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 1999, Lewinsohn et al. 2005, Santos et al. 2008). Inventory data help document distributions of species and decrease the Wallacean shortfall (Lomolino 2004, Whittaker et al. 2005). They provide data for studies in community ecology and biogeography (e.g., Robbins et al. 1996, Brown Jr. & Freitas 2000, Gonçalves-Souza et al. 2014, Zellweger et al. 2017). Scientifically undescribed taxa may be discovered (e.g., Biezanko & Mielke 1973, Núñez-Bustos 2008, Dolibaina et al. 2011, Lamas et al. 2021), thus decreasing the Linnean shortfall (Brown & Lomolino 1998). 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. 2003, Freitas et al. 2006, Santos et al. 2016).

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. 2009). 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. 2009) 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. 2008, Shirai et al. 2019), and its regional richness exceeds 2100 species (Brown Jr. & Freitas 2000). Despite these results, more information to better understand patterns of butterfly geographic distributions is needed (Francini et al. 2011, Iserhard et al. 2017).

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 2002). Mielke (1968) 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 1998). After that, Graciotim & Morais (2016) 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 2002, ICMBio 2018b). With further sampling, Santos et al. (2018) updated this list to 104 fruit-feeding nymphalid species. Considering the estimated richness of 700 butterfly species for Iguaçu NP (IBAMA 1999) and the 653 species recorded in Argentina’s Iguazú National Park (Núñez-Bustos 2009), 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 (1998); 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. 2013). 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. 2019). According to Google Earth (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 2018b): 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 2002, ICMBio 2018b) (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 2018b).

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. 2021b). 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), 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), 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.

Name	Phytophysiognomy	Present study	Mielke 1998	Municipality	Coordinates	Altitude
Pousada Eco Vine	MOF, OF	X		Santa Tereza do Oeste	25°03'S; 53°38'W	730 m
Research trail	MOF	X		Céu Azul	25°04'S; 53°39'W	750 m
Fazenda Rio Butu trail	MOF, FP	X		Céu Azul	25°05'S; 53°40'W	710 m
Linha Gonçalves Dias	MOF, OF	X		Santa Tereza do Oeste	25°06'S; 53°38'W	640 m
Abandoned quarry	SSFm	X		Céu Azul	25°06'S; 53°45'W	670 m
North of the municipality of Céu Azul	SSFm		X	Céu Azul	25°06'S; 53°46'W	680 m
Near the Gonçalves Dias River	MOF		X	Céu Azul	25°08'S; 53°39'W	600 m
Boa Vista trail	SSFm	X		Céu Azul	25°09'S; 53°47'W	650 m
Manoel Gomes trail	SSFm, FP	X	X	Céu Azul	25°10'S; 53°49'W	590 m
Jacutinga trail	SSFm	X		Céu Azul	25°14'S; 53°50'W	500 m
Middle course of the Floriano River	SSFm, FP		X	Céu Azul	25°17'S; 53°51'W	540 m
Benjamin Constant River	SSFs		X	Matelândia	25°23'S; 53°53'W	340 m
Bridge over the Silva Jardim River	OF		X	Matelândia	25°23'S; 53°56'W	280 m
Along the Pinheirinho River	SSFs		X	Matelândia	25°25'S; 53°54'W	290 m
Tributary of the Silva Jardim River	SSFs		X	Matelândia	25°26'S; 53°56'W	310 m
Along the Represa Grande River	SSFs		X	Serranópolis do Iguaçu	25°27'S; 54°07'W	220 m
Itaipu Binacional	SSFs, OF	X		Foz do Iguaçu	25°27'S; 54°36'W	120 m
Santa Maria Farm	SSFs, OF	X		Santa Terezinha de Itaipu	25°29'S; 54°22'W	330 m
Near the mouth of the Floriano River	SSFs	X	X	Céu Azul	25°30'S; 53°47'W	280 m
Estrada do Colono	SSFs	X	X	Serranópolis do Iguaçu	25°30'S; 54°01'W	270 m
Aparecidinha neighborhood	OF	X		Foz do Iguaçu	25°30'S; 54°30'W	270 m
Vietnã trail	SSFs, OF	X		Foz do Iguaçu	25°30'S; 54°34'W	200 m
Índio River	SSFs		X	São Miguel do Iguaçu	25°31'S; 54°16'W	210 m
Mouth of the Floriano River	SSFs	X		Capanema	25°32'S; 53°48'W	230 m
Linha Martins	SSFs	X		São Miguel do Iguaçu	25°32'S; 54°18'W	230 m
Estrada Velha de Guarapuava	SSFs, OF	X		São Miguel do Iguaçu	25°32'S; 54°21'W	270 m
Benjamin Constant River Waterfall	SSFs	X		Serranópolis do Iguaçu	25°33'S; 53°54'W	240 m
Cognópolis neighborhood	SSFs, OF	X		Foz do Iguaçu	25°33'S; 54°31'W	210 m
CITI Tênis	SSFs, OF	X		Foz do Iguaçu	25°33'S; 54°32'W	220 m
Iguaçu River lagoon	FP		X	Serranópolis do Iguaçu	25°34'S; 54°03'W	230 m
São João River basin	SSFs		X	Foz do Iguaçu	25°34'S; 54°25'W	220 m
Flora Éden	OF	X		Foz do Iguaçu	25°34'S; 54°30'W	200 m
Poço Preto	SSFs, FP	X	X	Foz do Iguaçu	25°35'S; 54°23'W	190 m
Cataratas Highway near Aquamania	OF	X		Foz do Iguaçu	25°35'S; 54°31'W	190 m
Marco das Três Fronteiras	SSFs, OF	X		Foz do Iguaçu	25°35'S; 54°35'W	140 m
Poço Preto trail	SSFs	X	X	Foz do Iguaçu	25°36'S; 54°26'W	240 m
Escola Parque trail	SSFs	X		Foz do Iguaçu	25°37'S; 54°28'W	210 m
Campinho trail	SSFs, OF	X		Foz do Iguaçu	25°37'S; 54°28'W	190 m
Represa trail	SSFs, FP	X		Foz do Iguaçu	25°37'S; 54°28'W	180 m
Canafístula trail	SSFs	X		Foz do Iguaçu	25°37'S; 54°28'W	190 m
Canzi Hotel trail	SSFs	X		Foz do Iguaçu	25°37'S; 54°29'W	170 m
Old power plant	SSFs	X		Foz do Iguaçu	25°37'S; 54°29'W	150 m
Monitoring trail	SSFs	X		Foz do Iguaçu	25°38'S; 54°26'W	250 m
Bananeiras trail	SSFs, OF	X		Foz do Iguaçu	25°39'S; 54°26'W	210 m
Macuco trail	SSFs	X		Foz do Iguaçu	25°39'S; 54°27'W	180 m
Higrômetro trail	SSFs	X		Foz do Iguaçu	25°41'S; 54°25'W	190 m
Cataratas trail	SSFs	X	X	Foz do Iguaçu	25°41'S; 54°26'W	180 m
Behind the Belmont Hotel	SSFs	X		Foz do Iguaçu	25°41'S; 54°26'W	220 m

