Abstract
Among the Atlantic Forest areas in Brazil, the portion north of the São Francisco River is the least studied regarding butterfly fauna. In this study, we present a list of species recorded in the Pedra Talhada Biological Reserve, built from five collection expeditions. A total of 241 species from seven families were identified, including 87 new records for the Alagoas state and 22 new records for the Atlantic Forest north of the São Francisco River. Notably, Scada karschina delicata Talbot, 1932, a critically endangered subspecies of high conservation interest, was recorded. This inventory of the Pedra Talhada Biological Reserve, a key conservation area in the region, is the first step to the conservation of local butterfly diversity.
Keywords
Borborema Plateau; Brejos de Altitude; Pernambuco Endemism Center; inventory; Insecta
Resumo
Dentre as áreas de Mata Atlântica no Brasil, a porção norte do rio São Francisco é a menos conhecida em relação à fauna de borboletas. Neste estudo, apresentamos uma lista das espécies registradas na Reserva Biológica de Pedra Talhada construída a partir de cinco expedições de coleta. No total, foram identificadas 241 espécies pertencentes a sete famílias, incluindo 87 novos registros para o estado de Alagoas e 22 novos registros para a Mata Atlântica ao norte do São Francisco. Destaca-se a presença de Scada karschina delicata Talbot, 1932, uma subespécie criticamente ameaçada de extinção, de relevante interesse para a conservação. Este inventário da Reserva Biológica de Pedra Talhada, uma área de conservação fundamental para a região, representa o primeiro passo para a preservação da diversidade local de borboletas.
Palavras-chave:
Planalto da Borborema; Brejos de Altitude; Centro de Endemismo de Pernambuco; levantamento; Insecta
Introduction
The Atlantic Forest is one of the most biodiverse regions on the planet and is also among the most threatened due to its history of deforestation and overexploitation (Myers et al. 2000). As a result, its landscape is highly fragmented, with only 8.5% of the remaining forest patches larger than 100 hectares (SOSMA & INPE 2024). The situation is even more critical in its portion north of the São Francisco River, known as the “Northern Atlantic Forest” or “Pernambuco Center of Endemism” (sensuBrown Jr. 1979), due to intense deforestation and the gap of knowledge about its biodiversity (Filho et al. 2023).
Knowledge about even well-studied organisms, such as butterflies, is still limited in the Northern Atlantic Forest (hereafter NAF), highlighting the scarcity of research on the local fauna (Santos et al. 2008, Shirai et al. 2019, Freitas et al. 2023). There are a few species lists for the five states of the NAF (Rio Grande do Norte, Paraíba, Pernambuco, Alagoas, and Sergipe), some being historical (D’Almeida 1935, Cardoso 1949, Kesselring & Ebert 1979) and others more recent (Nobre et al. 2008, Paluch et al. 2011, Kerpel et al. 2014, Rafael et al. 2017, Melo et al. 2019a, Medeiros et al. 2021, Gualberto et al. 2024), and among the recent published inventories, several are very partial or based on a low sampling effort. For the state of Alagoas, for example, except for the comprehensive inventory of Maceió and surroundings (Cardoso 1949, 1983), the remaining information is preliminary or restricted to some specific butterfly groups (Freitas 2003, Melo et al. 2019b, Godé & Nobre 2015, Bezerra et al. 2021).
Accessing the butterfly fauna in under-sampled regions is essential as insects play a crucial role in ecosystem services (Janzen 1987, Iserhard et al. 2018) and serve as important indicators of habitat integrity due to their sensitivity to changes and high specialization in specific resources (Brown Jr & Freitas 2000). Additionally, understanding species diversity and distribution is vital not only as a foundation for further research but also to support and guide conservation planning efforts (Santos et al. 2008, Freitas et al. 2023). Therefore, this study aimed to obtain a list of the butterfly species of the Pedra Talhada Biological Reserve, to enhance our understanding of the fauna in Alagoas and, consequently, of the Northern Atlantic Forest.
