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Butterflies (Lepidoptera: Papilionoidea) of Iguaçu National Park and surrounding areas in southern Brazil: a long-term survey, with six new records for the Brazilian fauna

As Borboletas (Lepidoptera: Papilionoidea) do Parque Nacional do Iguaçu e arredores no Sul do Brasil: um inventário de longo prazo, com seis novos registros para a fauna brasileira

Abstract

Iguaçu National Park is the second largest (1852.62 km²) protected area in the Atlantic Forest domain and harbors the largest area of semideciduous seasonal forest in Brazil. In this study, we present 795 subspecies and 787 species of butterflies that occur in this protected area and its surrounding areas, collected over 15 years and ten months using different non-standardized sampling methods. We also searched for additional records in the literature, entomological collections, and citizen science platforms on the internet. Among the sampled taxa, six are recorded for the first time in Brazil: Emesis orichalceus Stichel, 1916, Theope p. pakitza Hall & Harvey, 1998 (Riodinidae), Elbella v. viriditas (Skinner, 1920), Apaustus gracilis ssp. n. (Hesperiidae), Deltaya sp. n. (Nymphalidae), and Symbiopsis sp. n. (Lycaenidae). Another six are listed as endangered in lists of butterflies of conservation concern. The records for some species significantly increase previously documented distributions.

Keywords
Atlantic Forest; biodiversity; conservation; endangered species; Neotropical region

Resumo

O Parque Nacional do Iguaçu é a segunda maior Unidade de Conservação (1.852,62 km²) no domínio Mata Atlântica, abrigando a maior área de Floresta Estacional Semidecídua no Brasil. Neste estudo apresentamos uma lista com 795 subespécies e 787 espécies de borboletas que ocorrem nesta Unidade de Conservação e seus arredores, coligida ao longo de 15 anos e dez meses através do uso de diferentes métodos de amostragem não padronizados. Nós também procuramos por registros adicionais na literatura, coleções entomológicas e plataformas de ciência cidadã na internet. Dentre os táxons amostrados, seis são registrados pela primeira vez para o Brasil: Emesis orichalceus Stichel, 1916, Theope p. pakitza Hall & Harvey, 1998 (Riodinidae), Elbella v. viriditas (Skinner, 1920), Apaustus gracilis ssp. n. (Hesperiidae), Deltaya sp. n. (Nymphalidae) e Symbiopsis sp. n. (Lycaenidae). Outras seis espécies são consideradas ameaçadas de extinção em listas de borboletas de interesse para a conservação. Os registros de algumas espécies aumentam significativamente as suas distribuições previamente documentadas.

Palavras-chave
biodiversidade; conservação; espécies ameaçadas; Mata Atlântica; região Neotropical

Introduction

Species inventories document biodiversity by contributing to taxonomic, ecological, and biogeographical studies, and by providing foundational data for management plans and other conservation actions (Brown Jr. & Freitas 1999BROWN, J.R., K.S. & FREITAS, A.V.L. 1999. Lepidoptera. In Biodiversidade do Estado de São Paulo: síntese do conhecimento ao final do século XX, invertebrados terrestres (C.R.F. Brandão & E.M. Cancello, eds.). USP, São Paulo, p.226–243., Lewinsohn et al. 2005LEWINSOHN, T.M., FREITAS, A.V.L. & PRADO, P.I. 2005. Conservação de Invertebrados terrestres e seus hábitats no brasil. Megadiversidade 1(1):62–69., Santos et al. 2008SANTOS, E.C., MIELKE, O.H.H., & CASAGRANDE, M.M. 2008. Inventários de borboletas no Brasil: estado da arte e modelo de áreas prioritárias para pesquisa com vistas à conservação. Nat. Conserv. 6(2):68–90.). Inventory data help document distributions of species and decrease the Wallacean shortfall (Lomolino 2004LOMOLINO, M.V. 2004. Conservation Biogeography. In Frontiers of Biogeography: new directions in the Geography of Nature (M.V. Lomolino & L.R. Heaney, eds.). Sinauer Associates, Sunderland, p.293–296., Whittaker et al. 2005WHITTAKER, R.J., ARAÚJO, M.B., JEPSON, P.R., LADLE, R.J., WATSON, J.E.M. & WILLIS, K.J. 2005. Conservation Biogeography: assessment and prospect. Divers. Distrib. 11(1):3–23.). They provide data for studies in community ecology and biogeography (e.g., Robbins et al. 1996ROBBINS, R.K., LAMAS, G., MIELKE, O.H.H., HARVEY, D.J. & CASAGRANDE, M.M. 1996. Taxonomic composition and ecological structure of the species-rich butterfly community at Pakitza, Parque Nacional del Manu, Perú. In Manu: The biodiversity of Southeastern Perú (D.E. Wilson & A. Sandoval, eds.). Smithsonian Institution Press, Washington, D.C., p.217–252., Brown Jr. & Freitas 2000BROWN, J.R., K.S. & FREITAS, A.V.L. 2000. Atlantic Forest butterflies: indicators for landscape conservation. Biotropica 32(4b):934–956., Gonçalves-Souza et al. 2014GONÇALVES-SOUZA, T., ROMERO, G.Q. & COTTENIE, K. 2014. Metacommunity versus biogeography: a case study of two groups of Neotropical vegetation dwelling arthropods. PLoS ONE 9(12):1–20., Zellweger et al. 2017ZELLWEGER, F., ROTH, T., BUGMANN H. & BOLLMANN, K. 2017. Beta diversity of plants, birds and butterflies is close ly associated with climate and habitat structure. Global Ecol. Biogeogr. 26(8):898–906.). Scientifically undescribed taxa may be discovered (e.g., Biezanko & Mielke 1973BIEZANKO, C.M. & MIELKE O.H.H. 1973. Contribuição ao estudo faunístico dos Hesperiidae americanos. IV. Espécies do Rio Grande do Sul, Brasil, com notas taxonômicas e descrições de espécies novas. (Lepidoptera). Acta Biol. Parana. 2(1–4):51–102., Núñez-Bustos 2008NÚÑEZ-BUSTOS, E. 2008. Diversidad de mariposas diurnas en la Reserva Privada Yacutinga, Provincia de Misiones, Argentina (Lepidoptera: Hesperioidea y Papilionoidea). Trop. Lepid. Res. 18(2):78–87., Dolibaina et al. 2011DOLIBAINA, D.R., MIELKE, O.H.H. & CASAGRANDE, M.M. 2011. Borboletas (Papilionoidea e Hesperioidea) de Guarapuava e arredores, Paraná, Brasil: um inventário com base em 63 anos de registros. Biota Neotrop. 11(1):341–354. https://www.biotaneotropica.org.br/v11n1/en/abstract?inventory+bn00211012011 (last access on 12/08/2021)
https://www.biotaneotropica.org.br/v11n1...
, Lamas et al. 2021LAMAS, G., MCINNIS, M.L., BUSBY, R.C. & ROBBINS, R.K. 2021. The lycaenid butterfly fauna (Lepidoptera) of Cosñipata, Peru: annotated checklist, elevational patterns, and rarity. Insecta Mundi 0861:1–34.), thus decreasing the Linnean shortfall (Brown & Lomolino 1998BROWN, J.H. & LOMOLINO, M.V. 1998. Biogeography 2 ed. Sinauer Associates, Sunderland.). Additionally, since butterflies are good bioindicators, responding quickly to environmental changes, well-documented inventories provide the base data that allow early detection of such changes through monitoring (Freitas et al. 2003FREITAS, A.V.L., FRANCINI, R.B. & BROWN JR, K.S. 2003. Insetos como indicadores ambientais. In Métodos de estudo em Biologia da Conservação e manejo da vida silvestre (L. Cullen Jr., R. Rudran & C. Valladares-Pádua, orgs.). Fundação Boticário e Editora da UFPR, Curitiba, p.125–151., Freitas et al. 2006FREITAS, A.V.L., LEAL, I.R., UEHARA-PRADO, M. & IANNUZZI, L. 2006. Insetos como indicadores de conservação da paisagem. In Biologia da Conservação: Essências (C.F.D. Rocha, H.G. Bergallo, M.V. Sluys & M.A.S. Alves, eds.). Rima, São Carlos, p.357–384., Santos et al. 2016SANTOS, J.P., MARINI-FILHO, O.J., FREITAS, A.V.L. & UEHARA-PRADO, M. 2016. Monitoramento de borboletas: o papel de um indicador biológico na gestão de unidades de conservação. Biodivers. Bras. 6(1):87–99.).

The Atlantic Forest is one of the most important hotspots of biodiversity in the world, and one of the most threatened Brazilian domains, with only ~11.7% of its original vegetation cover (Ribeiro et al. 2009RIBEIRO, M.C., METZGER, J.P., MARTENSEN, A.C., PONZONI, F.J. & HIROTA, M.M. 2009. The Brazilian Atlantic Forest: how much is left, and how is the remaining forest distributed? Implications for conservation. Biol. Conserv. 142:1144–1156.). Its geographic extent in latitude (3°S to 31°S), longitude (35°W to 60°W) and elevation (0 to 3000 m) (Ribeiro et al. 2009RIBEIRO, M.C., METZGER, J.P., MARTENSEN, A.C., PONZONI, F.J. & HIROTA, M.M. 2009. The Brazilian Atlantic Forest: how much is left, and how is the remaining forest distributed? Implications for conservation. Biol. Conserv. 142:1144–1156.) makes it a diverse domain, with a wide range of climate regimes and environmental heterogeneity. Among all Brazilian domains, the Atlantic Forest has the most representative butterfly inventories (Santos et al. 2008SANTOS, E.C., MIELKE, O.H.H., & CASAGRANDE, M.M. 2008. Inventários de borboletas no Brasil: estado da arte e modelo de áreas prioritárias para pesquisa com vistas à conservação. Nat. Conserv. 6(2):68–90., Shirai et al. 2019SHIRAI, L.T., MACHADO, P.A., MOTA, L.L., ROSA, A.H.B. & FREITAS, A.V.L. 2019. DnB, the database of nymphalids in Brazil, with a checklist for standardized species lists. J. Lepid. Soc. 73(2):93–108.), and its regional richness exceeds 2100 species (Brown Jr. & Freitas 2000BROWN, J.R., K.S. & FREITAS, A.V.L. 2000. Atlantic Forest butterflies: indicators for landscape conservation. Biotropica 32(4b):934–956.). Despite these results, more information to better understand patterns of butterfly geographic distributions is needed (Francini et al. 2011FRANCINI, R.B., DUARTE, M., MIELKE, O.H.H., CALDAS, A. & FREITAS, A.V.L. 2011. Butterflies (Lepidoptera, Papilionoidea e Hesperioidea) of the “Baixada Santista” region, coastal São Paulo, southeastern Brazil. Rev. Bras. Entomol. 55(1):55–68., Iserhard et al. 2017ISERHARD, C.A., UEHARA-PRADO, M., MARINI-FILHO, O.J., DUARTE, M. & FREITAS, A.V.L. 2017. Fauna da Mata Atlântica: Lepidoptera – Borboletas. In Revisões em Zoologia: Mata Atlântica (E.L. de A. Monteiro-Filho & C.E. Conte, eds.). Editora da UFPR, Curitiba, p.57–102.).

