Ants (Hymenoptera: Formicidae) of the Parque Estadual São Camilo, an isolated Atlantic Forest remnant in western Paraná, Brazil

Natalia Ladino Rodrigo Machado Feitosa About the authors

ABSTRACT

We provide a list for the ants collected in the leaf litter, soil and vegetation of the Parque Estadual São Camilo, an important conservation unit of Atlantic Forest in Paraná, Brazil, and one of the oldest in the state. We report 108 species, of which eight species and two genera represent new records for Southern Brazil. Seven species are reported for the first time in Paraná. Our work is the first ant list for the western limit of the state, and reveals a surprisingly high number of species considering the extension of the study area. We highlight the importance of integrating different sampling techniques to explore ant diversity, and to conduct baseline studies in protected areas to document biodiversity.

KEY WORDS:
Biome; conservation; Malaise; pitfall; Winkler

INTRODUCTION

Originally, the Brazilian Atlantic Forest encompassed a 1.5 million km2 of the national territory, extending through most of the Brazilian coast and interior areas of Central and Southern regions (Ribeiro et al. 2011Ribeiro MC, Martensen AC, Metzger JP, Tabarelli M, Scaran F, Fortin M-J (2011) The Brazilian Atlantic Forest: a shrinking biodiversity hotspot. In: Zachos F, Habel J (Eds) Biodiversity hotspots: distribution and protection of conservation priority areas. Springer, Berlin, 405-433.). However, the biome has suffered with a drastic habitat loss and fragmentation, mainly derived from anthropogenic activities. Currently, the extension of the Atlantic Forest in Brazil is nearly 0.28-0.30 million km2, mainly represented by small, isolated and degraded fragments (Rosa et al. 2021aRosa MR, Brancallon PHS, Crouzeilles R, Tambosi LR, Piffer PR, Lenti FEB, Hirota M, Santiami E, Metzger JP (2021a) Hidden destruction of older forests threatens Brazil’s Atlantic Forest and challenges restoration programs. Science Advances 7(4): eabc4547. https://doi.org/10.1126/sciadv.abc4547
https://doi.org/10.1126/sciadv.abc4547...
). Some of those fragments are located in protected areas of the country (Rosa et al. 2021bRosa TF, Camarota F, Zuanon LA, Tito R, Maravalhas JB, Powell S, Vasconcelos HL (2021b) The effects of high-severity fires on the arboreal ant community of a Neotropical savanna. Oecologia 196: 951-961. https://doi.org/10.1007/s00442-021-04922-x
https://doi.org/10.1007/s00442-021-04922...
).

The state of Paraná, Southern Brazil, is covered by two of the six officially recognized Brazilian biomes (MMA 2021MMA (2021) Biomas. Ministério do Meio Ambiente, Brasília. https://antigo.mma.gov.br/biomas.html [Accessed: 04/01/2022]
https://antigo.mma.gov.br/biomas.html...
): the Atlantic Forest and the Cerrado. While the Cerrado is restricted to small, isolated fragments in the northern portion of the state (Oliveira and Feitosa 2021Oliveira AM, Feitosa RM (2021) Save the survivors: the remarkable ant diversity of the last protected fragment of savanna in Southern Brazil. Insectes Sociaux 68: 49-58. https://doi.org/10.1007/s00040-020-00804-2
https://doi.org/10.1007/s00040-020-00804...
), the Atlantic Forest occupies most of Paraná’s surface, including areas of semideciduous seasonal forests, ombrophilous forests with Araucaria and natural grasslands (Franco et al. 2021Franco W, Vasconcelos HL, Feitosa RM (2021) Patterns of Ant Diversity in the Natural Grasslands of Southern Brazil. Neotropical Entomology 50: 725-735. https://doi.org/10.1007/s13744-021-00886-y
https://doi.org/10.1007/s13744-021-00886...
). One of the conservation unities stablished to protect the Atlantic Forest in the state is the Parque Estadual São Camilo (hereafter PESC), created in 1990 and once named Reserva Biológica de São Camilo (IAT 2022aIAT (2022a) Instituto Água e Terra. https://www.iat.pr.gov.br/Pagina/Conheca-os-Parques-do-Parana
https://www.iat.pr.gov.br/Pagina/Conheca...
). The park covers an area of 3.85 km2 at the western limit of Paraná, near the border with Paraguay, representing only 0.0002% of the native fragments of the Atlantic Forest in the state (Cortez and Gonçalves 2015Cortez VG, Gonçalves RB (2015) Guia da biodiversidade de Palotina. Editora UFPR, Curitiba, 232 pp.). The PESC is also a strategic point to link other preserved areas in at least 15 municipalities at the southwestern of the state (IAT 2022aIAT (2022a) Instituto Água e Terra. https://www.iat.pr.gov.br/Pagina/Conheca-os-Parques-do-Parana
https://www.iat.pr.gov.br/Pagina/Conheca...
). Nevertheless, the published studies aimed to report the biological diversity of PESC have been focused on a few taxa (i.e. fungus, plants, bats, bees and wasps) (Ferreira and Cortez 2012Ferreira AJ, Cortez VG (2012) Lepiotoid Agaricaceae (Basidiomycota) from São Camilo State Park, Paraná State, Brazil. Mycosphere 3(6): 962-976. https://doi.org/10.5943/mycosphere/3/6/11
https://doi.org/10.5943/mycosphere/3/6/1...
, Dias and Cortez 2013Dias RL, Cortez VG (2013) New records of Pluteus (Agaricales) from Paraná State, Brazil. Mycosphere 4: 937-943. https://doi.org/10.5943/mycosphere/4/5/6
https://doi.org/10.5943/mycosphere/4/5/6...
, Alves and Cortez 2014Alves CR, Cortez VG (2014) Gasteroid Agaricomycetidae (Basidiomycota) from São Camilo State Park, Paraná, Brazil. Revista Brasileira de Biociências 12(1): 26. http://www.ufrgs.br/seerbio/ojs/index.php/rbb/article/view/2632
http://www.ufrgs.br/seerbio/ojs/index.ph...
, da Silva Cruz and Cortez 2015da Silva Cruz K, Cortez VG (2015) Xylaria (Xylariaceae, Ascomycota) in the Parque Estadual de São Camilo, Paraná, Brazil. Acta Biológica Paranaense 44: 1-4., Cortez and Gonçalves 2015Cortez VG, Gonçalves RB (2015) Guia da biodiversidade de Palotina. Editora UFPR, Curitiba, 232 pp., Oliveira and Gonçalves 2017Oliveira PS, Gonçalves RB (2017) Trap-nesting bees and wasps (Hymenoptera: Aculeata) in a Semidecidual Seasonal Forest fragment, southern Brazil. Papéis Avulsos de Zoologia 57: 149-156. https://doi.org/10.11606/0031-1049.2017.57.13
https://doi.org/10.11606/0031-1049.2017....
, Batista 2018Batista SC (2018) Assembleia de morcegos (Mammalia, Chiroptera) do Parque Estadual de São Camilo, Paraná, Brasil. MSc. Dissertation, Universidade Federal do Paraná, Brazil. Available online at: https://www.acervodigital.ufpr.br/handle/1884/55748?show=full
https://www.acervodigital.ufpr.br/handle...
, Silva-Filho et al. 2019Silva-Filho AGS, Seger C, Cortez VG (2019) Panaeolus (Agaricales) from Western Paraná state, South Brazil, with a description of a new species, Panaeolus sylvaticus. Edinburgh Journal of Botany 76(2): 297-309. https://doi:10.1017/S0960428619000064
https://doi:10.1017/S0960428619000064...
). In addition, anthropic pressures as hunting activities, animal traps and pesticides in the immediate surroundings, and irregular plant extraction represent a threat for the maintenance and preservation of the park’s biodiversity that need to be urgently addressed (IAT 2022bIAT (2022b) Instituto Água e Terra. http://www.iat.pr.gov.br/Pagina/Plano-de-Manejo-Parque-Estadual-de-Sao-Camilo
http://www.iat.pr.gov.br/Pagina/Plano-de...
).

