The butterflies (Lepidoptera, Papilionoidea) of the Parque Estadual Intervales and surroundings, São Paulo, Brazil

As borboletas (Lepidoptera, Papilionoidea) do Parque Estadual Intervales e arredores, São Paulo, Brasil

Abstract

The Global South has witnessed increasing sampling of its immense biological diversity during the past century. However, the diversity of many regions remains unknown, even at pristine and highly threatened places, such as in the Atlantic Forest; and for bioindicator, umbrella, and flagship groups. The present study reports the first butterfly list of the Parque Estadual Intervales, São Paulo, Brazil and surroundings, a key protected area in the last massive continuous of the Atlantic Forest. We compiled data from museums and four years of field work, under three sampling methods. We also aimed at providing resources to support conservation efforts by analyzing 27 years of climatic data (detailed in the Supplementary Material, in English and in Portuguese), discussing our results also for non-academics, and producing scientific outreach and educational material. A companion article dealt with the experiences of science outreach and capacity development, and illustrated the butterfly catalogue of the species sampled in the park. We found 312 species that sum to 2,139 records. The museum had 229 species (432 records), and we sampled 142 species (1,682 individuals), in a total effort of 36,679 sampling hours (36,432 trap and 247 net and observation hours). The richest families were Nymphalidae (148 species) and Hesperiidae (100 species). Most species were sampled exclusively by active methods (79.8%), but other sources (passive sampling, citizen science, etc.) also found unique records. We found the highest diversity metrics from January to May, and we demonstrated that winter months had less richness and abundance. We illustrated the 20 species common to all regions, and listed those that were found more than seven months in the year, as well as the most abundant species in trap sampling, with forest dwellers as well as species common to open and fragmented areas. The dominant species in our trap datasets was the iridescent white morpho, Morpho epistrophus (Fabricius, 1796), and we suggest it to become the park butterfly mascot.

Keywords
Species list; Paranapiacaba continuum; fish carrion bait; biodiversity knowledge shortfalls

Resumo

O Sul Global testemunhou crescente amostragem de sua imensa diversidade biológica durante o século passado. Entretanto, a diversidade de muitas regiões permanece desconhecida, mesmo em locais pristinos e altamente ameaçados, como na Mata Atlântica; e para grupos bioindicadores, guarda-chuva e emblemáticos. O presente estudo reporta a primeira lista de borboletas do Parque Estadual Intervales, São Paulo, Brasil e arredores, uma unidade de conservação chave no último maciço contínuo de Mata Atlântica. Compilamos dados de museus e quatro anos de campo, sob três métodos de amostragem. Também visamos oferecer recursos para apoiar os esforços de conservação, analisando 27 anos de dados climáticos (detalhados no Material Suplementar, em inglês e em português), discutindo nossos resultados numa linguagem também para não acadêmicos, e produzindo material de divulgação científica e educativos. Um artigo irmão tratou das experiências de divulgação científica e capacitação, e ilustrou o catálogo de borboletas das espécies amostradas no parque. Encontramos 312 espécies em 2.139 registros. O museu tinha 229 espécies (432 registros), e amostramos 142 espécies (1.682 indivíduos), em um esforço total de 36.679 horas de amostragem (36.432 armadilhas e 247 horas de rede e observação). As famílias mais ricas foram Nymphalidae (148 espécies) e Hesperiidae (100 espécies). A maioria das espécies foi amostrada exclusivamente por métodos ativos (79,8%), mas outras fontes (passiva, ciência cidadã, etc.) também encontraram registros únicos. Encontramos as maiores métricas de diversidade de janeiro a maio, e demonstramos que os meses de inverno tiveram menos riqueza e abundância. Ilustramos as 20 espécies comuns a todas as regiões, e listamos aquelas que foram encontradas em mais de sete meses no ano, bem como as espécies mais abundantes em armadilhas, com espécies florestais e também comuns em áreas abertas e fragmentadas. A espécie dominante em nossas armadilhas foi a morfo branca iridescente, Morpho epistrophus (Fabricius, 1796), e sugerimos que se torne a borboleta mascote do parque.

Palavras-chave
Lista de espécies; contínuo de Paranapiacaba; isca de peixe; deficit de conhecimento da biodiversidade

Introduction

Biodiversity studies are of central importance for the future, and inventories are one of their building blocks. Sampling distribution is, however, far from homogeneous, with most knowledge concentrated in the wealthiest, temperate, countries of the world (e.g.Girardello et al. 2019). The unequal sampling effort increases the relevance of carrying out inventories in the richest and megadiverse regions on Earth, usually found in the tropics, especially in those countries threatened by governments openly against science and the preservation of its environment (Myers et al. 2000, Alves et al. 2018, Andrade 2019, Angelo 2019, Tollefson 2019, Ferrante & Fearnside 2021, Hipólito et al. 2021). Despite an increase in sampling in the Global South during the past century (Girardello et al. 2019), it is remarkable that some regions with pristine natural environments remain unsampled, even at highly diverse and threatened regions, such as in the Atlantic Forest (Myers et al. 2000); and for biological groups that are relevant for conservation, such as bioindicator, umbrella, and flagship groups.

Bioindicator and umbrella groups are particularly useful for conservation planning and future attempts to understand what has been lost because they provide snapshots of the plant, animal or fungal species at a given time. The information they provide is more useful when inventories are performed according to best practices and standards. For example, to assess the fauna of a given location, it is important to use different sampling methods, to collect periodically (e.g. monthly) and for many years, to span different environments and altitudes, and to be careful with specimen and data curation (Santos et al. 2008, Shirai et al. 2019). That is rarely the case for several reasons, namely the financial and temporal availability of trained people, aggravated in places like South America (Elliot et al. 2018). However, in taxa that are easy to collect and identify even by non-specialists, approaches like citizen science and capacity development can be powerful allies to complement collection by scientists (Uehara-Prado et al. 2007, Davis 2015, Santos et al. 2016, Elliot et al. 2018, Mota et al. 2022, Shirai et al. 2022). It also helps when the bioindicator or umbrella group is aesthetically appealing, which is the case of most butterflies (Lepidoptera, Papilionoidea: Hesperiidae, Lycaenidae, Nymphalidae, Papilionidae, Pieridae, and Riodinidae).

Here we provide the first butterfly list of Intervales state park (Parque Estadual Intervales or PEI), a protected area located in the last massive continuous of the Atlantic Forest (Figure 1a). Despite the importance of this area, and associated environmental and social threats (see Methods), it is remarkable that we did not find any study sampling this bioindicator group, butterflies, although some works report the collection of focal taxa at the region (listed in Methods). The PEI boundaries were not determined for biological reasons, so we searched for inventories in the surrounding municipalities, all of which are in the state of São Paulo, Brazil (Figure 1a). The only published inventories near or at the PEI either collected all butterflies and did not report the species list (Brown & Freitas 1999) or had a species list but of a single tribe (Ithomiini, Shirai et al. 2017) or of a single guild on specific flowers (nectar-feeding butterflies on Lantana L. (Verbenaceae), Santos et al. 2015).

Figure 1.
Study area: (a) map of the Paranapiacaba Continuum: Intervales (PEI), Parque Estadual Carlos Botelho (PECB), Parque Estadual Turístico do Alto da Ribeira (PETAR), and Parque Estadual Nascentes do Paranapanema (PENaP), and Estação Ecológica Xitué (EEX); together with nearby Parque Estadual Cavernas do Diabo (PECD). The inset map shows the original distribution of the Atlantic Forest in light green and in dark green the remaining fragments, highlighting the importance of this Continuum, the largest continuous forest remanent; (b) map of sampling points by traps and net inside the PEI, showing some of its infrastructure; and representative images of canopy (above) and understory (below) traps of the PEI dataset: (c) “Minotauro” primary forest, (d) “Cidreira” secondary forest, and (e) “Guapiara” open road, with edge effect. (f) strategic traps set at PEI’s infrastructure for the PEI 1y dataset.

Figure 2.
Venn diagrams of (a) sampling methods (active versus passive), separating the sampling from the ZUEC museum and our study for the active search method; and (b) different regions (PEI, PENaP, AG) – the 20 species common to all regions illustrated in Figure 3.

We report the butterflies of the PEI and surroundings with data compiled from museums and four years of field work by us, under three sampling methods. Our aims were to contribute to the scientific community, but also to the local community, supporting conservation efforts in different ways. Here, we provide the butterfly list, discussing it academically but also in a language accessible for non-academics. We also analyzed 27 years of climatic data collected at the park but so far without any formal synthesis. This climatic profile can be reported by the protected area to support decision-makers to conserve the area with robust data, so we present it in the Supplementary Material both in English and in Portuguese. Some of the spread individuals returned to the park and are in display for visitors, along with informative posters with the results of this study (http://doi.org/10.5281/zenodo.5893810, press release available at https://www.infraestruturameioambiente.sp.gov.br/fundacaoflorestal/2022/02/parque-estadual-intervales-realiza-primeiro-plantio-coletivo-de-mudas/). Also, the “Butterflies of Intervales” outreach booklet is publicly available (http://doi.org/10.5281/zenodo.5068939). Finally, the experiences of science outreach and capacity development with the PEI staff, and an illustrated butterfly catalogue of species sampled here can be found in the companion publication (Shirai et al. 2022).

