Biota Neotropica 23(2): e20221453, 2023

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ISSN 1676-0611 (online edition)

Inventory

The butterflies (Lepidoptera, Papilionoidea) of the Parque Estadual Intervales and

surroundings, São Paulo, Brazil

Leila T. Shirai¹* , Renato O. Silva², Fernando M. S. Dias³, André L. C. Rochelle4 & André V. L. Freitas¹

Universidade Estadual de Campinas, Instituto de Biologia, Departamento de Biologia Animal

1

e Museu de Diversidade Biológica, 13083-862, Campinas, SP, Brasil.

2
Universidade de São Paulo, Museu de Zoologia, 04263-000, São Paulo, SP, Brasil.
3
Universidade Estadual de Londrina, Departamento de Biologia Animal e Vegetal,
Londrina, 86057-970, Londrina, PR, Brasil.
4
Universidade Estadual Paulista, Departamento de Biodiversidade, 13506-900, Rio Claro SP, Brasil.
*Corresponding author: [email protected]
SHIRAI, L.T., SILVA, R.O., DIAS, F.M.S., ROCHELLE, A.L.C., FREITAS, A.V.L. The butterflies (Lepidoptera,
Papilionoidea) of the Parque Estadual Intervales and surroundings, São Paulo, Brazil. Biota Neotropica
23(2): e20221453. https://doi.org/10.1590/1676-0611-BN-2022-1453

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.

As borboletas (Lepidoptera, Papilionoidea) do Parque Estadual Intervales e

arredores, São Paulo, Brasil

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

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Shirai L.T. et al.

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 diverse and threatened regions, such as in the Atlantic Forest (Myers
Biodiversity studies are of central importance for the future, and et al. 2000); and for biological groups that are relevant for conservation,
inventories are one of their building blocks. Sampling distribution is, such as bioindicator, umbrella, and flagship groups.
however, far from homogeneous, with most knowledge concentrated Bioindicator and umbrella groups are particularly useful for
in the wealthiest, temperate, countries of the world (e.g. Girardello conservation planning and future attempts to understand what has been
et al. 2019). The unequal sampling effort increases the relevance of lost because they provide snapshots of the plant, animal or fungal species
carrying out inventories in the richest and megadiverse regions on Earth, at a given time. The information they provide is more useful when
usually found in the tropics, especially in those countries threatened inventories are performed according to best practices and standards.
by governments openly against science and the preservation of its For example, to assess the fauna of a given location, it is important to
environment (Myers et al. 2000, Alves et al. 2018, Andrade 2019, use different sampling methods, to collect periodically (e.g. monthly)
Angelo 2019, Tollefson 2019, Ferrante & Fearnside 2021, Hipólito et al. and for many years, to span different environments and altitudes, and
2021). Despite an increase in sampling in the Global South during the to be careful with specimen and data curation (Santos et al. 2008,
past century (Girardello et al. 2019), it is remarkable that some regions Shirai et al. 2019). That is rarely the case for several reasons, namely
with pristine natural environments remain unsampled, even at highly the financial and temporal availability of trained people, aggravated in

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.