Insects play essential roles in the environment, mainly related to a high sensitivity to ecosystem alterations. Among the insects, ants (Hymenoptera, Formicidae) are a well-studied group in Brazil (Ramalho et al. 2020Ramalho M, Martins C, Moreau CS (2020) Myrmecology: majority of females only within the colony. Boletim do Museu Paraense Emílio Goeldi-Ciências Naturais 15(1): 17-26. https://doi.org/10.46357/bcnaturais.v15i1.241
https://doi.org/10.46357/bcnaturais.v15i...
). Biomass and dominance of ants in terrestrial ecosystems make them crucial in processes of bioturbation, nutrient cycling, decomposition, seed dispersal and energy flow. Moreover, their importance as biological indicators and pest regulators has also been long recognized (Elizalde et al. 2020Elizalde L, Arbetman M, Arnan X, Eggleton P, Leal IR, Lescano MN, Saez A, Werenkraut V, Pirk GI (2020) The ecosystem services provided by social insects: traits, management tools and knowledge gaps. Biological Reviews 95(5): 1418-1441. https://doi.org/10.1111/brv.12616
https://doi.org/10.1111/brv.12616...
, Ribas et al. 2012Ribas CR, Campos RBF, Schmidt FA, Solar RRC (2012) Ants as Indicators in Brazil: A Review with Suggestions to Improve the Use of Ants in Environmental Monitoring Programs. Psyche 2012: 636749. https://doi.org/10.1155/2012/636749
https://doi.org/10.1155/2012/636749...
).

One of the first steps to improve our understanding about biodiversity and how the ecosystems are being affected by anthropic activities is to identify the organisms inhabiting them (Souza-Campana et al. 2019Souza-Campana DR, Carvalho ACN, Canali M, Silva OGM, Morini MSC, Fujihara RT (2019) A rapid survey of ground-dwelling ants (Hymenoptera: Formicidae) in an urban park from state of São Paulo, Brazil. Brazilian Journal of Biology 80: 682-684. https://doi.org/10.1590/1519-6984.221831
https://doi.org/10.1590/1519-6984.221831...
). In that sense, baseline studies, as species lists, are a powerful tool to do so (Camacho and Vasconcelos 2015Camacho GP, Vasconcelos HL (2015) Ants of the Panga Ecological Station, a Cerrado reserve in central Brazil. Sociobiology 62(2): 281-295. https://doi.org/10.13102/sociobiology.v62i2.281-295
https://doi.org/10.13102/sociobiology.v6...
). We here provided a list of the ant species in the Parque Estadual São Camilo based on four sampling techniques. Our findings revealed a high local richness, also including new geographic records of species and genera for the Paraná and Southern Brazil.

MATERIAL AND METHODS

Study site

The study area was the Parque Estadual São Camilo, located at 8 km of the central area of the municipality of Palotina, Paraná state (see Supplementary file Movie 1). The park has an average elevation of about 250 m average above sea level, and a Cfa climate type according to Köppen classification (IAT 2022bIAT (2022b) Instituto Água e Terra. http://www.iat.pr.gov.br/Pagina/Plano-de-Manejo-Parque-Estadual-de-Sao-Camilo
http://www.iat.pr.gov.br/Pagina/Plano-de...
). The PESC is embedded in a broad matrix of anthropic use, and is covered by areas of submontane semideciduous seasonal forests interrupted by intermediate stages of plant succession, and pioneer formations with fluvio-lacustrine influence (IAT 2022bIAT (2022b) Instituto Água e Terra. http://www.iat.pr.gov.br/Pagina/Plano-de-Manejo-Parque-Estadual-de-Sao-Camilo
http://www.iat.pr.gov.br/Pagina/Plano-de...
) (Fig. 1).