Materials and Methods

1. Study area

The Parque Estadual Intervales (hereafter PEI, 41,704 ha) is part of the largest continuous of the Atlantic Forest at the Serra do Mar (1,109,546 ha, Ribeiro et al. 2009, inset of Figure 1a). The PEI is located within the Paranapiacaba Continuum (ca. 140,000 ha c.f. decrees) which, along with the contiguous ecological station Xitué and Parque Estadual Carlos Botelho (PECB), Parque Estadual Turístico do Alto da Ribeira (PETAR), and Parque Estadual Nascentes do Paranapanema (PENaP) (Figure 1a), is a UNESCO Biosphere Reserve and a World Heritage Site. This Continuum plays a central role in conserving one of the five hottest world hotspots (cf.Myers et al. 2000), due to its unique extension of continuous forest. For example, the Continuum is one of the last three areas capable of hosting viable populations of the biome’s top predator, the jaguar (Panthera onca (Linnaeus, 1758)), which is vulnerable to extinction (ICMBio 2018) due to hunting and habitat loss (Beisiegel & Nakano-Oliveira 2020)

It is estimated that 85% of the PEI still has well preserved old growth forest with very low levels of disturbance (Nisi 2006), which is predominantly dense montane ombrophilous forest (Leonel 2010). Intervales means “between valleys,” in reference to the hills that range from 140 m to 1,200 m. The complex landscape with vast intact forest hosts important remnants of wild life, for instances, 9.2% of the PEI fauna of 532 species preliminarily inventoried in the 1990’s was threatened (Nisi 2006). Particularly, several arthropods in the Red List (ICMBio 2018) were found at the calcareous caves of the region, some exclusively at the PEI. The pristine forest, the caves and the almost 400 species of birds (an international hotspot for birdwatching) are among the main attractions of the park. Illegal hunting, mining, and overexploitation of juçara palm (Euterpe edulis Mart.) are among the main threats to the life it preserves (Shirai et al. 2022).

The climate of the region can be classified as dry-winter subtropical highland, or Cwb in the Köppen-Geiger system (c.f.Beck et al. 2018), with rainy summers (December to March) and a noticeable, dry, winter (June to September). The PEI has, however, different stations with distinct climatic profiles. At the PEI main station (ca. 800 m.a.s.l), the annual precipitation is of 1,400 mm, with the summer having an average precipitation of 187 ± 30 mm, and the wettest month (January) having 208 mm (WorldClim 2.5’ data from 1970–2000, for 24°15’56” S 48°24’48” W, 841 m a.s.l., which is the coordinate of the “reception” in Figure 1b; Fick & Hijmans 2017, www.worldclim.org/data/worldclim21.html, see also Figure 2 of Leonel 2010). During winter, the average precipitation is 66 ± 21 mm, with the driest month (August) having 49 mm (WorldClim op. cit.). At another PEI station in the lowland (ca. 150 m a.s.l.), at Saibadela, the picture can be very different as its nonseasonal tropical rainy climate reaches 4,000 mm annual precipitation, with the rainy season always with > 200 mm/month, frequently > 400 mm/month, and the driest period (May to August) with > 100 mm/month (Morellato et al. 2000).

We had access to 27 years of maximum and minimum temperature and relative humidity data from the weather station located near the PEI reception (Figure 1b). Data collection began in April 1992 and, although it is collected until the present time, we analyzed it until June 2019. The weather station was visited three times a day (08h, 14h, and 20h), every day, resulting in 28,990 measurements (9,663 at 08h, 9,676 at 14h, 9,651 at 20h), spread in 9,722 days of 320 months of 27 years (Tables S2–S4). More details of the data and its analyses are provided in the Supplementary Material. This data showed high correlation with the temperature and humidity data we measured at the PEI trap sampling (Table S6).

2. Butterfly data sampling

This study joins datasets from museums (Museu de Diversidade Biológica of the University of Campinas, ZUEC-LEP and the Zoology Museum of the University of São Paulo, MZUSP, having also consulted the Coleção Entomológica Padre Jesus Santiago Moure, Zoology Department of the Federal University of Paraná, DZUP; and the National Museum at Rio de Janeiro, MNUFRJ) and sampling by the authors (referred by our initials) from different locations, under different methods.

In the Supplementary Material we describe the material and methods of each dataset (summarized in Table 1), including coordinates, method(s), collectors, specialists who identified the material, aim, sampling effort, and design. We organized them as follows: 1. museum data, and 2. sampling data by us (with PEI species fully illustrated in Shirai et al. 2022). We divided the museum data in: 1.1. PEI ZUEC dataset, and 1.2. PENaP dataset (Parque Estadual Nascentes do Paranapanema) – both datasets used the entomological net as the sampling method (communicated by the collectors). We divided the sampling data in: 2.1. PEI dataset; 2.2. PEI fish dataset (carrion bait test in the Atlantic Forest); 2.3. PEI 1y dataset (one-year monitoring for capacity development and science outreach); 2.4. AG dataset (Apiaí and Guapiara sampling); and 2.5. Other.

Table 1.
Summary of each dataset, with information of when they were sampled (month and year), how (method and details of trap studies), and sampling effort. Museum data: PEI ZUEC (Parque Estadual Intervales in the ZUEC collection) and PENaP (Parque Estadual Nascentes do Paranapanema). Sampled in this study: PEI, PEI fish (carrion bait test in the Atlantic Forest), PEI 1y (one-year monitoring for capacity development and science outreach), and AG (Apiaí and Guapiara). Full sampling data is in the Supplementary Material.

Dataset	Months	Year(s)	Sampling effort	Method(s)	No. Traps	Bait	Stratum(a)
PEI ZUEC	I, II, IV, VIII, XI, XII	1992, 2000–2003, 2007–2010, 2013	NA	net	NA	NA	NA
PENAP	III, IV	2012	NA	net	NA	NA	NA
PEI	III, IV, VI, VII	2016–2018	5,760 trap h; 76 net h	net, trap	30	banana	understory, canopy
PEI fish	III, XII	2018	3,456 trap h; 15 net h	trap	36	banana x fish	understory, canopy
PEI 1y	I to XII	2019	4,752 trap h	trap	11	banana	understory
AG	all but I, IV, VI, X	2014, 2017–2019	22,464 trap h; 156 net/obs h	net, trap, obs	48	banana	understory

Our sampling was carried out in the following periods: 2.1. PEI dataset (illustrated in Shirai et al. 2022) at 14 to 18.III.2016 (pilot), 06 to 13.III.2017, 07 to 12.IV.2017, 11 to 16.VI.2017, 21 to 23.VII.2017, 10 to 14.IV.2018, 26 to 29.VII.2018 with sampling effort of 5,760 trap hours (16 days * 30 traps * 12 trap hours/day), and ca. 76 net hours; 2.2. PEI fish dataset (07 to 12.III.2018 and 09 to 15.XII.2018) with sampling effort of 3,456 trap hours (8 days * 36 traps * 12 trap hours/day), and ca. 15 net hours; 2.3. PEI 1y dataset 14 to 17.I.2019, 18 to 21.II.2019, 18 to 21.III.2019, 15 to 18.IV.2019, 27 to 30.V.2019, 17 to 20.VI.2019, 16 to 19.VII.2019, 20 to 23.VIII.2019, 17 to 20.IX.2019, 14 to 17.X.2019, 25 to 28.XI.2019, 09 to 12.XII.2019 with sampling effort of 4,752 trap hours (36 days * 11 traps * 12 trap hours/day); and 2.4. AG dataset XII.2014, 06 to 19.VIII.2017, 18 to 27.XI.2017, 17 to 26.II.2018, 17 to 25.V.2018, 25.VIII to 03.IX.2018, 14 to 23.XII.2018, 08 to 17.III.2019, 02 to 12.VII.2019, 19 to 29.IX.2019 with sampling effort of 22,464 trap hours (39 days * 48 traps * 12 trap hours/day) and 156 hours of active search.

We classified species in subfamilies and used the phylogenetic hypotheses of Warren et al. (2008, 2009), Wahlberg et al. (2009, 2014), Seraphim et al. (2018), Li et al. (2019), and Dias et al. (2019). We based the identification of specimens on the specialized literature (e.g.Brown 1992, Uehara-Prado et al. 2004, Warren et al. 2016, Lamas 2022), and on the collections of the ZUEC-LEP and MZUSP. These are also the entomological collections where we housed the specimens collected by us, with material from LTS at Unicamp (ZUEC–LEP 10.018–10.070, 10.253–10.262, 11.312–11.445) and ROS at MZUSP. Part of the Unicamp material (~10%) has a pair of legs stored in EtOH and is available to the scientific community upon request.

We searched for information about the butterflies of the region by looking at the literature, asking specialists, and consulting a database with the state of the art of species lists in Brazil (Shirai et al. 2019). We found only three studies (Brown & Freitas 1999, Santos et al. 2015, Shirai et al. 2017), none of which had a species list for more than a tribe. Other studies reported the collection of focal taxa at the PEI (e.g. Forsterinaria pronophila (A. Butler, 1867) in Freitas & Peña 2006, Blepolenis bassus (C. Felder & R. Felder, 1867) in Penz et al. 2013, Taygetis ypthima (Hübner, [1821]) and T. rectifascia (Weymer, 1907) in Siewert et al. 2013, Godartiana muscosa (A. Butler, 1870) in Zacca et al. 2016, T. acuta Weymer, 1910 in Freitas 2017, T. yphtima in Uehara-Prado & Freitas 2019), and the referred specimens belong to ZUEC-LEP and ZUEC-AVLF private collection.

Except from ZUEC-LEP, other large butterfly collections in the country did not have material from the PEI: MZUSP (ROS pers. obs.); DZUP (O. H. H. Mielke, pers. comm.); the MNUFRJ probably did not have material from the region but even if it had, it would be reduced to ashes due to the fire in September 2018. In the 30 years and 300 research projects in the park, no foreign institution has had an official project that collected butterflies (T. B. Conforti, pers. comm.). Therefore, the museum data reported below belonged solely to ZUEC-LEP (now MZUSP has sampled material by ROS, and DZUP and MNRJ entomological collections also house PEI specimens donated by us).

