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 Zischkaia warreni Dias, Zacca & Dolibaina , sp. nov. urn:lsid:zoobank.org:act: F2CFEE62-2CC3-4FA5-BD05-D25F973A5691 Figs 1 , 3 M–P, 5 C, 7 G–H, 14 , 16 C Diagnosis Zischkaia warreni sp. nov. is distinguished from other species in the “ saundersii clade” by the absence of the VFW submedian line. Further distinctive characters include the rather prominent DHW submarginal and marginal line, reduced VHW ocellus in M 1 -M 2 combined with the ocellus in M 2 -M 3 being more than twice as large (in terms of diameter) and the median line and submarginal line being fused immediately after 2A in males. Additional characters are also discussed in the relevant sections of other species. Etymology This specific epithet is in honor of our good friend, Andrew D. Warren, a prominent lepidopterist with a broad knowledge of butterflies. The specific epithet is to be considered a latinized masculine noun in the genitive case. Type material examined Holotype BRAZIL • ♂ ; “// 2-XI-2000 , Represa Sanepar , Piraquara, PR [ Paraná , Brazil], Bizarro leg. // DZ 36.948 //BC-DZ Willmott 227 //”; DZUP . Paratypes ( 20 ♂♂ , 31 ♀♀ ) BRAZIL : Minas Gerais • 1 ♀ ; Itamonte ; [ 22°17′16″ S , 44°51′54″ W ]; Dec. 1957 ; Ebert leg.; DZ-36731 ; DZUP . – Paraná • 1 ♀ ; Balsa Nova , São Luís do Purunã ; [ 25°29′ S , 49°44′ W ]; 1000 m ; 21 Feb. 2001 ; G. Lamas leg.; MUSM-LEP-105660 ; MUSM • 3 ♀♀ ; Castro ; [ 24°47′ S , 50°1′ W ]; 950 m ; BMNH(E)-1718053 to BMNH(E)-1718054 , BMNH(E)-1718064 ; NHMUK • 4 ♀♀ , same collection data as for preceding; USNM • 1 ♂ , 1 ♀ ; same collection data as for preceding; E.D. Jones leg.; BMNH(E)-1718069 , BMNH(E)-1718061 ; NHMUK • 1 ♀ ; same collection data as for preceding but Sep. 1897 ; BMNH(E)-1718060 ; NHMUK • 1 ♀ ; Guarapuava , Serra da Esperança , N side BR 373, 4.3 rd km ENE of Guará ; [ 25°21’49.49”S 51°14’46.07”W ]; 1218 m ; 4 Feb. 2016 ; D. Dolibaina and A. Warren leg.; DD-380 • 2 ♂♂ , 3 ♀♀ ; same collection data as for preceding but 6 Feb. 2016 ; DD-375 to DD-379 • 7 ♂♂ , 3 ♀♀ , same collection data as for preceding but A. Warren leg.; ADW • 1 ♀ ; Piraquara , Mananciais da Serra ; [ 25°30′28″ S , 49°01′32″ W ]; 5 Nov. 1972 ; Mielke leg.; DZ-36739 ; DZUP • 1 ♂ ; Quatro Barras , Morro do Anhangava ; [ 25°23′18″ S , 49°0′14″ W ]; 1350 m ; 25 Mar. 2009 ; E. Carneiro leg.; DD-306 • 1 ♂ ; same collection data as for preceding but D. Dolibaina leg.; DD-305 • 1 ♀ ; Tijucas do Sul ; [ 25°55′ S , 49°12′ W ]; 850 m ; 13 Mar. 2004 ; G. Lamas leg.; MUSM-LEP-105659 ; MUSM • 1 ♀ ; Tijucas do Sul , Rincão ; [ 25°48′ S , 49°8′ W ]; 900 m ; 25 Feb. 1969 ; Mielke leg.; DZ-36719 ; DZUP • 1 ♂ , 1 ♀ ; Tijucas do Sul , Vossoroca ; 850 m ; 20 Mar. 1971 ; Mielke leg.; DZ-21214 , DZ-36669 ; DZUP • 2 ♂♂ , 1 ♀ ; same collection data as for preceding but 8 Mar. 1972 ; DZ-36659 , DZ-36679 , DZ-36709 ; DZUP • 1 ♂ ; União da Vitória ; [ 26°12′ S , 51°5′ W ]; 610 m ; Dec. 1919 – Jan. 1920 ; A. Hall leg.; BMNH(E)-10430703 ; NHMUK • 1 ♀ ; same collection data as for preceding; 750 m ; 7 Nov. 1985 ; O.H.H. Mielke and M.M. Casagrande leg.; DZ-36749 ; DZUP . – Rio de Janeiro • 1 ♂ ; Itatiaia , Serra Itatiaia , vertente SE ; [ 22°19′55″ S , 44°36′38″ W ]; 1300 m ; Feb. 1950 ; Ebert leg.; DZ-36639; DZUP . – Santa Catarina • 1 ♀ ; Campo Alegre , Serra do Quiriri , [ 26°1′34″ S , 48°59′2″ W ]; 1300 m ; 14 Nov. 2009 ; O.H.H. Mielke , E. Carneiro and Melo leg.; DZ-36538 ; DZUP • 1 ♀ ; same collection data as for preceding but 14 Nov. 2009 ; O. Mielke and E. Carneiro ; DD-304 • 1 ♂ , 1 ♀ ; Irani ; [ 27°01′20″ S , 51°54′15″ W ]; 1000 m ; 8 Nov. 1985 ; O.H.H. Mielke and M.M. Casagrande leg.; DZ-36689 , DZ-36699 ; DZUP • 1 ♀ ; Mafra ; [ 26°07′08″ S , 49°48′50″ W ]; 22–23 Feb. 1982 ; Mielke leg.; DZ-36729 ; DZUP • 1 ♂ ; São Bento do Sul ; [ 26°15′ S , 49°23′ W ]; 850 m ; 2 Feb. 1985 ; H. Miers leg.; FLMNH-MGCL-1036043 ; FLMNH • 1 ♂ ; Seara , Nova Teutonia ; [ 27°3′ S , 52°23′ W ]; 300–500 m ; Feb. 1973 ; F. Plaumann leg.; FLMNH-MGCL-1036044 ; FLMNH . – São Paulo • 2 ♀♀ ; Apiaí , Serra de Paranapiacaba ; [ 24°31′ S , 48°51′ W ]; Feb. 1972 ; Ebert leg.; DZ-36609 , DZ-36653 ; DZUP . – Not located • 1 ♀ ; ‘Brazil’; BMNH(E)-1205409 ; NHMUK . Description Male FOREWING LENGTH. 