Material and Methods
1. Study area
The Pedra Talhada Biological Reserve is located in northeastern Brazil, within the Atlantic Forest domain, spanning the states of Alagoas and Pernambuco. Most of the reserve lies in Alagoas, primarily within the municipality of Quebrangulo, with additional areas extending into Chã Preta (Alagoas) and Lagoa do Ouro (Pernambuco). It covers approximately 4,300 hectares and is geographically bounded at the coordinates 9°11’18”S 36°25’37”W (north), 9°13’51”S 36°23’52”W (east), 9°15’40”S 36°25’28”W (south) and 9°13’59”S, 36°27’33”W (west).
Situated in the “Agreste”, a transition zone between the coastal Atlantic Forest and the semiarid Caatinga of northeastern Brazil, this reserve features montane forest characteristics known as “Brejos de Altitude” or “Brejos Nordestinos” (Andrade-Lima 1982, Tabarelli & Santos 2004). Its vegetation is classified as seasonal semideciduous forest, with an average temperature of 22–24°C and an annual precipitation of 1,000–1,300 mm (Alvares et al. 2013). The reserve’s altitude ranges from 459 to 883 meters above sea level (Tscharner et al. 2015), encompassing a diverse array of microhabitats, including wetlands, dry areas, dense forests, and open fields (Figure 1). This environmental heterogeneity plays a crucial role in shaping the region’s rich biodiversity and ecological significance.
Landscapes and environments at Pedra Talhada Biological Reserve and surrounding areas, Quebrangulo, Alagoas. A-C) General view of Pedra Talhada and surrounding the forest; D) General view of the forest; E) Closer view of the forest; F) Open and flooded area; G) Interior view of the forest; H) Water stream inside the forest. Photos: A, B, D, E, and G courtesy of André César Lopes da Silva.
2. Sampling
The list was developed with data from five collection expeditions carried out from November 17 to 27, 2018; June 3 to 8, 2019; August 23 to 27, 2023; November 26 to December 3, 2023; and December 1 to 8, 2024. Consequently, the expeditions conducted in August, November, and December took place during the predominance of the dry season, whereas the June expedition occurred near the end of the rainy season (Marques et al. 2020).
Most of the sampling and observations occurred in the southern area of the reserve due to the availability of trails and the proximity to the research station. Sampling methods included the use of entomological nets during active searches, Van Someren-Rydon traps and light traps consisting of a white sheet and a 250 W mixed lamp.
Active searches involved trail walks conducted between 8:00 AM and 5:00 PM, totaling 252 hours of sampling effort. The trails, approximately 3 km long and ranging in altitude from 510 to 730 m, traversed a variety of microhabitats, with particular attention given to areas such as forest edges, clearings, flower fields, vegetation near water bodies and wetlands, where butterfly activity is highest (Brown Jr 1972, Freitas et al. 2021). Each trail was surveyed by at least three collectors, spaced apart to maximize coverage. Immature stages (eggs, caterpillars, and pupae) found in the field were reared in the laboratory to confirm identification with adults (see Greve et al. 2023). Photographic records of immature and adult stages were made available in the project “Borboletas da Rebio Pedra Talhada” on the iNaturalist citizen science platform (Kaminski 2025).
In two expeditions, six Van Someren-Rydon traps were used for four days each, and in two others, ten traps were fixed from six to ten days, totaling the sampling effort to 168 trap-days. The bait consisted of banana fermented with sugarcane juice for at least 48 hours before use (Freitas et al. 2014). During each expedition, traps were installed in the understory of the forest, at a height of 1.5 meters above the ground. The traps were set up on the first day and retrieved on the last. To sample nocturnal butterflies (Hedylidae), a white sheet with a mixed 250 W light was used between 17h30 to 22h00.
Collected specimens were deposited in three collections: 1) Entomological Collection at the Department of Systematics and Ecology (DSEC), Universidade Federal da Paraíba, João Pessoa, Paraíba, Brazil; 2) Entomological Collection of the Museu de História Natural da UFAL (MHNUFAL), Maceió, Alagoas, Brazil; 3) Zoological Collection of the Museu de Diversidade Biológica (ZUEC), Universidade Estadual de Campinas, Campinas, São Paulo, Brazil. Species identification was carried out using scientific literature, previously identified specimens in the DSEC - ZUEC collection, consultations with specialists, and online resources such as the Butterflies of America website (Warren et al. 2025).
