Zischkaia Forster 1964
Creators: name: Nakahara, Shinichi
name: Zacca, Thamara
name: Dias, Fernando M. S.
name: Dolibaina, Diego R.
name: Xiao, Lei
name: Espeland, Marianne
name: Casagrande, Mirna M.
name: Mielke, Olaf H. H.
name: Lamas, Gerardo
name: Huertas, Blanca
name: Kleckner, Kaylin
name: Willmott, Keith R.
Publisher: name: Zenodo
Year: 2019
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Lepidoptera
Nymphalidae
Zischkaia

description: Genus Zischkaia Forster, 1964
Zischkaia Forster, 1964: 116.
Type species
Euptychia fumata Butler, 1867: 109, pl. 12, fig. 14 (by original designation).
Systematics
Zischkaia is a member of a clade consisting of several species of Splendeuptychia Forster, 1964 (e.g., S. itonis (Hewitson, 1862), S. clementia (Butler, 1877)), Amphidecta Butler, 1867 and Rareuptychia Forster, 1964, based on the aforementioned large dataset including>2000 individuals representing> 420 species (unpubl. data). Although Zischkaia was not included in the analysis of Espeland et al. (2019), the clade containing these related species (the “ Amphidecta clade”) was well supported based on hybrid enrichment data incorporating 368 loci (Espeland et al. 2019). The “ Amphidecta clade” is sister to a large clade including some major euptychiine clades, such as the “ Pareuptychia clade”, “ Taygetis clade” and “ Splendeuptychia clade” (all sensu Peña et al. 2010), but the support for this relationship is low. Our molecular data based on the COI barcoding region strongly support Zischkaia itself as monophyletic (Fig. 1; SH-aLRT/UFBoot = 98.7/100), with the removal by Freitas et al. (2018) of Euptychia mima Butler, 1867 to Nhambikuara, a genus in the “ Pareuptychia clade”. Zischkaia can be divided into two moderate to well-supported clades based on our molecular data (Fig. 1; SH-aLRT/ UFBoot = 95.8/97; 91.3/93), which are also diagnosable by several morphological characters discussed below under the ‘Diagnosis’ section. Despite the morphological differences between the two clades, which are also highly supported based on molecular data, the overall wing patterns and habitats appear to be rather similar among species in both of these clades. Considering that generic names should ideally be informative for the recognition of monophyletic groups of similar appearing species, in addition to having potential value in predicting unknown biological traits, we here treat Zischkaia as representing both clades rather than treating them as distinct genera. The morphological diversity within Zischkaia, such as the presence or absence of wing androconial scales, is also seen within other euptychiine genera of similar species diversity, such as Taygetina Forster, 1964.
Diagnosis
Species of Zischkaia can be distinguished from all other genera of Euptychiina by the combination of the following characters: 1) absence of ocelli on the DFW, DHW and VFW (also true of some other euptychiines); 2) five or six ocelli on the VHW from Sc+R 1 or M 1 to 2A with the white pupils (often diffuse scales rather than a single spot) displaced distally from the center (Splendeuptychia doxes (Godart, [1824]) and relatives possess a superficially similar pupil, but it is displaced basally instead of distally); 3) VFW with submedian line restricted to discal cell or absent (also true of a number of other euptychiine genera); 4) inter-segmental membrane between seventh and eighth sternites not pleated, but folded posteriorly of ostium bursae with its sclerotized region forming a ‘scoop-like’ structure below lamella antevaginalis (projection more apparent in species of the “ pacarus clade”). This form of intersegmental membrane is rather unique among euptychiines, which often have this membrane pleated and expandable, although some species in the genus Euptychia Hübner, 1818 are somewhat similar in this respect.
