –   Rio de Janeiro• 1 ♀; Nova Iguaçu, Fazenda São Bernardino;  19 Sep. 1937; R.F. d’Almeidaleg.; DZ-36671; DZUP. –  Notlocated•  1 ♂; ‘ Brazil’; BMNH(E)-1205420; NHMUK.    Description  Male FOREWING LENGTH. 24–25 mm(n = 3). HEAD. Eyes naked, with whitish scales at base; labial palpi laterally dark reddish brown with whitish scales, ventrally dark brown and whitish lines, first segment about the length of the third segment and as thick as the second, third segment thin, second segment thicker and almost four times the length of the third segment, with long hair-like scales; antennae reddish dark brown dorsally with whitish scales at the base of each segment, ventrally naked, approximately two-fifths the forewing length, with about 46 segments (n = 5), distal segments forming a slender club. THORAX. Dorsally dark reddish brown; laterally and ventrally dark reddish brown with brownish gray scales. LEGS. Foreleg brownish gray with whitish scales; mid and hindleg femur brownish gray with whitish scales, tibiae and tarsi mostly brownish gray with whitish scales ventrally, with pair of tibial spurs present at distal end of tibia. ABDOMEN. Eighth tergite as stripe at base of eighth abdominal segment, in addition to presence of two distal broader patches. WING VENATION. Basal half of forewing Subcosta swollen; base of Cubitus swollen; forewing recurrent vein absent; origin of M 2towards M 1than M 3; hindwing humeral developed. WING SHAPE. Forewing triangular, costal margin rounded, outer margin straight, inner margin more or less straight, apex and tornus rounded; hindwing more or less rounded, humeral area expanded, costal margin slightly rounded, outer margin slightly crenulated, inner margin slightly sinuous, apex and tornus rounded, not very conspicuous. DORSAL FOREWING. Ground color variable, usually dark reddish brown; marginal area and apex conspicuously darker brown. DORSAL HINDWING. Ground color similar to forewing; submarginal and marginal line somewhat visible though transparency along the costal margin, specially near the tornus. VENTRAL FOREWING. Ground color brown, lighter than the dorsal forewing ground color; discal line reddish brown, noticeably fainter than other lines, running more or less obliquely from near the costal margin to 2A; postdiscal line dark reddish brown, variably developed in width, running obliquely from about M 1to 2A; submarginal line dark reddish brown, variably wavy, from the apex to the tornus along the outer margin, ending at about 2A; marginal line dark reddish brown, straight and narrower than submarginal line, from the apex to the tornus along the outer margin, ending at about 2A; fringe dark brown. VENTRAL HINDWING. Ground color similar to forewing; discal line dark reddish brown, almost straight, running from costal to the inner margin; postdiscal line similar to and more or less parallel to discal line, passing through the end of the discal cell at the origin of M 3, and posteriorly directed to the base of the wing at 2A-3A; submarginal band wavy, similar in color and width to the discal and postdiscal line, running from apex towards the tornus, occasionally fused with the posdiscal line near the costal margin and occasionally fused to the submedian line in 2A-3A; marginal line similar in color, but thinner than other lines, slightly wavy along the outer margin; submarginal ocelli from M 1to 2A, roughly oval, dark brown encircled by a marked orange ring and a subtler brown ring; pupil with scattered silver scales, ocelli at M 2-M 3, M 3-Cu 1, and Cu 1-Cu 2similar in size, somewhat elongate, those in M 1-M 2and Cu 2-2A underdeveloped, usually half the size of the other ocelli; scales of the areas between postdiscal and submarginal lines and submarginal and marginal lines with variable light purple to lilac tinge; fringe dark brownish. GENITALIA ( Fig. 5B). Tegumen somewhat flattened, anteriorly bulged, with a single dorsal dome-like projection near the base of the uncus; appendices angulares absent; uncus laterally flattened and straight, slightly longer than the length of the tegument in lateral view; brachia slightly shorter than the uncus, thicker at the base and narrowing to a point posteriorly; combination of ventral arms of the tegumen and dorsal arms of the saccus sinuous; saccus anterior projection tube-like, approximately the length of the tegumen; valva more or less triangular, stretched dorsal and posteriorly; basal half of valva larger, narrowing towards the apex, dorsal margin almost straight, ventral margin convex, apex broadly rounded; fultura inferior (i.e., juxta) strap-like; phallobase about one-third of phallus, rather straight; ductus ejaculatorius not discerned; aedeagus straight with manica covering approximately half, winglet absent, distal opening located ventrally where vesica is visible.  