Iguaçu National Park (hereafter Iguaçu NP) is a protected area in the Atlantic Forest domain that harbors the largest preserved area of semideciduous seasonal forest in Brazil (Urban 2002URBAN, T. 2002. Parque Nacional do Iguaçu: Caminho Aberto para a Vida. Rede Nacional Pró-Unidades de Conservação, Rede Verde de Informações Ambientais, Curitiba.). Mielke (1968)MIELKE, O.H.H. 1968. Contribuição ao estudo faunístico dos “Hesperiidae” brasileiros I. Resultados de uma excursão à Foz do Iguaçu, Paraná, Brasil, com notas taxonômicas (Lepidoptera). Atas Soc. Biol. Rio J. 12(2):73–78. was the first to publish a butterfly list for the region, with emphasis on Hesperiidae, and recorded 106 species in the municipality of Foz do Iguaçu. Decades later, a Rapid Ecological Assessment (REA), conducted to develop a management plan in this protected area, recorded 257 species of six families of butterflies (Mielke 1998MIELKE, O.H.H. 1998. Plano de Manejo do Parque Nacional do Iguaçu. Avaliação Ecológica Rápida. Caracterização da Entomofauna. Versão Revisada. IBAMA/FUPEF, Curitiba.). After that, Graciotim & Morais (2016)GRACIOTIM, C. & MORAIS, A.B.B. 2016. Borboletas frugívoras em florestas de mata atlântica do Parque Nacional do Iguaçu, Paraná, Brasil (Lepidoptera: Nymphalidae). SHILAP-Rev. Lepidopt. 44(173):115–128. published a list with 69 fruit-feeding nymphalid species for this region. The authors sampled using Van Someren-Rydon traps placed along two trails, which represented the two principal phytophysiognomies in the park (Urban 2002URBAN, T. 2002. Parque Nacional do Iguaçu: Caminho Aberto para a Vida. Rede Nacional Pró-Unidades de Conservação, Rede Verde de Informações Ambientais, Curitiba., ICMBio 2018bICMBIO 2018b. Plano de Manejo do Parque Nacional do Iguaçu. https://www.icmbio.gov.br/portal/images/stories/plano-de-manejo/plano_de_manejo_do_parna_do_iguacu_fevereiro_2018.pdf (last access on 16/09/2021)
https://www.icmbio.gov.br/portal/images/...
). With further sampling, Santos et al. (2018)SANTOS, J.P., FREITAS, A.V.L., BROWN JR., K.S., CARREIRA, J.Y.O, GUERATTO, P.E., ROSA, A.H.B., LOURENÇO, G.M., ACCACIO, G.M., UEHARA-PRADO, M., ISERHARD, C.A., RICHTER, A.G.K., ROMANOWSKI, H.P., MEGA, N.O., TEIXEIRA, M.O., MOSER, A., RIBEIRO, D.B., ARAUJO, P.F., FILGUEIRAS, B.K.C., MELO, D.H.A., LEAL, I.R., BEIRÃO, M.V., RIBEIRO, S.P., CAMBUÍ, E.C.B., VASCONCELOS, R.N., CARDOSO, M.Z., PALUCH, M., GREVE, R.R., VOLTOLINI, J.C., GALETTI, M., REGOLIN, A.L., SOUZA, T.S. & RIBEIRO, M.C. 2018. Atlantic butterflies: a data set of fruit feeding butterfly communities from the Atlantic forests. Ecology 99(12):2875. updated this list to 104 fruit-feeding nymphalid species. Considering the estimated richness of 700 butterfly species for Iguaçu NP (IBAMA 1999HALL, J.P.W. & K.R. WILLMOTT. 2010. Description of a new Lucillella species Riodinidae: Symmachiini) discovered in the eastern Andes of Ecuador using the single rope canopy access technique. J. Lepid. Soc. 64:139–146.) and the 653 species recorded in Argentina’s Iguazú National Park (Núñez-Bustos 2009NÚÑEZ-BUSTOS, E. 2009. Mariposas diurnas (Lepidoptera: Papilionoidea y Hesperioidea) del Parque Nacional Iguazú, Provincia de Misiones, Argentina. Trop. Lepid. Res. 19(2):71–81.), it appeared that Iguaçu NP was still under sampled. The purpose of this paper is to remedy this undersampling by producing an exhaustive list resulting from more than 15 years of sampling at Iguaçu NP and its surrounding areas, supplemented by records from the literature and from entomological collections. Our data are intended to contribute both to the taxonomy and ecology of Brazilian butterflies and to be used for conservation decisions in this emblematic National Park of the Atlantic Forest in Brazil.

Material and Methods

1. Study area

Iguaçu NP is a Brazilian protected area located in the state of Paraná, between the geographic coordinates of 25º05' and 25º41'S and 53º40' and 54º38'W (Datum WGS84) (Figure 1). The park area is 1852.62 km² with a 420 km perimeter. It encompasses six municipalities in the state of Paraná (Céu Azul, Capanema, Matelândia, Serranópolis do Iguaçu, São Miguel do Iguaçu and Foz do Iguaçu) and borders another four (Capitão Leônidas Marques, Lindoeste, Santa Tereza do Oeste and Santa Terezinha de Itaipu). The Iguaçu River separates Iguaçu NP in Brazil from its sister park in Argentina (Iguazú National Park). Five rivers, with their sources outside of the park, flow across the park to the Iguaçu River: Gonçalves Dias, São João, Represa Grande, Silva Jardim and Benjamin Constant. A sixth river, the Floriano River, is the only one situated almost entirely in the park. Geologically, Iguaçu NP is located on the third Paraná plateau, in the Paraná Basin.

Figure 1.
Map of Iguaçu National Park (in dark green) and its location in Brazil, with the main sampling sites; yellow circles = previous surveys by Mielke (1998MIELKE, O.H.H. 1998. Plano de Manejo do Parque Nacional do Iguaçu. Avaliação Ecológica Rápida. Caracterização da Entomofauna. Versão Revisada. IBAMA/FUPEF, Curitiba.); red triangles = sites surveyed in the present study.

The climate in the region is classified as humid subtropical (Cfa) according to Köppen, with mean temperatures below 18°C during the coldest month and above 22°C during the hottest month (Alvares et al. 2013ALVARES, C.A., STAPE, J.L., SENTELHAS, P.C., GONÇALVES, J.L.M & SPAROVEK, G. 2013. Köppen’s climate classification map for Brazil. Meteorol. Z. 22(6):711–728;). Annual precipitation varies from 1600 to 1800 mm, and the rains are concentrated mainly during hot months. The dry season is not well defined, and the frequency of frosts is low (Nitsche et al. 2019NITSCHE, P.R., CARAMORI, P.H., RICCE, W.S. & PINTO, L.F.D. 2019. Atlas climático do Estado do Paraná. Instituto Agronômico do Paraná, Londrina.). According to Google EarthGOOGLE EARTH. http://earth.google.com (last access on 13/06/2023)
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(http://earth.google.com), the elevation of the park varies from 120 m, at the Iguaçu River, to over 700 m at its northeastern extremity. The park is located in the Atlantic Forest domain and is composed of four phytophysiognomies (ICMBio 2018bICMBIO 2018b. Plano de Manejo do Parque Nacional do Iguaçu. https://www.icmbio.gov.br/portal/images/stories/plano-de-manejo/plano_de_manejo_do_parna_do_iguacu_fevereiro_2018.pdf (last access on 16/09/2021)
https://www.icmbio.gov.br/portal/images/...
): 1) the seasonal semideciduous forest (hereafter SSF), 2) the mixed ombrophilous forest (hereafter MOF), 3) the floodplain (hereafter FP), and 4) the open formations (hereafter OF). The SSF occupies 85% of the park area and has two subformations, the submontane (up to 400 m), and the montane (between 400 and 600 m). The MOF has a single subformation, the montane (above 600 m). The FP occurs in small areas, subject to extreme flooding conditions for long periods, and is generally occupied by herbaceous-shrub vegetation. The OF comprises all open vegetation formations of anthropogenic or natural origin (Urban 2002URBAN, T. 2002. Parque Nacional do Iguaçu: Caminho Aberto para a Vida. Rede Nacional Pró-Unidades de Conservação, Rede Verde de Informações Ambientais, Curitiba., ICMBio 2018bICMBIO 2018b. Plano de Manejo do Parque Nacional do Iguaçu. https://www.icmbio.gov.br/portal/images/stories/plano-de-manejo/plano_de_manejo_do_parna_do_iguacu_fevereiro_2018.pdf (last access on 16/09/2021)
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) (Figure 2). The surrounding areas of the park exhibit a heterogeneous landscape, characterized by forest fragments, extensive areas dedicated to agriculture and raising livestock, and urban areas (ICMBio 2018bICMBIO 2018b. Plano de Manejo do Parque Nacional do Iguaçu. https://www.icmbio.gov.br/portal/images/stories/plano-de-manejo/plano_de_manejo_do_parna_do_iguacu_fevereiro_2018.pdf (last access on 16/09/2021)
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).

Figure 2.
Landscapes and environments at Iguaçu National Park: A) semideciduous seasonal forest, 200 m; B) mixed ombrophilous forest, over 700 m, with the presence of several individuals of Brazilian pine Araucaria angustifolia (Bertol.) Kuntze (Araucariaceae); C) stony beach on the banks of the Iguaçu River; D) Blue River; E) a dense patch of tree ferns (Cyatheaceae spp.); F) close view of forest understory with young juçara palm Euterpe edulis Mart. (Arecaceae).

