Introduction

Neotropical Cerrado savannas cover extensive areas in northeastern Brazil, where they intermix with Caatinga dry forests, extending obliquely toward the southwest until reaching the boundaries of Pantanal and Chaco and, thereby, dividing two large, forested biomes, the Amazon and the Atlantic Forest (OLIVEIRA & MARQUIS, 2002). This region is considered an important biodiversity hotspot for combining a high rate of endemism and threats (MYERS et al., 2000). The area of Cerrado savannas has decreased in recent decades as a result of policies that prioritize the expansion of agricultural frontiers in Brazil (OLIVEIRA-FILHO, 2006; OLIVEIRA-FILHO & RATTER, 2002). Thus, effective public policies aiming at the conservation of Cerrado remnants are urgently needed to maintain biodiversity and, consequently, preserve ecosystem services (KLINK & MACHADO, 2005; STRASSBURG et al., 2017). For the proposal of new conservation actions and monitoring strategies, it is necessary to obtain information on Cerrado species, their geographical distributions and temporal patterns, and the ecology and natural history of groups occupying preserved Cerrado areas (BROWN & GIFFORD, 2002; CAVALCANTI & JOLY, 2002; PINHEIRO et al., 2010).

Biological inventories, that is, complete and well-represented lists of local species, are fundamental to the knowledge of the geographic distribution of organisms and development of environmental agendas, including conservation priorities and management programs (SANTOS et al., 2008; SILVEIRA et al., 2010). Insect inventories are prioritized because of the ecological relevance of these organisms to terrestrial and aquatic ecosystems. Insects are involved in several ecological processes, such as pollination, suppression of plant species through herbivory, predation of other insects and animals, and nutrient cycling; furthermore, insects are important components of virtually all food webs (MORSE, 1971; BARBOLA et al., 2007; POTTS et al., 2016; SCHOWALTER et al., 2018). Given such a close relationship with other organisms, several insect groups are used as indicators of anthropogenic disturbances (i.e., biological indicators), particularly Lepidoptera (BROWN, 1991; FREITAS et al., 2003; FREITAS et al., 2006).

Despite the growing knowledge of Brazilian Lepidoptera, inventories of Cerrado areas are still local, sparse, and underrepresented, mostly concentrated in the Central-West and Highlands (see BROWN & MIELKE 1967 a, b; BROWN & MIELKE 1968; PINHEIRO & ORTIZ 1992; EMERY et al., 2006; PINHEIRO & EMERY, 2006). Thus, there is a lack of information about butterflies occurring in southern Cerrado, transition zones between biomes, and elevated regions, in which Cerrado vegetation is replaced by natural, open, rocky montane vegetation, known locally as campo rupestre (see SOARES et al., 1999; GOZZI et al., 2012; NERY et al., 2014; PIRES et al., 2018; HENRIQUES et al., 2019; SOLDATI et al., 2019; FREITAS et al., 2021a). This survey provides an inventory of butterflies occurring in the Sempre Vivas National Park (SVNP), an area covered by Cerrado and campo rupestre vegetation in the north-central region of Minas Gerais State, Brazil.

Material and methods

The study was carried out in the Sempre Vivas National Park (SVNP) (17º55’S 43º47’W), a reserve that encompasses areas in the municipalities of Diamantina, Olhos D’Água, Bocaiúva, and Buenópolis, north-central Minas Gerais State, southeastern Brazil. The SVNP is located in the Serra do Espinhaço and extends over an area of approximately 124,156 ha, covered by a mosaic of different phytophysiognomies, such as riparian forest, seasonal semideciduous forest, several Cerrado physiognomies (such as Cerrado savannas sensu stricto, arboreal formations such as cerradão, and open vegetation such as campo limpo), and a high representation of campo rupestre. The climate is humid subtropical with dry winters and temperate summers (Cwb in the Köppen climate classification) (ÁLVARES et al., 2014). Altitudes range from 1200 to 1400 m a.s.l., and the annual precipitation is between 1300 and 1600 mm (ÁLVARES et al., 2014).