Two clades are recognized in Zischkaia, which we call the “ pacarus clade” and the “ saundersii clade”; the former can be distinguished from the latter by: 1) presence of androconial scales on DFW of males (absent in “ saundersii clade”) (Figs 2–3); 2) presence of a ‘tusk’-like projection (Fig. 4A) from the posterior region of the tegumen above the uncus (the projection of the tegumen appears as a ‘bulb’ in the “ saundersii clade” (Fig. 5A)); 3) uncus that is narrow and long, somewhat curving down, terminating in a small ‘bulb’ (rather than being straight and broad as in the “ saundersii clade”); 4) brachia curved dorsally (rather than straight as in the “ saundersii clade”); 5) valva being rather short in lateral view, with the apical point not extending beyond that of the uncus (the apical point extends beyond that of the uncus in the “ saundersii clade”); 6) tip of the anterior projection of the saccus extending further than the tegumen in lateral view (the tip of the anterior projection of the saccus does not extend beyond the tegumen in lateral view in the “ saundersii clade”); 7) fultura inferior (i.e., juxta) appearing as a thin strip in posterior view (Fig. 8A) (it appears as a well-developed plate in posterior view in the “ saundersii clade” (Fig. 8B)); 8) median region of the ductus bursae with a well-developed sclerotized half-ring (Fig. 6A) (absent in the “ saundersii clade” (Fig. 7A), or very reduced as in Z. saundersii and Z. josti sp. nov.).
The diagnostic characters provided above for the “ pacarus clade” can be used to separate all species in this clade from species in the “ saundersii clade”, and vice-versa. Therefore, species diagnoses focus on comparing similar species within the respective clade. Note that females of Z. amalda (Weymer, 1911), Z. argyrosflecha Nakahara, L. Miller & Huertas, sp. nov., Z. abanico Nakahara & Petit, sp. nov. and Z. arctoa Nakahara, sp. nov. are still unknown or unrecognized, thus the diagnostic characters provided for females of related species might not be applicable to these four species, and discovery of female specimens of these taxa would be extremely valuable. Similarly, the male is unknown or unrecognized for Z. josti Nakahara & Kleckner, sp. nov.
History of classification
Butler (1867 a) was the first to propose a systematic classification for Zischkaia. In his monograph of Euptychia (then used as a catch-all genus to include most euptychiine species), Butler divided the genus into seven groups (Division I to VII), and Division VI was the group relevant to Zischkaia. This division included E. saundersii Butler, 1867, E. mima Butler, 1867, E. fumata nomen nudum, E. pacarus (Godart, [1824]) and E. insignis Butler, 1867. The following diagnosis was given for Butler’s Division VI: “Upper surfaces brown, without marks; ventral forewings generally unmarked; [ventral] hindwings frequently with oval-shaped black ocelli, pupilled with silvery spots”. Subsequently, Butler (1877) proposed the “ E. pacarus group” and included E. saundersii, E. mima, E. fumata Butler, 1867, E. pacarus, E. insignis, E. peculiaris Butler, 1874 and E. erichtho Butler, 1867. The last taxon was described and placed in Division VII in Butler’s (1867 a) classification, but the name is currently regarded as a junior subjective synonym of Neonympha antonina C. Felder & R. Felder, 1867 (now placed in Erichthodes Forster, 1964) (Lamas 2004). Weymer (1911) proposed a similar group, named the “ pacarus group”, in which he included E. saundersii, E. mima, E. fumata, E. pacarus, E. peculiaris, E. insignis, E. erichtho and his E. amalda. Forster (1964) erected the genus Zischkaia by designating Euptychia fumata as the type species and stated: “In this newly erected genus I include some very similar in appearance, large, dorsally unicolorous brown species, which regarding the anatomical structure of the male Genitalia (figs 123– 125) correspond well with one another and are differentiated from all other ‘ Euptychia ’ species by the long, slender subunci and unique, unpaired outgrowths of the tegumen, which are located dorsally over the uncus”. Based on these diagnostic characters, in addition to the type species, Forster (1964) recognized Z. amalda and Z. saundersii in the genus, and figured the male genitalia for all three species. Lamas (2004) included four species, namely Z. amalda, Z. pacarus, Z. saundersii, and Z. mima, in addition to recognizing one undescribed species (described here as Z. arenisca sp. nov.). The checklist of Lamas (2004) also treated Euptychia fumata (described by Butler 1867 b: 109) as a junior subjective synomym of E. pacarus, and E. fumata (cited by Butler 1867a: 501) as a nomen nudum. Euptychia mima Butler, 1867, a taxon formerly placed in Zischkaia, was recently made the type species of Nhambikuara by Freitas et al. (2018).