2. Sampling

The faunal list is primarily the result of 15 years and ten months of sampling (September 2007 to July 2023) of adult butterflies by the first author, mainly using entomological nets, but in some situations, also using Van Someren-Rydon (hereafter VSR) traps and the Ahrenholz technique, which was especially useful for sampling Hesperiidae butterflies (Freitas et al. 2021bFREITAS, A.V.L., SANTOS, J.P., ROSA, A.H., ISERHARD, C.A., RICHTER, A., SIEWERT, R.R., GUERATTO, P.E., CARREIRA, J.Y.O. & LOURENÇO, G.M. 2021b. Sampling methods for butterflies (Lepidoptera). In Measuring arthropod biodiversity: a handbook of sampling methods (J.C. Santos & G.W. Fernandes, eds.). Springer International Publishing, Cham, p.101–123.). A digital camera was also used to record some species. Some immature stages were found, which were then raised in the laboratory until emergence.

In the collection with entomological net, many trails and environments were sampled, in all four phytophysiognomies and throughout the altitudinal range (Table 1). Following Brown Jr. & Freitas (2000)BROWN, J.R., K.S. & FREITAS, A.V.L. 2000. Atlantic Forest butterflies: indicators for landscape conservation. Biotropica 32(4b):934–956., special emphasis was given to sites with resources that attract butterflies, like flowering plants, plant exudates, sunny areas in the interior of the forest, mud puddles, fermented fruits, wet sand along trails or river margins, feces or urine from carnivores, and carcasses. Some sites were more intensively sampled, especially those in the municipality of Foz do Iguaçu. Field sampling was carried out at all seasons, at different times of the day, from 9:00 a.m. to 7:00 p.m., and in all microclimates or habitats within the park and in surrounding areas.

Table 1.
Sampled sites in the present study and in Mielke (1998)MIELKE, O.H.H. 1998. Plano de Manejo do Parque Nacional do Iguaçu. Avaliação Ecológica Rápida. Caracterização da Entomofauna. Versão Revisada. IBAMA/FUPEF, Curitiba., ordered from North to South. Nearby points were treated as a single point. In the case of trails, the altitude presented is the simple arithmetic mean between its highest and lowest point. SSFs = Seasonal Semideciduous Forest, subformation submontana; SSFm = Seasonal Semideciduous Forest, subformation montana; MOF = Mixed Ombrophilous Forest; FP = Floodplain; OF = Open formations.

In March 2013, VSR traps baited with a mixture of fermented banana and sugarcane juice were installed on two trails in the Iguaçu NP to sample fruit-feeding butterflies, following a protocol adapted from Uehara-Prado et al. (2005)UEHARA-PRADO, M., BROWN JR., K.S. & FREITAS, A.V.L. 2005. Biological traits of frugivorous butterflies in a fragmented and a continuous landscape in the South Brazilian Atlantic Forest. J. Lepid. Soc. 59(2):96–106.. These trails represent the two principal phytophysiognomies present in the park: 1) an area of SSF, located in the municipality of Foz do Iguaçu, and 2) an area of MOF, located in the municipality of Céu Azul. On each trail, ten traps were placed in line (starting at least 50 m from the forest edge) and with about 20 m between adjacent traps, alternating understory (~1.5 m above the ground) and canopy (~10 m above the ground). The traps remained suspended for two consecutive days on each trail (10 hours of effective sampling per day), resulting in a total sampling effort of 400 trap/hours.

Most Hesperiidae were deposited at the Entomological Collection Padre Jesus Santiago Moure in the Federal University of Paraná (DZUP/UFPR), and specimens of the other families were deposited at the Zoological Collection of the Museu de Diversidade Biológica in the University of Campinas (ZUEC/Unicamp). Some specimens of the genus Actinote were deposited at the Entomological Collection of the Museu de Zoologia e Paleontologia in the Federal University of Recôncavo da Bahia (MURB/UFRB).

3. Species list

We identified the specimens using taxonomic keys and species guides (e.g., Brown Jr. 1992BROWN, J.R., K.S. 1992. Borboletas da Serra do Japi: Diversidade, habitats, recursos alimentares e variação temporal. In História natural da Serra do Japi: Ecologia e preservação de uma área florestal no sudeste do Brasil (L.P.C. Morellato, org.). Editora da Unicamp/FAPESP, Campinas, p.142–187., Canals 2003CANALS, G.R. 2003. Mariposas de Misiones. L.O.L.A., Buenos Aires., D’Abrera 1984D’ABRERA, B.L. 1984. Butterflies of the Neotropical Region. Part. II. Danaidae, Ithomiidae Heliconidae and Morphidae. Hill House, Victoria., 1987D’ABRERA, B.L. 1987. Butterflies of the Neotropical Region. Part. IV. Nymphalidae (Partim). Hill House, Victoria., 1989D’ABRERA, B.L. 1989. Butterflies of the Neotropical Region. Part. V. Nymphalidae (Conc.) and Satyridae. Hill House, Victoria., 1994D’ABRERA, B.L. 1994. Butterflies of the Neotropical Region. Part. VI. Riodinidae. Hill House, Victoria., 1995D’ABRERA, B.L. 1995. Butterflies of the Neotropical Region. Part. VII. Lycaenidae. Hill House, Victoria., Tyler et al. 1994TYLER, H.A., BROWN JR., K.S. & WILSON, K.H. 1994. Swallowtail butterflies of the Americas. A study in biological dynamics, ecological diversity, biossystematics and conservation. Scientific Publishers, Gainesville., Klimaitis et al. 2018KLIMAITIS, J.F., NÚÑEZ-BUSTOS, E., KLIMAITIS, C.L. & GULLER, R.M. 2018. Mariposas de Argentina: Guía de identificación. Vazquez Mazzini Editores, Buenos Aires.). We also compared specimens to images of type specimens of American butterflies, available in the Butterflies of America database, on internet (Warren et al. 2016WARREN, A.D., DAVIS, K.J., STANGELAND, E.M., PELHAM, J.P., WILLMOTT, K.R. & GRISHIN, N.V. 2016. Illustrated Lists of American Butterflies. http://www.butterfliesofamerica.com (last access on 12/05/2022)
http://www.butterfliesofamerica.com...
), and to species maintained in reference collections (DZUP/UFPR and ZUEC/Unicamp). For some individuals, we examined the genitalia and, more rarely, we used barcoding sequences to determine the species. We also consulted specialists to help us determine specific groups of butterflies (see “Acknowledgements”). For recorded subspecies with sympatric distribution, we use an “x” to indicate that two phenotypes (and sometimes the putative hybrids) are present.

Besides sampling, we compiled data from the literature (Mielke 1968MIELKE, O.H.H. 1968. Contribuição ao estudo faunístico dos “Hesperiidae” brasileiros I. Resultados de uma excursão à Foz do Iguaçu, Paraná, Brasil, com notas taxonômicas (Lepidoptera). Atas Soc. Biol. Rio J. 12(2):73–78., Jenkins 1990JENKINS, D.W. 1990. Neotropical Nymphalidae. VIII. Revision of Eunica. Bulletin of the Allyn Museum, Florida., Casagrande & Mielke 1992CASAGRANDE, M.M. & MIELKE, O.H.H. 1992. Borboletas (Lepidoptera) Ameaçadas de Extinção no Paraná. Rev. Bras. Zool. 9(1/2):75–92., Anken 1994ANKEN, R.H. 1994. Neue taxa des genus Hermeuptychia Forster aus Brasilien (Lepidoptera, Satyridae). Entomol. Z. 104(14):283–291., D’Abrera 1995D’ABRERA, B.L. 1995. Butterflies of the Neotropical Region. Part. VII. Lycaenidae. Hill House, Victoria., Mielke 1998MIELKE, O.H.H. 1998. Plano de Manejo do Parque Nacional do Iguaçu. Avaliação Ecológica Rápida. Caracterização da Entomofauna. Versão Revisada. IBAMA/FUPEF, Curitiba., Hall & Harvey 2002HALL, J.P.W. & HARVEY, D.J. 2002. Basal subtribes of the Nymphidiini (Lepidoptera: Riodinidae): Phylogeny and myrmecophily. Cladistics 18(6):539–569., Callaghan 2010CALLAGHAN, C.J. 2010. A re-evaluation of the Aricoris constantius group with the recognition of three species (Lepidoptera: Riodinidae). Zoologia 27(3):395–401., Lourido 2011LOURIDO, G.M. 2011. Revisão taxonômica e análise filogenética da família Neotropical Hedylidae (Lepidoptera, Hedyloidea). Tese de doutorado, Instituto Nacional de Pesquisas da Amazônia, Manaus., Graciotim & Morais 2016GRACIOTIM, C. & MORAIS, A.B.B. 2016. Borboletas frugívoras em florestas de mata atlântica do Parque Nacional do Iguaçu, Paraná, Brasil (Lepidoptera: Nymphalidae). SHILAP-Rev. Lepidopt. 44(173):115–128.) for any of the ten municipalities in the study area, updating the nomenclature when necessary. We also examined scientific collections, where we found species from the study area that were not previously recorded for the park. These include DZUP at UFPR; two local collections in Foz do Iguaçu, the Ecomuseum of Itaipu and the Museum of Zoology in the Uniamérica Universitary Center; and two private collections, D.R. Dolibaina and A.D. Warren. We did not include species that could not be identified, as well as literature records for which we did not find voucher material or which may represent species misidentification. In addition, we searched for pictures of butterflies from any of the ten municipalities within and bordering Iguaçu NP on the citizen science platform iNaturalistINATURALIST. https://www.inaturalist.org (last access on 27/09/2022)
https://www.inaturalist.org...
(https://www.inaturalist.org), Google ImagesGOOGLE IMAGES. https://www.google.com/imghp?hl=EN (last access on 27/09/2022)
https://www.google.com/imghp?hl=EN...
(https://www.google.com/imghp?hl=EN) and FlickrFLICKR. https://www.flickr.com (last access on 27/09/2022)
https://www.flickr.com...
(https://www.flickr.com), but no additional records were obtained.