24–25 mm (n = 3). HEAD. Eyes naked, with whitish scales at base; labial palpi first segment with white and brownish long hair-like scales; second segment length almost twice as great as eye depth and covered with brown scales and hair-like scales laterally, with white scales and hair-like scales, dorsally with pale brownish long hair-like scales, ventrally with black and white hair-like scales, about 3–4 times as long as segment width; third segment about one-fourth of second segment in length and covered with black scales and hair-like scales, with slight patch of creamy-white scales laterally; antennae approximately two-fifth of forewing length, with ca 37 segments (n = 1), distal ca 15 composing club, club not prominent. THORAX. Dorsally, laterally and ventrally scattered with brownish gray scales. LEGS. Foreleg brownish, foretarsus and tibia similar in length, femur not examined; midleg and hindleg with femur creamy white ventrally, tibia and tarsus grayish dorsally, whitish to ocher ventrally, tarsus and tibia spined ventrally, and a pair of tibial spurs 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 2 towards M 1 than M 3 ; hindwing humeral developed. WING SHAPE. Forewing subtriangular, apex rounded, costal margin convex, outer margin slightly convex (almost straight), inner margin straight, but rounded towards thorax near base; hindwing slightly elongate, rounded, costal margin almost straight, angled towards thorax near base, outer margin slightly undulating, inner margin slightly concave near tornus, anal lobe convex, slightly round. DORSAL FOREWING. Ground color brownish, distally darker; androconial scales absent; trace of submarginal band indistinct. DORSAL HINDWING. Ground color similar to forewing, trace of submarginal and marginal band visible. VENTRAL FOREWING. Ground color light grayish brown, area below 2A paler; submedian line invisible; dark-brown narrow median line, extends from Radius to 2A, traversing wing outward with terminal end of median line almost reaching submarginal line; concolorous submarginal line extending from apex towards tornus, terminating around 2A; concolorous marginal line, narrower than submarginal line, extending from apex towards tornus, but terminates around 2A; fringe dark brownish. VENTRAL HINDWING. Ground color similar to forewing; regular dark-brown submedian line somewhat sigmoid, extending from costa to inner margin, terminal end curving basally along inner margin; median line almost parallel to submedian line, concolorous, similar in width, passing origin of M 3 , posterior and anterior end fused with submarginal line in 2A-3A and Rs-M 1 respectively; submarginal line extending from apex towards tornus, anterior and posterior end fused with median line in Rs-M 1 , jagged, posterior end slightly broadening and fused to median line in 2A-3A; marginal line, concolorous, undulate along outer margin, much thinner than submarginal line; submarginal ocelli from M 1 to 2A, roughly oval, pupil as silver scales placed distally, black central spot ringed with orangish ring then with thin dark brownish indistinct ring, those in M 2 -M 3 and M 3 -Cu 1 are both similar in size, those in M 1 -M 2 and Cu 2 -2A smaller, especially ocellus in M 1 -M 2 reduced, ocellus in Cu 1 -Cu 2 largest; fringe dark brownish. GENITALIA ( Fig. 5C ). Tegumen somewhat rounded in lateral view, dorsally curved in lateral view, domeshaped hump present posteriorly along dorsal margin; combination of ventral arms from tegumen and dorsal arms from saccus almost straight; appendices angulares present but rather inconspicuous; saccus shorter than uncus in length; uncus long and narrow, about twice as long as tegumen in length, with sparse hair-like setae, slightly hooked at posterior end; brachia long and narrow, shorter than uncus in length; fultura inferior (i.e., juxta) present as developed plate in posterior view; valva setose, basal half of valva roughly