We consulted ZUEC-LEP by the online database available at the SpeciesLink website (www.splink.org.br) at the very beginning and end of this study, filtering by the collection (ZUEC-LEP) and the municipalities of and around PEI (Capão Bonito, Eldorado, Guapiara, Iporanga, Ribeirão Grande, Sete Barras), returning results only for Ribeirão Grande and Sete Barras. We searched the Nymphalidae specimens with missing IDs in the database and checked every drawer of the collection to update the online database by including their identity.

3. Data treatment

We counted the number of individuals found at the ZUEC-LEP, those sampled by us under active searches and other sources (here referred as records), as well as under baited traps designs (here referred as abundance). The number of records and abundance are not comparable because specimens deposited in a museum, sampled with the net by different observers, or observational records (such as citizen science) do not reflect biological abundance. We also counted the number of species (richness), excluding genera not identified to the species level due to taxonomic issues (e.g., Hermeuptychia sp. Forster, 1964, Tan et al. 2021) or museum specimens identified as such (e.g., Actinote sp. Hübner, [1819], Adelpha sp. Hübner, [1819], etc); except from Autochton sp. Hübner, 1823, Ochlodes sp. Scudder, 1872, Emesis sp. Fabricius, 1807, and Mesene sp. E. Doubleday, 1847 because no other species of the same genus were in the list.

We reported the species lists by dataset (Table 2) and explored how many and which species belong to each dataset, or are shared among datasets, with Venn diagrams (using online tool Venny, https://bioinfogp.cnb.csic.es/tools/venny/). We also looked at the species assembly (presence/absence) by sampling method and by region. We then collapsed datasets in a single list to analyze richness (whole data, as well as PEI traps: PEI trap, PEI fish, and PEI 1y datasets) and abundance (only PEI traps) per month and per season. For PEI traps, we separated the lists by datasets and divided the diversity metrics by the number of traps and trap days.

Table 2.
Butterfly species of PEI and surroundings per dataset. Museum data: PEI ZUEC (Parque Estadual Intervales in the ZUEC collection) and PENaP (Parque Estadual Nascentes do Paranapanema). Sampled in this study: PEI, PEI fish (carrion bait test in the Atlantic Forest), PEI 1y (one-year monitoring for capacity development and science outreach), and AG (Apiaí and Guapiara). “Other” refers to citizen science, iNaturalist and Santos et al. (2015) sources. Numbers are either the number of records or the abundance (only for standardized method of baited traps), while “x” denotes presence. In the “Month caught” column, PENaP did not have specify collection dates, only III–IV. This table is available at 10.5281/zenodo.7429126.