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Butterflies of Intervales

places like South America (Elliot et al. 2018). However, in taxa that (Figure 1a), is a UNESCO Biosphere Reserve and a World Heritage Site.
are easy to collect and identify even by non-specialists, approaches This Continuum plays a central role in conserving one of the five hottest
like citizen science and capacity development can be powerful allies world hotspots (cf. Myers et al. 2000), due to its unique extension of
to complement collection by scientists (Uehara-Prado et al. 2007, continuous forest. For example, the Continuum is one of the last three
Davis 2015, Santos et al. 2016, Elliot et al. 2018, Mota et al. 2022, areas capable of hosting viable populations of the biome’s top predator,
Shirai et al. 2022). It also helps when the bioindicator or umbrella the jaguar (Panthera onca (Linnaeus, 1758)), which is vulnerable to
group is aesthetically appealing, which is the case of most butterflies extinction (ICMBio 2018) due to hunting and habitat loss (Beisiegel
(Lepidoptera, Papilionoidea: Hesperiidae, Lycaenidae, Nymphalidae, & Nakano-Oliveira 2020)
Papilionidae, Pieridae, and Riodinidae). It is estimated that 85% of the PEI still has well preserved old
Here we provide the first butterfly list of Intervales state park growth forest with very low levels of disturbance (Nisi 2006), which
(Parque Estadual Intervales or PEI), a protected area located in the is predominantly dense montane ombrophilous forest (Leonel 2010).
last massive continuous of the Atlantic Forest (Figure 1a). Despite Intervales means “between valleys,” in reference to the hills that
the importance of this area, and associated environmental and social range from 140 m to 1,200 m. The complex landscape with vast intact
threats (see Methods), it is remarkable that we did not find any study forest hosts important remnants of wild life, for instances, 9.2% of the
sampling this bioindicator group, butterflies, although some works PEI fauna of 532 species preliminarily inventoried in the 1990’s was
report the collection of focal taxa at the region (listed in Methods). threatened (Nisi 2006). Particularly, several arthropods in the Red List
The PEI boundaries were not determined for biological reasons, so we (ICMBio 2018) were found at the calcareous caves of the region, some
searched for inventories in the surrounding municipalities, all of which exclusively at the PEI. The pristine forest, the caves and the almost
are in the state of São Paulo, Brazil (Figure 1a). The only published 400 species of birds (an international hotspot for birdwatching) are
inventories near or at the PEI either collected all butterflies and did not among the main attractions of the park. Illegal hunting, mining, and
report the species list (Brown & Freitas 1999) or had a species list but overexploitation of juçara palm (Euterpe edulis Mart.) are among the
of a single tribe (Ithomiini, Shirai et al. 2017) or of a single guild on main threats to the life it preserves (Shirai et al. 2022).
specific flowers (nectar-feeding butterflies on Lantana L. (Verbenaceae), The climate of the region can be classified as dry-winter subtropical
Santos et al. 2015). highland, or Cwb in the Köppen-Geiger system (c.f. Beck et al. 2018),
We report the butterflies of the PEI and surroundings with data with rainy summers (December to March) and a noticeable, dry, winter
compiled from museums and four years of field work by us, under (June to September). The PEI has, however, different stations with
three sampling methods. Our aims were to contribute to the scientific distinct climatic profiles. At the PEI main station (ca. 800 m.a.s.l),
community, but also to the local community, supporting conservation the annual precipitation is of 1,400 mm, with the summer having
efforts in different ways. Here, we provide the butterfly list, discussing an average precipitation of 187 ± 30 mm, and the wettest month
it academically but also in a language accessible for non-academics. (January) having 208 mm (WorldClim 2.5’ data from 1970–2000, for
We also analyzed 27 years of climatic data collected at the park but so 24°15’56” S 48°24’48” W, 841 m a.s.l., which is the coordinate of
far without any formal synthesis. This climatic profile can be reported the “reception” in Figure 1b; Fick & Hijmans 2017, www.worldclim.
by the protected area to support decision-makers to conserve the area org/data/worldclim21.html, see also Figure 2 of Leonel 2010). During
with robust data, so we present it in the Supplementary Material both in winter, the average precipitation is 66 ± 21 mm, with the driest month
English and in Portuguese. Some of the spread individuals returned to (August) having 49 mm (WorldClim op. cit.). At another PEI station
the park and are in display for visitors, along with informative posters in the lowland (ca. 150 m a.s.l.), at Saibadela, the picture can be very
with the results of this study (http://doi.org/10.5281/zenodo.5893810, different as its nonseasonal tropical rainy climate reaches 4,000 mm
press release available at https://www.infraestruturameioambiente. annual precipitation, with the rainy season always with > 200 mm/
sp.gov.br/fundacaoflorestal/2022/02/parque-estadual-intervales-realiza- month, frequently > 400 mm/month, and the driest period (May to
primeiro-plantio-coletivo-de-mudas/). Also, the “Butterflies of August) with > 100 mm/month (Morellato et al. 2000).
Intervales” outreach booklet is publicly available (http://doi. We had access to 27 years of maximum and minimum temperature
org/10.5281/zenodo.5068939). Finally, the experiences of science and relative humidity data from the weather station located near the
outreach and capacity development with the PEI staff, and an illustrated PEI reception (Figure 1b). Data collection began in April 1992 and,
butterfly catalogue of species sampled here can be found in the although it is collected until the present time, we analyzed it until June
companion publication (Shirai et al. 2022). 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
Material and Methods at 14h, 9,651 at 20h), spread in 9,722 days of 320 months of 27 years
1. Study area (Tables S2–S4). More details of the data and its analyses are provided
in the Supplementary Material. This data showed high correlation
The Parque Estadual Intervales (hereafter PEI, 41,704 ha) is part with the temperature and humidity data we measured at the PEI trap
of the largest continuous of the Atlantic Forest at the Serra do Mar sampling (Table S6).
(1,109,546 ha, Ribeiro et al. 2009, inset of Figure 1a). The PEI is located
2. Butterfly data sampling
within the Paranapiacaba Continuum (ca. 140,000 ha c.f. decrees) which,
along with the contiguous ecological station Xitué and Parque Estadual This study joins datasets from museums (Museu de Diversidade
Carlos Botelho (PECB), Parque Estadual Turístico do Alto da Ribeira Biológica of the University of Campinas, ZUEC-LEP and the Zoology
(PETAR), and Parque Estadual Nascentes do Paranapanema (PENaP) Museum of the University of São Paulo, MZUSP, having also