Samplings took place at 10 sampling sites over two periods: (1) two expeditions of five consecutive days each in April 2010 and May 2011 and (2) four expeditions of five consecutive days each in October 2018 and January, February, and March 2019 (Table I). Samplings were carried out between 9:00 a.m. and 5:00 p.m. (about 8 h of sampling effort per day) by active search using entomological nets, according to the protocols proposed by BROWN (1972), FREITAS et al. (2003), and FREITAS et al. (2021b), totaling 240 h of sampling effort over the two periods. Additionally, baited traps (Van Someren-Rydon traps) were used to attract fruit-feeding butterflies, following a protocol adapted from FREITAS et al. (2014). Ten traps were placed 20 m away from each other in areas of open vegetation, forest, and forest edges and left in the field for five consecutive days per expedition, totaling 480 h of sampling effort per trap. The bait used was a mixture of fermented banana and sugarcane juice, prepared 48 h before sampling. Collections were restricted to individuals whose identification in the field was difficult or doubtful, in which case the collected specimens were taken to the laboratory for later identification and deposited in zoological collections. Species identification was carried out by authors with experience in the field, supported by specialized literature and comparisons with the illustrated list on the webpage “Butterflies of America” (WARREN et al., 2016) and specimens previously identified and deposited in museums. The classification used follows the checklist published by LAMAS (2004), modified according to WARREN et al. (2009), CONG et al. (2019), LI et al. (2019), and ZHANG et al. (2019) for Hesperiidae; SERAPHIM et al. (2018) for Riodinidae; WAHLBERG et al. (2009) for Nymphalidae; and ZACCA et al. (2018) for Euptychiina. Collection, storage, and preparation procedures were performed according to FREITAS et al. (2021c). Six families of diurnal butterflies were sampled (family Hedylidae was not included). Collected specimens were deposited in one of the following three scientific collections: (1) Coleção Zoológica do Museu de Diversidade Biológica da Universidade Estadual de Campinas (ZUEC), Campinas, São Paulo, Brazil; (2) Centro Nacional de Pesquisa e Conservação da Biodiversidade do Cerrado e Caatinga do Instituto Chico Mendes de Conservação da Biodiversidade (CECAT), Brasília, Distrito Federal, Brazil; or (3) Coleção do Laboratório de Zoologia, do Instituto Federal de Educação, Ciências e Tecnologia do Sul de Minas (CBVS), Campus Inconfidentes, Inconfidentes, Minas Gerais, Brazil.

Table I.–

Sampling sites, coordinates, and phytophysiognomies where butterfly individuals were collected in the Sempre Vivas National Park, Minas Gerais, southeastern Brazil.

Results

A total of 1181 individuals belonging to 238 species were collected/recorded, distributed in six families of Lepidoptera (Appendix I). The family with the highest species richness was Hesperiidae, with 73 species (30.7%), followed by Nymphalidae (n = 65, 27.3%), Lycaenidae (n = 42, 17.7%), Riodinidae (n = 37, 15.5%), Pieridae (n = 15, 6.3%), and Papilionidae (n = 6, 2.5%) (Appendix I).

Of the Lepidoptera species inventoried in this study, three are included in the Brazilian Red List of threatened fauna (MMA, 2014): (1) Magnastigma julia Nicolay, 1977 (Lycaenidae) (endangered); (2) Strymon ohausi (Spitz, 1933) (Lycaenidae) (endangered); and (3) Rhetus belphegor (Westwood, 1851) (Riodinidae) (critically endangered). At least five species are endemic to the Cerrado biome: (1) Cogia cerradicola (Mielke, 1967) (Hesperiidae); (2) Parides bunichusdiodorus (Hopffer, 1865) (Papilionidae); (3) Nhambikuara cerradensis Freitas, Barbosa & Zacca, 2018 (Nymphalidae); (4) Yphthimoides cipoensis Freitas, 2004 (Nymphalidae); and (5) Sertania jaibensis (Callaghan & Soares, 2001) (Riodinidae).