Pareuptychia summandosa (Gosse, 1880) was considered a junior synonym of P. ocirrhoe (Fabricius, 1776) (M.A. Marín, pers. comm.). In the case of Junonia, a species complex that has not yet been unveiled, the different sampled phenotypes were grouped under the single taxon Junonia e. evarete (Cramer, 1779). Following Freitas et al. (2014)FREITAS, A.V.L., ISERHARD, C.A., SANTOS, J.P., CARREIRA, J.Y.O., RIBEIRO, D.B., MELO, D.H.A., ROSA, A.H.B., MARINI-FILHO, O.J., ACCACIO, G.M. & UEHARA-PRADO, M. 2014. Studies with butterfly bait traps: an overview. Rev. Colomb. Entomol. 40(2):209–218., we consider as fruit-feeding species only the nymphalids belonging to the subfamilies Biblidinae (except Eubagini and Mestra), Charaxinae, Satyrinae (except non-feeding Brassolis and Dynastor), and Nymphalinae (including only Colobura, Historis and Smyrna).

We recognized seven butterfly families, including Hedylidae, following recent phylogenetic studies (Kawahara & Breinholt 2014KAWAHARA, A.Y. & BREINHOLT, J.W. 2014. Phylogenomics provides strong evidence for relationships of butterflies and moths. Proc. R. Soc. London, Ser. B 281(1788):1–8., Breinholt et al. 2018BREINHOLT, J.W., EARL, C., LEMMON, A.R., LEMMON, E.M., XIAO, L. & KAWAHARA, A.Y. 2018. Resolving relationships among the megadiverse butterflies and moths with a novel pipeline for Anchored Phylogenomics. Syst. Biol. 67(1):78–93., Espeland et al. 2018ESPELAND, M., BREINHOLT, J., WILLMOTT, K.R., WARREN, A.D., VILA, R., TOUSSAINT, E.F.A., MAUNSELL, S.C., ADUSE-POKU, K., TALAVERA, G., EASTWOOD, R., JARZYNA, M.A., GURALNICK, R., LOHMAN, D.J., PIERCE, N.E. & KAWAHARA, A.Y. 2018. A comprehensive and dated phylogenomic analysis of butterflies. Curr. Biol. 28(5):770–778.). For higher taxonomy within families, we followed Lamas (2004)LAMAS, G. 2004. Checklist: Part 4A. Hesperioidea – Papilionoidea. In Atlas of Neotropical Lepidoptera (J.B. Heppner, org.). vol. 5A. Association of Tropical Lepidoptera, Gainesville., updated for Nymphalidae (Wahlberg et al. 2009WAHLBERG, N., LENEVEU, J., KODANDARAMAIAH, U., PEÑA, C., NYLIN, S., FREITAS, A.V.L., & BROWER, A.V.Z. 2009. Nymphalid butterflies diversify following near demise at the Cretaceous/Tertiary boundary. Proc. R. Soc. London, Ser. B 276(1677):4295–4302.), Riodinidae (Seraphim et al. 2018SERAPHIM, N., KAMINSKI, L.A., DEVRIES, P.J., PENZ, C., CALLAGHAN, C.J., WAHLBERG, N., SILVA-BRANDÃO, K.L. & FREITAS, A.V.L. 2018. Molecular phylogeny and higher systematics of the metalmark butterflies (Lepidoptera: Riodinidae). Syst. Entomol. 43(2):407–425., Seraphim 2019SERAPHIM, N. 2019. Riodinidae Species Checklist: a preliminary species checklist for the Riodinidae. https://www2.ib.unicamp.br/labor/site/?page_id=805 (last access on 06/02/2022)
https://www2.ib.unicamp.br/labor/site/?p...
), and Hesperiidae (Li et al. 2019LI, W., CONG, Q., SHEN, J., ZHANG, J., HALLWACHS, W., JANZEN, D.H. & GRISHIN, N.V. 2019. Genomes of skipper butterflies reveal extensive convergence of wing patterns. Proc. Natl. Acad. Sci. U.S.A. 116:6232–6237., Zhang et al. 2019ZHANG, J., CONG, Q., SHEN, J., BROCKMANN, E. & GRISHIN, N.V. 2019. Genomes reveal drastic and recurrent phenotypic divergence in firetip skipper butterflies (Hesperiidae: Pyrrhopyginae). Proc. R. Soc. London, Ser. B. 286(1903):20190609., Cong et al. 2019CONG, Q., ZHANG, J., SHEN, J. & GRISHIN, N.V. 2019. Fifty new genera of Hesperiidae (Lepidoptera). Insecta Mundi 0731:1–56., Zhang et al. 2022ZHANG, J., CONG, Q., SHEN, J. & GRISHIN, N.V. 2022. Taxonomic changes suggested by the genomic analysis of Hesperiidae (Lepidoptera). Insecta Mundi 921:1–135.). For generic names, we followed Mielke et al. (2022)MIELKE, O.H.H., BROCKMANN, E. & MIELKE, C.G.C. 2022. Butterflies of the World 49. Hesperiidae II. New World Pyrrhopyginae, detailed text. Goecke & Evers, Keltern. and Brockmann et al. (2022)BROCKMANN, E., MIELKE, C.G.C. & MIELKE, O.H.H. 2022. Butterflies of the World 50. Hesperiidae III. New World Pyrrhopyginae, short text and plates. Goecke & Evers, Keltern. for the subfamily Pyrrhopyginae, Robbins et al. (2022)ROBBINS, R.K., CONG, Q., ZHANG, J., SHEN, J., BUSBY, R.C., FAYNEL, C., DUARTE, M., MARTINS, A.R.P., PRIETO, C., LAMAS, G. & GRISHIN, N.V. 2022. Genomics-based higher classification of the species-rich hairstreaks (Lepidoptera: Lycaenidae: Eumaeini). Syst. Entomol. 47(3):445–469. for the tribe Eumaeini and Espeland et al. (2023)ESPELAND, M., NAKAHARA, S., ZACCA, T., BARBOSA, E.P., HUERTAS, B., MARÍN, M.A., LAMAS, G., BENMESBAH, M., BRÉVIGNON, C., CASAGRANDE, M.M., FÅHRAEUS, C., GRISHIN, N., KAWAHARA, A.Y., MIELKE, O.H.H., MILLER, J.Y., NAKAMURA, I., NAVAS, V., PATRUSKY, B., PYRCZ, T.W., RICHARDS, L., TAN, D., TYLER, S., VILORIA, Á., WARREN, A.D., XIAO, L., FREITAS, A.V.L. & WILLMOTT, K.R. 2023. Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae). Syst. Biol. 1–73. for the subtribe Euptychiina. Taxonomy was also updated for the genera Morpho (Blandin 2007BLANDIN, P. 2007. The systematics of the genus Morpho Fabricius, 1807 (Lepidoptera Nymphalidae, Morphinae). Hillside Books, Canterbury., Pablos et al. 2021PABLOS, J.L., SILVA, A.K., SERAPHIM, N., MAGALDI, L.M., SOUZA, A.P., FREITAS, A.V.L. & SILVA-BRANDÃO, K.L. 2021. North-south and climate-landscape-associated pattern of population structure for the Atlantic Forest White Morpho butterflies. Mol. Phylogenet. Evol. 161(10):1–18.), Eryphanis (Penz 2008PENZ, C.M. 2008. Phylogenetic revision of Eryphanis Boisduval, with a description of a new species from Ecuador (Lepidoptera, Nymphalidae). Insecta Mundi 35:1–25.), Zaretis (Dias et al. 2018DIAS, F.M.S., JANZEN, D., HALLWACHS, W., CHACÓN, I., WILLMOTT, K., ORTIZ-ACEVEDO, E., MIELKE, O.H.H. & CASAGRANDE, M.M. 2018. DNA barcodes uncover hidden taxonomic diversity behind the variable wing patterns in the Neotropical butterfly genus Zaretis (Lepidoptera: Nymphalidae: Charaxinae). Zool. J. Linn. Soc.-Lond. XX:1–61.), Phoebis (Murillo-Ramos et al. 2018MURILLO-RAMOS, L., TORRES, R.H., NÚÑEZ, R. & AYAZO, R. 2018. New insights on the taxonomy and phylogenetic relationships of the Neotropical genus Phoebis (Pieridae: Coliadinae) revealed by molecular and morphological data. Zootaxa 4457(1):179–188.), Agraulis and Dryas (Núñez et al. 2022NÚÑEZ, R., WILLMOTT, K.R., ÁLVAREZ, Y., GENARO, J.A., PÉREZ‐ASSO, A.R., QUEJERETA, M., TURNER, T., MILLER, J.Y., BRÉVIGNON, C., LAMAS, G. & HAUSMANN, A. 2022. Integrative taxonomy clarifies species limits in the hitherto monotypic passion-vine butterfly genera Agraulis and Dryas (Lepidoptera, Nymphalidae, Heliconiinae). Syst. Entomol. 47:152–178.).

Results

We recorded 787 species (totalling 795 taxa including subspecies) from seven families of Papilionoidea at Iguaçu NP and its surrounding areas (Table 2), consisting of 339 (43.1%) Hesperiidae, 214 (27.2%) Nymphalidae, 115 (14.6%) Lycaenidae, 77 (9.8%) Riodinidae, 24 (3.0%) Pieridae, 17 (2.2%) Papilionidae, and 1 (0.1%) Hedylidae (Figure 3). Of these 795 taxa, 729 were sampled by the first author (including seven only by taking pictures of live specimens), 27 were obtained from the literature and 39 from scientific collections (see “Data Availability” for more details). From the guild of fruit-feeding nymphalids, we recorded a total of 112 species, representing 52.3% of Nymphalidae richness.