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Shirai L.T. et al.

consulted the Coleção Entomológica Padre Jesus Santiago Moure, Our sampling was carried out in the following periods: 2.1.
Zoology Department of the Federal University of Paraná, DZUP; and PEI dataset (illustrated in Shirai et al. 2022) at 14 to 18.III.2016
the National Museum at Rio de Janeiro, MNUFRJ) and sampling by (pilot), 06 to 13.III.2017, 07 to 12.IV.2017, 11 to 16.VI.2017, 21
the authors (referred by our initials) from different locations, under to 23.VII.2017, 10 to 14.IV.2018, 26 to 29.VII.2018 with sampling
different methods. effort of 5,760 trap hours (16 days * 30 traps * 12 trap hours/day),
In the Supplementary Material we describe the material and and ca. 76 net hours; 2.2. PEI fish dataset (07 to 12.III.2018 and
methods of each dataset (summarized in Table 1), including coordinates, 09 to 15.XII.2018) with sampling effort of 3,456 trap hours (8 days
method(s), collectors, specialists who identified the material, aim, * 36 traps * 12 trap hours/day), and ca. 15 net hours; 2.3. PEI 1y
sampling effort, and design. We organized them as follows: 1. museum dataset 14 to 17.I.2019, 18 to 21.II.2019, 18 to 21.III.2019, 15 to
data, and 2. sampling data by us (with PEI species fully illustrated in 18.IV.2019, 27 to 30.V.2019, 17 to 20.VI.2019, 16 to 19.VII.2019,
Shirai et al. 2022). We divided the museum data in: 1.1. PEI ZUEC 20 to 23.VIII.2019, 17 to 20.IX.2019, 14 to 17.X.2019, 25 to
dataset, and 1.2. PENaP dataset (Parque Estadual Nascentes do 28.XI.2019, 09 to 12.XII.2019 with sampling effort of 4,752 trap
Paranapanema) – both datasets used the entomological net as the hours (36 days * 11 traps * 12 trap hours/day); and 2.4. AG dataset
sampling method (communicated by the collectors). We divided the XII.2014, 06 to 19.VIII.2017, 18 to 27.XI.2017, 17 to 26.II.2018,
sampling data in: 2.1. PEI dataset; 2.2. PEI fish dataset (carrion bait 17 to 25.V.2018, 25.VIII to 03.IX.2018, 14 to 23.XII.2018, 08 to
test in the Atlantic Forest); 2.3. PEI 1y dataset (one-year monitoring 17.III.2019, 02 to 12.VII.2019, 19 to 29.IX.2019 with sampling
for capacity development and science outreach); 2.4. AG dataset (Apiaí effort of 22,464 trap hours (39 days * 48 traps * 12 trap hours/day)
and Guapiara sampling); and 2.5. Other. and 156 hours of active search.

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.

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, 1992, 2000–2003, NA net NA NA NA
XI, XII 2007–2010, 2013
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, 2014, 2017–2019 22,464 trap h; 156 net/ net, trap, obs 48 banana understory
VI, X obs h