Results and Discussion
In total, 241 butterfly species were recorded in the Pedra Talhada Biological Reserve (Table 1). The family with the highest species richness was Nymphalidae (n = 93, 38.59% out of total) followed by Hesperiidae (n = 80, 33.19%), Lycaenidae (n = 26, 11.79%), Riodinidae (n = 19, 7.88%), Pieridae (n = 17, 7.05%), Papilionidae (5 = 2.07%) and Hedylidae (n = 1, 0.41%). A considerable number of new records for Alagoas were found, totaling 87 out of the 241 identified species (36.1%). Of the 241 species recorded, 46 (19.1%) belong to the frugivorous guild and had the capture facilitated by Van Someren-Rydon traps. This proportion is consistent with similar studies in the region, such as Gualberto et al. (2024), which documented 47 frugivorous species, representing 22.17% of the total recorded.
Butterflies’ species list of Pedra Talhada Biological Reserve. (*) New records for the state of Alagoas; (**) new records for the NAF (Northern Atlantic Forest); (†) taxon present in NAF in an unpublished list by Heinz Ebert (see Freitas et al. 2023); (£), species illustrated in Figure 2.
The total richness of 241 species is comparable to that obtained in other well-sampled sites in the NAF (see table 4.1 in Freitas et al. 2023). There is substantial evidence, however, suggesting that the current list is likely to grow with additional sampling. The first is related to the total richness of each butterfly family. It is well known that in well-sampled sites in the Atlantic Forest, the richness of Hesperiidae should surpass that of Nymphalidae (Francini et al. 2011), contrary to the present study, where Nymphalidae is the most species-rich family. That means that at least 13 additional species of Hesperiidae (possibly more) could be present in the study site. The second is related to the absence of some common and widespread species, such as the Opsiphanes invirae (Hübner, [1808]) (Nymphalidae: Satyrinae) and Actinote pellenea auloeda Oberthür, 1917 (Nymphalidae: Heliconiinae), or species present in other similar localities in the northern Atlantic Forest, such as Callithomia lenea (Cramer, 1779) (Nymphalidae: Danainae), Amiga arnaca (Fabricius, 1776) (Nymphalidae: Satyrinae) and Dismorphia amphione (Cramer, 1779) (Pieridae: Dismorphiinae). Previous studies indicated that butterfly species richness can be predicted using specific subgroups that serve as surrogates for total richness (Beccaloni & Gaston 1995; Brown & Freitas 2000b). Following these studies, the predicted richness for the study site based on the Ithomiini (4.3–4.6% of total fauna) and Nymphalidae (25–29% of total fauna) as surrogates would be between 347 and 372 species in the first case, and 320 and 372 in the second. Based on the evidence presented, the Pedra Talhada Biological Reserve is likely to have around 300 species or more with some additional sampling effort. This level of biodiversity would be comparable to that of João Pessoa and Recife, which are the two most thoroughly studied sites in the NAF (Kesselring & Ebert 1982, Melo et al. 2019a).
Based on the present list, 22 species were added for the NAF. Although recent inventories in the region reported fewer new records, this situation is usual in regions with low biological information. For instance, in the inventory of Gualberto et al. (2024) in two conservation units in Paraíba, seven new records were added for the NAF. Similarly, Melo et al. (2019), listing 273 species from an urban park in Recife, recorded only two new occurrences.
It is also important to highlight the work of Godé and Nobre (2015), conducted in the same locality, which recorded 69 butterfly species for the reserve. Although preliminary, their study represented the first inventory of the area and, notably, included moth species, something rarely done in most regional Lepidoptera surveys. Their work also provided new records for the Northern Atlantic Forest, such as the species listed and illustrated as Pterourus sp.1 and Adelpha sp.1, which are treated here as Pterourus scamander grayi (Boisduval, 1836) and Adelpha erotia erotia (Hewitson, 1847), respectively.