Table 2.
List of the butterflies and moth-butterflies (Lepidoptera: Papilionoidea) of Iguaçu National Park and surrounding areas. Species richness for each major taxa are provided within parentheses. * Taxa recorded by the first author solely through photographs of live specimens; ** records from collections or literature (see “Data Availability” for more details); # new records for Brazil; the “x” indicates that phenotypes of two subspecies are present; superscript letters a = the first subspecies correspond to the predominant phenotype; b = very few specimens were collected to determine predominant phenotype; c = there is evidence suggesting that the two subspecies could in fact be two valid species; F = strictly fruit-feeding species (sensu Freitas et al. 2014).
Figure 3.
Sample of representative species present at Iguaçu National Park and surrounding areas representing six butterfly families (only Hedylidae is not illustrated): A) Pyrrhopygopsis s. socrates (Ménétriés, 1855); B) Dalla diraspes (Hewitson, 1877); C) Festivia cronion (Felder & Felder, 1867); D) Mysoria barcastus barta Evans, 1951 (Hesperiidae); E) Myscelia orsis (Drury, [1782]); F) Epityches eupompe (Geyer, [1832]); G) Argentaria libitina (Butler, 1870); H) Catagramma pygas thamyras Ménétriés, 1857 (Nymphalidae); I) Zizula cyna (Edwards, 1881); J) Cyanophrys herodotus (Fabricius, 1793); K) Heterosmaitia palegon (Cramer, [1780]); L) Evenus regalis (Cramer, [1775]) (Lycaenidae); M) Rhetus periander arthuriana (Sharpe, 1890); N) Alesa prema (Godart, [1824]); O) Caria p. plutargus (Fabricius, 1793); P) Ariconias glaphyra (Westwood, 1851) (Riodinidae); Q) Anteos clorinde (Godart, [1824]); R) Pyrisitia l. leuce (Boisduval, 1836); S) Itaballia demophile ssp.; T) Dismorphia amphione astynome (Dalman, 1823) (Pieridae); U) Heraclides hectorides (Esper, 1794); V) Protesilaus stenodesmus (Rothschild & Jordan, 1906); W) Mimoides protodamas (Godart, 1819); X) Parides agavus (Drury, 1782) (Papilionidae).

The eight species that had more than one recorded subspecies were: 1) Heliconius ethilla (Godart, 1819), 2) Hypothyris euclea (Godart, 1819), 3) Tithorea harmonia (Cramer, 1777) (Nymphalidae), 4) Ascia monuste (Linnaeus, 1764) (Pieridae), 5) Melanis aegates (Hewitson, 1874), 6) Rhetus periander (Cramer, 1777) (Riodinidae), 7) Saturnus reticulata (Plötz, 1883), and 8) Tisias lesueur (Latreille, [1824]) (Hesperiidae).

We also recorded several taxa that are typical from other domains, such as Amazon and Cerrado. Among typical Amazonian taxa (Díaz et al. 2014DÍAZ, S.D.R., DIAS, F.M.S., CASAGRANDE, M.M., MIELKE, O.H.H. & LAMAS, G. 2014. Notes on the geografic distribution and subspecific taxonomy of Sais rosalia (Cramer) (Lepidoptera, Nymphalidae, Ithomiini), including the first records in Paraguay. Rev. Bras. Entomol. 58(1):11–18., Busby et al. 2017BUSBY, R.C., FAYNEL, C., MOSER, A. & ROBBINS, R.K. 2017. Sympatric diversification in the upper Amazon: a revision of the Eumaeine genus Paraspiculatus (Lepidoptera: Lycaenidae). Smithson. Contrib. Zool. 649:1–65., Hall 2018HALL, J.P.W. 2018. A monograph of the Nymphidiina (Lepidoptera: Riodinidae: Nymphidiini): Phylogeny, Taxonomy, Biology, and Biogeography. Entomological Society of Washington, Washington, D.C.) are Sais rosalia rosalinde Weymer, 1890 (Nymphalidae), Paraspiculatus orobia (Hewitson, 1867) (Lycaenidae), and Pseudolivendula h. hemileuca (Bates, 1868) (Riodinidae). Among those typical from Cerrado (Mielke et al. 2008MIELKE, O.H.H., EMERY, E.O. & PINHEIRO, C.E.G. 2008. As borboletas Hesperiidae (Lepidoptera, Hesperioidea) do Distrito Federal, Brasil. Rev. Bras. Entomol. 52(2):283–288., Pinheiro et al. 2010PINHEIRO, C.E.G., MALINOV, I.K., EMERY, E.O. & SCHMIDT, K. 2010. Endemismos e conservação de borboletas (Lepidoptera: Papilionoidea e Hesperioidea) no bioma Cerrado. In Cerrado: Conhecimento científico quantitativo como subsídio para ações de conservação (I.R. Diniz, J. Marinho-Filho, R.B. Machado & R.B. Cavalcanti, eds.). Editora da UNB, Brasília, D.F., p.225–238., Freitas et al. 2021aFREITAS, A.V.L., MUNIZ, D.G., CARREIRA, J.Y.O. & OLIVEIRA, P.S. 2021a. Fruit-feeding butterfly assemblages in a Neotropical Savanna: Assessing phenological patterns using baited traps. J. Lepid. Soc. 75(2):88–103.) are Aguna squamalba Austin & Mielke, 1998, Blubella azeta giffordi (Mielke, 1995), Elbella intersecta losca (Evans, 1951), Staphylus melangon epicaste Mabile, 1903 (Hesperiidae), Amphidecta reynoldsi Sharpe, 1890, Brevioleria seba emyra (Haensch, 1905), Callicore s. sorana (Godart, [1824]), Fountainea glycerium cratais (Hewitson, 1874), Malaveria mimula (Hayward, 1954), Paryphthimoides p. poltys (Prittwitz, 1865), “Pharneuptychia” innocentia (Felder & Felder, 1867), Yphthimoides celmis (Godart, [1824]) (Nymphalidae), Symbiopsis lenitas (Druce, 1907) (Lycaenidae), and Rhetus periander arthuriana (Sharpe, 1890) (Riodinidae).

The records of six species at Iguaçu NP represent the southernmost point of their respective previously documented distributions. These are: 1) Camissecla vesper (Druce, 1909), 2) Ipidecla crepundia (Druce, 1909), 3) Nesiostrymon tristis (Lathy, 1926) (Lycaenidae), 4) Joiceya praeclarus Talbot, 1928 (for more details see Greve et al. 2013GREVE, R.R., CALLAGHAN, C.J., KAMINSKI, L.A. & FREITAS, A.V.L. 2013. The rediscovery of Joiceya praeclarus Talbot 1928 (Lepidoptera: Riodinidae), more than 80 years after its description. J. Lepid. Soc. 67(1):56–57.), 5) Theope p. pakitza Hall & Harvey, 1998 (Riodinidae), and 6) Eunica m. malvina Bates, 1864 (Nymphalidae).

Among the sampled taxa, six were recorded for the first time in Brazil (Figure 4), namely: 1) Apaustus gracilis ssp. n., 2) Elbella v. viriditas (Skinner, 1920) (Hesperiidae), 3) Deltaya sp. n. (Nymphalidae), 4) Symbiopsis sp. n. (Lycaenidae), 5) Emesis orichalceus Stichel, 1916 and 6) Theope p. pakitza (Riodinidae). The new taxa of Apaustus, Deltaya and Symbopsis were discovered in this study.

Figure 4.
Taxa recorded for the first time in Brazil (except for the undescribed species of Deltaya): A) an undescribed subspecies of Apaustus gracilis, male; B) Elbella v. viriditas (Skinner, 1920), female (Hesperiidae); C) an undescribed species of Symbiopsis, female (Lycaenidae); D) Emesis orichalceus Stichel, 1916, male (left) and female (right); E) Theope p. pakitza Hall & Harvey, 1998, male (left) and hitherto unknown female (right) (Riodinidae). Scale bar = 1 cm.

Six taxa were included in at least one endangered faunal list (Mielke & Casagrande 2004MIELKE, O.H.H. & CASAGRANDE, M.M. 2004. Borboletas. In Livro vermelho da fauna ameaçada no Estado do Paraná (S.B. Mikich & R.S. Bérnils, orgs.). Instituto Ambiental do Paraná, Curitiba, p.713–739., ICMBio 2018aICMBIO 2018a. Livro vermelho da fauna brasileira ameaçada de extinção: Volume VII – Invertebrados 1 ed. Instituto Chico Mendes de Conservação da Biodiversidade, Brasília, D.F.): 1) Zonia zonia diabo Mielke & Casagrande, 1998, 2) Ochropyge ruficauda (Hayward, 1932), 3) Passova passova practa Evans, 1951 (Hesperiidae), 4) Narope cyllarus Westwood, 1851 (Nymphalidae), 5) Alesa prema (Godart, [1824]), and 6) Joiceya praeclarus (Riodinidae).

Discussion

1. Species richness

The 787 species recorded at Iguaçu NP and its surrounding areas represent 22.2% of the 3549 butterfly species known in Brazil (Casagrande & Duarte 2022CASAGRANDE, M.M. & DUARTE, M. 2022. Lepidoptera. In Catálogo Taxonômico da Fauna do Brasil. PNUD. http://fauna.jbrj.gov.br/fauna/faunadobrasil/84 (last access on 29/07/2023)
http://fauna.jbrj.gov.br/fauna/faunadobr...
). Except for the family Hedylidae, which was not included in previous Papilionoidea inventories in the Atlantic Forest, the other 786 species make Iguaçu NP the richest known site in butterfly species among areas with Semideciduous Forest. The park is also among the three richest sites in butterfly species of Atlantic Forest domain, together with the municipality of Linhares, in the state of Espírito Santo (835 species) (Brown Jr. & Freitas 2000BROWN, J.R., K.S. & FREITAS, A.V.L. 2000. Atlantic Forest butterflies: indicators for landscape conservation. Biotropica 32(4b):934–956.), and Itatiaia National Park, in the states of Rio de Janeiro and Minas Gerais (914 species) (Zikán & Zikán 1968ZIKÁN, J.F. & ZIKÁN, W. 1968. Inseto-fauna do Itatiaia e da Mantiqueira. III. Lepidoptera. Pesq. Agropec. Bras. 3:45–109.).