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). 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. 1992, Canals 2003, D’Abrera 1984, 1987, 1989, 1994, 1995, Tyler et al. 1994, Klimaitis et al. 2018). We also compared specimens to images of type specimens of American butterflies, available in the Butterflies of America database, on internet (Warren et al. 2016), 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 1968, Jenkins 1990, Casagrande & Mielke 1992, Anken 1994, D’Abrera 1995, Mielke 1998, Hall & Harvey 2002, Callaghan 2010, Lourido 2011, Graciotim & Morais 2016) 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 iNaturalist (https://www.inaturalist.org), Google Images (https://www.google.com/imghp?hl=EN) and Flickr (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), 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 2014, Breinholt et al. 2018, Espeland et al. 2018). For higher taxonomy within families, we followed Lamas (2004), updated for Nymphalidae (Wahlberg et al. 2009), Riodinidae (Seraphim et al. 2018, Seraphim 2019), and Hesperiidae (Li et al. 2019, Zhang et al. 2019, Cong et al. 2019, Zhang et al. 2022). For generic names, we followed Mielke et al. (2022) and Brockmann et al. (2022) for the subfamily Pyrrhopyginae, Robbins et al. (2022) for the tribe Eumaeini and Espeland et al. (2023) for the subtribe Euptychiina. Taxonomy was also updated for the genera Morpho (Blandin 2007, Pablos et al. 2021), Eryphanis (Penz 2008), Zaretis (Dias et al. 2018), Phoebis (Murillo-Ramos et al. 2018), Agraulis and Dryas (Núñez et al. 2022).

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).

PAPILIONOIDEA (787)

PAPILIONIDAE (17)

Papilioninae (17)

Leptocircini (6)

Mimoides lysithous rurik (Eschscholtz, 1821)

Mimoides protodamas (Godart, 1819)

Neographium asius (Fabricius, 1781)

Protesilaus helios (Rothschild & Jordan, 1906)

Protesilaus protesilaus nigricornis (Staudinger, 1884)**

Protesilaus stenodesmus (Rothschild & Jordan, 1906)

Papilionini (5)

Heraclides anchisiades capys (Hübner, [1809])

Heraclides androgeus laodocus (Fabricius, 1793)

Heraclides astyalus astyalus (Godart, 1819)

Heraclides hectorides (Esper, 1794)

Heraclides thoas brasiliensis (Rothschild & Jordan, 1906)

Troidini (6)

Battus polydamas polydamas (Linnaeus, 1758)

Battus polystictus polystictus (Butler, 1874)

Parides agavus (Drury, 1782)

Parides anchises nephalion (Godart, 1819)

Parides bunichus perrhebus (Boisduval, 1836)**

Parides neophilus eurybates (Gray, [1853])

LYCAENIDAE (115)

Polyommatinae (3)

Hemiargus hanno (Stoll, [1790])

Leptotes cassius (Cramer, [1775])

Zizula cyna (Edwards, 1881)

Theclinae (112)

Eumaeini (112)

Allosmaitia strophius (Godart, [1824])

Arawacus dolylas (Cramer, [1777])

Arawacus ellida (Hewitson, 1867)

Arawacus meliboeus (Fabricius, 1793)

Arawacus separata (Lathy, 1926)

Arcas imperialis (Cramer, 1775)

Arzecla arza (Hewitson, 1874)

Arzecla ceromia (Hewitson, 1877)

Arzecla nubilum (Druce, 1907)**

Arzecla tucumanensis (Johnson & Kroenlein, 1993)

Atlides misma D’Abrera, 1995

Atlides polybe (Linnaeus, 1763)

Aubergina vanessoides (Prittwitz, 1865)

Badecla badaca (Hewitson, 1868)

Brangas getus (Fabricius, 1787)

Brangas moserorum Bálint & Faynel, 2008

Brangas neora (Hewitson, 1867)

Brevianta celelata (Hewitson, 1874)**

Caerofethra carnica (Hewitson, 1873)

Caerofethra eumorpha (Hayward, 1949)

Calycopis bellera (Hewitson, 1877)

Calycopis caulonia (Hewitson, 1877)

Calycopis gentilla (Schaus, 1902)

Calycopis sp.**

Camissecla vesper (Druce, 1909)

Celmia celmus (Cramer, [1775])

Chalybs chloris (Hewitson, 1877)

Chalybs hassan (Stoll, [1790])

Chlorostrymon simaethis (Drury, [1773])

Chlorostrymon telea (Hewitson, 1868)

Contrafacia catharina (Draudt, 1920)

Cupathecla cupentus (Stoll, [1781])

Cyanophrys acaste (Prittwitz, 1865)

Cyanophrys herodotus (Fabricius, 1793)

Cyanophrys pseudolongula (Clench, 1944)**

Cyanophrys remus (Hewitson, 1868)

Decussata empusa (Hewitson, 1867)

Denivia chaluma (Schaus, 1902)

Denivia deniva (Hewitson, 1874)**

Denivia hemon (Cramer, [1775])

Denivia lisus (Stoll, [1790])

Dicya dicaea (Hewitson, 1874)

Electrostrymon endymion (Fabricius, 1775)

Enos thara (Hewitson, 1867)

Erora biblia (Hewitson, 1868)

Erora sp.

Evenus latreillii (Hewitson, 1865)

Evenus regalis (Cramer, [1775])

Gargina gnosia (Hewitson, 1868)

Heterosmaitia marius (Lucas, 1857)

Heterosmaitia palegon (Cramer, [1780])

Heterosmaitia stagira (Hewitson, 1867)

Ignata brasiliensis (Talbot, 1928)

Ignata norax (Godman & Salvin, [1887])*

Ipidecla crepundia (Druce, 1909)

Ipidecla schausi (Godman & Salvin, [1887])

Janthecla aurora (Druce, 1907)

Janthecla rocena (Hewitson, 1867)

Kisutam syllis (Godman & Salvin, 1887)

Kolana chlamys (Druce, 1907)

Kolana ergina (Hewitson, 1867)

Kolana ligurina (Hewitson, 1874)

Laothus phydela (Hewitson, 1867)**

Lathecla mimula (Draudt, 1920)

Magnastigma hirsuta (Prittwitz, 1865)

Marachina asa (Hewitson, 1868)

Mercedes calus (Godart, [1824])

Michaelus ira (Hewitson, 1867)

Michaelus jebus (Godart, [1824])

Michaelus thordesa (Hewitson, 1867)

Ministrymon azia (Hewitson, 1873)

Ministrymon cruenta (Gosse, 1880)

Nesiostrymon calchinia (Hewitson, 1868)

Nesiostrymon tristis (Lathy, 1926)

Nicolaea torris (Druce, 1907)

Ocaria ocrisia (Hewitson, 1868)

Ocaria thales (Fabricius, 1793)

Ostrinotes sophocles (Fabricius, 1793)

Paiwarria aphaca (Hewitson, 1867)

Paiwarria venulius (Cramer, [1779])

Panthiades hebraeus (Hewitson, 1867)

Panthiades phaleros (Linnaeus, 1767)

Paraspiculatus orobia (Hewitson, 1867)

Parrhasius orgia (Hewitson, 1867)

Parrhasius polibetes (Stoll, [1781])

Pseudolycaena marsyas (Linnaeus, 1758)

Rekoa malina (Hewitson, 1867)

Rekoa meton (Cramer, [1779])

Semonina ares (Godman & Salvin, [1887])

Siderus eliatha (Hewitson, 1867)