Four taxa recorded in the present study are worthy of mentioning: the first is the threatened Scada karschina delicata Talbot, 1932 (Nymphalidae: Ithomiini). This subspecies, listed as Critically Endangered (Freitas et al. 2020, MMA 2022), is quite common at the study site, making this Conservation Unit an excellent opportunity for conserving this beautiful small butterfly. The second is Aeria olena tiumaFreitas & Rosa 2021. This recently described subspecies occurs in only seven localities in the NAF and will possibly be listed as threatened in future assessments (Freitas et al. 2021). The third is the recently described Actinote palmarina Freitas, Magaldi, Kaminski & Paluch, 2025, (Nymphalidae: Heliconiinae) a butterfly known from only two localities in the NAF. Fourth, the “American moth-butterfly” Macrosoma majormacula Lévêque, 2007, a rare representative of the poorly documented family Hedylidae. The latter was collected exclusively using a nocturnal light trap on two days across all expeditions, yielding four specimens, all males. This highlights the importance of this collection method for capturing these butterflies and emphasizes the need for its application in future faunal inventories.
It was also observed that during the dry season sampled (August, November and December months), S. karschina delicata and other several members of the tribe Ithomiini congregated in the so called ithomiine pockets (Collenette and Talbot 1928, Brown and Benson 1974, Pinheiro et al. 2008). These aggregations consist of limited physical spaces where various species seem to coexist in large abundance and appear to be associated with bodies of water, such as rivers and springs. In addition to S. karschina delicata, other ithomiines such as Dircenna dero (Hübner, 1823), Episcada hymenaea hymenaea (Prittwitz, 1865), Heterosais edessa (Hewitson, [1855]), Hypothyris ninonia daetina (Weymer, 1899), Ithomia spp. (Figure 2L), Scada reckia (Hübner, [1808]), Napeogenes inachia grazielaeFreitas, 2020 (Figure 2I) and Mechanitis nesaea Hübner, [1820]) were also observed in these same pockets.
Sample of representative butterflies taxa present at Pedra Talhada Biological Reserve and surrounding areas, Quebrangulo, Alagoas. A) Mesosemia minos tetrophthalma Stichel, 1915; B) Eurybia gonzaga Dolibaina, Dias, Mielke & Casagrande, 2014; C) Chamaelimnas tircis tircis C. Felder & R. Felder, 1865, (Riodinidae); D) Strephonota tephraeus (Geyer, 1837); E) Denivia hemon (Cramer, 1775); F) Strymon bubastus bubastus (Stoll, 1780), (Lycaenidae); G) Myscelia orsis (Drury, 1782); H) Junonia evarete evarete (Cramer, 1779); I) Napeogenes inachia grazielaeFreitas, 2020; J) Hypothyris euclea laphria (E. Doubleday, 1847); K) Scada karschina delicata Talbot, 1932; L) Ithomia sp.1; M) Pierella kesselringi Zacca, Siewert & Paluch, 2016; N) Magneuptychia libye (Linnaeus, 1767); O) Caligo illioneus (Cramer, 1775) (Nymphalidae); P) Cecropterus (Thorybes) dorantes dorantes (Stoll, 1790) (Hesperiidae); Q) Eurema (Abaeis) elvina (Godart, 1819) R) Eurema (Eurema) phiale paula (Röber, 1909) (Pieridae). Photos: A, E, J, O, and R courtesy of André César Lopes da Silva.
The findings of this study highlight the critical role of inventories in advancing butterfly research and conservation efforts. By providing essential data on species distribution, abundance, and habitat preferences, such surveys contribute to the delineation of occurrence areas, support biogeographical studies, and offer valuable material for taxonomic and systematic research. These efforts are vital for informing conservation strategies, managing biodiversity hotspots, and preserving the ecological integrity of areas like the NAF. As such, this study not only expands our understanding of butterfly diversity in Alagoas but also serves as a foundation for future research and conservation initiatives in the region.
Acknowledgments
We thank the fire brigade team of the REBIO Pedra Talhada for their support throughout the collection period. We thank Anita Studer and Luis Freitas in the name of Nordesta Association and to Jaílton Fernandes and all the staff of the Pedra Talhada Biological Reserve. We also thank Adalberto D. Medeiros and Olaf H. H. Mielke for his assistance in identifying some Hesperiidae, Robert K. Robbins for his assistance in identifying some Lycaenidae, and Fernando Dias for helping with the Charaxinae. LAK thanks Bruna S. S. Santos, Josivânia G. Silva and Rawelly O. Gonçalves for help in the fieldwork. We thank André César Lopes da Silva for the photos of live butterflies and environment. LAK acknowledges support from Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq 175508/2023-2) and Fundação de Amparo à Pesquisa do Estado de Alagoas (FAPEAL APQ2022021000087). AHBR thanks Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) - Finance code 001 and Conselho Nacional de Desenvolvimento Científico e Tecnológico – (CNPq grant 151203/2024-5). EPB thanks the Brazilian Research Council – Conselho Nacional de Desenvolvimento Científico e Tecnológico – (CNPq grant 174863/2023-3) for a post-doc fellowship. JYOC thanks the Brazilian Research Council – Conselho Nacional de Desenvolvimento Científico e Tecnológico – (CNPq grant 141443/2018-9). AVLF thanks the CNPq (grants 304291/2020-0 and 421248/2017-3) and the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP, grant 2021/03868-8). The present study is registered in the SISGEN (A8E47EE).