Discussion

Although species richness was relatively low in the SVNP compared with other Neotropical sites (see BROWN 2005 and the discussion below), the present list can be considered representative of the SVNP Lepidoptera assemblage for the following reasons: (1) expeditions were well spaced in time, including periods characteristically known to have high species richness in the region and occurrence of univoltine species (see BROWN, 1972), (2) the team was composed of researchers with vast experience in butterfly inventories, and (3) the richness of Hesperiidae species was greater than that of Nymphalidae species, a characteristic also observed in complete inventories of Lepidoptera in the Atlantic Forest and Cerrado (FRANCINI et al., 2011; ISERHARD et al., 2017). In summary, although many additions of species are expected in all six families represented here, this list may adequately represent richness patterns of the SVNP Rhopalocera assemblage.

Inventories in forested biomes, such as the Atlantic Forest and Amazon, generally comprise more than 500 species (e. g., BROWN JR. & FREITAS, 2002; BROWN, 2005; MIELKE et al., 2010; FRANCINI et al., 2011; BELTRAMI et al., 2014; GARCIA-SALIK et al., 2014; FREITAS et al., 2016), as do inventories carried out in central Cerrado areas (EMERY et al., 2006; MIELKE et al., 2008). The species richness recorded here is comparable to that obtained in relatively wellsampled areas of southern Cerrado (CASAGRANDE et al., 2012), sites in ecotone with the Atlantic Forest (also known as transition zones) (BROWN & MIELKE, 1968; SOLDATI et al., 2019), and in similar areas of campo rupestre (PIRES et al., 2018; HENRIQUES et al., 2019). Some groups were clearly undersampled, and their richness would likely increase with some additional visits. For example, the subfamily Ithomiini (Nymphalidae: Danainae) had only three sampled species, but, according to other lists from the Cerrado, this number could increase to 15 or more. A good strategy is to visit riparian forests during the dry season, when Ithomiini species can usually be found in large groups in humid, shaded environments. Other poorly sampled groups were two genera of the family Nymphalidae, Adelpha (Limenitidinae) and Actinote (Heliconiinae), both with only one sampled species. These two genera are usually represented by eight or more species in similar habitats; the number of sampled species could increase with additional visits in March and April.

Finally, it is worth noting that, although richness was not very high, three threatened species were recorded in the SVNP, namely M. julia, S. ohausi, and R. belphegor, all associated with open formations of Cerrado and campo rupestre (FREITAS & MARINI-FILHO, 2011; KAMINSKI et al., 2015; ICMBio, 2018). By contrast, no threatened species were identified in the Baixada Santista region, São Paulo State, where more than 500 species were recorded (FRANCINI et al., 2011). Furthermore, some endemic species of the Cerrado were recorded, such as Sertania jaibensis, which was previously observed in only two localities (CALLAGHAN & SOARES, 2001; KAMINSKI et al., 2017). The Serra do Espinhaço region is very peculiar and important to several fauna and flora species, extending over a small area (compared to the Cerrado as a whole) from Minas Gerais to Bahia States. Thus, we highlight the importance of preserved areas in unique landscapes such as campo rupestre in the SVNP. This area, belonging to the Cerrado domain, preserves and maintains various populations of endangered and endemic species in Brazil, representing a crucial and singular ecosystem to the biodiversity of Brazilian elevated regions.

Taxonomic composition

As previously mentioned, the SVNP list reflects the richness distribution of Brazilian species lists, with Hesperiidae being more represented than Nymphalidae (BROWN & FREITAS, 1999). Such a family composition is evidence that the inventory constituted a representative sampling effort (FRANCINI et al., 2011). This is due to the fact that Nymphalidae has a greater proportion of species that are common and easily detectable, leading to a high accumulation of species in the first expeditions. By contrast, Hesperiidae, Lycaenidae, and Riodinidae show a slower rate of species accumulation, given that these individuals are generally furtive and low in abundance (BROWN & FREITAS, 2000; ISERHARD et al., 2013, 2017a). Thus, these groups demand greater collection effort and time to obtain a well-represented inventory (e.g., BROWN & FREITAS, 2000: Table 3; LAMAS et al., 2021). Inventories carried out with low sampling efforts tend to show greater Nymphalidae than Hesperiidae richness, although local richness is usually greater in the latter family.