Family	Subfamily	Tribe	Taxon	Species description	PEI ZUEC	PENaP	PEI	AG	PEI	PEI fish	PEI 1y	AG	other	Month caught
					entomological net				baited traps
Hesperiidae	Eudaminae		Autochton sp.	NA	x									NA
Hesperiidae	Eudaminae		Cecropterus albimargo	(Mabille, 1876)	1									I
Hesperiidae	Eudaminae		Cecropterus zarex	(Hübner, 1818)	2		1							I, III
Hesperiidae	Eudaminae		Oechydrus chersis	(Herrich-Schäffer, 1869)			1							IV
Hesperiidae	Eudaminae		Perichares seneca	(Latreille, [1824])			1							VII
Hesperiidae	Eudaminae		Phanus australis	L. Miller, 1965	2									I
Hesperiidae	Eudaminae		Phocides pialia	(Hewitson, 1857)	x									NA
Hesperiidae	Eudaminae		Polygonus savigny	(Latreille, [1824])	1									I
Hesperiidae	Eudaminae		Spicauda procne	(Plötz, 1881)			1							III
Hesperiidae	Eudaminae		Spicauda simplicius	(Stoll, 1790)			2							III, IV
Hesperiidae	Eudaminae		Spicauda teleus	(Hübner, 1821)		4								III–IV, IV
Hesperiidae	Eudaminae		Telegonus alardus	(Stoll, 1790)	1									I
Hesperiidae	Eudaminae		Typhedanus aziris	(Hewitson, 1867)	1									XII
Hesperiidae	Eudaminae		Typhedanus stylites	(Herrich-Schäffer, 1869)			1			1				XII
Hesperiidae	Eudaminae		Urbanus esmeraldus	(A. Butler, 1877)	2									I
Hesperiidae	Eudaminae		Urbanus pronta	Evans, 1952			3							III
Hesperiidae	Hesperiinae		Anthoptus epictetus	(Fabricius, 1793)	3									XII
Hesperiidae	Hesperiinae		Callimormus rivera	(Plötz, 1882)	2									XII
Hesperiidae	Hesperiinae		Carystoides sicania	(Hewitson, 1876)	x									NA
Hesperiidae	Hesperiinae		Cobalopsis brema	E. Bell, 1959	1									I
Hesperiidae	Hesperiinae		Cobalopsis vorgia	(Schaus, 1902)	1									I
Hesperiidae	Hesperiinae		Conga immaculata	(E. Bell, 1930)			1							XII
Hesperiidae	Hesperiinae		Corticea lysias	Evans, 1955	x									NA
Hesperiidae	Hesperiinae		Corticea oblinita	(Mabille, 1891)			1							III
Hesperiidae	Hesperiinae		Decinea dama	(Herrich-Schäffer, 1869)						1				III
Hesperiidae	Hesperiinae		Decinea decinea	(Hewitson, 1876)	1									I
Hesperiidae	Hesperiinae		Enosis schausI	O. Mielke & Casagrande, 2002	1									I
Hesperiidae	Hesperiinae		Eutocus matildae	(Hayward, 1941)	x									NA
Hesperiidae	Hesperiinae		Eutychide physcella	(Hewitson, 1866)	x									NA
Hesperiidae	Hesperiinae		Lamponia lamponia	(Hewitson, 1876)	x									NA
Hesperiidae	Hesperiinae		Lerema duroca	Evans, 1955	x									NA
Hesperiidae	Hesperiinae		Levina levina	(Plötz, 1884)	1									XII
Hesperiidae	Hesperiinae		Libra aligula	(Schaus, 1902)	2									II
Hesperiidae	Hesperiinae		Lucida lucia	(Capronnier, 1874)	x									NA
Hesperiidae	Hesperiinae		Lucida ranesus	(Schaus, 1902)	1									II
Hesperiidae	Hesperiinae		Lucida schmithI	(E. Bell, 1930)	1									XII
Hesperiidae	Hesperiinae		Lychnuchoides ozias	(Hewitson, 1878)	x									NA
Hesperiidae	Hesperiinae		Lychnuchus celsus	(Fabricius, 1793)	1	3								I, IV
Hesperiidae	Hesperiinae		Metron oropa	(Hewitson, 1877)	3									II, XII
Hesperiidae	Hesperiinae		Miltomiges cinnamomea	(Herrich-Schäffer, 1869)	1	3	4							I, III, IV, VII
Hesperiidae	Hesperiinae		Mnasitheus ritans	(Schaus, 1902)			3							IV, VII, XII
Hesperiidae	Hesperiinae		Moeris remus	(Fabricius, 1798)		1								IV
Hesperiidae	Hesperiinae		Naevolus orius	(Mabille, 1883)	x									NA
Hesperiidae	Hesperiinae		Nastra lurida	(Herrich-Schäffer, 1869)	1									XII
Hesperiidae	Hesperiinae		Niconiades cydia	(Hewitson, 1876)	4									I
Hesperiidae	Hesperiinae		Niconiades merenda	(Mabille, 1878)	1									I
Hesperiidae	Hesperiinae		Ochlodes sp.	NA	x									NA
Hesperiidae	Hesperiinae		Papias phainis	Godman, 1900									x	XII
Hesperiidae	Hesperiinae		Perichares philetes	(Gmelin, [1790])	x									NA
Hesperiidae	Hesperiinae		Pheraeus fastus	(Hayward, 1939)									x	NA
Hesperiidae	Hesperiinae		Pheraeus odilia	(Plötz, 1884)	1									XII
Hesperiidae	Hesperiinae		Polites vibex	(Geyer, 1832)									x	XII
Hesperiidae	Hesperiinae		Pompeius pompeius	(Latreille, [1824])	x									NA
Hesperiidae	Hesperiinae		Psoralis coyana	(Schaus, 1902)			1							VII
Hesperiidae	Hesperiinae		Psoralis stacara	(Schaus, 1902)		3	1							IV, VII
Hesperiidae	Hesperiinae		Quasimellana nicomedes	(Mabille, 1883)	2									I
Hesperiidae	Hesperiinae		Saliana esperI	Evans, 1955	x									NA
Hesperiidae	Hesperiinae		Saliana saladin	Evans, 1955	x									NA
Hesperiidae	Hesperiinae		Sodalia dimassa	(Hewitson, 1876)	1									XII
Hesperiidae	Hesperiinae		Talides sergestus	(Cramer, 1775)	x									NA
Hesperiidae	Hesperiinae		Tirynthia conflua	(Herrich-Schäffer, 1869)	2									I
Hesperiidae	Hesperiinae		Vehilius clavicula	(Plötz, 1884)	1	2	2							III, III–IV, XII
Hesperiidae	Hesperiinae		Vehilius stictomenes	(A. Butler, 1877)	1									I
Hesperiidae	Hesperiinae		Vettius diana	(Plötz, 1886)	x									NA
Hesperiidae	Hesperiinae		Vettius lucretius	(Latreille, [1824])	x									NA
Hesperiidae	Hesperiinae		Vettius phyllus	Evans, 1955	x									NA
Hesperiidae	Hesperiinae		Vettius umbrata	(Erschoff, 1876)	2		3							II, VII
Hesperiidae	Hesperiinae		Vinius letis	(Plötz, 1883)	1									XII
Hesperiidae	Hesperiinae		Zariaspes mys	(Hübner, [1808])		5								III–IV, IV
Hesperiidae	Hesperiinae		Zenis jebus	(Plötz, 1882)	x									NA
Hesperiidae	Pyrginae		Achlyodes mithridates	(Fabricius, 1793)	1									I
Hesperiidae	Pyrginae		Anastrus obscurus	Hübner, [1824]	x									NA
Hesperiidae	Pyrginae		Anastrus ulpianus	Poey, 1832	x									NA
Hesperiidae	Pyrginae		Bolla catharina	(E. Bell, 1937)	1									I
Hesperiidae	Pyrginae		Burnsius orcus	(Stoll, 1780)		1		x						IV, XII
Hesperiidae	Pyrginae		Diaeus lacaena	(Hewitson, 1869)			1							XII
Hesperiidae	Pyrginae		Diaeus sp.	NA		1								III–IV
Hesperiidae	Pyrginae		Ebrietas infanda	(A. Butler, 1877)			1							III
Hesperiidae	Pyrginae		Gindanes brebisson	(Latreille, [1824])	x									NA
Hesperiidae	Pyrginae		Heliopetes (Heliopetes) arsalte	(Linnaeus, 1758)		1								IV
Hesperiidae	Pyrginae		Heliopetes (Heliopetes) ochroleuca	J. Zikán, 1938	1		1							VII, XII
Hesperiidae	Pyrginae		Heliopetes (Heliopyrgus) americanus	(Blanchard, 1852)			1							III
Hesperiidae	Pyrginae		Heliopetes (Leucoscirtes) omrina	(A. Butler, 1870)		1								IV
Hesperiidae	Pyrginae		Heliopetes alana	(Reakirt, 1868)	x									NA
Hesperiidae	Pyrginae		Mylon maimon	(Fabricius, 1775)	1									I
Hesperiidae	Pyrginae		Nisoniades castolus	(Hewitson, 1878)			1							III
Hesperiidae	Pyrginae		Noctuana diurna	(A. Butler, 1870)	1									I
Hesperiidae	Pyrginae		Pythonides lancea	(Hewitson, 1868)	1								x	I
Hesperiidae	Pyrginae		Quadrus cerialis	(Stoll, 1782)	1									XII
Hesperiidae	Pyrginae		Theagenes dichrous	(Mabille, 1878)			1							VII
Hesperiidae	Pyrginae		Trina geometrina	(C. Felder & R. Felder, 1867)	x									NA
Hesperiidae	Pyrginae		Viola violella	(Mabille, 1898)									x	XII
Hesperiidae	Pyrginae		Xenophanes tryxus	(Stoll, 1780)		5	3							III, III–IV, VII
Hesperiidae	Pyrginae		Zera hyacinthinus	(Mabille, 1877)	x									NA
Hesperiidae	Pyrginae		Zera zera	(A. Butler, 1870)	x									NA
Hesperiidae	Pyrrhopyginae		Elbella lamprus	(Hopffer, 1874)	x									NA
Hesperiidae	Pyrrhopyginae		Mimoniades (Mahotis) versicolor	(Latreille, [1824])	2									I
Hesperiidae	Pyrrhopyginae		Myscelus amystis	(Hewitson, 1867)	1									XII
Hesperiidae	Pyrrhopyginae		Pyrrhopyge charybdis	Westwood, 1852	x									NA
Hesperiidae	Tagiadinae		Celaenorrhinus eligius	(Stoll, 1781)	1									I
Hesperiidae	Tagiadinae		Celaenorrhinus similis	Hayward, 1933	x									NA
Lycaenidae	Polyommatinae		Elkalyce cogina	(Schaus, 1902)	1									I
Lycaenidae	Polyommatinae		Hemiargus hanno	(Stoll, 1790)		1								IV
Lycaenidae	Polyommatinae		Leptotes cassius	(Cramer, 1775)			1							III
Lycaenidae	Polyommatinae		Zizula cyna	(W. H. Edwards, 1881)		3								III–IV, IV
Lycaenidae	Theclinae		Arawacus meliboeus	(Fabricius, 1793)		1								III–IV
Lycaenidae	Theclinae		Arzecla arza	(Hewitson, 1874)	x									NA
Lycaenidae	Theclinae		Aubergina vanessoides	(Prittwitz, 1865)	x									NA
Lycaenidae	Theclinae		Brangas silumena	(Hewitson, 1867)	x									NA
Lycaenidae	Theclinae		Calycopis janeirica	(C. Felder, 1862)	x									NA
Lycaenidae	Theclinae		Chalybs janias or C. chloris	NA	x									NA
Lycaenidae	Theclinae		Evenus satyroides	(Hewitson, 1865)	x									NA
Lycaenidae	Theclinae		Janthecla flosculus	(H. Druce, 1907)			1							III
Lycaenidae	Theclinae		Laothus phydela	(Hewitson, 1867)		1								IV
Lycaenidae	Theclinae		Ocaria thales	(Fabricius, 1793)	x									NA
Lycaenidae	Theclinae		Ostrinotes empusa	(Hewitson, 1867)			1							XII
Lycaenidae	Theclinae		Rekoa palegon	(Cramer, 1780)	x									NA
Lycaenidae	Theclinae		Strephonota elika	(Hewitson, 1867)				x						XII
Lycaenidae	Theclinae		Theritas hemon	(Cramer, 1775)		1								III–IV
Lycaenidae	Theclinae		Theritas lisus	(Stoll, 1790)	x									NA
Nymphalidae	Apaturinae		Doxocopa laurentia	(Godart, [1824])			1							IV
Nymphalidae	Apaturinae		Doxocopa zunilda	(Godart, [1824])									x	NA
Nymphalidae	Biblidinae		Catonephele acontius	(Linnaeus, 1771)				1	1					II, III
Nymphalidae	Biblidinae		Catonephele numilia	(Cramer, 1775)				1	2	1		1		III, IV
Nymphalidae	Biblidinae		Diaethria clymena	(Cramer, 1775)				7	1	3		4		II, III, V, XII
Nymphalidae	Biblidinae		Diaethria eluina	(Hewitson, [1855])					1					III
Nymphalidae	Biblidinae		Dynamine athemon	(Linnaeus, 1758)	x									NA
Nymphalidae	Biblidinae		Dynamine tithia	(Hübner, 1823)	x									NA
Nymphalidae	Biblidinae		Ectima thecla	(Fabricius, 1796)				2				3		II, IX
Nymphalidae	Biblidinae		Epiphile orea	(Hübner, [1823])			1	4	2	2	1	1		II to V, IX, XI, XII
Nymphalidae	Biblidinae		Haematera pyrame	(Hübner, [1819])				2				1		II
Nymphalidae	Biblidinae		Hamadryas amphinome	(Linnaeus, 1767)				2		1		1		II, XII
Nymphalidae	Biblidinae		Hamadryas epinome	(C. Felder & R. Felder, 1867)				41				17		II, V, IX, XI
Nymphalidae	Biblidinae		Hamadryas februa	(Hübner, [1823])	1			2						II
Nymphalidae	Biblidinae		Hamadryas feronia	(Linnaeus, 1758)	x									NA
Nymphalidae	Biblidinae		Hamadryas fornax	(Hübner, [1823])			1	3				1		II, III, XI
Nymphalidae	Biblidinae		Myscelia orsis	(Drury, 1782)		2	1	33	1			18		II to V, VIII, IX, XI
Nymphalidae	Biblidinae		Temenis laothoe	(Cramer, 1777)				2		2		1		V, XII
Nymphalidae	Charaxinae		Archaeoprepona amphimachus	(Fabricius, 1775)	1			2	1			1		I to III, V
Nymphalidae	Charaxinae		Archaeoprepona chalciope	(Hübner, [1823])				5		2				II, III, XII
Nymphalidae	Charaxinae		Archaeoprepona demophon	(Linnaeus, 1758)	2			4				1		I, II, XI
Nymphalidae	Charaxinae		Consul fabius	(Cramer, 1776)	1			1						I, II
Nymphalidae	Charaxinae		Fountainea ryphea	(Cramer, 1775)			1	2	10	6	2	2		II to VI, XII
Nymphalidae	Charaxinae		Hypna clytemnestra	(Cramer, 1777)	1		1				1			I, III, V
Nymphalidae	Charaxinae		Memphis acidalia	(Hübner, [1819])		1			1		1			III–IV, IV, V
Nymphalidae	Charaxinae		Memphis appias	(Hübner, [1825])			2	5	51	8	1			II to IV, XII
Nymphalidae	Charaxinae		Memphis moruus	(Fabricius, 1775)					4		1			II, III
Nymphalidae	Charaxinae		Memphis otrere	(Hübner, [1825])				4	2	2		2		II to IV, XI
Nymphalidae	Charaxinae		Siderone galanthis	(Cramer, 1775)							1			IV
Nymphalidae	Charaxinae		Zaretis strigosus	(Gmelin, [1790])				2	3	1	3			II to V, XI
Nymphalidae	Danainae		Aeria olena	Weymer, 1875	x									NA
Nymphalidae	Danainae		Callithomia lenea	(Cramer, 1779)	1		3							I, III
Nymphalidae	Danainae		Danaus erippus	(Cramer, 1775)			8							III, IV, VII, XII
Nymphalidae	Danainae		Danaus gilippus	(Cramer, 1775)		1							x	IV
Nymphalidae	Danainae		Dircenna dero	(Hübner, 1823)	2									I
Nymphalidae	Danainae		Episcada hymenaea	(Prittwitz, 1865)			1							III
Nymphalidae	Danainae		Episcada philoclea	(Hewitson, [1855])	x									NA
Nymphalidae	Danainae		Episcada striposis	Haensch, 1909	x									NA
Nymphalidae	Danainae		Episcada sylvo	(Geyer, 1832)	2									I
Nymphalidae	Danainae		Epityches eupompe	(Geyer, 1832)	5	2	52						x	I, III, IV, VI, VII, XII
Nymphalidae	Danainae		Hypoleria adasa	(Hewitson, [1855])	x									NA
Nymphalidae	Danainae		Hypothyris ninonia	(Hübner, [1806])			4			1	1			III to VI
Nymphalidae	Danainae		Ithomia agnosia	Hewitson, [1855]			1							IV
Nymphalidae	Danainae		Ithomia drymo	Hübner, 1816	4	4	6	x	2				x	I, III, IV, VI, VII, XII
Nymphalidae	Danainae		Ithomia lichyI	R.F. d’Almeida, 1939			3							III
Nymphalidae	Danainae		Lycorea halia	(Hübner, 1816)									x	NA