Data Availability
All the data is reported in the main text as Table 1.
References
-
ALVARES, C.A., STAPE, J.L., SENTELHAS, P.C., & DE MORAES GONÇALVES, J.L. (2013). Köppen’s climate classification map for Brazil. Meteorologische Zeitschrift, 22(6), 711–728. https://doi.org/10.1127/0941-2948/2013/0507
» https://doi.org/10.1127/0941-2948/2013/0507 - ANDRADE-LIMA, D. (1982). Present day forest refuges in Northeastern Brazil. In G. T. Prance (Ed.), Biological diversification in the tropics (pp. 245–254). Columbia University Press.
-
BECCALONI, G.W., & GASTON, K.J. (1995). Predicting the species richness of Neotropical forest butterflies: Ithomiinae (Lepidoptera: Nymphalidae) as indicators. Biological Conservation, 71(1), 77–86. https://doi.org/10.1016/0006-3207(94)00023-J
» https://doi.org/10.1016/0006-3207(94)00023-J -
BEZERRA, L.T., GONZAGA, E.P., LANA, M.D., BREDA, M.O., & DUARTE, M. (2021). Butterflies (Lepidoptera: Papilionoidea) in the Arboretum of Alagoas State, Brazil. Brazilian Journal of Animal and Environmental Research, 4(3), 3248–3268. https://doi.org/10.34188/bjaerv4n3-037
» https://doi.org/10.34188/bjaerv4n3-037 - BROWN Jr, K.S. (1972). Maximizing daily butterfly counts. Journal of the Lepidopterists’ Society, 26, 183–196.
- BROWN Jr, K.S. (1979). Ecologia geográfica e evolução nas florestas Neotropicais. Universidade Estadual de Campinas, Campinas, 265 p.
-
BROWN Jr, K.S., & FREITAS, A.V.L. (2000). Atlantic Forest butterflies: Indicators for landscape conservation. Biotropica, 32(4b), 934–956. https://doi.org/10.1646/0006-3606(2000)032[0934:AFBIFL]2.0.CO;2
» https://doi.org/10.1646/0006-3606(2000)032[0934:AFBIFL]2.0.CO;2 - CARDOSO, A. (1949). Lepidópteros de Alagoas. Revista de Entomologia, 20(1), 427–436.
- CARDOSO, A. (1983). Apreciações sobre a fauna de lepidópteros de Alagoas. Anais da Sociedade Nordestina de Zoologia, 1(1), 41–46.
-
COLLENETTE, C.L., & TALBOT, G. (1928). Observations on the bionomics of the Lepidoptera of Matto Grosso, Brazil. Transactions of the Entomological Society of London, 76(2), 391–416. https://doi.org/10.1111/j.1365-2311.1929.tb01413.x
» https://doi.org/10.1111/j.1365-2311.1929.tb01413.x - D’ALMEIDA, R.F. (1935). Lista dos lepidópteros capturados pelo Dr. RV Ihering no Nordeste do Brasil. Revista de Entomologia, 5(3), 326–328.
- FILHO, G.A.P., FRANÇA, F.G.R., ALVES, R.R.N., & VASCONCELLOS, A. (eds.). (2023). Animal Biodiversity and Conservation in Brazil’s Northern Atlantic Forest. Springer, Cham.