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Butterflies of Intervales

We classified species in subfamilies and used the phylogenetic (e.g., Actinote sp. Hübner, [1819], Adelpha sp. Hübner, [1819], etc);
hypotheses of Warren et al. (2008, 2009), Wahlberg et al. (2009, except from Autochton sp. Hübner, 1823, Ochlodes sp. Scudder, 1872,
2014), Seraphim et al. (2018), Li et al. (2019), and Dias et al. Emesis sp. Fabricius, 1807, and Mesene sp. E. Doubleday, 1847 because
(2019). We based the identification of specimens on the specialized no other species of the same genus were in the list.
literature (e.g. Brown 1992, Uehara-Prado et al. 2004, Warren et al. We reported the species lists by dataset (Table 2) and explored
2016, Lamas 2022), and on the collections of the ZUEC-LEP and how many and which species belong to each dataset, or are shared
MZUSP. These are also the entomological collections where we among datasets, with Venn diagrams (using online tool Venny, https://
housed the specimens collected by us, with material from LTS at bioinfogp.cnb.csic.es/tools/venny/). We also looked at the species
Unicamp (ZUEC–LEP 10.018–10.070, 10.253–10.262, 11.312– assembly (presence/absence) by sampling method and by region.
11.445) and ROS at MZUSP. Part of the Unicamp material (~10%) We then collapsed datasets in a single list to analyze richness (whole
has a pair of legs stored in EtOH and is available to the scientific data, as well as PEI traps: PEI trap, PEI fish, and PEI 1y datasets) and
community upon request. abundance (only PEI traps) per month and per season. For PEI traps,
We searched for information about the butterflies of the region by we separated the lists by datasets and divided the diversity metrics by
looking at the literature, asking specialists, and consulting a database the number of traps and trap days.
with the state of the art of species lists in Brazil (Shirai et al. 2019). Lastly, focusing on baited traps (PEI traps and AG traps), to
We found only three studies (Brown & Freitas 1999, Santos et al. understand how the richness distribution match what has been found
2015, Shirai et al. 2017), none of which had a species list for more in other trap studies in the Atlantic Forest, we used a database of
than a tribe. Other studies reported the collection of focal taxa at the Nymphalidae species lists in Brazil called DnB (Shirai et al. 2019,
PEI (e.g. Forsterinaria pronophila (A. Butler, 1867) in Freitas & Peña downloaded from https://doi.org/10.5281/zenodo.2561408), filtering
2006, Blepolenis bassus (C. Felder & R. Felder, 1867) in Penz et al. studies that exclusively used baited traps, exclusively at the Atlantic
2013, Taygetis ypthima (Hübner, [1821]) and T. rectifascia (Weymer, Forest. That is, we did not include studies of the DnB database that
1907) in Siewert et al. 2013, Godartiana muscosa (A. Butler, 1870) in used e.g. active searches or active search plus traps; or done in other
Zacca et al. 2016, T. acuta Weymer, 1910 in Freitas 2017, T. yphtima biomes or the Atlantic Forest plus other biomes, because the lists would
in Uehara-Prado & Freitas 2019), and the referred specimens belong not be strictly comparable. The DnB study used the same source for
to ZUEC-LEP and ZUEC-AVLF private collection. phylogenetic arrangement as ours (Wahlberg et al. 2009). In relation
Except from ZUEC-LEP, other large butterfly collections in to species identities, we filtered species considered valid by Lamas
the country did not have material from the PEI: MZUSP (ROS (2004) which, despite being outdated, standardizes the taxonomy to a
pers. obs.); DZUP (O. H. H. Mielke, pers. comm.); the MNUFRJ single source. To counterbalance this compromise, we also consulted
probably did not have material from the region but even if it had, it the largest database of fruit-feeding butterflies in the Atlantic Forest
would be reduced to ashes due to the fire in September 2018. In the (Santos et al. 2018), that has an updated taxonomy. Using Santos et al.
30 years and 300 research projects in the park, no foreign institution (2018) classification of tribes, we compared our totals per tribe with
has had an official project that collected butterflies (T. B. Conforti, their data (Figure 3 of Santos et al. 2018).
pers. comm.). Therefore, the museum data reported below belonged
solely to ZUEC-LEP (now MZUSP has sampled material by ROS, Results
and DZUP and MNRJ entomological collections also house PEI
specimens donated by us). In total, we found 312 species for PEI and surroundings (Table 2)
We consulted ZUEC-LEP by the online database available at the that sum to 2,139 records (1,605 in active search, 509 in traps, 25 other).
SpeciesLink website (www.splink.org.br) at the very beginning and The museum contributed with 432 records for 229 species. We sampled
end of this study, filtering by the collection (ZUEC-LEP) and the 1,682 specimens belonging to 142 species, in a total effort of 36,679
municipalities of and around PEI (Capão Bonito, Eldorado, Guapiara, sampling hours (36,432 trap and 247 net and observation hours, Table 1).
Iporanga, Ribeirão Grande, Sete Barras), returning results only for The richest families were Nymphalidae (148 species), the single
Ribeirão Grande and Sete Barras. We searched the Nymphalidae family caught by all sampling methods, and Hesperiidae (100),
specimens with missing IDs in the database and checked every drawer followed by Riodinidae (23 species), Lycaenidae (19 species), Pieridae
of the collection to update the online database by including their identity. (14 species), and Papilionidae (8 species). Within nymphalids, the
richest subfamily was Satyrinae (58 species), followed by Danainae
3. Data treatment
(23), Heliconiinae (18), Biblidinae (16), Nymphalinae (13), Charaxinae
We counted the number of individuals found at the ZUEC-LEP, (12), Limenitidinae (6), and Apaturinae (2). Within skippers, the richest
those sampled by us under active searches and other sources (here subfamily was Hesperiinae (54 species), followed by Pyrginae (24),
referred as records), as well as under baited traps designs (here Eudaminae (16), Pyrrhopyginae (4), and Tagiadinae (2). None of the
referred as abundance). The number of records and abundance are species were among the 63 threatened (that is, critically endangered,
not comparable because specimens deposited in a museum, sampled endangered, or vulnerable) butterfly species (ICMBio 2018, updated
with the net by different observers, or observational records (such as by Augusto H. B. Rosa).
citizen science) do not reflect biological abundance. We also counted The great majority of species was sampled exclusively by active
the number of species (richness), excluding genera not identified to methods (79.8%), with 18.3% having been collected by both active
the species level due to taxonomic issues (e.g., Hermeuptychia sp. and passive methods, and only 1.9% sampled in traps only (Figure 2a).
Forster, 1964, Tan et al. 2021) or museum specimens identified as such Additionally, the citizen science, iNaturalist and Santos et al. (2015)