Female FOREWING LENGTH. 26–27 mm(n = 6). Similar to male except as follows: foretarsus divided into five tarsomers; wings ground color generally lighter; forewing more rounded and broader, outer margin usually slightly convex; hindwing outer margin more crenulated; dorsal hindwing submarginal and marginal lines more noticeable seen through transparency. FEMALE ABDOMEN AND GENITALIA ( Fig. 7E–F): membranous areas between seventh and eight sternite folded forming the sclerotized antrum attached to the lamella antevaginalis near the ostium bursae; lamella antevaginalis sclerotized; lamella postvaginalis absent; eighth abdomen segment with a lateral sclerotized area; ductus bursae with a sclerotized area of about a third of its length near the ostium bursae; ductus seminalis origin close to the ostium bursae; corpus bursae oval, with a pair of parallel and dorsal signa, signa narrow and slightly shorter than the length of the corpus bursae.   Variation The silvery blue patch between the median and submarginal lines in VHW can vary in intensity, and the coloration of the ocellar ring is also variable, being more yellowish in some specimens.  Distribution( Fig. 14) This species is known to date from southeastern Brazil, from the states of Espírito Santo, Rio de Janeiroand Minas Gerais.    Remarks Three barcoded specimens of  Z. mielkeorum sp. nov.(BC-DZ-225, 248, 249) form a well supported clade (SH-aLRT/UF Boot = 99.8/95), with two individuals of  Z. josti sp. nov.(MUSM-LEP-68763, 18704) ( Fig. 1). Despite the small sampling size (n = 5) for these two species, the existence of a ‘barcoding gap’ between these two taxa (see corresponding section of  Z. josti sp. nov.), combined with the presence of wing pattern and genitalia differences that separate these two species, leads us to consider them as distinct species. The known ranges of these species are approximately 3000 kmapart, reinforcing this taxonomic decision. In fact, no known area of sympatry exists between any species pair in the “  saundersiiclade”, which thus appears to have undergone allopatric speciation. However, this taxonomic treatment will result in accepting  Z. mielkeorum sp. nov.as a paraphyletic group. Although it is not an acceptable taxonomic practice to name non-monophyletic groups, we argue in this case based on the fact that BC-DZ-249 is missing first 80 or so base pairs of its sequence. This missing information might have resulted in separation from the sequence generated from the individual from the same locality (BC-DZ-248, from Res. Ecologica Sooretama, Espírito Santo, Brazil) and, in fact, grouped together with BC-DZ-225, which is a specimen from a different site ( Minas Gerais, Brazil). It is also worth noting that a number of butterfly species are recovered as paraphyletic, including species in some of the best studied butterfly genus  HeliconiusKluk, 1780(e.g.,  H. erato(Linnaeus, 1758)as in Kozak et al.2015). Clearly, more data based on more individuals is needed in order to draw a firm conclusion as to taxonomic status of these two populations, and our treatment herein is a mere provisional hypothesis based on available information. 2425040156 2013-02-19 2013-02-26 2013-02-19 DZUP Casagrande Brazil Brasil -19.056944 Res 21 -40.147224 Sooretama 51 52 1 1 Espirito Santo holotype 2425040111 1969-10-06 DZUP C. - C. Elias Brazil -18.566668 Conceicao da Barra 1275 -39.75 51 52 1 1 Espirito Santo 2425040164 [486,1381,723,752] 1981-09 DZUP Elias Brazil -19.383333 Linhares 1272 -40.066666 51 52 1 1 Espirito Santo 2425040001 2013-02-19 2013-02-26 2013-02-19 DZUP O. H. H. Mielke & M. M. Casagrande Brazil 100 -19.056944 Reserva Ecologica de Sooretama 21 -40.147224 Sooretama 51 52 3 3 Espirito Santo 2425040097 1970-03-25 1970-03-29 1970-03-25 DZUP Ebert Brazil -19.933332 Santa Teresa 1270 -40.6 51 52 1 1 Espirito Santo 2425040061 2007-02-11 DZUP O. H. H. Mielke & M. M. Casagrande Brazil 600 12 km S of Teofilo Otoni 51 52 1 1 Minas Gerais 2425040127 1937-09-19 DZUP R. F. d'Almeida & Not Brazil Fazenda Sao Bernardino Nova Iguacu 51 52 1 1 Rio de Janeiro 2425040144 [467,1001,951,978] NHMUK Brazil Brazil' 51 52 1 1 Rio de Janeiro