trapezoidal in lateral view, ventral margin convex in middle, dorsal margin almost straight except for margin basal to costa angles down to meet ventral margin, distal half narrow and somewhat elongate with rounded apex; costa subtriangular in lateral view, projecting towards appendices angulares; phallobase about one-third of phallus, rather straight; ductus ejaculatorius coming out higher than anterior end of coecum; 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 first and second subsegments apparently fused; forewing more rounded and broad; ground color of both wing surfaces paler, posterior end of VFW discal line do not come close to submarginal line as in male. VHW area between postdiscal and submarginal line whitish (around ocelli). Fig. 14. Distribution maps for taxa of Zischkaia Forster, 1964 . FEMALE ABDOMEN AND GENITALIA ( Fig. 7 G–H): inter-segmental membrane between seventh and eighth tergite not pleated, but expanded with posterior edge forming a smooth, curving sclerotized band anterior to ostium bursae that seamlessly borders the broad, sclerotized plate of lamella antevaginalis, which narrows ventrally to encircle ostium bursae and is indented throughout with the edges forming raised lip; eighth abdominal segment lateral side with sclerotized plate and with spiracle near dorsal margin, ventrally connected to lamella antevaginalis as sclerotized stripe, fused below ostium bursae as ‘arm’; ductus bursae membranous; origin of ductus seminalis close to ostium bursae; corpus bursae roughly oval in dorsal view, extending to third abdominal segment, with two signa in middle, parallel to each other. Variation The VFW submarginal and marginal lines are more or less fused in cell Cu 2 in some specimens, whereas they are placed rather further apart in other specimens. Fig. 15. Habitats of species of Zischkaia Forster, 1964 . A . Climbing bamboo species ( Poaceae , likely Chusquea Kunth ) that was common along the Sumak Yaku mine track, Zamora-Chinchipe, Ecuador, where Z. arenisca Nakahara, Willmott & Hall , sp. nov. was common (photo by Keith Willmott). B . Ridge top at San Roque ridge, Zamora-Chinchipe, Ecuador, where females of Z. arenisca sp. nov. were found (photo by Keith Willmott). C . View of the ridge line 500 m west of the Río Abanico (Morona-Santiago, Ecuador) collecting site (photo by Andrew Neild). Distribution ( Fig. 14 ) This species is known to date from southeastern to southern Brazil , in the states of Rio de Janeiro , Minas Gerais , São Paulo , Paraná and Santa Catarina . The geographic distribution of Z. warreni sp. nov. is apparently confined to areas associated with plateaus with Araucaria forest in middle to high elevations of southern Brazil , not extending beyond Serra Geral in the west. This distributional pattern is distinct from its potentially sympatric species Z. pacarus , which is also found in low elevation areas in the Paraná drainage and in the Atlantic coastal zone. Fig. 16. Habitats of species of Zischkaia Forster, 1964 . A . Habitat at the type locality of Z. josti Nakahara & Kleckner , sp. nov. , Kavanayen, Venezuela (photo by Bernhard Jost). B . Adult in situ at Serra da Esperança, Guarapuava, Paraná, Brazil (photo by Andrew Warren). C . Habitat of Z. warreni Dias, Zacca & Dolibaina , sp. nov. , Serra da Esperança, Guarapuava, Paraná, Brazil (photo by Andrew Warren). Remarks The evidence from molecular data (see Fig. 1 and Table 4 ) and morphological distinctiveness documented above leaves no doubt as to its specific status. Adults of Z. warreni sp. nov. are similar in behavior to Z. pacarus , flying in association with the bamboo Merostachys ( Poaceae : Bambusoideae ) ( Fig. 16A, C ). The males’ flight is faster than that of the females, and individuals are commonly found engaged in patrolling flight. Both sexes are never found away from the bamboo and they are more active between the middle of the morning and the beginning of the afternoon. The restricted habitat of Z. warreni sp. nov. in Araucaria forests suggests that they likely use Merostachys as the caterpillar host plant.