Nymphalidae	Danainae		Mechanitis lysimnia	(Fabricius, 1793)	2	1	4	x						I, III, III–IV, VI, XII
Nymphalidae	Danainae		Melinaea ethra	(Godart, 1819)	x									NA
Nymphalidae	Danainae		Melinaea ludovica	(Cramer, 1780)	2									I
Nymphalidae	Danainae		Oleria aquata	(Weymer, 1875)	1		1							I, III
Nymphalidae	Danainae		Placidina euryanassa	(C. Felder & R. Felder, 1860)			1							III
Nymphalidae	Danainae		Pseudoscada erruca	(Hewitson, 1855)	6		2	x						I, III, XII
Nymphalidae	Danainae		Pteronymia carlia	Schaus, 1902	4	3	5							I, III, IV, XII
Nymphalidae	Heliconiinae		Actinote carycina	Jordan, 1913			1							IV
Nymphalidae	Heliconiinae		Actinote dalmeidaI	Francini, 1996	1									XII
Nymphalidae	Heliconiinae		Actinote genitrix	R.F. d’Almeida, 1922	2									XII
Nymphalidae	Heliconiinae		Actinote melanisans	Oberthür, 1917			1							XII
Nymphalidae	Heliconiinae		Actinote parapheles	Jordan, 1913	1									XII
Nymphalidae	Heliconiinae		Actinote pyrrha	(Fabricius, 1775)		1	5	x						IV, XII
Nymphalidae	Heliconiinae		Actinote sp.	NA	4	10							x	III–IV, IV, XI, XII
Nymphalidae	Heliconiinae		Agraulis vanillae	(Linnaeus, 1758)	x									NA
Nymphalidae	Heliconiinae		Dione juno	(Cramer, 1779)	x									NA
Nymphalidae	Heliconiinae		Dryadula phaetusa	(Linnaeus, 1758)									x	NA
Nymphalidae	Heliconiinae		Dryas iulia	(Fabricius, 1775)	1		5						x	I, III, IV, VI
Nymphalidae	Heliconiinae		Eueides aliphera	(Godart, 1819)		2								III–IV
Nymphalidae	Heliconiinae		Eueides isabella	(Stoll, 1781)			1	x						IV, XII
Nymphalidae	Heliconiinae		Eueides pavana	Ménétriés, 1857		1								IV
Nymphalidae	Heliconiinae		Heliconius besckeI	(Ménétriés, 1857)	3	1	8	x						I, III, IV, VII, XII
Nymphalidae	Heliconiinae		Heliconius erato	(Linnaeus, 1758)	3	1	10							I, III, IV, XII
Nymphalidae	Heliconiinae		Heliconius ethilla	(Godart, 1819)	1		4							III, VII, XII
Nymphalidae	Heliconiinae		Heliconius sara	(Fabricius, 1793)	1	2	1							III, III–IV, XII
Nymphalidae	Heliconiinae		Philaethria wernickeI	(Röber, 1906)		1	3						x	III, IV
Nymphalidae	Limenitidinae		Adelpha cocala	(Cramer, 1779)	1									I
Nymphalidae	Limenitidinae		Adelpha gavina	Fruhstorfer, 1915	x									NA
Nymphalidae	Limenitidinae		Adelpha lycorias	(Godart, [1824])	x									NA
Nymphalidae	Limenitidinae		Adelpha mythra	(Godart, [1824])	1									I
Nymphalidae	Limenitidinae		Adelpha serpa	(Boisduval, 1836)	1			x						I, XII
Nymphalidae	Limenitidinae		Adelpha sp.	NA	1	3							x	III–IV, IV, XI
Nymphalidae	Limenitidinae		Adelpha syma	(Godart, [1824])			1							IV
Nymphalidae	Nymphalinae		Anartia amathea	(Linnaeus, 1758)	1	1	10	x					x	I, III, IV, XII
Nymphalidae	Nymphalinae		Eresia lansdorfI	(Godart, 1819)		1	1	x						III, III–IV, XII
Nymphalidae	Nymphalinae		Eresia perna	Hewitson, 1852	x									NA
Nymphalidae	Nymphalinae		Historis odius	(Fabricius, 1775)	1									I
Nymphalidae	Nymphalinae		Hypanartia bella	(Fabricius, 1793)		2	3							III–IV, XII
Nymphalidae	Nymphalinae		Hypanartia lethe	(Fabricius, 1793)	x									NA
Nymphalidae	Nymphalinae		Junonia evarete	(Cramer, 1779)			1	x						IV, XII
Nymphalidae	Nymphalinae		Ortilia ithra	(W. F. Kirby, 1900)		1								III–IV
Nymphalidae	Nymphalinae		Siproeta epaphus	(Latreille, [1813])	1			x						I, XII
Nymphalidae	Nymphalinae		Smyrna blomfildia	(Fabricius, 1781)	2				2					I, III
Nymphalidae	Nymphalinae		Tegosa claudina	(Eschscholtz, 1821)	1	1	3							III, IV, XII
Nymphalidae	Nymphalinae		Tegosa sp.	NA			2							III
Nymphalidae	Nymphalinae		Telenassa teletusa	(Godart, [1824])	3	4		x						I, III–IV, XII
Nymphalidae	Nymphalinae		Vanessa braziliensis	(Moore, 1883)	1		1							VII, XI
Nymphalidae	Satyrinae	BrassolinI	Blepolenis bassus	(C. Felder & R. Felder, 1867)	5						1			I, II
Nymphalidae	Satyrinae	BrassolinI	Blepolenis batea	(Hübner, [1821])	1	2	3		2		3	3		I to III, III–IV
Nymphalidae	Satyrinae	BrassolinI	Blepolenis catharinae	(Stichel, 1902)				1				1		II
Nymphalidae	Satyrinae	BrassolinI	Caligo arisbe	Hübner, [1822]								1		II
Nymphalidae	Satyrinae	BrassolinI	Caligo beltrao	(Illiger, 1801)	1								x	XI
Nymphalidae	Satyrinae	BrassolinI	Caligo brasiliensis	(C. Felder, 1862)	x									NA
Nymphalidae	Satyrinae	BrassolinI	Catoblepia amphirhoe	(Hübner, [1825])				1						II
Nymphalidae	Satyrinae	BrassolinI	Dasyophthalma creusa	(Hübner, [1821])			1	8				4		II, III
Nymphalidae	Satyrinae	BrassolinI	Dasyophthalma rusina	(Godart, [1824])		1		1			1			II, IV
Nymphalidae	Satyrinae	BrassolinI	Eryphanis reevesiI	(E. Doubleday, [1849])	1			16	2		1	2		II, III, V, X to XII
Nymphalidae	Satyrinae	BrassolinI	Narope cyllene	C. Felder & R. Felder, 1859				2	2	1		1		II to IV, XII
Nymphalidae	Satyrinae	BrassolinI	Opoptera aorsa	(Godart, [1824])	1		3							III, XII
Nymphalidae	Satyrinae	BrassolinI	Opoptera sulcius	(Staudinger, 1887)		1	2	12	1			2		II to IV
Nymphalidae	Satyrinae	BrassolinI	Opsiphanes cassiae	(Linnaeus, 1758)	1									I
Nymphalidae	Satyrinae	BrassolinI	Opsiphanes invirae	(Hübner, [1808])	1			4						I
Nymphalidae	Satyrinae	HaeterinI	Pierella lamia	(Sulzer, 1776)	1									I
Nymphalidae	Satyrinae	HaeterinI	Pierella nereis	(Drury, 1782)	1									XII
Nymphalidae	Satyrinae	MorphinI	Antirrhea archaea	Hübner, [1822]	x									NA
Nymphalidae	Satyrinae	MorphinI	Morpho aega	(Hübner, [1822])	3	1	2	80	1			10		all but VI, VII, IX, X
Nymphalidae	Satyrinae	MorphinI	Morpho anaxibia	(Esper, [1801])		2	1							III–IV, IV
Nymphalidae	Satyrinae	MorphinI	Morpho epistrophus	(Fabricius, 1796)			5	125	37	24	4	9	x	II to IV
Nymphalidae	Satyrinae	MorphinI	Morpho helenor	(Cramer, 1776)		1	3	46				3		II, III, III–IV, XI
Nymphalidae	Satyrinae	MorphinI	Morpho hercules	(Dalman, 1823)		1								IV
Nymphalidae	Satyrinae	SatyrinI	Archeuptychia cluena	(Drury, 1782)	1									VIII
Nymphalidae	Satyrinae	SatyrinI	Capronnieria galesus	(Godart, [1824])	2	2					2			I, III, IV
Nymphalidae	Satyrinae	SatyrinI	Carminda griseldis	(Weymer, 1911)	2							1		I, II, XII
Nymphalidae	Satyrinae	SatyrinI	Carminda paeon	(Godart, [1824])	2	5	5	7	2	1		8		all but VI, VII, X
Nymphalidae	Satyrinae	SatyrinI	Cissia eous	(A. Butler, 1867)		3								III–IV
Nymphalidae	Satyrinae	SatyrinI	Cissia phronius	(Godart, [1824])	4	3	6	10	1		7	4		I, III to V, IX, XII
Nymphalidae	Satyrinae	SatyrinI	Eteona tisiphone	(Boisduval, 1836)				12	6		1	7		II to V, IX, XI
Nymphalidae	Satyrinae	SatyrinI	Euptychoides castrensis	(Schaus, 1902)	3	2		16				8		I, II, III–IV, XI, XII
Nymphalidae	Satyrinae	SatyrinI	Forsterinaria necys	(Godart, [1824])	1		4	115	10	2		12		all but VI, X
Nymphalidae	Satyrinae	SatyrinI	Forsterinaria pronophila	(A. Butler, 1867)	1		1	44		3		11		I to III, V, VIII, IX, XI
Nymphalidae	Satyrinae	SatyrinI	Forsterinaria quantius	(Godart, [1824])	4	2		52				2		all but III, VI, VII, X
Nymphalidae	Satyrinae	SatyrinI	Godartiana muscosa	(A. Butler, 1870)	2		4	133	3			41		all but VII, X
Nymphalidae	Satyrinae	SatyrinI	Hermeuptychia aff. hermes	NA			10		1					III, IV, VI, VII, XII
Nymphalidae	Satyrinae	SatyrinI	Hermeuptychia hermes	(Fabricius, 1775)	2			33				14		I, II, XII
Nymphalidae	Satyrinae	SatyrinI	Hermeuptychia sp.	NA			11				3			III to V
Nymphalidae	Satyrinae	SatyrinI	Moneuptychia soter	(A. Butler, 1877)	1			3	1		4	2		I, III to VI, XI
Nymphalidae	Satyrinae	SatyrinI	Pareuptychia ocirrhoe	(Fabricius, 1776)	1									IV
Nymphalidae	Satyrinae	SatyrinI	Paryphthimoides grimon	(Godart, [1824])	1									IV
Nymphalidae	Satyrinae	SatyrinI	Paryphthimoides poltys	(Prittwitz, 1865)	x									NA
Nymphalidae	Satyrinae	SatyrinI	Praepedaliodes phanias	(Hewitson, 1862)			1					5		II, III, V
Nymphalidae	Satyrinae	SatyrinI	Praepedaliodes sp.	NA	1									II
Nymphalidae	Satyrinae	SatyrinI	Pseudodebis celia	(Cramer, 1779)				2	1	1				III, XI
Nymphalidae	Satyrinae	SatyrinI	Pseudodebis euptychidia	(A. Butler, 1868)	5		3		8		10			all but II, VII, VIII
Nymphalidae	Satyrinae	SatyrinI	Splendeuptychia aff. boliviensis	NA				6				18		V, VIII, IX, XI
Nymphalidae	Satyrinae	SatyrinI	Splendeuptychia ambra	(Weymer, [1911])				11				2		XI
Nymphalidae	Satyrinae	SatyrinI	Splendeuptychia sp.	NA	1									XII
Nymphalidae	Satyrinae	SatyrinI	Taygetina kerea	(A. Butler, 1869)	x									NA
Nymphalidae	Satyrinae	SatyrinI	Taygetis acuta	Weymer, 1910				1			1			II, V
Nymphalidae	Satyrinae	SatyrinI	Taygetis mermeria	(Cramer, 1776)			2		3					III, VI
Nymphalidae	Satyrinae	SatyrinI	Taygetis rectifascia	(Weymer, 1907)	14							1		I, II, XII
Nymphalidae	Satyrinae	SatyrinI	Taygetis rufomarginata	Staudinger, 1888	x									NA
Nymphalidae	Satyrinae	SatyrinI	Taygetis sosis	Hopffer, 1874	x									NA
Nymphalidae	Satyrinae	SatyrinI	Taygetis sp.	NA	2									I
Nymphalidae	Satyrinae	SatyrinI	Taygetis tripunctata	Weymer, 1907	x									NA
Nymphalidae	Satyrinae	SatyrinI	Taygetis ypthima	(Hübner, [1821])	12					1	3			I to III, VI, XII
Nymphalidae	Satyrinae	SatyrinI	Yphthimoides affinis	(A. Butler, 1867)	1									I
Nymphalidae	Satyrinae	SatyrinI	Yphthimoides ochracea	(A. Butler, 1867)	1									IV
Nymphalidae	Satyrinae	SatyrinI	Yphthimoides ordinaria	A.V.L. Freitas, L. Kaminski & O.H.H. Mielke, 2012	1									II
Nymphalidae	Satyrinae	SatyrinI	Zischkaia pacarus	(Godart, [1824])								1		NA
Papilionidae	Papilioninae		Eurytides bellerophon	(Dalman, 1823)	1									I
Papilionidae	Papilioninae		Heraclides thoas	(Rothschild & Jordan, 1906)	x									NA
Papilionidae	Papilioninae		Mimoides lysithous	(Hübner, [1821])	1								x	I
Papilionidae	Papilioninae		Mimoides protodamas	(Godart, 1819)	1									I
Papilionidae	Papilioninae		Parides agavus	(Drury, 1782)			2							III
Papilionidae	Papilioninae		Parides anchises	(Linnaeus, 1758)	2									I
Papilionidae	Papilioninae		Parides proneus	(Hübner, [1831])	2									I
Papilionidae	Papilioninae		Pterourus scamander	(Boisduval, 1836)		1								IV
Pieridae	Coliadinae		Eurema albula	(Cramer, 1775)	1	5	5	x						III, IV, VI, XII
Pieridae	Coliadinae		Phoebis argante	(Fabricius, 1775)		3		x						III–IV, IV, XII
Pieridae	Coliadinae		Phoebis philea	(Linnaeus, 1763)	2	1	2	x					x	I, III, IV, XII
Pieridae	Coliadinae		Phoebis sennae	(Linnaeus, 1758)		1								IV
Pieridae	Coliadinae		Pyrisitia nise	(Cramer, 1775)		1								III–IV
Pieridae	Coliadinae		Rhabdodryas trite	(Linnaeus, 1758)	1	1	1	x					x	III, IV, XII
Pieridae	Dismorphiinae		Dismorphia amphione	(Cramer, 1779)			1							III
Pieridae	Dismorphiinae		Dismorphia crisia	(Drury, 1782)			2							III
Pieridae	Dismorphiinae		Dismorphia thermesia	(Godart, 1819)	1	1								I, III–IV
Pieridae	Dismorphiinae		Enantia clarissa	(Weymer, 1895)				x						XII
Pieridae	Dismorphiinae		Pseudopieris nehemia	(Boisduval, 1836)	1									XII
Pieridae	Pierinae		Archonias brassolis	(Fabricius, 1776)	1									I
Pieridae	Pierinae		Melete lycimnia	(Cramer, 1777)		2								IV
Pieridae	Pierinae		Pereute swainsonI	(G. Gray, 1832)			4						x	III
Riodinidae	Riodininae		Adelotypa bolena	(A. Butler, 1867)	x								x	NA
Riodinidae	Riodininae		Ancyluris aulestes	(Saunders, 1850)	x								x	NA
Riodinidae	Riodininae		Calospila apotheta	(H. Bates, 1868)	x									NA
Riodinidae	Riodininae		Emesis sp.	NA			1							III
Riodinidae	Riodininae		Eurybia carolina	Godart, [1824]			1							IV
Riodinidae	Riodininae		Eurybia molochina	Stichel, 1910	x									NA
Riodinidae	Riodininae		Eurybia pergaea	(Geyer, 1832)	1	2								IV, XII
Riodinidae	Riodininae		Euselasia hygenius	(Stoll, 1787)	x									NA
Riodinidae	Riodininae		Ionotus alector	(Geyer, 1837)	x									NA
Riodinidae	Riodininae		Leucochimona icare	(Hübner, [1819])			1							III
Riodinidae	Riodininae		Mesene sp.	NA	1									I
Riodinidae	Riodininae		Mesosemia mayi	Lathy, 1958	x									NA
Riodinidae	Riodininae		Mesosemia odice	(Godart, [1824])	4	1	1							I, IV, VI, XII
Riodinidae	Riodininae		Mesosemia rhodia	(Godart, [1824])	4									XII
Riodinidae	Riodininae		Metacharis ptolomaeus	(Fabricius, 1793)			1							III
Riodinidae	Riodininae		Napaea elisae	(J. Zikán, 1952)	x									XII
Riodinidae	Riodininae		Napaea nepos	(Fabricius, 1793)	1									XII
Riodinidae	Riodininae		Panara soana	Hewitson, 1875	x									NA
Riodinidae	Riodininae		Rhetus periander	(Cramer, 1777)			1							III
Riodinidae	Riodininae		Rhetus arcius	(Linnaeus, 1763)									x	NA
Riodinidae	Riodininae		Stichelia bocchoris	(Hewitson, 1876)	x									NA
Riodinidae	Riodininae		Symmachia aconia	Hewitson, 1876	x									NA
Riodinidae	Riodininae		Voltinia cebrenia	(Hewitson, [1873])	x									NA
				Total n	230 + 74	128	288	866 + 19	165	64	53	227	25
				Total richness	195	60	92	64	31	20	21	40	23