Figure 1
Map showing the collection points indicated by yellow circles at Parque Estadual São Camilo, Palotina, Paraná state, Southern Brazil. The photos show the main environments explored, and the green filled polygons depict the vegetation cover of the park. Photos: N. Ladino, 8-11/12/2020.

Experimental design

We established five sampling points at PESC at the semidecidual forest areas, two in the south limits of the park and three adjacent to the visitor’s trail and the administrative center (Table 1, Fig. 1). We collected the ants under the collection permit IAT 21.19/15.713.531-7, at the second week of December 2020. Four techniques were employed, including non-baited epigean pitfall traps, Winkler extractors of leaf litter, free hand-collection in vegetation and soil strata, and a flight interception Malaise trap. Our study was a rapid survey, the sampling was not standardized, and there were no replicates. All the techniques, except by the Malaise trap, were employed in the first point, Winkler and hand-collection in the second point, Malaise and hand-collection in the third point, hand-collection in the fourth point, and pitfall and Winkler in the fifth point. The pitfall traps consisted in 25 plastic 200 ml cups containing a solution of water and li quid unscented soap, with the opening positioned at the ground level. We kept the traps in the field for 48 hours, after that, we removed them and sorted the ants (Lasmar et al. 2017Lasmar CJ, Queiroz ACM, Rabello AM, Feitosa RM, Canedo-Júnior EO, Schmidt FA, Cuissi RG, Ribas CR (2017) Testing the effect of pitfall-trap installation on ant sampling. Insectes Sociaux 64: 445-451. https://doi.org/10.1007/s00040-017-0558-7
https://doi.org/10.1007/s00040-017-0558-...
). We randomly selected areas to implement the Winkler technique as described by Fisher (1999Fisher BL (1999) Improving inventory efficiency: a case study of leaf-litter ant diversity in Madagascar. Ecological Applications 9(2): 714-731. https://doi.org/10.1890/1051-0761(1999)009[0714:IIEACS]2.0.CO;2), except by the ant sorting, which was carried out with ants still alive in white trails. After collecting 31 sifted litter samples, we transported them to our field lab at the PESC administrative center, and sorted the specimens until ant activity ceased. We performed hand-collection of ants in soil and vegetation strata. The Malaise trap was installed adjacent to the visitor’s trail of the park, and kept in field for 72 hours (Montgomery et al. 2021Montgomery GA, Belitz MW, Guralnick RP, Tingley MW (2021) Standards and Best Practices for Monitoring and Benchmarking Insects. Frontiers in Ecology and Evolution 8: 579193. https://doi.org/10.3389/fevo.2020.579193
https://doi.org/10.3389/fevo.2020.579193...
). All samples were stored in vials with ethanol at 100%, and transported to the Laboratório de Sistemática e Biologia de Formigas at the Universidade Federal do Paraná, where the ants were separated in morphospecies, dry-mounted, pinned, and labelled.

Table 1
Collection points at Parque Estadual São Camilo, Palotina, PR. Brazil.

Ant identification

We firstly identified the ants to the genus level using Baccaro et al. (2015Baccaro FB, Feitosa RM, Fernández F, Fernandes IO, Izzo TJ, Souza JLP, Solar RRC (2015) Guia para os gêneros de formigas do Brasil. INPA, Manaus, 388 pp. https://doi.org/10.5281/zenodo.32912
https://doi.org/10.5281/zenodo.32912...
). For the identification of species, we used specific taxonomic publications (Gonçalves 1961Gonçalves CR (1961) O gênero Acromyrmex no Brasil (Hym. Formicidae). Studia Entomologica 4: 113-180., Watkins 1976Watkins JF (1976) The identification and distribution of New World Army Ants. Baylor University Press, Waco, Texas, 109 pp., Kugler and Brown 1982Kugler C, Brown WL (1982) Revisionary and other studies on the ant genus Ectatomma, including the description of two new species. Search Agriculture 24: 1-8., Mackay and Mackay 2010Mackay WP, Mackay EE (2010) Systematics and biology of the New World ants of the genus Pachycondyla (Hymenoptera: Formicidae). Edwin Mellen Press, Lewiston, 642 pp., Longino 2013Longino JT (2013) A revision of the ant genus Octostruma Forel 1912 (Hymenoptera, Formicidae). Zootaxa 3699(1): 1-61. https://doi.org/10.11646/ZOOTAXA.3699.1.1
https://doi.org/10.11646/ZOOTAXA.3699.1....
, Johnson and Cover 2015Johnson RA, Cover SP (2015) A taxonomic revision of the seed-harvester ant genus Pogonomyrmex (Hymenoptera: Formicidae) on Hispaniola. Zootaxa 3972(2): 19. https://doi.org/10.11646/zootaxa.3972.2.5
https://doi.org/10.11646/zootaxa.3972.2....
, Ortiz et al. 2019Ortíz-Sepúlveda CM, Van Bocxlaer B, Meneses AD, Fernández F (2019) Molecular and morphological recognition of species boundaries in the neglected ant genus Brachymyrmex (Hymenoptera: Formicidae): toward a taxonomic revision. Organisms Diversity & Evolution 19(3): 447-542. https://doi.org/10.1007/s13127-019-00406-2
https://doi.org/10.1007/s13127-019-00406...
, Oliveira et al. 2021Oliveira AM, Powell S, Feitosa RM (2021) A taxonomic study of the Brazilian turtle ants (Formicidae: Myrmicinae: Cephalotes). Revista Brasileira de Entomologia 65(3): e20210028. https://doi.org/10.1590/1806-9665-RBENT-2021-0028
https://doi.org/10.1590/1806-9665-RBENT-...
). For the identification of Rogeria species we consulted Otávio Guilherme Morais Silva (Museu Paraense Emílio Goeldi). For the identification of Odontomachus and Pachycondyla species we consulted unpublished keys provided by Eder Cleyton Barbosa de França and Frederico Rottgers Marcineiro (Universidade Federal do Paraná). For the species of Pheidole we used the interactive key as implemented in the Lucid platform, provided by John T. Longino (https://sites.google.com/site/newworldpheidole/home), along with Wilson’s monography (2003Wilson EO (2003) Pheidole in the New World: a dominant, hyperdiverse ant genus. Harvard University Press, Cambridge, vol. 1, 820 pp.), and AntWeb (2022AntWeb (2022) AntWeb. v. 8.75.3. California Academy of Science. Available online at: Available online at: http://www.antweb.org [Accessed: 04/01/2022]
http://www.antweb.org...
). We listed the unidentified ants as morphospecies and deposited vouchers of all collected specimens at the Entomological Collection Padre Jesus Santiago Moure (DZUP), at Universidade Federal do Paraná, Curitiba, Brazil. The new records for Paraná state and Southern Brazil were determined by the revision of taxonomic works and ant surveys carried out in this region, together with the information provided by Antmaps.org (Janicki et al. 2016Janicki J, Narula N, Ziegler M, Guénard B, Economo EP (2016) Visualizing and interacting with large-volume biodiversity data using client-server web-mapping applications: The design and implementation of antmaps.org. Ecological Informatics 32: 185-193.).