Among the records that may be added in the future to Iguaçu NP are the taxa that have been recorded exclusively at Iguazú National Park (hereafter Iguazú NP), Yacutinga Private Reserve (hereafter Yacutinga PR) and Surucuá Private Reserve (hereafter Surucuá PR), three neighboring protected areas in Argentina (Núñez-Bustos 2008NÚÑEZ-BUSTOS, E. 2008. Diversidad de mariposas diurnas en la Reserva Privada Yacutinga, Provincia de Misiones, Argentina (Lepidoptera: Hesperioidea y Papilionoidea). Trop. Lepid. Res. 18(2):78–87., 2009NÚÑEZ-BUSTOS, E. 2009. Mariposas diurnas (Lepidoptera: Papilionoidea y Hesperioidea) del Parque Nacional Iguazú, Provincia de Misiones, Argentina. Trop. Lepid. Res. 19(2):71–81., 2019NÚÑEZ-BUSTOS, E. 2019. Nuevos registros de mariposas diurnas para Misiones, Argentina (Lepidoptera: Papilionoidea), con notas toxonómicas. Historia Natural (Tercera Serie) 9(1):105–113., Klimaitis et al. 2018KLIMAITIS, J.F., NÚÑEZ-BUSTOS, E., KLIMAITIS, C.L. & GULLER, R.M. 2018. Mariposas de Argentina: Guía de identificación. Vazquez Mazzini Editores, Buenos Aires., Núñez-Bustos & Penco 2020NÚÑEZ-BUSTOS, E. & PENCO, F.C. 2020. Macrosoma hedylaria (Warren, 1894): Un Nuevo Registro de Hedylidae Guenée, 1858 de Misiones, Argentina (Lepidoptera: Papilionoidea). Historia Natural (Tercera Serie) 10(2):199–203., Núñez-Bustos in preparation). Together, these protected areas in Argentina and the Iguaçu NP in Brazil, form a practically continuous forest, composed by the same vegetal formation (except by MOF, which does not occur in the Argentinian parks), separated by the Iguaçu River. As such, it is reasonable to expect that these areas share a similar fauna. At least 77 species have been recorded only in the Argentinian parks, but all are likely to occur at Iguaçu NP. These records include 1 Papilionidae, 9 Lycaenidae, 8 Riodinidae, 15 Nymphalidae, 8 Pieridae, 35 Hesperiidae and 1 Hedylidae (Table 3), resulting in a list with 864 species of butterflies. Relevantly, D’Abrera (1987)D’ABRERA, B.L. 1987. Butterflies of the Neotropical Region. Part. IV. Nymphalidae (Partim). Hill House, Victoria. mentioned that a tourist recorded Prepona claudina annetta (Gray, 1832) (at the time, Agrias claudina claudianus Staudinger, [1885]) in February 1987 at the Argentinian Falls, so it is possible that this species occurs in the Brazilian park as well. Among other species with possible occurrence in Iguaçu NP, we cite the Papilionidae Pterourus cleotas (Gray, 1832) and the Nymphalidae Splendeuptychia tupinamba Freitas, Huertas & Rosa 2021. P. cleotas has specimens deposited in DZUP/UFPR from Cascavel, a municipality with approximately the same elevation and a few kilometers from Santa Tereza do Oeste, in the Northern border of Iguaçu NP. The species S. tupinamba was recently described and recorded at Yacutinga PR (Misiones, Argentina) (Rosa et al. 2021ROSA, A.H.B., HUERTAS, B., WILLMOTT, K.R., BARBOSA, E.P., MACHADO, P.A., MIELKE, O.H.H., CANAAN, C.H.P. & FREITAS, A.V.L. 2021. Fifty years without a name: a new species of Splendeuptychia Forster (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa 5061(1):95–114.).

Table 3.
Species that are likely to occur in Iguaçu NP and may be included in future lists, based on records from neighboring protected areas in Argentina (Iguazú NP, Yacutinga PR and Surucuá PR) (Núñez-Bustos 2008NÚÑEZ-BUSTOS, E. 2008. Diversidad de mariposas diurnas en la Reserva Privada Yacutinga, Provincia de Misiones, Argentina (Lepidoptera: Hesperioidea y Papilionoidea). Trop. Lepid. Res. 18(2):78–87., 2009NÚÑEZ-BUSTOS, E. 2009. Mariposas diurnas (Lepidoptera: Papilionoidea y Hesperioidea) del Parque Nacional Iguazú, Provincia de Misiones, Argentina. Trop. Lepid. Res. 19(2):71–81., 2019NÚÑEZ-BUSTOS, E. 2019. Nuevos registros de mariposas diurnas para Misiones, Argentina (Lepidoptera: Papilionoidea), con notas toxonómicas. Historia Natural (Tercera Serie) 9(1):105–113., Klimaitis et al. 2018KLIMAITIS, J.F., NÚÑEZ-BUSTOS, E., KLIMAITIS, C.L. & GULLER, R.M. 2018. Mariposas de Argentina: Guía de identificación. Vazquez Mazzini Editores, Buenos Aires., Núñez-Bustos & Penco 2020NÚÑEZ-BUSTOS, E. & PENCO, F.C. 2020. Macrosoma hedylaria (Warren, 1894): Un Nuevo Registro de Hedylidae Guenée, 1858 de Misiones, Argentina (Lepidoptera: Papilionoidea). Historia Natural (Tercera Serie) 10(2):199–203., Rosa et al. 2021ROSA, A.H.B., HUERTAS, B., WILLMOTT, K.R., BARBOSA, E.P., MACHADO, P.A., MIELKE, O.H.H., CANAAN, C.H.P. & FREITAS, A.V.L. 2021. Fifty years without a name: a new species of Splendeuptychia Forster (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa 5061(1):95–114., Núñez-Bustos in preparation). Species richness for each family are provided within parenthesis.

2. Taxonomic composition

Brown Jr. & Freitas (2000)BROWN, J.R., K.S. & FREITAS, A.V.L. 2000. Atlantic Forest butterflies: indicators for landscape conservation. Biotropica 32(4b):934–956. suggested that Nymphalidae generally represent 25% to 29% of the total species in a well-sampled inventory in the Neotropical region. At Iguaçu NP, the Nymphalidae make up 27.2% of the butterfly fauna, in full accord with predictions. Since the Nymphalidae can be satisfactorily documented with low sampling effort and their proportion in the butterfly fauna is relatively consistent, the richness of this family is an accurate surrogate for the total butterfly species richness. Alternatively, Robbins et al. (1996)ROBBINS, R.K., LAMAS, G., MIELKE, O.H.H., HARVEY, D.J. & CASAGRANDE, M.M. 1996. Taxonomic composition and ecological structure of the species-rich butterfly community at Pakitza, Parque Nacional del Manu, Perú. In Manu: The biodiversity of Southeastern Perú (D.E. Wilson & A. Sandoval, eds.). Smithsonian Institution Press, Washington, D.C., p.217–252. suggested that Neotropical faunas consist of approximately a third Papilionidae + Pieridae + Nymphalidae (255 species at Iguaçu NP), a third Lycaenidae + Riodinidae (192 species at Iguaçu NP), and a third Hesperiidae (339 species at Iguaçu NP). This prediction was not especially accurate at Iguaçu NP. Further, even though the proportional richness of Hesperiidae greatly exceeded a third, we consider it likely that most species to be discovered in the future will belong to this family. The reason is that they are mostly inconspicuous, small-sized and fast-flying species that make them difficult to sample (Brown Jr. 1992BROWN, J.R., K.S. 1992. Borboletas da Serra do Japi: Diversidade, habitats, recursos alimentares e variação temporal. In História natural da Serra do Japi: Ecologia e preservação de uma área florestal no sudeste do Brasil (L.P.C. Morellato, org.). Editora da Unicamp/FAPESP, Campinas, p.142–187., Brown Jr. & Freitas 1999BROWN, J.R., K.S. & FREITAS, A.V.L. 1999. Lepidoptera. In Biodiversidade do Estado de São Paulo: síntese do conhecimento ao final do século XX, invertebrados terrestres (C.R.F. Brandão & E.M. Cancello, eds.). USP, São Paulo, p.226–243.).

At greater detail, the higher species richness of Hesperiidae (43.1%) than Nymphalidae (27.2%) is consistent in well-detailed Neotropical inventories (Brown Jr. & Freitas 2000BROWN, J.R., K.S. & FREITAS, A.V.L. 2000. Atlantic Forest butterflies: indicators for landscape conservation. Biotropica 32(4b):934–956.) and may be a good indicator of “representativity” of butterfly sampling in the region (Francini et al. 2011FRANCINI, R.B., DUARTE, M., MIELKE, O.H.H., CALDAS, A. & FREITAS, A.V.L. 2011. Butterflies (Lepidoptera, Papilionoidea e Hesperioidea) of the “Baixada Santista” region, coastal São Paulo, southeastern Brazil. Rev. Bras. Entomol. 55(1):55–68.). Furthermore, the predominance of Lycaenidae over Riodinidae is another expected pattern in butterfly assemblages in the southern Atlantic Forest (Francini et al. 2011FRANCINI, R.B., DUARTE, M., MIELKE, O.H.H., CALDAS, A. & FREITAS, A.V.L. 2011. Butterflies (Lepidoptera, Papilionoidea e Hesperioidea) of the “Baixada Santista” region, coastal São Paulo, southeastern Brazil. Rev. Bras. Entomol. 55(1):55–68.) and seems to be related to a positive association between Riodinidae richness and local mean temperature (Brown Jr. 2005BROWN, J.R., K.S. 2005. Geological, evolutionary and ecological bases of the diversification of Neotropical butterflies: implications for conservation. In Tropical rainforests: Past, present and future (E. Bermingham, C.W. Dick & C. Moritz, eds.). University of Chicago Press, Chicago, p.166–201.). Assemblages in northern locations of the Atlantic Forest and in the Amazon Basin, accordingly, show greater riodinid species richness.

So far, there are no long-term butterfly inventories carried out in locations in the Atlantic Forest that include Hedylidae. Thus, we do not know the representativeness of this family in the total butterfly fauna in this domain. In the Brazilian butterfly fauna, Hedylidae represents 0.6% of the total (Lourido & Duarte 2023LOURIDO, G.M. & DUARTE, M. 2023. Hedylidae. in Catálogo Taxonômico da Fauna do Brasil. PNUD. http://fauna.jbrj.gov.br/fauna/faunadobrasil/1735 (last access on 25/05/2023)
http://fauna.jbrj.gov.br/fauna/faunadobr...
). Assuming that the same proportion applies to the Atlantic Forest, we could expect the occurrence of up to four additional species of this family in Iguaçu NP, suggesting that it may be underrepresented in the present inventory.