-
FREITAS, A.V.L. (2003). Lista preliminar das espécies de borboletas na Usina Serra Grande. Relatório com a primeira listagem das borboletas da mata de Coimbra, na Usina Serra Grande, AL. Available at: https://labbor.ib.unicamp.br/wp-content/uploads/2020/08/RT-Serra-Grande-AL-Relatório-preliminar.pdf. Accessed 03 Set 2025
» https://labbor.ib.unicamp.br/wp-content/uploads/2020/08/RT-Serra-Grande-AL-Relatório-preliminar.pdf. Accessed 03 Set 2025 -
FREITAS, A.V.L., SANTOS, J.P., ROSA, A.H.B., MELO, D.H.A., NOBRE, C.E.B., SANTOS, L.N., & CARDOSO, M.Z. (2023). The Butterfly Fauna of the Northern Atlantic Forest. In: FILHO, G.A.P., FRANÇA, F.G.R., ALVES, R.R.N., & VASCONCELLOS, A. (eds.). Animal Biodiversity and Conservation in Brazil’s Northern Atlantic Forest. Springer, Cham, p.33-46. https://doi.org/10.1007/978-3-031-21287-1_4
» https://doi.org/10.1007/978-3-031-21287-1_4 -
FREITAS, A.V.L. (2004). Lista preliminar das espécies de borboletas na RPPN Frei Caneca, Jaqueira, PE. Relatório Técnico com a primeira listagem das borboletas na RPPN Frei Caneca, Jaqueira, PE. Unicamp, Campinas. Available at: http://www.cepan.org.br (last access on 15/08/2025)
» http://www.cepan.org.br - FREITAS, A.V.L., ISERHARD, C.A., SANTOS, J.P., CARREIRA, J.Y.O., RIBEIRO, D.B., MELO, D.H.A., ROSA, A.H.B., MARINI-FILHO, O.J., ACCACIO, G.M., & UEHARA-PRADO, M. (2014). Studies with butterfly bait traps: an overview. Rev. Colomb. Entomol. 40(2):203–212.
-
FREITAS, A.V.L., ROSA, A.H.B., NOBRE, C.E.B., MELO, D.H.A., MOTA, L.L., SILVA-BRANDÃO, K.L., MACHADO, P.A., & CARREIRA, J.Y. O. (2020). Immature stages, natural history, systematics and conservation of two atan endangered Neotropical butterfly: the case of Scada karschina delicata (Nymphalidae: Ithomiini). Neotrop. Entomol. 49(5):685–695. https://doi.org/10.1007/s13744-020-00797-4
» https://doi.org/10.1007/s13744-020-00797-4 -
FREITAS, A.V.L., ROSA, A.H.B., BROWN, K.S., Jr., NOBRE, C.E.B., MELO, D.H.A., & KERPEL, S.M. (2021). A new and rare Ithomiini from northeastern Brazil (Lepidoptera: Nymphalidae: Danainae). J. Lepid. Soc. 75:44–48. https://doi.org/10.18473/lepi.75i1.a6
» https://doi.org/10.18473/lepi.75i1.a6 -
FREITAS, A.V.L., SANTOS, J.P., ROSA, A.H.B., ISERHARD, C.A., RICHTER, A., SIEWERT, R.R., GUERATTO, P.E., CARREIRA, J.Y.O., & LOURENÇO, G.M. (2021). Sampling methods for butterflies (Lepidoptera). p. 101–123. In: SANTOS, J.C. & FERNANDES, G.W. (eds.). Measuring Arthropod Biodiversity. Springer, Cham. xvii + 600 p. ISBN 978-3-030-53225-3. https://doi.org/10.1007/978-3-030-53226-0_5
» https://doi.org/10.1007/978-3-030-53226-0_5 - GODÉ, L., & NOBRE, C.E.B. (2015). Inventário XIX da Reserva Biológica de Pedra Talhada: Borboletas e Mariposas (Lepidoptera). In: STUDER, A., NUSBAUMER, L. & SPICHIGER, R. (eds.). Biodiversidade da Reserva Biológica de Pedra Talhada (Alagoas, Pernambuco - Brasil). Boissiera 68:683–692.