Distribution and natural history
All confirmed records for Zischkaia are exclusively east of the Andes (but see also discussion under Z. abanico sp. nov.), where species are found from sea level up to about 1600 m. The known distributions of a number of species are highly restricted, with typically only a single species occurring in a particular locality, although label data suggest local sympatry between Z. warreni Dias, Zacca & Dolibaina, sp. nov. and Z. pacarus in southeastern Brazil, and Zischkaia chullachaki Nakahara & Zacca, sp. nov. and Z. saundersii in southeastern Peru. Perhaps notably, these two cases concern species in the two different clades, with no known cases of sympatry in members of the same clade. There are a few published observations on the behavior of Zischkaia, which are typically found in forest in close proximity to patches of bamboo (Poaceae: Bambusoideae), their likely hostplant; Brown (1992) reported the larvae of Z. pacarus using “bamboo”, as did Freitas in Beccaloni et al. (2008) (see also natural history notes under Z. arenisca sp. nov.).
Species accounts
Zischkaia Forster, 1964
amalda (Weymer, 1911)
arctoa Nakahara, sp. nov.
chullachaki Nakahara & Zacca, sp. nov.
baku Zacca, Dolibaina & Dias, sp. nov.
arenisca Nakahara, Willmott & Hall, sp. nov.
argyrosflecha Nakahara, L. Miller & Huertas, sp. nov.
pacarus (Godart, [1824])
= fumata (Butler, 1867a), nom. nud.
= fumata (Butler, 1867b)
abanico Nakahara & Petit, sp. nov.
saundersii (Butler, 1867)
josti Nakahara & Kleckner, sp. nov.
mielkeorum Dolibaina, Dias & Zacca, sp. nov.
warreni Dias, Zacca & Dolibaina, sp. nov.
Key to species of Zischkaia
DFW and DHW androconial scales present in males; projection from posterior region of tegumen above uncus ‘tusk’-like; uncus narrow and long, somewhat curving down, terminating in a small ‘bulb’ in lateral view; brachia curved dorsally; valva rather short in lateral view, apical point does not extend beyond that of uncus; tip of anterior projection of saccus extends further than tegumen in lateral view; fultura inferior (i.e., juxta) appears as a thin strip in posterior view; developed sclerotized halfring in median region of ductus bursae present........................................................ Z. pacarus clade
DFW and DHW androconial scales absent in males; projection from posterior region of tegumen above uncus ‘bulb’-like; uncus appearing straight in lateral view; valva apical point extends beyond that of uncus; tip of anterior projection of saccus does not extend beyond tegumen in lateral view; fultura inferior (i.e., juxta) appears as a developed plate in posterior view; developed sclerotized half-ring in median region of ductus bursae absent or reduced........................................... Z. saundersii clade
“ pacarus clade”:
1. VHW ocelli elongated (ocelli in VHW cells M 2 -M 3, M 3 -Cu 1, and Cu 1 -Cu 2 occupying more than half space between submedian line and submarginal line)....................................................................... 2
– VHW ocelli not elongated, rather circular (ocelli in VHW cells M 2 -M 3, M 3 -Cu 1 and Cu 1 -Cu 2 occupying less than half space between submedian line and submarginal line) ............................... 3
2. Prominent ocellus absent in VHW Rs-M 1; lack of silverish-purple scales on the basal side of VHW ocelli (distad of submedian line); one winglet on aedeagus; aedeagus winglet obtuseangled; ventral margin of apical process of valva not concave; saccus longer than ventral margin of valva ........................................................................ Z. arenisca Nakahara, Willmott & Hall, sp. nov.