By using VSR traps, 27 species of fruit-feeding nymphalids were sampled (out of a total of 112 sampled from this guild), of which four were sampled exclusively by this method: Amphidecta pignerator simplicia Weymer, 1910, A. reynoldsi, Moneuptychia castrensis (Schaus, 1902), and Yphthimoides yphthima (Felder & Felder, [1867]). In the present study, VSR traps were used in a complementary way, and despite the low sampling effort employed in this method, it was effective in recording species that were not found in the active sampling with entomological nets. This may be related to the habits of these species (Freitas et al. 2021bFREITAS, A.V.L., SANTOS, J.P., ROSA, A.H., ISERHARD, C.A., RICHTER, A., SIEWERT, R.R., GUERATTO, P.E., CARREIRA, J.Y.O. & LOURENÇO, G.M. 2021b. Sampling methods for butterflies (Lepidoptera). In Measuring arthropod biodiversity: a handbook of sampling methods (J.C. Santos & G.W. Fernandes, eds.). Springer International Publishing, Cham, p.101–123.). For example, among the species that were sampled exclusively by traps, A. pignerator simplicia and A. reynoldsi have crepuscular habits, while Y. yphthima was sampled in a canopy trap.

Active searching for immature stages (especially on cold or rainy days, unfavorable for sampling adults) has also proven useful in adding species to the list. Whenever possible, the collected immatures were raised in the laboratory until the emergence of the adult for secure identification. The adults of some of these reared species were never seen in the field, such as the hesperid Thracides c. cleanthes (Latreille, [1824]), with eggs found on Syagrus romanzoffiana (Cham.) Glassman (Arecaceae); the nymphalid Opsiphanes cassiae crameri Felder & Felder, 1862, with a pupa found on exotic Heliconia rostrata Ruiz & Pav. (Heliconiaceae); the nymphalid Dynastor darius ictericus Stichel, 1904, whose pupa with signs of parasitoidism was found on Ananas bracteatus (Lindl.) Schult. & Schult.f. (Bromeliaceae); the riodinid Napaea eucharila (Bates, 1867), with larvae found on several Bromeliaceae species; and the riodinid Hyphilaria thasus (Stoll, 1780), with larvae found on several native and exotic Orchidaceae species, similar to that reported by Núñez-Bustos (2008)NÚÑEZ-BUSTOS, E. 2008. Diversidad de mariposas diurnas en la Reserva Privada Yacutinga, Provincia de Misiones, Argentina (Lepidoptera: Hesperioidea y Papilionoidea). Trop. Lepid. Res. 18(2):78–87., who raised a larva found on Octomeria pinicola Barb. Rodr. A remarkable case was that of the nymphalid Selenophanes cassiope guarany Casagrande, 1992, whose immature stages were previously unknown and could then be described (Shirai et al. 2016SHIRAI, L.T., BARBOSA, E.P., GREVE. R.R., MAGALDI, L.M., NASCIMENTO, A.R. & FREITAS, A.V.L. 2016. Natural history of Selenophanes cassiope guarany (Lepidoptera: Nymphalidae: Brassolini): an integrative approach, from molecules to ecology. Ann. Entomol. Soc. Am. 110(2):145–159.) after a female was observed ovipositing on leaves of S. romanzoffiana at dusk. Certain species may be more likely to be found in their immature stages for several reasons. Some spend most of their life cycle as immatures, especially those whose adults do not feed (e.g., Urich & Emmel 1991URICH, F.C. & EMMEL, T.C. 1991. Life histories of Neotropical butterflies from Trinidad 5. Dynastor darius darius (Lepidoptera: Nymphalidae: Brassolinae). Trop. Lep. 2(2):145–149.). Others have adults that fly mostly in the canopy or are active for a restricted time of day (DeVries 1997DEVRIES, P.J. 1997. The butterflies of Costa Rica. Vol. 2. Riodinidae. Princeton University Press, Princeton., Hall & Willmott 2010HALL, J.P.W. & K.R. WILLMOTT. 2010. Description of a new Lucillella species Riodinidae: Symmachiini) discovered in the eastern Andes of Ecuador using the single rope canopy access technique. J. Lepid. Soc. 64:139–146.). D. darius fits into more than one of these explanations. In the adult stage, this is a crepuscular species that lives relatively little time and does not feed, which is also why it is not sampled in VSR traps. In a study conducted in the Cerrado domain (Silva et al. 2011SILVA, N.A.P., DUARTE, M., DINIZ, I.R. & MORAIS, H.C. 2011. Host plants of Lycaenidae on inflorescences in the central Brazilian Cerrado. J. Res. Lepid. 44:95–105.), a survey focused on immature stages of Lycaenidae resulted in six new species records for a heavily sampled locality such as the Federal District (central Brazil), suggesting that this methodology has been underestimated in butterfly inventories.

Some taxa were only recorded above 500 meters elevation, where the montane subformation of SSF and its ecotone with MOF occur. These taxa include Amphidecta reynoldsi, Archaeoprepona chalciope (Hübner, [1823]), Diaethria e. eluina (Hewitson, [1855]), Dynamine meridionalis Röber, 1915, Eteona tisiphone (Boisduval, 1836), Heliconius besckei (Ménétriés, 1857), Cisandina lea (Cramer, 1777), Morpho aega (Hübner, [1822]), Opoptera a. aorsa (Godart, [1824]) (Nymphalidae), Mimoides protodamas (Godart, 1819), Neographium asius (Fabricius, 1781) (Papilionidae), Catocyclotis malca (Schaus, 1902), Emesis fastidiosa Ménétriés, 1855, Euselasia eusepus (Hewitson, [1853]), Euselasia zara (Westwood, 1851), Rhetus periander eleusinus Stichel, 1910 (Riodinidae), Gorgopas petale (Mabille, 1888), and Thespieus x. xarippe (Butler, 1870) (Hesperiidae). Although subspecies R. periander eleusinus and R. periander arthuriana fly together in other regions, in Iguaçu NP they were recorded separately – one individual of R. periander eleusinus in the highest part of the park whereas R. periander arthuriana was very common in low areas.

Among the recorded species, six were included in the Brazil Red Book of Threatened Species of Fauna (ICMBio 2018aICMBIO 2018a. Livro vermelho da fauna brasileira ameaçada de extinção: Volume VII – Invertebrados 1 ed. Instituto Chico Mendes de Conservação da Biodiversidade, Brasília, D.F.) and/or in the Paraná State Red Book of Threatened Species of Fauna (Mielke & Casagrande 2004)MIELKE, O.H.H. & CASAGRANDE, M.M. 2004. Borboletas. In Livro vermelho da fauna ameaçada no Estado do Paraná (S.B. Mikich & R.S. Bérnils, orgs.). Instituto Ambiental do Paraná, Curitiba, p.713–739.. These species are Zonia zonia diabo, Ochropyge ruficauda, Passova passova practa (Hesperiidae), Narope cyllarus (Nymphalidae), Alesa prema and Joiceya praeclarus (Riodinidae). Greve et al. (2013)GREVE, R.R., CALLAGHAN, C.J., KAMINSKI, L.A. & FREITAS, A.V.L. 2013. The rediscovery of Joiceya praeclarus Talbot 1928 (Lepidoptera: Riodinidae), more than 80 years after its description. J. Lepid. Soc. 67(1):56–57. suggested changing the conservation status of J. praeclarus to DD (“Data Deficient”) due to the new record for Foz do Iguaçu during the inventory. This record increased the species geographic distribution, which was previously known only from the type locality, in Mato Grosso, Brazil. In the case of Passova passova practa, Dolibaina et al. (2010)DOLIBAINA, D.R., CARNEIRO, E., DIAS, F.M.S., MIELKE, O.H.H. & CASAGRANDE, M.M. 2010. Registros inéditos de borboletas (Papilionoidea e Hesperioidea) ameaçadas de extinção para o Estado do Paraná, Brasil: novos subsídios para reavaliação dos critérios de ameaça. Biota Neotrop. 10(3):75–81 https://www.biotaneotropica.org.br/v10n3/pt/abstract?article+bn01210032010 (last access on 12/08/2021)
https://www.biotaneotropica.org.br/v10n3...
suggested removing the species from the Red List of Paraná, because new records of it in the study area and the abundance of larvae on host plant suggest that the rarity of this species is an adult sampling artifact. Z. zonia diabo and J. praeclarus were recorded only in surrounding areas of Iguaçu NP, in small patches of secondary forest, which suggests that these species may be more environmentally tolerant than previously thought. However, even though these species have not yet been detected inside the Iguaçu NP, they certainly also occur there, reinforcing the importance of this protected area for its conservation, because populations that occur in small forest fragments tend to be much more unstable and subject to local extinctions by stochastic factors or anthropogenic disturbances (Brown Jr. 1992)BROWN, J.R., K.S. 1992. Borboletas da Serra do Japi: Diversidade, habitats, recursos alimentares e variação temporal. In História natural da Serra do Japi: Ecologia e preservação de uma área florestal no sudeste do Brasil (L.P.C. Morellato, org.). Editora da Unicamp/FAPESP, Campinas, p.142–187..

Six species had their known distribution significantly increased after the present study: Camissecla vesper, Ipidecla crepundia, Nesiostrymon tristis (Lycaenidae), Joiceya praeclarus, Theope p. pakitza (Riodinidae), and Eunica m. malvina (Nymphalidae). The three species of Lycaenidae had previously been recorded in Rio de Janeiro state (Duarte et al. 2010DUARTE, M., ROBBINS, R.K., FREITAS, A.V.L., BROWN JR., K.S., MONTEIRO, R.F., CASAGRANDE, M.M. MIELKE, O.H.H., NASCIMENTO, M.S. & ALVES, T.G. 2010. Borboletas da Mata Atlântica do Estado do Rio de Janeiro: Lycaenidae (Lepidoptera). Arq. Mus. Nac. 67(3-4):291–302.). Theope p. pakitza had been known only from a Peruvian male (Hall 1998HALL, J.P.W. 1998. Six new species in the “foliorum group” of Theope (Lepidoptera: Riodinidae). Lambillionea 98(4):562–573.), and its female is illustrated here for the first time (Figure 4E). In the case of Eunica m. malvina, we recorded a female in March 2016 landed on a high leaf of Luehea divaricata Mart. & Zucc. (Malvaceae) at the margin of the Iguaçu River. Otherwise, in the Atlantic Forest this species occurs in the states of Bahia, Espírito Santo, Rio de Janeiro, Minas Gerais and São Paulo Paulo ( Jenkins 1990JENKINS, D.W. 1990. Neotropical Nymphalidae. VIII. Revision of Eunica. Bulletin of the Allyn Museum, Florida. , Santos et al. 2018SANTOS, J.P., FREITAS, A.V.L., BROWN JR., K.S., CARREIRA, J.Y.O, GUERATTO, P.E., ROSA, A.H.B., LOURENÇO, G.M., ACCACIO, G.M., UEHARA-PRADO, M., ISERHARD, C.A., RICHTER, A.G.K., ROMANOWSKI, H.P., MEGA, N.O., TEIXEIRA, M.O., MOSER, A., RIBEIRO, D.B., ARAUJO, P.F., FILGUEIRAS, B.K.C., MELO, D.H.A., LEAL, I.R., BEIRÃO, M.V., RIBEIRO, S.P., CAMBUÍ, E.C.B., VASCONCELOS, R.N., CARDOSO, M.Z., PALUCH, M., GREVE, R.R., VOLTOLINI, J.C., GALETTI, M., REGOLIN, A.L., SOUZA, T.S. & RIBEIRO, M.C. 2018. Atlantic butterflies: a data set of fruit feeding butterfly communities from the Atlantic forests. Ecology 99(12):2875.). In the case of the distributions previously documented for the riodinids J. praeclarus and T. p. pakitza, the increase was over 1000 km, showing that even in a well-studied group like butterflies, the Wallacean shortfall is still present.