Strephonota ambrax (Westwood, 1852)

Strephonota elika (Hewitson, 1867)*

Strephonota jactator (Druce, 1907)**

Strephonota tephraeus (Geyer, 1837)

Strymon astiocha (Prittwitz, 1865)

Strymon bazochii (Godart, [1824])

Strymon bubastus (Stoll, 1780)

Strymon cestri (Reakirt, 1867)

Strymon eurytulus (Hübner, [1819])

Strymon megarus (Godart, [1824])

Strymon mulucha (Hewitson, 1867)

Strymon rufofusca (Hewitson, 1877)

Strymon ziba (Hewitson, 1868)

Symbiopsis sp. n.#

Symbiopsis lenitas (Druce, 1907)

Symbiopsis strenua (Hewitson, 1877)

Theclopsis gargara (Hewitson, 1868)

Thepytus thyrea (Hewitson, 1867)

Theritas triquetra (Hewitson, 1865)*

Tmolus cydrara (Hewitson, 1868)

Tmolus echion (Linnaeus, 1767)

Ziegleria hesperitis (Butler & Druce, 1872)

RIODINIDAE (77)

Nemeobiinae (6)

Euselasiini (6)

Euselasia eucerus (Hewitson, 1872)

Euselasia eusepus (Hewitson, 1853)

Euselasia hygenius occulta Stichel, 1919

Euselasia marica Stichel, 1919

Euselasia satyroides Lathy, 1926

Euselasia zara (Westwood, 1851)

Riodininae (71)

Eurybiini (11)

Alesa prema (Godart, [1824])

Eurybia halimede passercula Stichel, 1915

Eurybia pergaea (Geyer, 1832)

Hyphilaria thasus (Stoll, 1780)

Ionotus alector (Geyer, 1837)

Leucochimona icare matatha (Hewitson, 1873)

Mesosemia friburguensis Schaus, 1902**

Mesosemia odice (Godart, [1824])

Mesosemia rhodia (Godart, [1824])

Napaea eucharila (Bates, 1867)

Perophthalma tullius (Fabricius, 1787)

Helicopini (1)

Anteros formosus (Cramer, [1777])

Emesidini (9)

Emesis cerea cerea (Linnaeus, 1767)

Emesis diogenia Prittwitz, 1865

Emesis fastidiosa Ménétriés, 1855

Emesis mandana (Cramer, 1780)

Emesis melancholica Stichel, 1916

Emesis neemias neemias Hewitson, 1872

Emesis ocypore ocypore (Geyer, 1837)

Emesis orichalceus Stichel, 1916#

Emesis russula Stichel, 1910

Nymphidiini (22)

Adelotypa bolena (Butler, 1867)

Ariconias glaphyra (Westwood, 1851)

Aricoris indistincta (Lathy, 1932)**

Aricoris monotona (Stichel, 1910)**

Aricoris signata (Stichel, 1910)

Catocyclotis malca (Schaus, 1902)

Catocyclotis sejuncta (Stichel, 1910)

Joiceya praeclarus Talbot, 1928

Juditha azan azan (Westwood, [1851])

Juditha molpe (Hübner, [1808])

Juditha odites praeclarum (Bates, 1866)

Nymphidium lisimon (Stoll, 1790)

Pseudolivendula hemileuca hemileuca (Bates, 1868)

Synargis calyce (Felder & Felder, 1862)

Synargis ethelinda (Hewitson, 1870)**

Synargis paulistina (Stichel, 1910)

Synargis aff. regulus (Fabricius, 1793)

Synargis regulus (Fabricius, 1793)

Teenie argiella (Bates, 1868)

Theope pakitza pakitza Hall & Harvey, 1998#

Theope terambus (Godart, [1824])

Theope thestias Hewitson, 1860

Riodinini (23)

Ancyluris rubrofilum Stichel, 1909

Barbicornis basilis mona Westwood, 1851

Calephelis aymaran McAlpine, 1971

Caria marsyas Godman, 1903

Caria plutargus plutargus (Fabricius, 1793)

Chalodeta theodora (Felder & Felder, 1862)

Chamaelimnas briola doryphora Stichel, 1910

Chorinea amazon amazon (Saunders, 1859)

Detritivora brasilia (Harvey & Hall, 2002)

Inkana incoides (Schaus, 1902)

Lasaia agesilas agesilas (Latreille, [1809])

Lasaia arsis Staudinger, 1888

Lasaia oileus Godman, 1903

Melanis aegates albugo (Stichel, 1910) x Melanis aegates limbata (Stichel, 1925)b

Melanis marathon charon (Butler, 1874)

Melanis xenia xenia (Hewitson, 1853)

Notheme erota angellus Stichel, 1910

Parcella amarynthina (Felder & Felder, [1865])

Pheles atricolor atricolor (Butler, 1871)

Rhetus periander arthuriana (Sharpe, 1890) x Rhetus periander eleusinus Stichel, 1910a

Riodina lycisca lycisca (Hewitson, [1853])

Riodina lysippoides Berg, 1882

Syrmatia nyx (Hübner, [1817])

Symmachiini (5)

Mesene aff. cyneus (Hewitson, 1874)

Mesene aff. simplex Bates, 1868

Mesene monostigma monostigma (Erichson, [1849])

Mesene pyrippe sanguilenta Stichel, 1910**

Pirascca sagaris satnius (Dalman, 1823)

NYMPHALIDAE (214)

Apaturinae (5)

Doxocopa agathina vacuna (Godart, [1824])

Doxocopa kallina (Staudinger, 1886)

Doxocopa laurentia laurentia (Godart, [1824])

Doxocopa linda mileta (Boisduval, 1870)

Doxocopa zunilda zunilda (Godart, [1824])

Biblidinae (40)

Ageroniini (7)

Ectima thecla thecla (Fabricius, 1796)F

Hamadryas amphinome amphinome (Linnaeus, 1767)F

Hamadryas arete (Doubleday, 1847)*F

Hamadryas epinome (Felder & Felder, 1867)F

Hamadryas februa februa (Hübner, [1823])F

Hamadryas feronia feronia (Linnaeus, 1758)F

Hamadryas fornax fornax (Hübner, [1823])F

Biblidini (2)

Biblis hyperia nectanabis (Frühstorfer, 1909)F

Mestra hersilia apicalis (Staudinger, 1886)**

Callicorini (8)

Callicore hydaspes (Drury, 1782)F

Callicore sorana sorana (Godart, [1824])F

Catagramma pygas thamyras Ménétriés, 1857F

Catagramma pyracmon pyracmon (Godart, [1824])F

Diaethria candrena candrena (Godart, [1824])F

Diaethria clymena janeira (C. Felder, 1862)F

Diaethria eluina eluina (Hewitson, [1855])F

Haematera pyrame pyrame Hübner, [1819]F

Catonephelini (9)

Catonephele acontius caeruleus Jenkins, 1985F

Catonephele numilia penthia (Hewitson, 1852)F

Cybdelis phaesyla (Hübner, [1831])**F

Eunica eburnea Frühstorfer, 1907F

Eunica eurota dolores (Prittwitz, 1871)**F

Eunica malvina malvina Bates, 1864F

Eunica margarita (Godart, [1824])F

Eunica tatila bellaria Frühstorfer, 1908F

Myscelia orsis (Drury, [1782])F

Epiphilini (5)