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Table 2. Butterfly species of PEI and surroundings per dataset. Museum data: PEI ZUEC (Parque Estadual Intervales in the ZUEC collection) and PENaP (Parque Es-
tadual 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 collec-
tion dates, only III–IV. This table is available at 10.5281/zenodo.7429126.
PEI PEI PEI
PENaP PEI AG PEI AG Month
Family Subfamily Tribe Taxon Species description ZUEC fish 1y other
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, 1 I
2002
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
Continue...
...Continuation

PEI PEI PEI

PENaP PEI AG PEI AG Month
Family Subfamily Tribe Taxon Species description ZUEC fish 1y other
caught
entomological net baited traps
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) (Linnaeus, 1758) 1 IV
arsalte
Hesperiidae Pyrginae Heliopetes (Heliopetes) J. Zikán, 1938 1 1 VII, XII
ochroleuca
Hesperiidae Pyrginae Heliopetes (Heliopyrgus) (Blanchard, 1852) 1 III
americanus
Hesperiidae Pyrginae Heliopetes (Leucoscirtes) (A. Butler, 1870) 1 IV
omrina
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, x NA
1867)
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) (Latreille, [1824]) 2 I
versicolor
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. NA x NA
chloris
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
Continue...
...Continuation

PEI PEI PEI

PENaP PEI AG PEI AG Month
Family Subfamily Tribe Taxon Species description ZUEC fish 1y other
caught
entomological net baited traps
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, 41 17 II, V, IX, XI
1867)
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 (Fabricius, 1775) 1 2 1 1 I to III, V
amphimachus
Nymphalidae Charaxinae Archaeoprepona (Hübner, [1823]) 5 2 II, III, XII
chalciope
Nymphalidae Charaxinae Archaeoprepona (Linnaeus, 1758) 2 4 1 I, II, XI
demophon
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, 1 III
1860)
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
Continue...
...Continuation

PEI PEI PEI

PENaP PEI AG PEI AG Month
Family Subfamily Tribe Taxon Species description ZUEC fish 1y other
caught
entomological net baited traps
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, 5 1 I, II
1867)
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. NA 10 1 III, IV, VI,
hermes VII, XII

Continue...
...Continuation

PEI PEI PEI

PENaP PEI AG PEI AG Month
Family Subfamily Tribe Taxon Species description ZUEC fish 1y other
caught
entomological net baited traps
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. NA 6 18 V, VIII, IX,
boliviensis 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 & 1 II
O.H.H. Mielke, 2012
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 + 165 64 53 227 25
19
15

Total richness 195 60 92 64 31 20 21 40 23

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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).

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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.
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 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
euptychidia (A. Butler, 1868). Except from E. orea and F. quantius, line, and abundance, in solid line, per trap and per trap day) reporting them by
the remaining seven species also ranked among the 15 most abundant (b) month, and (c) by season.
species the trap sampling.
Among trap datasets, we caught 505 individuals of 63 species.
The richest and most abundant subfamily was Satyrinae, followed by Discussion
Charaxinae, Biblidinae, and Nymphalinae (Table 3). We found that
51.3% of the individuals belong to seven species: Morpho epistrophus Butterflies are excellent model organisms for scientific research
(Fabricius, 1796) (74), Memphis appias (Hübner, [1825]) (60), (reviewed in e.g. Brown & Freitas 1999, Santos et al. 2008, Sourakov &
G. muscosa (44), F. necys (24), Fountainea ryphea (Cramer, 1775) Shirai 2020) and for conservation purposes (see Introduction). However,
(20), M. orsis (19), and P. euptychidia (18). On average, we caught 0.19 even considering they are among the best-studied insects, important
individuals/trap/day in the PEI (0.34), PEI-fish (0.22), PEI 1y (0.13) and gaps in basic information still exist, such as for species description and
AG (0.09) datasets. The comparison between PEI habitats and vertical mapping species distributions (Linnean and Wallacean shortfalls c.f.
stratification using the PEI and PEI fish datasets can be found in the Hortal et al. 2015). That results in the exclusion of butterflies, as well
Supplementary Material. as other invertebrates, from studies like those that established hotspots

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Shirai L.T. et al.