Lastly, focusing on baited traps (PEI traps and AG traps), to understand how the richness distribution match what has been found in other trap studies in the Atlantic Forest, we used a database of Nymphalidae species lists in Brazil called DnB (Shirai et al. 2019, downloaded from https://doi.org/10.5281/zenodo.2561408), filtering studies that exclusively used baited traps, exclusively at the Atlantic Forest. That is, we did not include studies of the DnB database that used e.g. active searches or active search plus traps; or done in other biomes or the Atlantic Forest plus other biomes, because the lists would not be strictly comparable. The DnB study used the same source for phylogenetic arrangement as ours (Wahlberg et al. 2009). In relation to species identities, we filtered species considered valid by Lamas (2004) which, despite being outdated, standardizes the taxonomy to a single source. To counterbalance this compromise, we also consulted the largest database of fruit-feeding butterflies in the Atlantic Forest (Santos et al. 2018), that has an updated taxonomy. Using Santos et al. (2018) classification of tribes, we compared our totals per tribe with their data (Figure 3 of Santos et al. 2018).

Figure 3.
Butterfly species common to all sampling regions (intersection of PEI, PENaP, AG datasets in Figure 2). These are likely the species that any person will see at the PEI and surroundings. Sizes are proportional to their real size, with dorsal wing surfaces shown spread, and ventral surfaces tilted similar to the butterfly’s resting position. The net symbol indicates species caught exclusively with the entomological net, while all others were caught with the net and traps. A more complete illustrated guide can be found in the companion paper (Shirai et al. 2022).