A second pattern observed here was the relative proportions of Lycaenidae and Riodinidae. Previous studies suggested that Lycaenidae predominate over Riodinidae in colder sites of the Atlantic Forest, including southern localities and highlands (ISERHARD & ROMANOWSKI, 2004; ISERHARD et al., 2010; BELLAVER et al., 2012). Riodinidae, by contrast, is more frequent than Lycaenidae in lowlands, northern regions, and the interior of the Atlantic Forest (FRANCINI et al., 2011). Therefore, the SVNP inventory, carried out in a mountainous region with elevations of up to 1600 m a.s.l., is in accordance with this pattern, given that Lycaenidae was better represented than Riodinidae. The reasons for this pattern are unknown; however, warmer climates seem to favor sampling of the latter in relation to the former. According to BROWN (2005), several Amazon sites, including some that exhibit the highest butterfly species richness, follow this pattern, with reports of a significant positive relationship between Riodinidae richness and mean annual temperature. In the current study, the difference in richness between the two families was small, but it is expected that with more collections, the relative proportion between Lycaenidae and Riodinidae may change.

Conclusion

Given the importance of knowledge about Neotropical Lepidoptera, the inclusion of wellrepresented inventories is crucial to cover the lack of information on the occurrence and distribution of Lepidoptera in Brazil. This study represents an important contribution to such knowledge, including Lepidoptera that occur in montane sites within the Cerrado biome, where savannas are replaced by campo rupestre. In general, these habitats are spatially restricted and extremely sensitive to human disturbances and climate change, in addition to harboring several endemic and endangered species. Thus, knowledge of the diversity of these ecosystems is essential to support future conservation actions, including monitoring and management of protected areas (as in the present case), and to identify and propose priority areas for conservation. This survey has great relevance and is in accordance with the aims proposed by the National Plan for the Conservation of Threatened Lepidoptera, a great effort involving several researchers and the ICMBio Environmental Agency to increase the knowledge of Lepidoptera in all Brazilian biomes. Thus, this list holds valuable information not only for the management of the SVNP but also, combined with other campo rupestre inventories, for the consolidation of a more robust body of information to support actions aimed at the conservation of Cerrado biodiversity in Brazil.

Acknowledgments

To Olaf Mielke, Thamara Zacca, Eduardo Barbosa, Tiago Barbosa, Gustavo Accacio, Eurides Furtado, Curtis Callaghan, Diego Dolibaina, and Fernando Dias for assisting in species identification; to Olaf Mielke for critically reading and contributing to the improvement of the manuscript; to the entire team of the zoology lab of IFSULDEMINAS (Campus Inconfidentes, Minas Gerais) for assisting with fieldwork and providing logistical support; to IFSULDEMINAS for supporting and financing our transport; and to ICMBio’s team of brigade members and SVNP directory for assisting in the logistics of collection campaigns. We also thank CNPq/FAPs/MMA (Edital 47/2010) for creating SISBIOTA Brazil, encouraging the organization of RedeLep (Rede Nacional de Pesquisa e Conservação de Lepidópteros) (563332/2010-7). AHBR thanks the Brazilian Federal Agency for Support and Evaluation of Graduate Education (CAPES, Finance Code 001) for his scholarship; LAK was supported by CNPq (163119/2013-9) and FAPESP (10/51340-8); CAI was supported by CNPq, FAPESP (2011/08433-8), and INCT (Institutos Nacionais de Ciência e Tecnologia) in Ecology, Evolution, and Biodiversity Conservation (EECBio) through MCTIC/CNPq (process no. 465610/2014-5) and the Goiás State Research Foundation; AVLF is grateful to FAPESP (Biota-Fapesp 2011/50225-3), CNPq (421248/2017-3, 304291/2020-0), and NSF (DEB-1256742). Collection licenses: SISBIO 63930-1 and 53016. This study is registered in SISGEN (A5334BA).

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Appendix I.–

List of Papilionoidea species recorded in the Sempre Vivas National Park, Minas Gerais,southeastern Brazil. The total number of species in each taxon is indicated in parentheses.

Notas de autor

Autor para la correspondencia / Corresponding author augustohbrosa@hotmail.com