Data treatment

We elaborated a dataset including specimens records by subfamily, genus, species, collection point, and sampling technique. The dataset was analyzed in the R environment with the R Base package (R Core Team 2020R Core Team (2020) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, v. 3.6.3. https://www.R-project.org/), and the graph relating the number of species by genus in each subfamily was made with the ggplot2 package (Wickham 2016Wickham H (2016) ggplot2: Elegant Graphics for Data Analysis. Springer, Cham, 260 pp. https://doi.org/10.1007/978-3-319-24277-4
https://doi.org/10.1007/978-3-319-24277-...
). Our list was compared with other records for ants in the region (see Golias et al. 2018Golias HC, Lopes J, Delabie JHC, de Azevedo F (2018) Diversity of ants in citrus orchards and in a forest fragment in Southern Brazil. EntomoBrasilis 11(1): 01-08. https://doi.org/10.12741/ebrasilis.v11i1.703
https://doi.org/10.12741/ebrasilis.v11i1...
, Holdefer et al. 2017Holdefer DR, Lutinski JA, Garcia FRM (2017) Does organic management of agroecosystems contribute to the maintenance of the richness of ants? Semina: Ciências Agrárias 38(6): 3455-3468. https://doi.org/10.5433/1679-0359.2017v38n6p3455
https://doi.org/10.5433/1679-0359.2017v3...
, Franco and Feitosa 2018Franco W, Feitosa RM (2018) First standardized inventory of ants (Hymenoptera: Formicidae) in the natural grasslands of Paraná: New records for Southern Brazil. Papéis Avulsos de Zoologia 58: e20185812. https://doi.org/10.11606/1807-0205/2018.58.12
https://doi.org/10.11606/1807-0205/2018....
, Martins et al. 2019Martins MFO, Thomazini MJ, Baretta D, Brown GG, da Rosa MG, Zagatto MR, Santos A, Nadolny HS, Cardoso GBX, Niva CC, Bartz ML, Feitosa RM (2019) Accessing the subterranean ant fauna (Hymenoptera: Formicidae) in native and modified subtropical landscapes in the Neotropics. Biota Neotropica 20: e20190782. https://doi.org/10.1590/1676-0611-BN-2019-0782
https://doi.org/10.1590/1676-0611-BN-201...
, Franco et al. 2021Franco W, Vasconcelos HL, Feitosa RM (2021) Patterns of Ant Diversity in the Natural Grasslands of Southern Brazil. Neotropical Entomology 50: 725-735. https://doi.org/10.1007/s13744-021-00886-y
https://doi.org/10.1007/s13744-021-00886...
, Oliveira and Feitosa 2021Oliveira AM, Feitosa RM (2021) Save the survivors: the remarkable ant diversity of the last protected fragment of savanna in Southern Brazil. Insectes Sociaux 68: 49-58. https://doi.org/10.1007/s00040-020-00804-2
https://doi.org/10.1007/s00040-020-00804...
). We elaborated a map of the study area in Brazil with the collection points established at PESC in QGIS v.3.16.7 (QGIS 2021QGIS (2021) QGIS Geographic Information System. QGIS Association. http://www.qgis.org
http://www.qgis.org...
), the vegetation cover was retrieved from IAT (2022bIAT (2022b) Instituto Água e Terra. http://www.iat.pr.gov.br/Pagina/Plano-de-Manejo-Parque-Estadual-de-Sao-Camilo
http://www.iat.pr.gov.br/Pagina/Plano-de...
). Also, provided a time-lapse video derived from aerial images of changes in land cover over time in Palotina, obtained in Google Earth Pro, v. 7.3.4.8248, as supplementary material (see Supplementary file Movie 1). High-resolution photos from new ant genera records were elaborated with a Zeiss Axiocam 305 camera coupled to a Zeiss Stereo Discovery V20 stereomicroscope. Parameters of brightness and contrast were enhanced in Adobe Photoshop CC2014, and the figure plates were elaborated in Adobe Illustrator 2020.

RESULTS

We recorded 108 species at PESC, distributed in 41 genera and eight subfamilies (Table 2, Fig. 2). Ten species were exclusively collected with pitfall traps, 29 with Winkler extractors, three with Malaise and 12 by hand-collection, from which seven were solely found in vegetation. Pheidole was the richest genus with 27 species (25.2%), followed by Solenopsis with eight (7.5%) and Camponotus with seven (6.5%). We reported the species Rasopone ferruginea (Smith, 1858), Thaumatomyrmex mutilatus Mayr, 1887, Pheidole lucretii Santschi, 1923, Brachymyrmex santschii Menozzi, 1927 and Carebara brevipilosa Fernández, 2004 for the first time in Paraná; while we firstly recorded Anochetus bispinosus (Smith, 1858), Rogeria scobinataKugler, 1994Kugler C (1994) A revision of the ant genus Rogeria with description of the sting apparatus (Hymenoptera: Formicidae). Journal of Hymenoptera Research 3:17-89., Solenopsis iheringi Forel, 1908, Strumigenys hyphata (Brown, 1953), and Wasmannia iheringi for Southern Brazil (Figs 3-11).