Results

In total, we found 312 species for PEI and surroundings (Table 2) that sum to 2,139 records (1,605 in active search, 509 in traps, 25 other). The museum contributed with 432 records for 229 species. We sampled 1,682 specimens belonging to 142 species, in a total effort of 36,679 sampling hours (36,432 trap and 247 net and observation hours, Table 1).

The richest families were Nymphalidae (148 species), the single family caught by all sampling methods, and Hesperiidae (100), followed by Riodinidae (23 species), Lycaenidae (19 species), Pieridae (14 species), and Papilionidae (8 species). Within nymphalids, the richest subfamily was Satyrinae (58 species), followed by Danainae (23), Heliconiinae (18), Biblidinae (16), Nymphalinae (13), Charaxinae (12), Limenitidinae (6), and Apaturinae (2). Within skippers, the richest subfamily was Hesperiinae (54 species), followed by Pyrginae (24), Eudaminae (16), Pyrrhopyginae (4), and Tagiadinae (2). None of the species were among the 63 threatened (that is, critically endangered, endangered, or vulnerable) butterfly species (ICMBio 2018, updated by Augusto H. B. Rosa).

The great majority of species was sampled exclusively by active methods (79.8%), with 18.3% having been collected by both active and passive methods, and only 1.9% sampled in traps only (Figure 2a). Additionally, the citizen science, iNaturalist and Santos et al. (2015) sources (“other” dataset, not included in Figure 2a due to the uncertain methodology) contributed with eight species not found by any of the active or passive methods: the skippers Papias phainis Godman, 1900, Pheraeus fastus (Hayward, 1939), Polites vibex (Geyer, 1832), and Viola violella (Mabille, 1898); the nymphalids Doxocopa zunilda (Godart, [1824]), Lycorea halia (Hübner, 1816), Dryadula phaetusa (Linnaeus, 1758); and the metalmark Rhetus arcius (Linnaeus, 1763).

The comparison by region (Figure 2b) showed, expectedly, that the majority was collected at the focal region (PEI, with 64.5% being exclusive). The 20 species common to all regions (Figure 3) included both forest dwellers and open area species.

We found the highest diversity in warmer months (Figure 4), as can be seen in the total richness (Figure 4a, using all data but not standardized by sampling effort). The information of which month each species was sampled (last column of Table 2) should, however, be used with caution because it does not necessarily reflect the true phenology of species due to sampling bias, such as the overrepresentation of summer months. Also, the number of individuals is subject to methodological bias, so looking at the PEI trap data, where we can compare abundance, March, April, and May outstood in both richness and abundance (Figure 4b). We would not biologically interpret the results in September since it seems an effect of statistic inflation: only 5 individuals of 4 species were sampled but, as they were caught only the PEI 1y dataset (3 days in 11 traps), the metrics end up higher than other datasets that sampled for more days in a higher number of traps.

Figure 4.
Diversity metrics for (a) all data, under the three sampling methods (absolute number of species) by month; and for PEI traps (richness, in dashed line, and abundance, in solid line, per trap and per trap day) reporting them by (b) month, and (c) by season.

As we sampled in every month of the year, we can demonstrate that winter months have lower richness compared to any other season in the whole dataset. We found in June: 13 species, July: 16 species, August: nine species, September: 13 species (Figure 4a); which is much lower compared to remaining seasons (average ± standard deviation 142.3 ± 47.2). This result can also be observed in the comparable data of baited traps (Figure 4b-c). Lastly, 15 out of the 19 Brassolini and Morphini species were reported exclusively from December to April.

Despite our sampling bias, we found that nine species should, with some certainty, be observed most of the year (species caught in seven-10 months in the year, Table 2): biblidines Epiphile orea (Hübner, [1823]) and Myscelia orsis (Drury, 1782), and satyrines Carminda paeon (Godart, [1824]), Forsterinaria necys (Godart, [1824]), F. pronophila (A. Butler, 1867), F. quantius (Godart, [1824]), Godartiana muscosa (A. Butler, 1870), Morpho aega (Hübner, [1822]), and Pseudodebis euptychidia (A. Butler, 1868). Except from E. orea and F. quantius, the remaining seven species also ranked among the 15 most abundant species the trap sampling.

Among trap datasets, we caught 505 individuals of 63 species. The richest and most abundant subfamily was Satyrinae, followed by Charaxinae, Biblidinae, and Nymphalinae (Table 3). We found that 51.3% of the individuals belong to seven species: Morpho epistrophus (Fabricius, 1796) (74), Memphis appias (Hübner, [1825]) (60), G. muscosa (44), F. necys (24), Fountainea ryphea (Cramer, 1775) (20), M. orsis (19), and P. euptychidia (18). On average, we caught 0.19 individuals/trap/day in the PEI (0.34), PEI-fish (0.22), PEI 1y (0.13) and AG (0.09) datasets. The comparison between PEI habitats and vertical stratification using the PEI and PEI fish datasets can be found in the Supplementary Material.

Table 3.
Richness and abundance of trap studies: “Our study” sums the trap individuals from the PEI, PEI fish, PEI 1y and AG datasets; “DnB” refers to “Database of nymphalids in Brazil” (Shirai et al. 2019), with species lists exclusively caught with traps at the Atlantic Forest (numbers are sums of 40 studies, see Material and Methods); and “Atl butterflies” refers to the “Atlantic Butterflies” database (Santos et al. 2018), the largest database for fruit-feeding butterflies. The number of individuals refers to abundance (our study) or to the number of records (= presence) in the literature (for DnB and Atl butterflies). Nymphalidae classification follows Wahlberg et al. 2009 (subfamilies) and Santos et al. 2018 (tribes).

Taxa	Richness			Abundance (our study)/records
	Our study	DnB	Atl butterflies	Our study	DnB	Atl butterflies
Biblidinae	12	45	63	66	513	2336
Ageroniini	4	11	11	23	208	639
Biblidini	0	1	2	0	29	136
Callicorini	3	11	11	10	81	448
Epicaliini (Catonephilini)	3	5	6	24	79	244
Epiphilini	2	6	6	9	61	272
Eubagini	0	3	12	0	5	362
Eunicini	0	8	15	0	50	235
Charaxinae	11	30	34	107	331	1010
Anaeini	8	19	20	102	190	656
Preponini	3	11	14	5	141	354
Nymphalinae	1	5	5	2	100	268
Coeini	1	5	5	2	100	268
Satyrinae	37	126	176	328	897	3434
Brassolini	9	28	36	28	244	881
Haeterini	0	4	5	0	17	79
Morphini	3	12	9	88	73	354
Satyrini	25	82	126	212	563	2120
Total	61	206	278	503	1841	7048

Discussion

Butterflies are excellent model organisms for scientific research (reviewed in e.g.Brown & Freitas 1999, Santos et al. 2008, Sourakov & Shirai 2020) and for conservation purposes (see Introduction). However, even considering they are among the best-studied insects, important gaps in basic information still exist, such as for species description and mapping species distributions (Linnean and Wallacean shortfalls c.f.Hortal et al. 2015). That results in the exclusion of butterflies, as well as other invertebrates, from studies like those that established hotspots (Myers et al. 2000) or other global conservation efforts (Bossart & Carlton 2002, Barua et al. 2012), even considering that butterflies are a bioindicator, flagship, and umbrella group. In this study, we aimed at filling this gap by inventorying butterfly diversity in a region within the Paranapiacaba Continuum, a key network of protected areas for the preservation of the Atlantic Forest hotspot. We also provided several resources to aid conservation efforts (see Introduction), and a discussion for both academics and non-academics.

Most of butterfly richness is in the tropics. While Nymphalidae diversified in the Asian, African and American tropics, several lineages of Hesperiidae and Riodinidae adaptive radiations happened exclusively in the Neotropics (Toussaint et al. 2018, Seraphim et al. 2018), with metalmarks being mainly Amazonian. Thus, it is not surprising that the richest families we found were Nymphalidae and Hesperiidae, frequently reported as the most diverse in the Atlantic Forest (Brown & Freitas 2000, Francini et al. 2011). Hesperiidae is probably the richest butterfly family in Brazil and the fact that we caught them less than Nymphalidae probably reflects the suggestion of Francini et al. (2011): in relatively complete inventories, the small and inconspicuous Hesperiidae are better sampled, but in short term studies, Nymphalidae appears with higher richness because they are easily captured, with both net and traps. Lycaenidae and Riodinidae, despite having less species in the Atlantic Forest than the two families above, equally suffer in shorter inventories by being harder to catch. Moreover, most Pieridae species we caught are found in open areas across the country but important species are expected to be caught only at the higher altitudes in the park, above 800 m a.s.l. (Francini et al. 2011).