Epiphile hubneri Hewitson, 1861F

Epiphile orea orea (Hübner, [1823])F

Nica flavilla flavilla (Godart, [1824])F

Pyrrhogyra neaerea arge Gosse, 1880F

Temenis laothoe meridionalis Ebert, 1965F

Eubagini (9)

Dynamine aerata (Butler, 1877)

Dynamine agacles agacles (Dalman, 1823)

Dynamine artemisia artemisia (Fabricius, 1793)

Dynamine athemon athemaena (Hübner, [1824])

Dynamine coenus coenus (Fabricius, 1793)

Dynamine meridionalis Röber, 1915

Dynamine myrrhina (Doubleday, 1849)

Dynamine postverta postverta (Cramer, 1779)

Dynamine tithia tithia (Hübner, 1823)

Charaxinae (13)

Anaeini (8)

Consul fabius drurii (Butler, 1874)F

Fountainea glycerium cratais (Hewitson, 1874)F

Fountainea ryphea phidile (Geyer, 1837)F

Hypna clytemnestra huebneri Butler, 1866F

Memphis acidalia victoria (Druce, 1877)F

Memphis moruus stheno (Prittwitz, 1865)F

Zaretis hurin Dias, 2018F

Zaretis strigosus (Gmelin, [1790])F

Preponini (5)

Archaeoprepona chalciope (Hübner, [1823])F

Archaeoprepona demophon thalpius (Hübner, [1814])F

Archaeoprepona demophoon demophoon (Hübner, [1814])F

Prepona laertes (Hübner, [1811])F

Prepona pylene laertides Staudinger, 1898F

Cyrestinae (3)

Cyrestini (3)

Marpesia chiron marius (Cramer, [1779])

Marpesia petreus petreus (Cramer, [1776])

Marpesia zerynthia zerynthia Hübner, [1823]**

Danainae (24)

Danaini (5)

Danaus eresimus plexaure (Godart, 1819)*

Danaus erippus (Cramer, [1775])

Danaus gilippus gilippus (Cramer, [1775])

Lycorea halia discreta Haensch, 1909

Lycorea ilione ilione (Cramer, [1775])

Ithomiini (19)

Aeria olena olena Weymer, 1875

Brevioleria seba emyra (Haensch, 1905)

Callithomia lenea methonella (Weymer, 1875)

Dircenna dero celtina Burmeister, 1878

Episcada carcinia Schaus, 1902

Episcada hymenaea hymenaea (Prittwitz, 1865)

Epityches eupompe (Geyer, [1832])

Hypothyris euclea laphria (Doubleday, 1847) x Hypothyris euclea nina (Haensch, 1905)a

Ithomia agnosia zikani d’Almeida, 1940

Ithomia lichyi lichyi d’Almeida, 1939**

Mcclungia cymo salonina (Hewitson, 1855)

Mechanitis lysimnia lysimnia (Fabricius, 1793)

Methona themisto themisto (Hübner, 1818)

Placidina euryanassa (Felder & Felder, 1860)

Pseudoscada erruca (Hewitson, 1855)

Pteronymia carlia Schaus, 1902

Sais rosalia rosalinde Weymer, 1890

Thyridia psidii cetoides (Rosenberg & Talbot, 1914)

Tithorea harmonia pseudethra Butler, 1873 x Tithorea harmonia pseudonyma Staudinger, 1894a

Heliconiinae (20)

Acraeini (7)

Actinote carycina Jordan, 1913

Actinote discrepans d’Almeida, 1958

Actinote genitrix genitrix d’Almeida, 1922**

Actinote melanisans Oberthür, 1917

Actinote pellenea pellenea Hübner, [1821]

Actinote pyrrha pyrrha (Fabricius, 1775)

Actinote sp.

Argynnini (2)

Euptoieta hegesia meridiania Stichel, 1938

Euptoieta hortensia (Blanchard, 1852)

Heliconiini (11)

Agraulis maculosa (Stichel, [1908])

Dione juno juno (Cramer, 1779)

Dione moneta moneta Hübner, [1825]

Dryadula phaetusa (Linnaeus, 1758)

Dryas alcionea (Cramer, 1779)

Eueides aliphera aliphera (Godart, 1819)

Eueides isabella dianasa (Hübner, [1806])

Heliconius besckei (Ménétriés, 1857)

Heliconius erato phyllis (Fabricius, 1775)

Heliconius ethilla polychrous Felder & Felder, 1865 x Heliconius ethilla narcaea (Godart, 1819)a

Philaethria wernickei (Röber, 1906)**

Libytheinae (1)

Libytheana carinenta carinenta (Cramer, [1777])

Limenitidinae (18)

Limenitidini (18)

Adelpha abia (Hewitson, 1850)

Adelpha calliphane Frühstorfer, 1915

Adelpha epizygis epizygis Frühstorfer, 1915

Adelpha falcipennis Frühstorfer, 1915

Adelpha gavina Frühstorfer, 1915**

Adelpha iphicleola leucates Frühstorfer, 1915

Adelpha iphiclus iphiclus (Linnaeus, 1758)**

Adelpha lycorias lycorias (Godart, [1824])

Adelpha malea goyama Schaus, 1902

Adelpha melona pseudarete Frühstorfer, 1915

Adelpha mythra (Godart, [1824])

Adelpha plesaure phliassa (Godart, [1824])

Adelpha serpa serpa (Boisduval, [1836])

Adelpha syma (Godart, [1824])

Adelpha thesprotia (Felder & Felder, [1867])

Adelpha thessalia indefecta Frühstorfer, 1913

Adelpha thoasa gerona (Hewitson, 1867)

Adelpha zea (Hewitson, 1850)

Nymphalinae (22)

Coeini (1)

Historis odius dious Lamas, 1995F

Junoniini (1)

Junonia evarete evarete (Cramer, 1779)

Melitaeini (9)

Anthanassa hermas hermas (Hewitson, 1864)

Chlosyne lacinia saundersi (Doubleday, [1847])

Eresia lansdorfi (Godart, 1819)

Ortilia dicoma (Hewitson, 1864)

Ortilia ithra (Kirby, 1900)

Ortilia orthia (Hewitson, 1864)

Ortilia velica durnfordi (Godman & Salvin, 1878)

Tegosa claudina (Eschscholtz, 1821)

Telenassa teletusa teletusa (Godart, [1824])**

Nymphalini (7)

Colobura dirce dirce (Linnaeus, 1758)F

Hypanartia bella (Fabricius, 1793)

Hypanartia lethe (Fabricius, 1793)

Smyrna blomfildia blomfildia (Fabricius, 1781)F

Vanessa braziliensis (Moore, 1883)

Vanessa carye (Hübner, [1812])

Vanessa myrinna (Doubleday, 1849)

Victorinini (4)

Anartia amathea roeselia (Eschscholtz, 1821)

Anartia jatrophae jatrophae (Linnaeus, 1763)

Siproeta epaphus trayja Hübner, [1823]

Siproeta stelenes meridionalis (Frühstorfer, 1909)

Satyrinae (68)

Brassolini (17)