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).

Richness Abundance (our study)/records

Taxa
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

(Myers et al. 2000) or other global conservation efforts (Bossart & we caught are found in open areas across the country but important
Carlton 2002, Barua et al. 2012), even considering that butterflies are species are expected to be caught only at the higher altitudes in the
a bioindicator, flagship, and umbrella group. In this study, we aimed park, above 800 m a.s.l. (Francini et al. 2011).
at filling this gap by inventorying butterfly diversity in a region within Focusing on nymphalids, a review of species lists in Brazil (Shirai
the Paranapiacaba Continuum, a key network of protected areas for the et al. 2019) listed the presence of 162-315 butterfly species in the top-
preservation of the Atlantic Forest hotspot. We also provided several 10 richest places in the country. The corresponding biomes (Amazon,
resources to aid conservation efforts (see Introduction), and a discussion Atlantic Forest and one in the Cerrado) rank among the richest because
for both academics and non-academics. 1) they are biologically rich but also, and importantly, 2) they were well-
Most of butterfly richness is in the tropics. While Nymphalidae sampled (Shirai et al. 2019). The most complete dataset of a subset of
diversified in the Asian, African and American tropics, several lineages Nymphalidae, fruit-feeding butterflies, listed 279 species for the whole
of Hesperiidae and Riodinidae adaptive radiations happened exclusively Atlantic Forest (Santos et al. 2018), which is a substantial increment
in the Neotropics (Toussaint et al. 2018, Seraphim et al. 2018), with over previously available data, such as the 88–127 fruit-feeding species
metalmarks being mainly Amazonian. Thus, it is not surprising that found in the same biome (Brown 2005). We were able to record 148
the richest families we found were Nymphalidae and Hesperiidae, nymphalids (61 fruit-feeding), but these numbers as well as our total of
frequently reported as the most diverse in the Atlantic Forest (Brown 312 butterfly species are certainly not the true richness of the PEI and
& Freitas 2000, Francini et al. 2011). Hesperiidae is probably the surroundings. The richness in the region could easily surpass 500 species
richest butterfly family in Brazil and the fact that we caught them less or more, as compared to similar areas in the same region (Francini
than Nymphalidae probably reflects the suggestion of Francini et al. et al. 2011). More sampling, and sampling by different strategies (like
(2011): in relatively complete inventories, the small and inconspicuous comparing to an estimate of the “true richness” by maximized sampling,
Hesperiidae are better sampled, but in short term studies, Nymphalidae Uehara-Prado et al. 2007), are necessary to reflect the diversity of this
appears with higher richness because they are easily captured, with both region. Despite the attempt to combine different datasets to enhance
net and traps. Lycaenidae and Riodinidae, despite having less species in this inventory, some obvious gaps remained, such as 1) a thorough
the Atlantic Forest than the two families above, equally suffer in shorter expedition to the other PEI station at the East lowlands (Saibadela),
inventories by being harder to catch. Moreover, most Pieridae species that has a different climate and altitude (Morellato et al. 2000); 2) the