-
GREVE, R.R., CARNEIRO, E., MIELKE, O.H.H., ROBBINS, R.K., CALLAGHAN, C.J., & FREITAS, A.V.L. (2023). Butterflies (Lepidoptera: Papilionoidea) of Iguaçu National Park and surrounding areas in southern Brazil: a long-term survey, with six new records for the Brazilian fauna. Biota Neotropica 23(3): e20231487. https://doi.org/10.1590/1676-0611-BN-2023-1487 (last access on 15/08/2025)
» https://doi.org/10.1590/1676-0611-BN-2023-1487 -
GUALBERTO, E.P., MEDEIROS, A.D., & KERPEL, S.M. (2024). Butterflies of two Atlantic Forest conservation units from Paraíba state, northeast of Brazil. EntomoBrasilis 17:e1068. https://doi.org/10.12741/ebrasilis.v17.e1068
» https://doi.org/10.12741/ebrasilis.v17.e1068 - ISERHARD, C.A., UEHARA-PRADO, M., MARINI-FILHO, O.J., DUARTE, M., & FREITAS, A.V.L. (2018). Fauna da Mata Atlântica: Lepidoptera-borboletas. In: MONTEIRO-FILHO, E.L.A. & CONTE, C.E. (eds.). Revisões em Zoologia: Mata Atlântica. Ed. UFPR, Curitiba, p.57-102.
-
JANZEN, D.H. (1987). Insect diversity of a Costa Rican dry forest: why keep it, and how? Biological Journal of the Linnean Society, v. 30, n. 4, p. 343–356. https://doi.org/10.1111/j.1095-8312.1987.tb00307.x
» https://doi.org/10.1111/j.1095-8312.1987.tb00307.x -
KAMINSKI, L.A. (2025). Borboletas da Rebio Pedra Talhada. https://www.inaturalist.org/projects/borboletas-da-rebio-pedra-talhada (last access on 15/08/2025)
» ttps://www.inaturalist.org/projects/borboletas-da-rebio-pedra-talhada - KERPEL, S.M., ZACCA, T., NOBRE, C.E.B., FERREIRA JÚNIOR, A., ARAÚJO, M.X., & FONSECA, A. (2014). Borboletas do Semiárido: conhecimento atual e contribuições do PPBio. In: BRAVO, F. & CALOR, A. (eds.). Artrópodes do Semiárido: Biodiversidade e Conservação. Printmídia, p.245–267.
- KESSELRING, J., & EBERT, H. (1982). Relação de Borboletas encontradas na “Mata do Buraquinho”, João Pessoa, estado da Paraíba, Brasil. Rev. Nordestina Biol. 2(1):105–118.
-
MARQUES, T.V., MENDES, K., MUTTI, P., MEDEIROS, S., SILVA, L., PEREZ-MARIN, A.M., CAMPOS, S., LÚCIO, P.S., LIMA, K., DOS REIS, J., et al. (2020). Environmental and biophysical controls of evapotranspiration from Seasonally Dry Tropical Forests (Caatinga) in the Brazilian Semiarid. Agric. For. Meteorol. 287, 107957. https://doi.org/10.1016/j.agrformet.2020.107957.
» https://doi.org/10.1016/j.agrformet.2020.107957 -
MEDEIROS, A., GUALBERTO, E.P., RODRIGUES, R.P., & KERPEL, S.M. (2021). Butterflies (Lepidoptera: Papilionoidea) of the Restinga de Cabedelo National Forest, Paraíba State, Brazil. EntomoBrasilis 14:e970. https://doi.org/10.12741/ebrasilis.v14.e970
» https://doi.org/10.12741/ebrasilis.v14.e970 -
MELO, D.H.A., DUARTE, M., MIELKE, O.H.H., ROBBINS, R.K., & FREITAS, A.V.L. (2019a). Butterflies (Lepidoptera: Papilionoidea) of an urban park in northeastern Brazil. Biota Neotropica. 19(1): e20180614. http://dx.doi.org/10.1590/1676-0611-BN-2018-0614 (last access on 15/08/2025)
» https://doi.org/10.1590/1676-0611-BN-2018-0614 -
MELO, D.H.A., FILGUEIRAS, B.K.C., ISERHARD, C.A., IANNUZZI, L., FREITAS, A.V.L., & LEAL, I.R. (2019b). Effect of habitat loss and fragmentation on fruit-feeding butterflies in the Brazilian Atlantic Forest. Can. J. Zool. 97(7):588–596. https://doi.org/10.1139/cjz-2018-0202
» https://doi.org/10.1139/cjz-2018-0202 - Ministério do Meio Ambiente (MMA). (2022). Lista oficial da fauna brasileira ameaçada de extinção - Anexo II à Portaria Nº 148, de 7 de junho de 2022. Diário Oficial da União, Brasília, 08 de junho de 2022.