Blepolenis batea batea (Hübner, [1821])F

Brassolis sophorae vulpeculus Stichel, 1902

Caligo beltrao (Illiger, 1801)F

Caligo illioneus pampeiro Frühstorfer, 1904F

Catoblepia amphirhoe (Hübner, [1825])**F

Catoblepia berecynthia unditaenia (Frühstorfer, 1907)F

Dynastor darius ictericus Stichel, 1904

Eryphanis automedon automedon (Cramer, [1775])F

Eryphanis reevesii reevesii (Doubleday, [1849])F

Narope cyllarus Westwood, 1851**F

Narope cyllastros Doubleday, [1849]F

Narope panniculus Stichel, 1904F

Opoptera aorsa aorsa (Godart, [1824])F

Opsiphanes cassiae crameri Felder & Felder, 1862F

Opsiphanes invirae remoliatus Frühstorfer, 1907F

Opsiphanes quiteria meridionalis Staudinger, 1887F

Selenophanes cassiope guarany Casagrande, 1992F

Melanitini (1)

Manataria hercyna hercyna (Hübner, [1821])F

Morphini (6)

Antirrhea archaea Hübner, [1822]F

Morpho aega (Hübner, [1822])F

Morpho anaxibia (Esper, [1801])F

Morpho epistrophus titei (Le Moult & Réal, 1962)F

Morpho helenor achillides Felder & Felder, 1867F

Morpho portis thamyris (Felder & Felder, 1867)**F

Satyrini (44)

Amphidecta pignerator simplicia Weymer, 1910F

Amphidecta reynoldsi Sharpe, 1890F

Argentaria hygina (Butler, 1877)F

Argentaria libitina (Butler, 1870)F

Argentaria pagyris (Godart, [1824])F

Caeruleuptychia helena (Anken, 1994)**F

Capronnieria galesus (Godart, [1824])F

Carminda griseldis (Weymer, 1911)F

Carminda paeon (Godart, [1824])F

Cisandina lea (Cramer, 1777)F

Cissia eous (Butler, 1867)F

Cissia phronius (Godart, [1824])F

Deltaya ocypete (Fabricius, 1776)F

Deltaya pallema (Schaus, 1902)F

Deltaya sp. n.#F

Eteona tisiphone (Boisduval, [1836])F

Forsterinaria necys (Godart, [1824])F

Forsterinaria quantius (Godart, [1824])F

Godartiana muscosa (Butler, 1870)F

Hermeuptychia atalanta (Butler, 1867)F

Hermeuptychia gisella (Hayward, 1957)F

Malaveria affinis (Butler, 1867)F

Malaveria grimon (Godart, [1824])**F

Malaveria mimula (Hayward, 1954)F

Modestia sylvina (Felder & Felder, 1867)F

Moneuptychia castrensis (Schaus, 1902)F

Moneuptychia soter (Butler, 1877)F

Pareuptychia ocirrhoe interjecta (d’Almeida, 1952)F

Paryphthimoides poltys poltys (Prittwitz, 1865)F

Paryphthimoides terrestris grevei Zacca, Casagrande & Mielke 2020

“Pharneuptychia” innocentia (Felder & Felder, 1867)**F

Posttaygetis penelea (Cramer, [1777])F

Praepedaliodes phanias (Hewitson, 1862)F

Pseudodebis euptychidia (Butler, 1868)F

Taguaiba ypthima (Hübner, [1821])F

Taygetina kerea (Butler, 1869)F

Taygetis acuta Weymer, 1910**F

Taygetis laches (Fabricius, 1793)F

Taygetis rufomarginata rufomarginata Staudinger, 1888F

Taygetis tripunctata Weymer, 1907F

Yphthimoides celmis (Godart, [1824])F

Yphthimoides ordinaria Freitas, Kaminski & Mielke, 2012F

Yphthimoides yphthima (Felder & Felder, [1867])F

Zischkaia pacarus (Godart, [1824])F

PIERIDAE (24)

Coliadinae (14)

Anteos clorinde (Godart, [1824])

Anteos menippe (Godart, [1818])

Eurema albula sinoe (Godart, 1819)

Eurema deva deva (Doubleday, 1847)

Eurema elathea flavescens (Chavannes, 1850)

Phoebis argante argante (Fabricius, 1775)

Phoebis marcellina (Cramer, [1779])

Phoebis neocypris neocypris (Hübner, [1823])

Phoebis philea philea (Linnaeus, 1763)

Phoebis statira statira (Cramer, [1777])

Phoebis trite banksi (Breyer, 1939)

Phoebis wallacei wallacei (Felder & Felder, 1862)

Pyrisitia leuce leuce (Boisduval, 1836)

Pyrisitia nise tenella (Boisduval, 1836)

Dismorphiinae (5)

Dismorphia amphione astynome (Dalman, 1823)

Dismorphia astyocha Hübner, [1831]

Dismorphia thermesia thermesia (Godart, 1819)

Enantia lina psamathe (Fabricius, 1793)

Pseudopieris nehemia nehemia (Boisduval, 1836)

Pierinae (5)

Pierini (5)

Ascia monuste orseis (Godart, 1819) x Ascia monuste automate (Burmeister, 1878)a

Glutophrissa drusilla drusilla (Cramer, 1777)

Itaballia demophile ssp.*

Leptophobia aripa balidia (Boisduval, 1836)

Melete lycimnia paulista Frühstorfer, 1908

HESPERIIDAE (339)

Eudaminae (68)

Entheini (1)

Phanus australis Miller, 1965

Phocidini (11)

Bungalotis astylos (Cramer, 1780)

Bungalotis midas (Cramer, 1775)

Dyscophellus damias (Plötz, 1882)**

Nascus broteas (Cramer, 1780)

Nascus paulliniae (Sepp, [1842])

Nascus phocus (Cramer, [1777])

Phocides charon (Felder & Felder, 1859)

Phocides metrodorus metron Evans, 1952**

Phocides pigmalion hewitsonius (Mabille, 1883)

Phocides polybius phanias (Burmeister, 1880)

Phocides thermus valgus (Mabille, 1883)

Eudamini (54)

Aguna asander asander (Hewitson, 1867)

Aguna glaphyrus (Mabille, 1888)

Aguna megaeles megaeles (Mabille, 1888)

Aguna metophis (Latreille, [1824])

Aguna squamalba Austin & Mielke, 1998

Astraptes aulus (Plötz, 1881)

Astraptes enotrus (Stoll, [1781])

Astraptes janeira (Schaus, 1902)

Autochton neis (Geyer, 1832)*

Autochton reflexus (Mabille & Boullet, 1912)

Cecropterus dorantes (Stoll, [1790])

Cecropterus doryssus albicuspis (Herrich-Schäffer, 1869)

Cecropterus rica (Evans, 1952)

Cecropterus virescens (Mabille, 1877)

Cecropterus zarex (Hübner, 1818)

Cephise cephise (Herrich-Schäffer, 1869)

Chioides catillus catillus (Cramer, 1779)

Codatractus aminias (Hewitson, 1867)

Ectomis caunus (Herrich-Schäffer, 1869)

Ectomis octomaculata (Sepp, [1844])

Ectomis perniciosus (Herrich-Schäffer, 1869)

Epargyreus clavicornis clavicornis (Herrich-Schäffer, 1869)

Epargyreus exadeus (Cramer, [1780])

Epargyreus pseudexadeus Westwood, 1852

Epargyreus socus socus (Hübner, [1825])

Narcosius parisi (Williams, 1927)

Polygonus leo pallida Röber, 1925

Polygonus savigny savigny (Latreille, [1824])

Proteides mercurius mercurius (Fabricius, 1787)

Spathilepia clonius (Cramer, 1775)

Spicauda procne (Plötz, 1880)

Spicauda simplicius (Stoll, [1790])

Spicauda teleus (Hübner, 1821)

Telegonus alardus alardus (Stoll, 1790)

Telegonus anaphus anaphus (Cramer, [1777])

Telegonus chalco (Hübner, 1823)

Telegonus cretatus adoba (Evans, 1952)

Telegonus creteus siges (Mabille, 1903)

Telegonus elorus (Hewitson, 1867)**

Telegonus fulgor (Hayward, 1939)

Telegonus sp.