Among the pairs of subspecies with sympatric distribution sampled, some produce intermediate forms, natural hybrids, such as the pairs nymphalids Heliconius ethilla polychrous Felder & Felder, 1865 x H. ethilla narcaea (Godart, 1819), Hypothyris euclea laphria (Doubleday, 1847) x H. euclea nina (Haensch, 1905), and Tithorea harmonia pseudethra Butler, 1873 x T. harmonia pseudonyma Staudinger, 1894. Such phenomenon is common where endemism zones meet, as shown by Brown Jr. (1982)BROWN, J.R., K.S. 1982. Historical and ecological factors in the biogeography of aposematic Neotropical butterflies. Am. Zool. 22:453–471.. However, other two pairs of subspecies, the hesperids Saturnus reticulata reticulata (Plötz, 1883) x S. reticulata conspicuus (Bell, 1941) and 8) Tisias lesueur lesueur (Latreille, [1824]) x T. lesueur canna Evans, 1955, actually correspond to distinct species (Mielke et al. in preparation). In other cases, such as the pairs riodinids Melanis aegates albugo (Stichel, 1910) x M. aegates limbata (Stichel, 1925) and Rhetus periander arthuriana (Sharpe, 1890) x R. periander eleusinus Stichel, 1910, further studies are needed to define the real status of these taxa. These pairs of sympatric subspecies recorded, as well as several taxa typical of other domains, is remarkable. Biogeographically, Iguaçu NP is situated in a transitional area between endemism zones (Brown Jr.& Mielke 1968BROWN, J.R., K.S. & MIELKE, O.H.H. 1968. Lepidoptera of the central Brazil plateau. III. partial list for the Belo Horizonte area, showing the character of the southeastern “blend zone”. J. Lepid. Soc. 22(3):147–157., Brown Jr. 1982BROWN, J.R., K.S. 1982. Historical and ecological factors in the biogeography of aposematic Neotropical butterflies. Am. Zool. 22:453–471.), which may help explain these findings. In addition, the riparian forest of the Paraná basin functions as an ecological corridor for species with a more northerly distribution, facilitating their dispersal to the region, which seems to be the southern limit of distribution for many butterfly species.

For those species that were only found as literature records or without vouchers, these records were not included. For example, there is a citation of Caligo b. brasiliensis (C. Felder, 1862) in an unpublished study, which was mentioned in a former version of the management plan of Iguaçu NP (IBAMA 1999IBAMA 1999. Plano de manejo do Parque Nacional do Iguaçu. Instituto Brasileiro de Meio Ambiente e dos Recursos Naturais Renováveis, Brasília, D.F.), but the species was not listed in the present study because the voucher specimen was not located. Also, all records for this species from iNaturalist were rejected because the available pictures did not correspond to natural observations in Foz do Iguaçu, but to individuals raised in captivity in the butterfly center of “Parque das Aves”, a private institution neighboring Iguaçu NP. Furthermore, the record of Taygetis virgilia (Cramer, 1776) (Nymphalidae) cited in IBAMA (1999)IBAMA 1999. Plano de manejo do Parque Nacional do Iguaçu. Instituto Brasileiro de Meio Ambiente e dos Recursos Naturais Renováveis, Brasília, D.F. was rejected because the voucher specimen could not be located at DZUP. However, there are records for this species in Iguazú National Park (Misiones, Argentina) (Núñez-Bustos 2009NÚÑEZ-BUSTOS, E. 2009. Mariposas diurnas (Lepidoptera: Papilionoidea y Hesperioidea) del Parque Nacional Iguazú, Provincia de Misiones, Argentina. Trop. Lepid. Res. 19(2):71–81.), so it is likely that future studies may record it in the Brazilian Park as well. Finally, four species of fruit-feeding nymphalids reported in Graciotim & Morais (2016)GRACIOTIM, C. & MORAIS, A.B.B. 2016. Borboletas frugívoras em florestas de mata atlântica do Parque Nacional do Iguaçu, Paraná, Brasil (Lepidoptera: Nymphalidae). SHILAP-Rev. Lepidopt. 44(173):115–128. were also not included, because: 1) the records of Taygetis sylvia Bates, 1866 and Yphthimoides ochracea (Butler, 1867) are possible misidentifications; and 2) based on nomenclatural updates (Espeland et al. 2023), Hermeuptychia hermes (Fabricius, 1775) and H. fallax (Felder & Felder, 1862) probably refer to H. atalanta (Butler, 1867) and H. gisella (Hayward, 1957), respectively.

Conclusion

Iguaçu NP and surrounding areas exhibit a heterogenous butterfly fauna with elements from different regions. Its species richness is high for a subtropical zone distant from the coast. Besides the three undescribed taxa recorded in this study, the number of other taxa originally described from specimens collected in the region, both in the Brazilian and Argentinian parks, is remarkable. Some examples are Ochropyge ruficauda, Atlides misma D’Abrera, 1995, Caeruleuptychia helena (Anken, 1994), and Agara pardalina yacutinga (Mielke & Casagrande, 2011). Furthermore, several unexpected records suggest that Iguaçu NP is a fertile field for new findings, such as the rediscoveries of Joiceya praeclarus and T. p. pakitza. The new records of these species represent a significant expansion in their geographic distributions, since J. praeclarus was known only from state of Mato Grosso in Brazil (over 1000 km) and had not been observed for 80 years, while T. p. pakitza was known only from Manú National Park in Peru (over 2000 km).

We recommend that future studies focus sampling efforts in MOF areas and their ecotone with SSF, which are the most preserved and least explored areas due to difficult accessibility. Considering the groups with high potential to represent new records for the park and its surroundings, we suggest that further studies aim at extensive collections using VSR traps for fruit-feeding Nymphalidae, the Ahrenholz technique for Hesperiidae, and light traps for mostly nocturnal Hedylidae. Due to the seasonality and short flight period of adults of certain species, some univoltine, these samplings must cover all months of the year, including during winter, in order to increase the chance of their detection.

Acknowledgments

RRG thanks the access to scientific collections allowed by Dayane Rossa and Adriane Guerino at Uniamérica Universitary Center, and Isabela da Costa Moreira at Itaipu Ecomuseum (technical reserve). This author is also thankful to all administration of Iguaçu NP, especially to Ivan Baptiston, Jorge Pegoraro, Pedro Fogaça, Apolônio Rodrigues, Raphael Xavier, Marina da Silva, Cíntia Mazon, Lucimara Frederico and Thais Oliveira. For assistance during fieldwork, RRG thanks Diego Dolibaina, Andrew Warren and Lucas Kaminski (and their help with species identification); Jessica Arruda and Thiago Farias (and their help with the Van Someren-Rydon traps); Márcia dos Anjos, Rafael Franco and Maximiano Duarte (and their help in many occasions). Ezequiel Núñez-Bustos has been in constant contact, sharing information and unpublished records for Argentine protected areas neighboring Iguaçu NP, in addition to fruitful discussions about the identity of several taxa that occur on both sides of the border. Roberto Leimig revised the first draft of this manuscript, giving valuable suggestions, as well as helping to identify several plant species. He also shared his knowledge about the plant physiognomies that occur in the Iguaçu NP and surrounding areas. For assistance in mounting numerous specimens, RRG thanks the collaborators of ZUEC/Unicamp. RRG also thanks Fabiane Annibale, who translated the manuscript and made several helpful suggestions. For their help with administrative matters, RRG thanks Luísa Mota, Augusto Rosa, Thiego Cotrim and Celso Paula Junior. For her valuable assistance in improving the quality of some of the figures that illustrate this paper, RRG thanks Valesca Ferreira. This author also thanks Chico Mendes Institute for Biodiversity Conservation (ICMBio) for the collection permit (SISBIO number 34730). AVLF thanks São Paulo Research Foundation (FAPESP 2012/50260-6, 2013/50297-0 and 2021/03868-8), Brazilian National Council for Scientific and Technological Development Pq-1A grant (CNPq 304291/2020-0), RedeLep-SISBIOTA-Brasil/CNPq (563332/2010-7) and National Science Foundation (DEB 1256742). We are all thankful for the assistance received to identify specimens: Márlon Paluch (Actinote), Fernando Dias (Fountainea, Memphis and Zaretis), Ricardo Siewert (Telemiades), Thamara Zacca, Eduardo Barbosa (several Euptychiina), Jorge Bizarro (Doxocopa) and Noemy Seraphim (Hermeuptychia). We are also grateful to Thamara Zacca and an anonymous reviewer for their careful reading of our manuscript and their many insightful comments and suggestions. Brazilian butterfly species are registered under SISGEN (A3EC5CC).

Data availability

A MS Excel file containing additional information on the taxa recorded from the literature and from entomological collections is available at: https://doi.org/10.48331/scielodata.NPDRGN

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Edited by

Associate Editor
Gustavo Graciolli

Publication Dates

  • Publication in this collection
    15 Sept 2023
  • Date of issue
    2023

History

  • Received
    08 Mar 2023
  • Accepted
    01 Aug 2023
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