Figure 2
Number of ant species per genus recorded in the Parque Estadual São Camilo, Palotina, PR, Brazil.

Table 2
List of ants of the Parque Estadual São Camilo, Palotina, PR, Brazil, discriminated by collection point and sampling technique: pitfall (P), Winkler (W), Malaise (M), and hand-collection (H; S: soil, V: vegetation). New records for the Paraná state are indicated by one asterisk (*), and new records for Southern Brazil are indicated by two asterisks (**).

DISCUSSION

We present here the first list of the ants occurring in the Parque Estadual São Camilo, Paraná, increasing the number of species formerly recorded for the state. We found that less than 30 species listed in the PESC are shared with other areas of the state based on previous studies on ant diversity in Paraná (Holdefer et al. 2017Holdefer DR, Lutinski JA, Garcia FRM (2017) Does organic management of agroecosystems contribute to the maintenance of the richness of ants? Semina: Ciências Agrárias 38(6): 3455-3468. https://doi.org/10.5433/1679-0359.2017v38n6p3455
https://doi.org/10.5433/1679-0359.2017v3...
, Franco and Feitosa 2018Franco W, Feitosa RM (2018) First standardized inventory of ants (Hymenoptera: Formicidae) in the natural grasslands of Paraná: New records for Southern Brazil. Papéis Avulsos de Zoologia 58: e20185812. https://doi.org/10.11606/1807-0205/2018.58.12
https://doi.org/10.11606/1807-0205/2018....
, Golias et al. 2018Golias HC, Lopes J, Delabie JHC, de Azevedo F (2018) Diversity of ants in citrus orchards and in a forest fragment in Southern Brazil. EntomoBrasilis 11(1): 01-08. https://doi.org/10.12741/ebrasilis.v11i1.703
https://doi.org/10.12741/ebrasilis.v11i1...
, Martins et al. 2019Martins MFO, Thomazini MJ, Baretta D, Brown GG, da Rosa MG, Zagatto MR, Santos A, Nadolny HS, Cardoso GBX, Niva CC, Bartz ML, Feitosa RM (2019) Accessing the subterranean ant fauna (Hymenoptera: Formicidae) in native and modified subtropical landscapes in the Neotropics. Biota Neotropica 20: e20190782. https://doi.org/10.1590/1676-0611-BN-2019-0782
https://doi.org/10.1590/1676-0611-BN-201...
, Franco et al. 2021Franco W, Vasconcelos HL, Feitosa RM (2021) Patterns of Ant Diversity in the Natural Grasslands of Southern Brazil. Neotropical Entomology 50: 725-735. https://doi.org/10.1007/s13744-021-00886-y
https://doi.org/10.1007/s13744-021-00886...
, Oliveira and Feitosa 2021Oliveira AM, Feitosa RM (2021) Save the survivors: the remarkable ant diversity of the last protected fragment of savanna in Southern Brazil. Insectes Sociaux 68: 49-58. https://doi.org/10.1007/s00040-020-00804-2
https://doi.org/10.1007/s00040-020-00804...
).

Different works have cited the advantages of combining sampling techniques for rapid ant surveys (e.g. Delabie et al. 2020Delabie J, Koch E, Dodonov P, Caitano B, DaRocha W, Jahyny B, Leponce M, Majer J, Mariano C (2020) Sampling and Analysis Methods for Ant Diversity Assessment. In: Santos JC, Fernandes GW (Eds) Measuring Arthropod Biodiversity. Springer, Cham, 13-54. https://doi.org/10.1007/978-3-030-53226-0_2
https://doi.org/10.1007/978-3-030-53226-...
, Salata et al. 2020Salata S, Kalarus K, Borowiec L, Trichas A, Kujawa K (2020) How estimated ant diversity is biased by the sampling method? A case study of Crete: a Mediterranean biodiversity hotspot. Biodiversity and Conservation 29(9): 3031-3050. https://doi.org/10.1007/s10531-020-02014-5
https://doi.org/10.1007/s10531-020-02014...
). This approach has a positive impact on the reported richness, collection of different ant guilds, and the comprehension of the community structure of ants (Delabie et al. 2020Delabie J, Koch E, Dodonov P, Caitano B, DaRocha W, Jahyny B, Leponce M, Majer J, Mariano C (2020) Sampling and Analysis Methods for Ant Diversity Assessment. In: Santos JC, Fernandes GW (Eds) Measuring Arthropod Biodiversity. Springer, Cham, 13-54. https://doi.org/10.1007/978-3-030-53226-0_2
https://doi.org/10.1007/978-3-030-53226-...
, Salata et al. 2020Salata S, Kalarus K, Borowiec L, Trichas A, Kujawa K (2020) How estimated ant diversity is biased by the sampling method? A case study of Crete: a Mediterranean biodiversity hotspot. Biodiversity and Conservation 29(9): 3031-3050. https://doi.org/10.1007/s10531-020-02014-5
https://doi.org/10.1007/s10531-020-02014...
). Implementing the four sampling techniques applied here was crucial to achieve the results we present, once ants establish their nests and forage in different substrates (Soares and Schoereder 2001Soares SM, Schoereder JH (2001) Ant-nest distribution in a remnant of tropical rainforest in southeastern Brazil. Insectes Sociaux 48(3): 280-286. https://doi.org/10.1007/PL00001778
https://doi.org/10.1007/PL00001778...
). Among the outcomes of using these sampling techniques we outstand that the epigean pitfall and Winkler techniques were effective to reveal ants inhabiting or foraging in the soil, litter and rotten wood. On the other hand, species of Cephalotes and Nesomyrmex, two exclusively arboreal ant genera listed here, were obtained only by hand-collection, once we did not use any technique to explore the arboreal ant fauna. Moreover, the single species of Neivamyrmex and Neoponera marginata (Roger, 1861), two taxa known by their nomadic habits, were also exclusively captured by hand-collection. In fact, trails of N. marginata were widely spread along the PESC visitors trail, but we fail to record the species at the collection points established (see Supplementary file Movie 2). Finally, the Malaise trap allowed us to register a great representativity of ant queens and males. Otherwise, those individuals would have been difficult to capture, since most techniques aimed to collect ants on our study are focused on workers.