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most inaccessible and pristine region of the PEI, at the South, as well as The best months to collect in South Eastern Brazil are March to
the few but important high altitude sites in the park; 3) the contiguous May (end of the rainy season, Ebert 1970, Brown 1972, 1992, Freitas
protected areas PECB, PETAR and Xitué (see Figure 1); and 4) more et al. 2014) but other seasonal intervals are also relevant. Here, we
sampling with different baits. chose when to collect aiming to span different seasons and months,
The use of different sampling methods certainly adds to more complementing the effort from museum data at the same time as
comprehensive inventories (e.g. Clench 1949, 1979, Brown 1972, revisiting good months (like December and March) and we indeed
Brown & Freitas 2000, Iserhard et al. 2013, Freitas et al. 2014, Checa found the highest diversity metrics in warmer months, particularly
et al. 2018). For example, most of the species we found were sampled from March to May (Figure 4).
by active methods (Figure 2a), collected by highly experienced people, We thus reinforce a previous suggestion (Brown 1972, Uehara-
during different years. However, despite having caught only about a Prado et al. 2007) that the best sampling months in the Atlantic
fifth (63/312 species) of the total richness with traps, Siderone galanthis Forest should be extended to the period between December to May
(Cramer 1775) was among the six trap-exclusive species (Figure 2a), (summer plus end of rainy season). The summer months of December
caught by a recently trained PEI guide (“PEI 1y” dataset). Decinea to February are mandatory for tribes like Morphini and Brassolini that
dama (Herrich-Schäffer 1869) was also among the trap-exclusive only fly in this period (Freitas et al. 2014). Here, we found 79% of
species, only caught with fish carrion (“PEI fish” dataset). It is also brassolines and morphines within the months of December and April.
worth mentioning that the citizen science approaches and a field course However, winter months should not be completely disregarded since
experiment (“other” dataset) contributed with eight species not found some species were exclusively caught in this season and, in the case
with neither active nor passive methods. of the PEI, it is the only season when an Epityches eupompe (Geyer,
Including efforts of other places than the PEI also enhanced the 1832) aggregation can be observed (Shirai et al. 2017, https://youtu.
inventory for the region adding, in our case, 33 species (20 exclusive of be/bUO4kpYS2uo).
PENaP, 11 exclusive of AG, plus two in common, Figure 2b) not found For Brazilian rainforests standards, the PEI is chilly, partly because
at the park, that had much more sampling effort (Table 1). Several of it has a lot of forest and in high altitudes, but also because it has a cold
the 20 species common to all regions (Figure 2b) are associated with winter: the annual average temperature ranged from 15.1 to 19.2°C
open or fragmented areas; we illustrated them (Figure 3) because they (Table S5), similar to what is reported in other sources (17.5°C from
are, aside from beautiful, colorful and diverse, likely the species that WorldClim and 17–18°C from Leonel 2010). More importantly, the
any person will observe at the region. temperature is seasonal, with a warm summer and a cold winter (Figure
An interesting statistic, both biologically and in terms of planning, is S1, Table S5): maximum temperatures of the warmest month (26–27°C)
the number of individuals caught per trap per day, as the 0.5 individuals/ and minimum temperature of the coldest month (7.5–11°C), also like
trap/day reported in an Amazonian terra firme site (Ribeiro & Freitas the WorldClim data.
2012). Traps at open and/or fragmented habitats tend to catch more The dominant species in our trap datasets was Morpho epistrophus
individuals, but not species (e.g. Uehara-Prado et al. 2007). We caught (15%), the only Morpho species in the Atlantic Forest that is iridescent
0.19 individuals/trap/day, but the highest values were not necessarily at white (Pablos et al. 2021, Figure 5) – caught on traps with black mesh
the open, disturbed or fragmented sites, such as PEI 1y (0.13) and AG (see Freitas et al. 2014). Although we caught them in traps only from
(0.09) datasets, but rather at the PEI (0.34) and PEI-fish (0.22). It would February to April (many of which with worn wings, LTS pers. obs.), this
be interesting to gather more data like this to uncover differences, for large butterfly is visible anywhere in the park since November. Butterflies
instances, between environments and biomes. are good flagship invertebrates because of their aesthetic appeal which, in

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.

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Shirai L.T. et al.