-
MYERS, N., MITTERMEIER, R.A., MITTERMEIER, C.G., DA FONSECA, G.A.B., & KENT, J. (2000). Biodiversity hotspots for conservation priorities. Nature 403(6772):853–858. https://doi.org/10.1038/35002501
» https://doi.org/10.1038/35002501 -
NOBRE, C.E.B., Schlindwein, C., & Mielke, O.H.H. (2008). The butterflies (Lepidoptera: Papilionoidea and Hesperioidea) of the Catimbau National Park, Pernambuco, Brazil. Zootaxa 1751(1):35–45. https://doi.org/10.11646/zootaxa.1751.1.3
» https://doi.org/10.11646/zootaxa.1751.1.3 -
PALUCH, M., MIELKE, O.H.H., NOBRE, C.E.B., CASAGRANDE, M.M., MELO, D.H.A., & FREITAS, A.V.L. (2011). Butterflies (Lepidoptera: Papilionoidea and Hesperioidea) of the Parque Ecológico João Vasconcelos Sobrinho, Caruaru, Pernambuco, Brazil. Biota Neotrop. 11(4):229–238. https://doi.org/10.1590/S1676-06032011000400020
» https://doi.org/10.1590/S1676-06032011000400020 -
PINHEIRO, C.E., MEDRI, Í.M., & SALCEDO, A.K.M. (2008). Why do the ithomiines (Lepidoptera, Nymphalidae) aggregate? Notes on a butterfly pocket in central Brazil. Rev. Bras. Entomol. 52:610–614. https://doi.org/10.1590/S0085-56262008000400012
» https://doi.org/10.1590/S0085-56262008000400012 - RAFAEL, J.A., OLIVEIRA, M.L., SOUZA, U.R.M., GALILEO, M.H.M., SILVA, A.S., MELLO, F.V., … LIMEIRA-DE-OLIVEIRA, F. (2017). Insetos dos Parques Nacionais de Sete Cidades e de Ubajara. In: Pesquisas em Unidades de Conservação no Domínio da Caatinga: Subsídios à Gestão. UFC, p.301–388.
- SANTOS, E.C.D., MIELKE, O.H.H., & CASAGRANDE, M.M. (2008). Butterfly inventories in Brazil: the state of the art and the priority-areas model for research aiming at conservation. Nat. Conserv. 6(2):178–200.
-
SHIRAI, L.T., MACHADO, P.A., MOTA, L.L., ROSA, A.H.B., & FREITAS, A.V.L. (2019). DnB, the database of nymphalids in Brazil, with a checklist for standardized species list. J. Lepid. Soc. 73(2):93–108. https://doi.org/10.18473/lepi.73i2.a4
» https://doi.org/10.18473/lepi.73i2.a4 - SOS MATA ATLÂNTICA & INPE. (2024). Atlas dos Remanescentes Florestais da Mata Atlântica no período 2022–2023. Fundação SOS Mata Atlântica.
- TABARELLI, M., & SANTOS, A.M.M. (2004). Uma breve descrição sobre a história natural dos brejos nordestinos. In: PORTO, K.C., CABRAL, J.J.P. & TABARELLI, M. (eds.). Brejos de Altitude em Pernambuco e Paraíba: História Natural, Ecologia e Conservação. Ministério do Meio Ambiente, p.39–58.
- TSCHARNERT, T., DUDA, G.P., OLIVEIRA, SILVA, C.M.S., NUSBAUMER, L., & FILHO, A.F.S. (2015). Parâmetros abióticos da Reserva Biológica de Pedra Talhada. In: STUDER, A., NUSBAUMER, L. & SPICHIGER, R. (eds.). Biodiversidade da Reserva Biológica de Pedra Talhada Alagoas, Pernambuco, Brasil. Boissiera, p.39–58.
- WARREN, A.D., DAVIS, K.J., STANGELAND, E.M., PELHAM, J.P., & GRISHIN, N.V. (2025). Illustrated lists of American butterflies. Available at: (last access: August 2025).