Telegonus talus (Cramer, [1777])

Telemiades aff. squanda Evans, 1953

Telemiades amphion (Geyer, 1832)

Telemiades antiope (Plötz, 1882)

Telemiades atlantiope Siewert, Mielke & Casagrande 2020

Telemiades brazus Bell, 1949

Telemiades laogonus (Hewitson, 1876)

Telemiades marpesus (Hewitson, 1876)

Urbanus esma Evans, 1952**

Urbanus esmeraldus (Butler, 1877)

Urbanus esta Evans, 1952

Urbanus pronta Evans, 1952

Urbanus proteus proteus (Linnaeus, 1758)

Oileidini (2)

Cogia calchas (Herrich-Schäffer, 1869)

Cogia undulatus (Hewitson, 1867)

Tagiadinae (1)

Celaenorrhinini (1)

Celaenorrhinus similis Hayward, 1933

Hesperiinae (187)

Pericharini (8)

Lycas argentea (Hewitson, [1866])

Orses cynisca (Swainson, [1821])

Oz ozias (Hewitson, 1878)

Perichares adela (Hewitson, 1867)

Perichares aurina Evans, 1955

Perichares lotus (Butler, 1870)

Perichares metallica (Riley, 1921)

Perichares seneca seneca (Latreille, [1824])

Hesperiini (179)

Adlerodea mineira Mielke, 1968**

Adlerodea modesta Hayward, 1940

Adlerodea petrovna (Schaus, 1902)

Adlerodea subpunctata subpunctata (Hayward, 1940)

Aides duma duma Evans, 1955

Alerema simplex (Bell, 1930)

Anatrytone perfida (Möschler, 1879)

Anthoptus epictetus (Fabricius, 1793)

Anthoptus insignis (Plötz, 1882)**

Apaustus gracilis ssp. n.#

Artines aepitus (Geyer, [1832])

Artines aquilina (Plötz, 1882)

Artines liege Medeiros, Mielke & Casagrande 2019

Artonia artona (Hewitson, 1868)

Callimormus diaeses Schaus, 1902**

Callimormus rivera (Plötz, 1882)**

Callimormus saturnus (Herrich-Schäffer, 1869)

Callimormus simplicius Hayward, 1939**

Calpodes esperi esperi (Evans, 1955)

Calpodes ethlius (Stoll, [1782])

Calpodes longirostris (Sepp, [1840])

Calpodes saladin catha (Evans, 1955)**

Calpodes salius (Cramer, [1775])

Cantha roraimae (Bell, 1932)

Carystoides basoches (Latreille, [1824])

Carystus hylaspes (Stoll, 1781)

Carystus lota (Hewitson, 1877)

Cobalopsis catocala (Herrich-Schäffer, 1869)

Cobalopsis cocalus (Hayward, 1939)

Cobalopsis nero (Herrich-Schäffer, 1869)

Cobalopsis obscurior (Hayward, 1934)

Cobalopsis valerius (Möschler, 1879)

Cobalus virbius virbius (Cramer, [1777])

Conga chydaea (Butler, 1877)

Conga iheringii (Mabille, 1891)

Contrastia distigma (Plötz, 1882)**

Corticea corticea (Plötz, 1882)

Corticea lysias potex Evans, 1955

Corticea sp.**

Cumbre meridionalis (Hayward, 1934)

Cyclosma altama (Schaus, 1902)

Cymaenes alumna (Butler, 1877)

Cymaenes cavalla Evans, 1955

Cymaenes gisca Evans, 1955

Cymaenes lepta (Hayward, 1939)

Cymaenes loxa Evans, 1955

Cymaenes tripunctata (Latreille, [1824])

Cynea bistrigula (Herrich-Schäffer, 1869)

Cynea cannae (Herrich-Schäffer, 1869)

Cynea diluta (Herrich-Schäffer, 1869)

Cynea irma (Möschler, 1879)

Cynea melius (Geyer, 1832)

Damas clavus (Herrich-Schäffer, 1869)

Decinea dama (Herrich-Schäffer, 1869)

Decinea denta pruda Evans, 1955

Dion uza uza (Hewitson, 1877)

Ebusus ebusus ebusus (Cramer, [1780])

Eprius obrepta (Kivirikko, 1936)

Eprius repens Evans, 1955

Euphyes leptosema (Mabille, 1891)

Euphyes subferrugineus subferrugineus (Hayward, 1934)

Eutocus vetulus matildae (Hayward, 1941)

Eutus mubevensis (Bell, 1932)

Eutychide olympia (Plötz, 1882)

Eutychide physcella (Hewitson, [1866])

Evansiella cordela (Plötz, 1882)

Gallio carasta (Schaus, 1902)

Gufa fusca (Hayward, 1940)

Haza hazarma (Hewitson, 1877)

Hedone vibex catilina (Plötz, 1886)

Hylephila phyleus phyleus (Drury, [1773])

Koria kora (Hewitson, 1877)

Lamponia elegantula (Herrich-Schäffer, 1869)

Lento krexoides (Hayward, 1940)

Lerema compta (Butler, 1877)

Lerema geisa (Möschler, 1879)

Lerodea eufala eufala (W. H. Edwards, 1869)

Lucida lucia (Capronnier, 1874)

Methionopsis cinnamomea (Herrich-Schäffer, 1869)

Methionopsis ina (Plötz, 1882)

Metrocles dejongi (Austin, 2008)

Metrocles hyboma (Plötz, 1886)

Metrocles sp.

Metron noctis (Kaye, 1914)

Metron oropa (Hewitson, 1877)

Mnaseas inca Bell, 1930

Mnasicles hicetaon Godman, [1901]**

Mnasicles remus (Fabricius, 1798)

Mnasitheus gemignanii (Hayward, 1940)

Mnasitheus submetallescens (Hayward, 1940)

Mnasitheus sucova (Schaus, 1902)**

Mnestheus silvaticus Hayward, 1940**

Moeris striga (Geyer, 1832)

Mucia zygia (Plötz, 1886)

Naevolus orius (Mabille, 1883)

Nastra celeus vetus (Mielke, 1969)

Neoxeniades scipio scipio (Fabricius, 1793)

Niconiades caeso (Mabille, 1891)

Niconiades linga Evans, 1955**

Niconiades merenda (Mabille, 1878)

Niconiades nikko Hayward, 1948**

Niconiades xanthaphes Hübner, [1821]

Nyctelius nyctelius nyctelius (Latreille, [1824])

Oligoria aff. locutia (Hewitson, 1876)

Oligoria aff. percosius (Godman, 1900)

Oligoria lucifer (Hübner, [1831])