The ants recorded in PESC where collected inside a fragment of submontane semidecidual seasonal forest in an advanced stage of recovery (IAT 2022bIAT (2022b) Instituto Água e Terra. http://www.iat.pr.gov.br/Pagina/Plano-de-Manejo-Parque-Estadual-de-Sao-Camilo
http://www.iat.pr.gov.br/Pagina/Plano-de...
, Cortez and Gonçalves 2015Cortez VG, Gonçalves RB (2015) Guia da biodiversidade de Palotina. Editora UFPR, Curitiba, 232 pp.). This forest type was once considered one of the richest in Brazil in terms of wood volume per unit of area due to a fertile soil derived from basalt (IAT 2022bIAT (2022b) Instituto Água e Terra. http://www.iat.pr.gov.br/Pagina/Plano-de-Manejo-Parque-Estadual-de-Sao-Camilo
http://www.iat.pr.gov.br/Pagina/Plano-de...
). Also, the high humidity and wide range substrates for nesting and forage as mold, leaf litter and well-developed roots provide conditions that favor the establishment of ant colonies (Inkotte et al. 2022Inkotte J, Bomfim B, da Silva SC, Valadão MBX, da Rosa MG, Viana RB, Rios Polliana DA, Gatto A, Pereira RS (2022) Linking soil biodiversity and ecosystem function in a Neotropica savana. Applied Soil Ecology 169: 104209. https://doi.org/10.1016/j.apsoil.2021.104209
https://doi.org/10.1016/j.apsoil.2021.10...
). Ants in these environments are more prone to be captured with Winkler extractors, which could explain the higher number of exclusive species obtained with this technique in our study. The prevalence of species of Pheidole, Solenopsis and Camponotus in our list is an expected outcome. Together, these taxa account for nearly 40% of the number of ant species collected here and correspond to genera known by their high diversity worldwide. Pheidole is the richest ant genera, with around 150 species recognized in Brazil (Casadei-Ferreira et al. 2020Casadei-Ferreira A, Economo EP, Feitosa RM (2020) Additions to the taxonomy of Pheidole (Hymenoptera: Formicidae) from the southern grasslands of Brazil. Revista Brasileira de Entomologia 64(4): e20200068. https://doi.org/10.1590/1806-9665-RBENT-2020-0068
https://doi.org/10.1590/1806-9665-RBENT-...
). Camponotus and Solenopsis include species commonly sampled in ant inventories, and stand out for their diversity and complex taxonomy, particularly in South America (Pacheco et al. 2013Pacheco JA, Mackay WP, Lattke J (2013) The systematics and biology of the New World thief ants of the genus Solenopsis (Hymenoptera: Formicidae). Edwin Mellen Press, New York, 501 pp., Mackay 2019Mackay WP (2019) New World Carpenter Ants of the Hyperdiverse genus Camponotus. Lambert Academic Publishing, El Paso, vol. 1, 412 pp.).