– Prominent ocellus present in VHW Rs-M 1; two winglets on aedeagus; aedeagus winglet an acuteangled triangle; ventral margin of apical process of valva concave; saccus shorter than ventral margin of valva ..................................................... Z. argyrosflecha Nakahara, L. Miller & Huertas, sp. nov.
3. Apical process of valva somewhat rectangular with a convex distal margin .................................... 7
– Apical process of valva somewhat subtriangular ............................................................................. 4
4. Adults small (forewing length 21–23 mm); male dorsal androconial scales rather indistinct; from Andean foothills................................................................................................................................. 5
– Adults large (forewing length around 25 mm); male dorsal androconial scales distinct; from Amazon Basin .................................................................................................................................................. 6
5. Apical process of valva somewhat elongate; ventral margin of apical process of valva convex; from Cordillera de la Costa (Venezuela) ......................................................... Z. arctoa Nakahara, sp. nov.
– Apical process of valva not elongate; ventral margin of apical process of valva almost straight or concave; from Andean foothills of Peru and Bolivia ................................ Z. amalda (Weymer, 1911)
6. Posterior projection of tegumen rather straight; winglet of aedeagus reduced, almost absent; lamella antevaginalis developed as a tube around ductus bursae .. Z. baku Zacca, Dolibaina & Dias, sp. nov.
– Posterior projection of tegumen curved; winglet of aedeagus prominent, clearly visible; lamella antevaginalis not developed as a tube around ductus bursae .............................................................................. Z. chullachaki Nakahara & Zacca, sp. nov.
7. Southern and southwestern Brazil to northeastern Argentina ................. Z. pacarus (Godart, [1824]) – Tropical Andes of Colombia and Ecuador ............................... Z. abanico Nakahara & Petit, sp. nov.
“ saundersii clade”:
1. VHW marked with prominent blue/grayish shading between submedian line and marginal line; signa long, extending almost entire length of corpus bursae .................. Z. saundersii (Butler, 1867)
– VHW not marked with prominent blue/grayish shading between submedian line and marginal line; signa not extending across the entire corpus bursae.......................................................................... 2
2. DHW submarginal and marginal line clearly visible; ocellus in VHW M 1 -M 2 often less than half size of that in VHW M 2 -M 3; median line and submarginal line fused immediately after 2A in males ............................................................................ Z. warreni Dias, Zacca & Dolibaina, sp. nov.
– DHW submarginal and marginal line almost invisible; ocellus in VHW M 1 -M 2 often about half size (or larger) of that in VHW M 2 -M 3; median line and submarginal line not fused immediately after 2A in males ............................................................................................................................................. 3
3. VHW ocelli in M 1 -M 2 and/or Cu 2 -2A appearing as incomplete ocelli with black central area and/or pupil indistinct; small sclerotized region absent at one-third distance from ostium bursae to corpus bursae; from southeastern Brazil .......................... Z. mielkeorum Dolibaina, Dias & Zacca, sp. nov.
– VHW ocelli in M 1 -M 2 and/or CuP-2A appearing as complete ocellus with black central area and/or pupil rather distinct and clearly visible; small sclerotized region present at one-third distance from ostium bursae to corpus bursae; from Guianas....................... Z. josti Nakahara & Kleckner, sp. nov.
description: Published as part of 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., 2019, Revision of the poorly known Neotropical butterfly genus Zischkaia Forster, 1964 (Lepidoptera, Nymphalidae, Satyrinae), with descriptions of nine new species, pp. 1-67 in European Journal of Taxonomy 551 on pages 11-20, DOI: 10.5852/ejt.2019.551, http://zenodo.org/record/3476676