Onophas columbaria distigma (Bell, 1930)

Orthos hyalinus (Bell, 1930)

Oxynthes corusca (Herrich-Schäffer, 1869)

Panoquina fusina viola Evans, 1955

Panoquina hecebolus (Scudder, 1872)

Panoquina lucas lucas (Fabricius, 1793)

Panoquina ocola ocola (Edwards, 1863)

Papias phainis Godman, 1900

Paracarystus evansi Hayward, 1938

Paracarystus hypargyra (Herrich-Schäffer, 1869)

Parphorus decora (Herrich-Schäffer, 1869)

Parphorus pseudecorus (Hayward, 1934)

Peba verames (Schaus, 1902)

Phemiades pohli pohli (Bell, 1932)

Pheraeus fastus (Hayward, 1939)

Pheraeus perpulcher (Hayward, 1934)

Phlebodes fuldai (Bell, 1930)

Polites premnas (Wallengren, 1860)

Pompeius amblyspila (Mabille, 1897)

Pompeius pompeius (Latreille, [1824])

Propapias sipariana (Kaye, 1925)

Psoralis arva (Evans, 1955)

Psoralis stacara (Schaus, 1902)

Psoralis umbrata (Erschoff, 1876)

Pyrrhopygopsis socrates socrates (Ménétriés, 1855)

Quasimellana eulogius (Plötz, 1882)

Quasimellana meridiani (Hayward, 1934)

Saturnus reticulata reticulata (Plötz, 1883) x Saturnus reticulata conspicuus (Bell, 1941)c

Saturnus saturnus saturnus (Fabricius, 1787)

Sodalia coler (Schaus, 1902)

Sodalia dimassa (Hewitson, 1876)

Sodalia sodalis (Butler, 1877)**

Synapte aff. silius (Latreille, [1824])

Synapte malitiosa antistia (Plötz, 1882)

Talides riosa Evans, 1955

Thargella caura occulta (Schaus, 1902)

Thargella evansi Biezanko & Mielke, 1973

Thespieus aspernatus Draudt, 1923

Thespieus dalman (Latreille, [1824])

Thespieus ethemides (Burmeister, 1878)

Thespieus lutetia (Hewitson, [1866])**

Thespieus xarippe xarippe (Butler, 1870)

Thracides cleanthes cleanthes (Latreille, [1824])

Tigasis arita (Schaus, 1902)

Tigasis perloides (Plötz, 1882)

Tigasis serra (Evans, 1955)

Tirynthia conflua (Herrich-Schäffer, 1869)

Tirynthoides virilis (Riley, 1929)

Tisias lesueur lesueur (Latreille, [1824])** x Tisias lesueur canna Evans, 1955c

Tricrista roppai (Mielke, 1980)**

Troyus diversa diversa (Herrich-Schäffer, 1869)

Troyus phyllides (Röber, 1925)

Turesis complanula (Herrich-Schäffer, 1869)

Vacerra bonfilius bonfilius (Latreille, [1824])

Vacerra caniola elva Evans, 1955

Vacerra evansi Hayward, 1938

Vehilius inca (Scudder, 1872)

Vehilius stictomenes stictomenes (Butler, 1877)

Vettius phyllus prona Evans, 1955**

Vidius similis Mielke, 1980

Vidius vidius (Mabille, 1891)

Vinius letis (Plötz, 1883)**

Vinius pulcherrimus Hayward, 1934

Vinius tryhana istria Evans, 1955

Virga austrinus (Hayward, 1934)

Virga silvanus (Hayward, 1947)

Vistigma bryanti (Weeks, 1906)

Vistigma xanthobasis (Hayward, 1939)

Xeniades chalestra corna Evans, 1955

Xeniades orchamus (Cramer, [1777])

Zariaspes mys (Hübner, [1808])

Zenis jebus jebus (Plötz, 1882)

Zenis minos (Latreille, [1824])**

Heteropterinae (1)

Heteropterini (1)

Dalla diraspes (Hewitson, 1877)

Pyrginae (62)

Achlyodini (10)

Achlyodes busirus rioja Evans, 1953

Aethilla echina coracina Butler, 1870

Eantis thraso (Hübner, [1807])

Milanion leucaspis (Mabille, 1878)

Ouleus fridericus riona Evans, 1953

Pythonides lancea (Hewitson, 1868)

Quadrus cerialis (Stoll, [1782])

Quadrus u-lucida mimus (Mabille & Boullet, 1917)

Zera hyacinthinus servius (Plötz, 1884)

Zera tetrastigma erisichthon (Plötz, 1884)

Carcharodini (20)

Bolla atahuallpai (Lindsey, 1925)

Bolla catharina (Bell, 1937)**

Gorgopas petale (Mabille, 1888)

Incisus incisus (Mabille, 1878)

Nisoniades bipuncta (Schaus, 1902)

Nisoniades castolus (Hewitson, 1878)

Nisoniades macarius (Herrich-Schäffer, 1870)

Nisoniades maura (Mabille & Boullet, 1917)

Pachyneuria inops (Mabille, 1877)

Pellicia costimacula costimacula Herrich-Schäffer, 1870

Pellicia dimidiata zamia Plötz, 1882**

Pellicia ranta rancida Evans, 1953**

Perus minor (Schaus, 1902)

Polyctor polyctor polyctor (Prittwitz, 1868)

Staphylus ascalon (Staudinger, 1876)

Staphylus chlorocephala (Latreille, [1824])

Staphylus insignis Mielke, 1980

Staphylus melangon melangon (Mabille, 1883)

Viola minor (Hayward, 1933)

Xispia satyrus (Jörgensen, 1935)

Erynnini (16)

Camptopleura auxo (Möschler, 1879)

Chiomara mithrax (Möschler, 1879)

Chiothion asychis autander (Mabille, 1891)

Cycloglypha thrasibulus thrasibulus (Fabricius, 1793)

Ebrietas anacreon anacreon (Staudinger, 1876)

Echelatus sempiternus simplicior (Möschler, 1877)

Festivia cronion (Felder & Felder, 1867)

Gesta gesta (Herrich-Schäffer, 1863)

Gorgythion begga begga (Prittwitz, 1868)

Gorgythion beggina escalophoides Evans, 1953

Grais stigmaticus stigmaticus (Mabille, 1883)

Helias phalaenoides palpalis (Latreille, [1824])

Hoodus pelopidas (Fabricius, 1793)

Mylon maimon (Fabricius, 1775)

Sostrata bifasciata bifasciata (Ménétriés, 1829)

Timochares trifasciata trifasciata (Hewitson, 1868)

Pyrgini (16)

Anisochoria sublimbata Mabille, 1883

Antigonus erosus (Hübner, [1812])

Antigonus nearchus (Latreille, [1817])

Burnsius orcus (Stoll, [1780])

Burnsius orcynoides (Giacomelli, 1928)

Canesia canescens pallida (Röber, 1925)

Heliopetes alana (Reakirt, 1868)

Heliopetes arsalte (Linnaeus, 1758)

Heliopetes libra Evans, 1944

Heliopetes ochroleuca Zikán, 1938

Heliopetes omrina (Butler, 1870)

Heliopetes petrus (Hübner, [1819])**

Paches liborius areta (Evans, 1953)

Spioniades artemides (Stoll, [1782])