turn, relates to species attributes of size and brightness/color (Barua et al. M. C. Aguiar, André R. Nascimento, and Joel Lastra-Valdés for their help
2012). The iridescent white color of M. epistrophus might seem dull to during field sampling; Patrício A. Salazar for designing and performing the
some visitors, but the fact that tourists in Neotropical forests are somewhat preliminary analysis of the carrion test with LTS; Jessie P. dos Santos for
used to seeing, and being amused by, the blue morphos, might raise interest teaching LTS how to use the slingshot; Wesley R. Silva for kindly lending
to their phylogenetic relationship, enhancing the color feature. Another his car in the first field trip; Thamara Zacca for identifying many Satyrinae,
important aspect of butterflies as invertebrate flagships is the fact that they Ronaldo B. Francini for identifying the Actinote; Augusto H. B. Rosa and
are harmless. The elegant flight of M. epistrophus does not incite danger Olaf H. H. Mielke for museum information and access; Marcelo Duarte
or disgust like wasps or mosquitoes might do. Although caterpillars are for allowing LTS to access the MZUSP collection; Tamara M. C. Aguiar,
not always understood as an early stage of butterflies (Barua et al. 2012), Artur N. Furegatti, and Michela Borges for their assistance at the ZUEC-
they can trigger fear or dislike because of damage to crops. M. epistrophus LEP collection; Almir C. Almeida for allowing with such enthusiasm
harmless caterpillars are large, gregarious, bright red, and visible in many to use his photographs; and Renato R. Ramos, Eduardo P. Barbosa
Inga Mill. tree trunks near the PEI reception (Figure 5). The contrasting and Junia Y. O. Carreira for remote assistance during the COVID-19
effect of beauty and fear/disgust of these caterpillars could be used as lockdown. LTS acknowledges FAPESP (14/23504-7) for a fellowship,
an opportunity: the particularly skilled, enthusiastic and charismatic and ALCR thanks CNPq (166036/2020-0). AVLF thanks (FAPESP)
PEI staff could educate visitors about holometabolous life cycle, what (2021/03868-8) and the Conselho Nacional de Desenvolvimento
makes a caterpillar harmful or harmless, what are aposematic colors, Científico e Tecnológico (CNPq) (421248/2017-3, 304291/2020-0).
and warn visitors about moth caterpillars found at PEI (like Megalopyge This study was done under environmental licenses ICMBio 8585-1
Hübner 1820) that actually sting. Thus, the easy identification, large size, (sol. 10438), IBAMA 02001.000480/2014-38 (820/2017), and COTEC
omnipresence in the park, relationship with famous blue morphos, elegant 260108-004.611/2016. The SisGen registry is AE4A636. All authors
flight, and inciting caterpillars make M. epistrophus eligible to become declare no conflict of interest.
the emblematic invertebrate mascot of the PEI.
Overexploitation of resources (like the juçara palm), illegal hunting Associate Editor
and mining are permanent issues that endanger threatened species (such
as the jaguar, Beisiegel & Nakano-Oliveira 2020) and a threatened Gustavo Graciolli
biome (Tabarelli et al. 2005, Maxwell et al. 2016, ICMBio 2018).
Even in the largest and strongest continuous of Atlantic Forest, we Author Contributions
face the danger of (irreversibly) losing species and natural resources.
More involvement and investment in the area would result in much Leila T. Shirai: contribution to data analysis and interpretation;
more for the people and the planet than e.g. the end goals of several contribution to critical revision, adding intellectual content; read and
“perverse subsidies” (Myers 1998, Tabarelli et al. 2005). For example, approved the manuscript; substantial contribution to the conception
investing in a geographically wider and temporally longer sampling of and design of the study; contribution to data collection; contribution
butterflies, together with more science outreach seminars and material, to manuscript preparation.
capacity development and teaching in local schools are ideas that need Renato O. Silva: contribution to data analysis and interpretation;
dedicated, almost exclusive, time of trained people but, surprisingly, contribution to critical revision, adding intellectual content; read and
they do not require a substantial financial investment (see Shirai et al. approved the manuscript; substantial contribution to the conception and
2022). Another better idea, though, would be to invest in bioliteracy design of the study; contribution to data collection.
(c.f. Janzen 2010), which is a strong chance to save tropical diversity. Fernando M. S. Dias: contribution to data analysis and interpretation;
contribution to critical revision, adding intellectual content; read and
Supplementary Material approved the manuscript.
André L. C. Rochelle: contribution to data analysis and
The following online material is available for this article: interpretation; contribution to critical revision, adding intellectual
content; read and approved the manuscript.
A – Full description of our data. André V. L. Freitas: contribution to data analysis and interpretation;
B – Climatic data from the weather station at the PEI, in English contribution to critical revision, adding intellectual content; read and
and in Portuguese. approved the manuscript; substantial contribution to the conception
C – PEI trap datasets: habitats and vertical stratification in the PEI. and design of the study.

Acknowledgments Conflicts of Interest

This work is dedicated to Benedito Amaral, a highly skilled citizen The authors declare that they have no conflict of interest related to
scientist and fine observer of the natural world. We would like to thank: the publication of this manuscript
Benedito Amaral, Thiago B. Conforti, Eliseu C. de Paula, Faustino A.
Ribeiro, José Floido, Mara C. F. Paiva, Irene A. Ribeiro, Zarife O. Mora, Ethics
and the remaining PEI staff for their outstanding contribution; Benedito
Amaral, Massuo J. Kato, Lydia F. Yamaguchi, Mariana A. Stanton, This study did not involve human beings and/or clinical trials that
Dimitre Ivanov, Sidneia C. C. do Nascimento, Adilson R. Moreira, Tamara should be approved by one Institutional Committee.

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Data Availability DAVIS, A.K. 2015. Conservation of monarch butterflies (Danaus plexippus)
could be enhanced with analyses and publication of citizen science tagging
data. Insect Conserv. Divers. 8:103–106.
The primary data is reported in the main text as Table 2 and
DIAS, F.M.S., SIEWERT, R.R., FREITAS, A.V.L., LAMAS, G., MAGALDI,
is also available at the public repository https://doi.org/10.5281/ L.M., MIELKE, O.H.H. and CASAGRANDE, M.M. 2019. An integrative
zenodo.7429126, with the metadata in the readme file available at approach elucidates the systematics of Sea Hayward and Cybdelis Boisduval
https://doi.org/10.5281/zenodo.7439430. (Lepidoptera: Nymphalidae: Biblidinae). Syst. Entomol. 44:226–250.
EBERT, H. 1970. On the frequency of butterflies in Eastern Brazil, with a list of
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