Among the new records made here, our discovery of C. brevipilosa, R. ferruginea and T. mutilatus in the PESC filled the occurrence gap for these three genera in Southern Brazil. The southernmost occurrence for this species before ours was recorded in Misiones, Argentina (Hanisch et al. 2018Hanisch PE, Suarez AV, Tubaro PL, Paris CI (2018) Co-occurrence patterns in a subtropical ant community revealed by complementary sampling methodologies. Environmental Entomology 47(6): 1402-1412. https://doi.org/10.1093/ee/nvy143
https://doi.org/10.1093/ee/nvy143...
). Rasopone ferruginea was already known from Santa Catarina and Rio Grande do Sul, but has never been reported to Paraná until now (Cantarelli et al. 2015Cantarelli EB, Fleck MD, Granzotto F, Corassa JN, D’Avila M (2015) Diversidade de formigas (Hymenoptera:Formicidae) da serrapilheira em diferentes sistemas de uso do solo. Ciência Florestal 25(3): 607-616. https://doi.org/10.5902/1980509819612
https://doi.org/10.5902/1980509819612...
, Martins et al. 2019Martins MFO, Thomazini MJ, Baretta D, Brown GG, da Rosa MG, Zagatto MR, Santos A, Nadolny HS, Cardoso GBX, Niva CC, Bartz ML, Feitosa RM (2019) Accessing the subterranean ant fauna (Hymenoptera: Formicidae) in native and modified subtropical landscapes in the Neotropics. Biota Neotropica 20: e20190782. https://doi.org/10.1590/1676-0611-BN-2019-0782
https://doi.org/10.1590/1676-0611-BN-201...
). This species is exclusively Neotropi cal, and the southernmost limit of its distribution is Formosa, Argentina (Leponce et al. 2004Leponce M, Theunis L, Delabie JH, Roisin Y (2004) Scale dependence of diversity measures in a leaf-litter ant assemblage. Ecography 27(2): 253-267. https://doi.org/10.1111/j.0906-7590.2004.03715.x
https://doi.org/10.1111/j.0906-7590.2004...
, Theunis et al. 2005Theunis L, Gilbert M, Roisin Y, Leponce M (2005) Spatial structure of litter-dwelling ant distribution in a subtropical dry forest. Insectes Sociaux 52(4): 366-377. https://doi.org/10.1007/s00040-005-0822-0
https://doi.org/10.1007/s00040-005-0822-...
). The unprecedent record of T. mutilatus was rather expected in Paraná, since neighboring states also reported its presence (Brandão et al. 1991Brandão CRF, Diniz JLM, Tomotake EM (1991) Thaumatomyrmex strips millipedes for prey: a novel predatory behaviour in ants, and the first case of sympatry in the genus (Hymenoptera: Formicidae). Insectes Sociaux 38(4): 335-344. https://doi.org/10.1007/BF01241869
https://doi.org/10.1007/BF01241869...
, Kempf 1975Kempf WW (1975) A revision of the Neotropical ponerine ant genus Thaumatomyrmex Mayr (Hymenoptera: Formicidae). Studia Entomologica 18: 95-126., Ulysséa et al. 2011Ulysséa MA, Cereto CE, Rosumek FB, da Silva RR, Lopes BC (2011) Updated list of ant species (Hymenoptera, Formicidae) recorded in Santa Catarina State, southern Brazil, with a discussion of research advances and priorities. Revista Brasileira de Entomologia 55(4): 603-611. https://doi.org/10.1590/S0085-56262011000400018
https://doi.org/10.1590/S0085-5626201100...
, Demetrio et al. 2017Demétrio MF, Silvestre R, Souza PR, Aoki C (2017) Checklist of ants (Hymenoptera, Formicidae) from Mato Grosso do Sul, Brazil. Iheringia, Série Zoologia 107: e2017126. https://doi.org/10.1590/1678-4766e2017126
https://doi.org/10.1590/1678-4766e201712...
). Outside Brazil, the southernmost records of T. mutilatus include Paraguay (Wild 2007Wild AL (2007) A catalogue of the ants of Paraguay (Hymenoptera: Formicidae). Zootaxa 1622(1): 1-55. https://doi.org/10.11646/ZOOTAXA.1622.1.1
https://doi.org/10.11646/ZOOTAXA.1622.1....
) and Misiones, Argentina (Hanisch et al. 2015Hanisch PE, Calcaterra LA, Leponce M, Achury R, Suarez AV, Silva RR, Paris CI (2015) Check list of ground-dwelling ant diversity (Hymenoptera: Formicidae) of the Iguazú National Park with a comparison at regional scale. Sociobiology 62(2): 213-227.). Before our work, A. bispinosus was only known from the states of Central and Northern Brazil (Brown 1978Brown WL (1978) Contributions toward a reclassification of the Formicidae. Part VI. Ponerinae, tribe Ponerini, subtribe Odontomachiti. Section B. Genus Anochetus and bibliography. Studia Entomologica 20: 549-638.). Thus, our finding constitutes the southernmost record for the distribution of this species in South America. Similarly, S. hyphata was also known so far by a few states Central-North Brazil. Despite the few records in Brazil, the occurrence of W. iheringi, S. iheringi and R. scobinata was not surprising, since these species have been collected in São Paulo, a state that borders Paraná to the north (Kugler 1994Kugler C (1994) A revision of the ant genus Rogeria with description of the sting apparatus (Hymenoptera: Formicidae). Journal of Hymenoptera Research 3:17-89., Longino and Fernández 2007Longino JT, Fernández F (2007) Taxonomic review of the genus Wasmannia. Memoirs of the American Entomological Institute 80: 271-289., Pacheco et al. 2013Pacheco JA, Mackay WP, Lattke J (2013) The systematics and biology of the New World thief ants of the genus Solenopsis (Hymenoptera: Formicidae). Edwin Mellen Press, New York, 501 pp.). Also, S. iheringi and R. scobinata have been reported in Paraguay and states of north Argentina (Kugler 1994Kugler C (1994) A revision of the ant genus Rogeria with description of the sting apparatus (Hymenoptera: Formicidae). Journal of Hymenoptera Research 3:17-89., Wild 2007Wild AL (2007) A catalogue of the ants of Paraguay (Hymenoptera: Formicidae). Zootaxa 1622(1): 1-55. https://doi.org/10.11646/ZOOTAXA.1622.1.1
https://doi.org/10.11646/ZOOTAXA.1622.1....
, Pacheco et al. 2013Pacheco JA, Mackay WP, Lattke J (2013) The systematics and biology of the New World thief ants of the genus Solenopsis (Hymenoptera: Formicidae). Edwin Mellen Press, New York, 501 pp., Hanisch et al. 2015Hanisch PE, Calcaterra LA, Leponce M, Achury R, Suarez AV, Silva RR, Paris CI (2015) Check list of ground-dwelling ant diversity (Hymenoptera: Formicidae) of the Iguazú National Park with a comparison at regional scale. Sociobiology 62(2): 213-227., 2018Hanisch PE, Suarez AV, Tubaro PL, Paris CI (2018) Co-occurrence patterns in a subtropical ant community revealed by complementary sampling methodologies. Environmental Entomology 47(6): 1402-1412. https://doi.org/10.1093/ee/nvy143
https://doi.org/10.1093/ee/nvy143...
). Some works published in the last decade have reported the presence of Brachymyrmex santschii in Southern Brazil, but solely in Santa Catarina (da Silva 1999da Silva RR (1999) Formigas (Hymenoptera: Formicidae) do oeste de Santa Catarina: histórico das coletas e lista atualizada das espécies do Estado de Santa Catarina. Biotemas 12(2): 75-100. https://doi.org/10.5007/%25x
https://doi.org/10.5007/%25x...
, Favretto et al. 2013Favretto MA, dos Santos EB, Geuster CJ (2013) Entomofauna do oeste do estado de Santa Catarina, Sul do Brasil. EntomoBrasilis 6(1): 42-63. https://doi.org/10.12741/ebrasilis.v6i1.271
https://doi.org/10.12741/ebrasilis.v6i1....
, Ulysséa et al. 2011Ulysséa MA, Cereto CE, Rosumek FB, da Silva RR, Lopes BC (2011) Updated list of ant species (Hymenoptera, Formicidae) recorded in Santa Catarina State, southern Brazil, with a discussion of research advances and priorities. Revista Brasileira de Entomologia 55(4): 603-611. https://doi.org/10.1590/S0085-56262011000400018
https://doi.org/10.1590/S0085-5626201100...
). Nevertheless, the most recent taxonomic revision of Brachymyrmex report this species for Costa Rica and Panamá (Ortíz-Sepúlveda et al. 2019Ortíz-Sepúlveda CM, Van Bocxlaer B, Meneses AD, Fernández F (2019) Molecular and morphological recognition of species boundaries in the neglected ant genus Brachymyrmex (Hymenoptera: Formicidae): toward a taxonomic revision. Organisms Diversity & Evolution 19(3): 447-542. https://doi.org/10.1007/s13127-019-00406-2
https://doi.org/10.1007/s13127-019-00406...
). Finally, the species Pheidole lucrettii was only known from Santa Catarina state until our work (Ulysséa et al. 2011Ulysséa MA, Cereto CE, Rosumek FB, da Silva RR, Lopes BC (2011) Updated list of ant species (Hymenoptera, Formicidae) recorded in Santa Catarina State, southern Brazil, with a discussion of research advances and priorities. Revista Brasileira de Entomologia 55(4): 603-611. https://doi.org/10.1590/S0085-56262011000400018
https://doi.org/10.1590/S0085-5626201100...
), and we here expand its distribution to the north.

