Revision of the poorly known Neotropical butterfly genus Zischkaia Forster, 1964 (Lepidoptera, Nymphalidae, Satyrinae), with descriptions of nine new species Author Nakahara, Shinichi A7240997-442C-41C7-9B20-D8D4AE31AED9 McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA. & Department of Entomology and Nematology, University of Florida, Gainesville, Florida 32611, USA. & Departamento de Entomología, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru. snakahara@ufl.edu Author Zacca, Thamara EBE99273-B02D-4A17-AD52-5BB3DFBA6C85 Departamento de Biologia Animal and Museu de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas, Campinas, São Paulo, Brazil. zacca.butterfly@gmail.com Author Dias, Fernando M. S. FA0950D9-DC58-4230-8180-F26CB5788E55 Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil. fernandomsdias@yahoo.com.br Author Dolibaina, Diego R. 0A762FB6-AF41-4A09-9DF8-6F3954BA8956 Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil. dirodrido@gmail.com Author Xiao, Lei 4233F498-4D30-428A-B2A4-A6A76DC5FCA4 McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA. lxiao@floridamuseum.ufl.edu Author Espeland, Marianne 00D6F9F9-3902-4A8B-846F-720AB32922A6 Arthropoda Department, Zoological Research Museum Alexander Koenig, Adenauer Allee 160, 53113 Bonn, Germany. marianne.espeland@gmail.com Author Casagrande, Mirna M. Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil. mibras@ufpr.br Author Mielke, Olaf H. H. Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil. omhesp@ufpr.br Author Lamas, Gerardo Departamento de Entomología, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru. glamasm@unmsm.edu.pe Author Huertas, Blanca Life Sciences Department, Natural History Museum, London, UK. B.Huertas@nhm.ac.uk Author Kleckner, Kaylin McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA. & Department of Entomology and Nematology, University of Florida, Gainesville, Florida 32611, USA. kaylin.kleckner@ufl.edu Author Willmott, Keith R. McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA. kwillmott@flmnh.ufl.edu text European Journal of Taxonomy 2019 2019-09-13 551 1 67 journal article 10.5852/ejt.2019.551 98f587bb-3baf-4961-89a0-5679c0dcf731 2118-9773 3476676 C3C851C3-0F12-412C-A15B-56F0F263CD00 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 . Fig. 1. Likelihood tree with the best log-likelihood score (-LnL = -2536.1613) given our data based on DNA ‘barcodes’, depicting Zischkaia as a well-supported clade. Support values are represented by SHaLRT/UF Bootstrap. Abbreviations: HT = Holotype; NT = Neotype. 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. Fig. 2. Zischkaia “ pacarus clade”, wing plate. A–B . Z. amalda ( Weymer, 1911 ) , lectotype, ♂, dorsal and ventral views. C–D . Z. arctoa Nakahara , sp. nov. , paratype, ♂ (in RFC, collected in 1975), dorsal and ventral views. E–F . Z. chullachaki Nakahara & Zacca , sp. nov. , holotype, ♂, dorsal and ventral views. G–H . Z. chullachaki sp. nov. , paratype, ♀ (DZ 36. 508), dorsal and ventral views. I–J . Z. baku Zacca, Dolibaina & Dias , sp. nov. , holotype, ♂, dorsal and ventral views. K–L . Z. baku sp. nov. , paratype, ♀ (DZ 36. 404), dorsal and ventral views. M–N . Z. arenisca Nakahara, Willmott & Hall , sp. nov. , holotype, ♂, dorsal and ventral views. O–P . Z. arenisca sp. nov. , paratype, ♀ (FLMNH-MGCL 297320), dorsal and ventral views. Q–R . Z. argyrosflecha Nakahara, L. Miller & Huertas , sp. nov. , holotype, ♂, dorsal and ventral views. S–T . Z. pacarus (Godart, [1824]), neotype, ♂, dorsal and ventral views. U–V . Z. pacarus , ♀ (DZ 36. 