Focusing on nymphalids, a review of species lists in Brazil (Shirai et al. 2019) listed the presence of 162-315 butterfly species in the top-10 richest places in the country. The corresponding biomes (Amazon, Atlantic Forest and one in the Cerrado) rank among the richest because 1) they are biologically rich but also, and importantly, 2) they were well-sampled (Shirai et al. 2019). The most complete dataset of a subset of Nymphalidae, fruit-feeding butterflies, listed 279 species for the whole Atlantic Forest (Santos et al. 2018), which is a substantial increment over previously available data, such as the 88–127 fruit-feeding species found in the same biome (Brown 2005). We were able to record 148 nymphalids (61 fruit-feeding), but these numbers as well as our total of 312 butterfly species are certainly not the true richness of the PEI and surroundings. The richness in the region could easily surpass 500 species or more, as compared to similar areas in the same region (Francini et al. 2011). More sampling, and sampling by different strategies (like comparing to an estimate of the “true richness” by maximized sampling, Uehara-Prado et al. 2007), are necessary to reflect the diversity of this region. Despite the attempt to combine different datasets to enhance this inventory, some obvious gaps remained, such as 1) a thorough expedition to the other PEI station at the East lowlands (Saibadela), that has a different climate and altitude (Morellato et al. 2000); 2) the most inaccessible and pristine region of the PEI, at the South, as well as the few but important high altitude sites in the park; 3) the contiguous protected areas PECB, PETAR and Xitué (see Figure 1); and 4) more sampling with different baits.

The use of different sampling methods certainly adds to more comprehensive inventories (e.g. Clench 1949, 1979, Brown 1972, Brown & Freitas 2000, Iserhard et al. 2013, Freitas et al. 2014, Checa et al. 2018). For example, most of the species we found were sampled by active methods (Figure 2a), collected by highly experienced people, during different years. However, despite having caught only about a fifth (63/312 species) of the total richness with traps, Siderone galanthis (Cramer 1775) was among the six trap-exclusive species (Figure 2a), caught by a recently trained PEI guide (“PEI 1y” dataset). Decinea dama (Herrich-Schäffer 1869) was also among the trap-exclusive species, only caught with fish carrion (“PEI fish” dataset). It is also worth mentioning that the citizen science approaches and a field course experiment (“other” dataset) contributed with eight species not found with neither active nor passive methods.

Including efforts of other places than the PEI also enhanced the inventory for the region adding, in our case, 33 species (20 exclusive of PENaP, 11 exclusive of AG, plus two in common, Figure 2b) not found at the park, that had much more sampling effort (Table 1). Several of the 20 species common to all regions (Figure 2b) are associated with open or fragmented areas; we illustrated them (Figure 3) because they are, aside from beautiful, colorful and diverse, likely the species that any person will observe at the region.

An interesting statistic, both biologically and in terms of planning, is the number of individuals caught per trap per day, as the 0.5 individuals/trap/day reported in an Amazonian terra firme site (Ribeiro & Freitas 2012). Traps at open and/or fragmented habitats tend to catch more individuals, but not species (e.g. Uehara-Prado et al. 2007). We caught 0.19 individuals/trap/day, but the highest values were not necessarily at the open, disturbed or fragmented sites, such as PEI 1y (0.13) and AG (0.09) datasets, but rather at the PEI (0.34) and PEI-fish (0.22). It would be interesting to gather more data like this to uncover differences, for instances, between environments and biomes.

The best months to collect in South Eastern Brazil are March to May (end of the rainy season, Ebert 1970, Brown 1972, 1992, Freitas et al. 2014) but other seasonal intervals are also relevant. Here, we chose when to collect aiming to span different seasons and months, complementing the effort from museum data at the same time as revisiting good months (like December and March) and we indeed found the highest diversity metrics in warmer months, particularly from March to May (Figure 4).

We thus reinforce a previous suggestion (Brown 1972, Uehara-Prado et al. 2007) that the best sampling months in the Atlantic Forest should be extended to the period between December to May (summer plus end of rainy season). The summer months of December to February are mandatory for tribes like Morphini and Brassolini that only fly in this period (Freitas et al. 2014). Here, we found 79% of brassolines and morphines within the months of December and April. However, winter months should not be completely disregarded since some species were exclusively caught in this season and, in the case of the PEI, it is the only season when an Epityches eupompe (Geyer, 1832) aggregation can be observed (Shirai et al. 2017, https://youtu.be/bUO4kpYS2uo).

For Brazilian rainforests standards, the PEI is chilly, partly because it has a lot of forest and in high altitudes, but also because it has a cold winter: the annual average temperature ranged from 15.1 to 19.2°C (Table S5), similar to what is reported in other sources (17.5°C from WorldClim and 17–18°C from Leonel 2010). More importantly, the temperature is seasonal, with a warm summer and a cold winter (Figure S1, Table S5): maximum temperatures of the warmest month (26–27°C) and minimum temperature of the coldest month (7.5–11°C), also like the WorldClim data.

The dominant species in our trap datasets was Morpho epistrophus (15%), the only Morpho species in the Atlantic Forest that is iridescent white (Pablos et al. 2021, Figure 5) – caught on traps with black mesh (see Freitas et al. 2014). Although we caught them in traps only from February to April (many of which with worn wings, LTS pers. obs.), this large butterfly is visible anywhere in the park since November. Butterflies are good flagship invertebrates because of their aesthetic appeal which, in turn, relates to species attributes of size and brightness/color (Barua et al. 2012). The iridescent white color of M. epistrophus might seem dull to some visitors, but the fact that tourists in Neotropical forests are somewhat used to seeing, and being amused by, the blue morphos, might raise interest to their phylogenetic relationship, enhancing the color feature. Another important aspect of butterflies as invertebrate flagships is the fact that they are harmless. The elegant flight of M. epistrophus does not incite danger or disgust like wasps or mosquitoes might do. Although caterpillars are not always understood as an early stage of butterflies (Barua et al. 2012), they can trigger fear or dislike because of damage to crops. M. epistrophus harmless caterpillars are large, gregarious, bright red, and visible in many Inga Mill. tree trunks near the PEI reception (Figure 5). The contrasting effect of beauty and fear/disgust of these caterpillars could be used as an opportunity: the particularly skilled, enthusiastic and charismatic PEI staff could educate visitors about holometabolous life cycle, what makes a caterpillar harmful or harmless, what are aposematic colors, and warn visitors about moth caterpillars found at PEI (like Megalopyge Hübner 1820) that actually sting. Thus, the easy identification, large size, omnipresence in the park, relationship with famous blue morphos, elegant flight, and inciting caterpillars make M. epistrophus eligible to become the emblematic invertebrate mascot of the PEI.

Figure 5.
The emblematic white morpho, Morpho epistrophus, adult and caterpillar, that we suggest becoming the PEI invertebrate mascot. Images used with permission of the author Almir Cândido Almeida.

Overexploitation of resources (like the juçara palm), illegal hunting and mining are permanent issues that endanger threatened species (such as the jaguar, Beisiegel & Nakano-Oliveira 2020) and a threatened biome (Tabarelli et al. 2005, Maxwell et al. 2016, ICMBio 2018). Even in the largest and strongest continuous of Atlantic Forest, we face the danger of (irreversibly) losing species and natural resources. More involvement and investment in the area would result in much more for the people and the planet than e.g. the end goals of several “perverse subsidies” (Myers 1998, Tabarelli et al. 2005). For example, investing in a geographically wider and temporally longer sampling of butterflies, together with more science outreach seminars and material, capacity development and teaching in local schools are ideas that need dedicated, almost exclusive, time of trained people but, surprisingly, they do not require a substantial financial investment (see Shirai et al. 2022). Another better idea, though, would be to invest in bioliteracy (c.f.Janzen 2010), which is a strong chance to save tropical diversity.

Acknowledgments

This work is dedicated to Benedito Amaral, a highly skilled citizen scientist and fine observer of the natural world. We would like to thank: Benedito Amaral, Thiago B. Conforti, Eliseu C. de Paula, Faustino A. Ribeiro, José Floido, Mara C. F. Paiva, Irene A. Ribeiro, Zarife O. Mora, and the remaining PEI staff for their outstanding contribution; Benedito Amaral, Massuo J. Kato, Lydia F. Yamaguchi, Mariana A. Stanton, Dimitre Ivanov, Sidneia C. C. do Nascimento, Adilson R. Moreira, Tamara M. C. Aguiar, André R. Nascimento, and Joel Lastra-Valdés for their help during field sampling; Patrício A. Salazar for designing and performing the preliminary analysis of the carrion test with LTS; Jessie P. dos Santos for teaching LTS how to use the slingshot; Wesley R. Silva for kindly lending his car in the first field trip; Thamara Zacca for identifying many Satyrinae, Ronaldo B. Francini for identifying the Actinote; Augusto H. B. Rosa and Olaf H. H. Mielke for museum information and access; Marcelo Duarte for allowing LTS to access the MZUSP collection; Tamara M. C. Aguiar, Artur N. Furegatti, and Michela Borges for their assistance at the ZUEC-LEP collection; Almir C. Almeida for allowing with such enthusiasm to use his photographs; and Renato R. Ramos, Eduardo P. Barbosa and Junia Y. O. Carreira for remote assistance during the COVID-19 lockdown. LTS acknowledges FAPESP (14/23504-7) for a fellowship, and ALCR thanks CNPq (166036/2020-0). AVLF thanks (FAPESP) (2021/03868-8) and the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) (421248/2017-3, 304291/2020-0). This study was done under environmental licenses ICMBio 8585-1 (sol. 10438), IBAMA 02001.000480/2014-38 (820/2017), and COTEC 260108-004.611/2016. The SisGen registry is AE4A636. All authors declare no conflict of interest.

Supplementary Material

The following online material is available for this article:

A – Full description of our data.

B – Climatic data from the weather station at the PEI, in English and in Portuguese.

C – PEI trap datasets: habitats and vertical stratification in the PEI.

Data Availability

The primary data is reported in the main text as Table 2 and is also available at the public repository https://doi.org/10.5281/zenodo.7429126, with the metadata in the readme file available at https://doi.org/10.5281/zenodo.7439430.

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Edited by

Associate Editor

Gustavo Graciolli

Publication Dates

• Publication in this collection
  10 July 2023
• Date of issue
  2023

History

• Received
  16 Dec 2022
• Accepted
  01 May 2023