We reported an important contribution to the knowledge of ants from Southern Brazil, and added taxonomic novelties for the ant diversity in Paraná (Table 2, Fig. 2, Figs 3-11). Still, we are certain that the ant diversity of the region remains underestimated. The PESC represents a refuge for biodiversity, and one of the last forest fragments of considerable size in Palotina (Cortez and Gonçalves 2015Cortez VG, Gonçalves RB (2015) Guia da biodiversidade de Palotina. Editora UFPR, Curitiba, 232 pp.). Considering this, a greater sampling effort and the publication of taxonomic works could lead to more species reported to this area. Our results are evidence of the historical lack of ant surveys in the state, which can be related to the low amount or even the absence of ant specialists based in Paraná institutions so far (Andrade and Feitosa 2020Andrade PL, Feitosa RM (2020) Coleção Entomológica Padre Jesus Santiago Moure: um novo centro de referência para a formação de sistematas de formigas (Hymenoptera: Formicidae). Boletim do Museu Paraense Emílio Goeldi, Ciências Naturais 15(1): 277-288. https://doi.org/10.46357/bcnaturais.v15i1.244
https://doi.org/10.46357/bcnaturais.v15i...
). Complementary to this, the current scenario of systematic dismantling of environmental policies enhanced by the current Brazilian government, suggest a disheartening panorama for the maintenance of ecosystem services within the country biomes (Lira et al. 2021Lira PK, Portela RCQ, Tambosi LR (2021) Land-Cover Changes and an Uncertain Future: Will the Brazilian Atlantic Forest Lose the Chance to Become a Hopespot? In: Marques MCM, Grelle CEV (Eds) The Atlantic Forest. Springer, Cham , 233-251.).

Figures 3-11
Full-face, lateral and dorsal view of the new genera records for the Paraná state and Southern Brazil according to the survey carried out in the Parque Estadual São Camilo, Palotina: (3-5) Carebara brevipilosa; (6-8) Rasopone ferruginea; (9-11) Thaumatomyrmex mutilatus. Scale: 0.2 mm.

ACKNOWLEDGMENTS

We are indebted with the park direction and employees for the permission and support to conduct the collection activities, particularly to Valdivino Martins dos Anjos. We thank Edilson Caron (UFPR, Palotina) for logistics assistance, the Programa de Apoio à Pós-graduação (PROAP, Ciências Biológicas, Entomologia) of the Universidade Federal do Paraná, and the Conselho Nacional de Desenvolvimento Científico e Tecnológico for financial support to RMF (CNPq: 301495/2019-0). We thank to anonymous reviewers for the comments of previous versions of this manuscript. We also thank Andrés Ramírez-Mejía (Instituto de Ecología Regional, CONICET) for guidance in the use of the ggplot2 R package.

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ADDITIONAL NOTES

  • Zoobank register

    http://zoobank.org/97A3CDE2-8A4D-4DAA-B5CB-C3E291CD7BD5
  • How to cite this article

    Ladino N, Feitosa RM (2022) Ants (Hymenoptera: Formicidae) of the Parque Estadual São Camilo, an isolated Atlantic Forest remnant in western Paraná, Brazil. Zoologia (Curitiba) 39: e22001. https://doi.org/10.1590/S1984-4689.v39.e22001
  • Published by

    Sociedade Brasileira de Zoologia at Scientific Electronic Library Online

Supplementary material 1

Supplementary file Movie 1. Evolution of the surrounding area of Parque Estadual São Camilo along time, with details of vegetation covering and urban expansion.

Authors: N Ladino, RM Feitosa.

Data type: Geographical data.

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Link: https://doi.org/10.1590/S1984-4689.v39.e22001

Supplementary material 2

Supplementary file Movie 2. Record of a nest migration event of Neoponera marginata (Formicidae: Ponerinae) in one of the trails of Parque Estadual São Camilo.

Authors: N Ladino, RM Feitosa.

Data type: Species data.

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Link: https://doi.org/10.1590/S1984-4689.v39.e22001

Publication Dates

  • Publication in this collection
    20 May 2022
  • Date of issue
    2022

History

  • Received
    04 Jan 2022
  • Accepted
    22 Feb 2022
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