909), dorsal and ventral views. W–X . Z. abanico Nakahara & Petit , sp. nov. , holotype, ♂, dorsal and ventral views. Scale bar: 1 cm. 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) . Fig. 3. Zischkaia “ saundersii clade”, wing plate. A–B . Z. saundersii ( Butler, 1867 ) , ♂ (DZ 37. 029), dorsal and ventral views. C–D . Z. saundersii , ♀ (DZ 36. 969), dorsal and ventral views. E–F . Z. josti Nakahara & Kleckner , sp. nov. , holotype, ♀, dorsal and ventral views. G–H . Z. josti sp. nov. , paratype, ♀ (USNM ENT 00233844), dorsal and ventral views. I–J . Z. mielkeorum Dolibaina, Dias & Zacca , sp. nov. , holotype, ♂, dorsal and ventral views. K–L . Z. mielkeorum sp. nov. , paratype, ♀ (DZ 36. 618), dorsal and ventral views. M–N . Z. warreni Dias, Zacca & Dolibaina , sp. nov. , holotype, ♂, dorsal and ventral views. O–P . Z. warreni sp. nov. , paratype, ♀ (DZ 36. 508), dorsal and ventral views. Scale bar: 1 cm. Fig. 4. Zischkaia “ pacarus clade”, male genitalia. A . Z. amalda ( Weymer, 1911 ) , lectotype, ♂, lateral view, with phallus separated (winglet on aedeagus shown inside rectangle) (Genitalic vial: SN-17- 21). B . Z. arctoa Nakahara , sp. nov. , ♂, lateral view, with phallus separated (Genitalic vial: SN-17-9). C . Z. chullachaki Nakahara & Zacca , sp. nov. , ♂, lateral view, with phallus separated (Genitalic vial: DZ 21. 235). D . Z. baku Zacca, Dolibaina & Dias , sp. nov. , ♂, lateral view, with phallus separated (Genitalic vial: DZ 5. 573). E . Z. arenisca Nakahara, Willmott & Hall , sp. nov. , ♂, lateral view, with phallus separated (Genitalic vial: SN-17-16). F . Z. argyrosflecha Nakahara, L. Miller & Huertas , sp. nov. ♂, lateral view, with phallus separated (two winglets on aedeagus shown inside rectangle) (Genitalic vial: M-9142 ♂, Lee D. Miller). G . Z. pacarus (Godart, [1824]), ♂, lateral view, with phallus separated (Genitalic vial: DZ 5. 578). H . Z. abanico Nakahara & Petit , sp. nov. , ♂, lateral view, with phallus separated (Genitalic vial: 575/19.07.2017, J. Lorenc-Brudecka, MZUJ). Fig. 5. Zischkaia “ saundersii clade”, male genitalia. A . Z. saundersii ( Butler, 1867 ) , ♂, lateral view, with phallus separated (Genitalic vial: DZ 21. 221). B . Z. mielkeorum Dolibaina, Dias & Zacca , sp. nov. , ♂, lateral view, with phallus separated (Genitalic vial: DZ 21. 200). C . Z. warreni Dias, Zacca & Dolibaina , sp. nov. , ♂, lateral view, with phallus separated (Genitalic vial DZ 21. 214). Fig. 6. Zischkaia “ pacarus clade”, female genitalia. A–B . Z. chullachaki Nakahara & Zacca , sp. nov. , ♀, lateral and ventral views (Genitalic vial DZ 36. 508). C–D . Z. baku Zacca, Dolibaina & Dias , sp. nov. , ♀, lateral and ventral views (Genitalic vial: DZ 5. 571). E–F . Z. arenisca Nakahara, Willmott & Hall , sp. nov. , ♀, lateral and ventral views (Genitalic vial: SN-17-174). G–H . Z. pacarus (Godart, [1824]), ♀, lateral and ventral views (Genitalic vial: DZ 36. 731). Fig. 7. Zischkaia “ saundersii clade”, female genitalia. A–B . Z. saundersii ( Butler, 1867 ) , ♀, lateral and ventral views (Genitalic vial: DZ 36. 969). C–D . Z. josti Nakahara & Kleckner , sp. nov. , ♀, lateral and ventral views (Genitalic vial: 2018-002, D.J. Harvey). E–F . Z. mielkeorum Dolibaina, Dias & Zacca , sp. nov. , ♀, lateral and ventral views (Genitalic vial: DZ 36. 688). G–H . Z. warreni Dias, Zacca & Dolibaina , sp. nov. , ♀, lateral and ventral views (Genitalic vial: DZ 36. 731). 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. ). Fig. 8. Fultura inferior (i.e., juxta). A . Zischkaia pacarus (Godart, [1824]) (DZ 5. 578), in ventral view. B . Z. saundersii ( Butler, 1867 ) (DZ 5. 580), in posterior view. 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.