Trina geometrina geometrina (Felder & Felder, [1867])

Xenophanes tryxus (Stoll, [1780])

Pyrrhopyginae (20)

Zoniini (1)

Zonia zonia diabo Mielke & Casagrande, 1998**

Passovini (6)

Agara epimachia edix (Evans, 1951)

Agara pardalina yacutinga (Mielke & Casagrande, 2011)

Granila paseas (Hewitson, 1857)**

Myscelus epigona Herrich-Schäffer, 1869

Passova passova practa Evans, 1951

Passova polemon (Hopffer, 1874)

Pyrrhopygini (13)

Blubella adonis (Bell, 1931)

Blubella azeta giffordi (Mielke, 1995)

Elbella intersecta losca Evans, 1951

Elbella lamprus lamprus (Hopffer, 1874)

Elbella viriditas viriditas (Skinner, 1920)#

Jematus gnetus brevipennis (Schaus, 1902)

Mysoria barcastus barta Evans, 1951

Ochropyge ruficauda (Hayward, 1932)**

Parelbella ahira extrema (Röber, 1925)

Pyrrhopyge attis subnubilus Hayward, 1935

Pyrrhopyge charybdis charybdis Westwood, 1852**

Pyrrhopyge pelota Plötz, 1879**

Santea antias (Felder & Felder, 1859)**

HEDYLIDAE (1)

Hedylinae (1)

Macrosoma zikani (Prout, 1932)**

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. 2014, Busby et al. 2017, Hall 2018) 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. 2008, Pinheiro et al. 2010, Freitas et al. 2021a) 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. 2013), 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 2004, ICMBio 2018a): 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 2022). 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 2000), and Itatiaia National Park, in the states of Rio de Janeiro and Minas Gerais (914 species) (Zikán & Zikán 1968).

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 2008, 2009, 2019, Klimaitis et al. 2018, Núñez-Bustos & Penco 2020, 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) 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. 2021).

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 2008, 2009, 2019, Klimaitis et al. 2018, Núñez-Bustos & Penco 2020, Rosa et al. 2021, Núñez-Bustos in preparation). Species richness for each family are provided within parenthesis.

Family	Species
Papilionidae (1)	Protesilaus telesilaus vitellus (Frühstorfer, 1907)
Lycaenidae (9)	Contrafacia imma (Prittwitz, 1865) Cyanophrys bertha (Jones, 1912) Kolana sp. n. Ministrymon una (Hewitson, 1873) Nicolaea cupa (Druce, 1907) Nicolaea opaliana (Hayward, 1967) Olynthus ophelia (Hewitson, 1867) Paraspiculatus hannelore (Bálint & Moser, 2001) Strephonota dindymus (Cramer, 1775) (cited as Strephonota sphinx (Fabricius, 1775))
Riodinidae (8)	Baeotis hisbon (Cramer, 1775) Chadia cadytis (Hewitson, 1866) Emesis fatimella fatimella Westwood, 1851 Eurybia misellivestis Stichel, 1910 Exoplisia aphanis (Stichel, 1910) Ithomiola orpheus (Westwood, 1851) Melanis smithiae smithiae (Westwood, 1851) Theope cmielkei Jauffret & Jauffret, 2009
Nymphalidae (15)	Actinote brylla Oberthür, 1917 Adelpha cytherea aea (Felder & Felder, 1867) Caligo martia (Godart, [1824]) Heterosais edessa (Hewitson, [1855]) Ithomia drymo Hübner, 1816 Memphis appias (Hübner, [1825]) Memphis otrere (Hübner, [1825]) Morpho hercules diadema Frühstorfer, 1905 Morpho menelaus coeruleus (Perry, 1810) Penetes pamphanis Doubleday, [1849] Pharneuptychia phares (Godart, [1824]) Phystis simois variegata (Hewitson, 1864) Splendeuptychia ambra (Weymer, [1911]) Splendeuptychia tupinamba Freitas, Huertas & Rosa 2021 Taygetis virgilia (Cramer, 1776)
Pieridae (8)	Colias lesbia lesbia (Fabricius, 1775) Enantia clarissa (Weymer, 1895) Eurema phiale paula (Röber, 1909) Ganyra phaloe endeis (Godart, 1819) Glennia pylotis (Godart, 1819) Hesperocharis paranensis paranensis Schaus, 1898 Perrhybris pamela eieidias Hübner, [1821] Pieriballia viardi molione (Frühstorfer, 1908)
Hesperiidae (35)	Anatrytone mella (Godman, 1900) Blubella blanda (Evans, 1951) Callimormus radiola pusillus Hayward, 1934 Carystus phorcus claudianus (Latreille, [1824]) Conga urqua (Schaus, 1902) Corticea noctis (Plötz, 1882) Cycloglypha caeruleonigra Mabille, 1903 Cynea popla Evans, 1955 Cynea robba nippa Evans, 1955 Diaeus variegata (Plötz, 1884) Dyscophellus porcius doriscus (Hewitson, 1867) Ectomis orphne (Plötz, 1881) Epargyreus tmolis (Burmeister, 1875) Jember menechmus menechmus (Mabille, 1878) Justinia maculata (Bell, 1930) Libra aligula decia (Hayward, 1948) Lindra brasus brasus (Mielke, 1968) Mahotis versicolor (Latreille, [1824]) Mimadia fallax fida (Evans, 1951) Nastra ethologus (Hayward, 1934) Nicephellus nicephorus (Hewitson, 1876) Nyctelius paranensis (Schaus, 1902) Olafia roscius roscius (Hopffer, 1874) Pellicia hersilia Hayward, 1939 Pellicia najoides Hayward, 1933 Phanes rezia (Plötz, 1882) Pheraeus odilia odilia (Plötz, 1884) Phlebodes campo campo (Bell, 1947) (misidentified as Thoon taxes Godman, 1900) Porphyrogenes vulpecula vulpecula (Plötz, 1882) Pseudosarbia phoenicicola Berg, 1897 Pythonides jovianus fabricii Kirby, 1871 Salatis salatis (Stoll, 1782) Staphylus melaina (Hayward, 1947) Staphylus musculus (Burmeister, 1875) Talides sergestus (Cramer, [1775])
Hedylidae (1)	Macrosoma hedylaria (Warren, 1894)

2. Taxonomic composition

Brown Jr. & Freitas (2000) 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) 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. 1992, Brown Jr. & Freitas 1999).

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 2000) and may be a good indicator of “representativity” of butterfly sampling in the region (Francini et al. 2011). Furthermore, the predominance of Lycaenidae over Riodinidae is another expected pattern in butterfly assemblages in the southern Atlantic Forest (Francini et al. 2011) and seems to be related to a positive association between Riodinidae richness and local mean temperature (Brown Jr. 2005). 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 2023). 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. 2021b). 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), 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. 2016) 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 1991). Others have adults that fly mostly in the canopy or are active for a restricted time of day (DeVries 1997, Hall & Willmott 2010). 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. 2011), 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 2018a) and/or in the Paraná State Red Book of Threatened Species of Fauna (Mielke & Casagrande 2004). 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) 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) 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).

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. 2010). Theope p. pakitza had been known only from a Peruvian male (Hall 1998), 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 1990 , Santos et al. 2018). 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). 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 1968, Brown Jr. 1982), 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 1999), 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) 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 2009), 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) 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