DOI: 10.13102/sociobiology.v67i4.5741Sociobiology 67(4): 514-525 (December, 2020) Open access journal: http://periodicos.uefs.br/ojs/index.php/sociobiology ISSN: 0361-6525 Introduction Leaf-cutting ants are known in Brazil as saúvas, the genus Atta Fabricius (tribe Attini, subfamily Myrmicinae). Atta occurs throughout the Neotropical region (Mariconi, 1970) and can cause severe losses to the forest and agricultural sector of Latin American countries (Mariconi, 1970; Fowler et al., 1989; Jaffé, 1993). However, leaf-cutting ants are ecosystem engineers as they move massive amounts of soil during nest construction and remove foliage, which may change plant composition, acting as herbivores in natural systems (Farji-Brener & Illes, 2000; Urbas et al., 2007; Costa et al., 2008; Meyer et al., 2011; Leal et al., 2014; Stephan et al., 2015). In agricultural areas these alteration of the soil and Abstract Leaf-cutting ants are widely distributed in Brazil, particularly species of the genus Atta. We therefore described the occurrence of leaf-cutting and grass-cutting ant species of the genus Atta. Five routes comprising some of the main highways in the South, Southeast, Midwest, and North of Brazil were sampled, in addition to ants received from other 82 municipalities, composing 300 municipalities sampled. This is the first comprehensive study of Atta in Brazil. The following species and subespecies were found: A. laevigata, A. capiguara, A. sexdens rubropilosa, A. sexdens piriventris, A. sexdens sexdens, and A. cephalotes. Atta laevigata and A. capiguara were the species present in the largest number of the Brazilian municipalities sampled. Atta sexdens piriventris was only recorded in the southern region of Brazil. Atta bisphaerica presented lower expansion than A. capiguara. Atta cephalotes and Atta opaciceps are species of very restricted occurrence. Southeastern region was characterized by the occurrence of A. capiguara and A. laevigata. Atta laevigata exhibited a generalized pattern of occurrence in the Midwest and North. Our study contributes to a better understanding of the so far unknown occurrence of leaf-cutting and grass-cutting ants within Brazil. Sociobiology An international journal on social insects LC Forti1, JS Rando2, RS Camargo1, AA Moreira3, MA Castellani3, SA Leite3, KKA Sousa1, N Caldato1 Article History Edited by Evandro Nascimento Silva, UEFS, Brazil Received 30 July 2020 Initial acceptance 07 September 2020 Final acceptance 12 October 2020 Publication date 28 December 2020 Keywords Attini, Brazilian biomes, grass-cutting ants, leaf-cutting ants. Corresponding author Aldenise Alves Moreira https://orcid.org/0000-0002-2362-9624 Departamento de Fitotecnia e Zootecnia Universidade Estadual do Sudoeste da Bahia 45083-300, Vitória da Conquista-BA, Brasil. E-Mail: aldenise.moreira@gmail.com the attack on plants led to the classification as pest insects (Fowler et al., 1989). According to Mariconi (1970), leaf-cutting ants show a wide geographic distribution, occurring from the south of the United States (latitude 33o N) to central Argentina (latitude 44o S). The geographic distribution, frequency and density of Attini ants in certain habitats is related to the environmental conditions such as type of vegetation, soil type, cultivation systems, climate change (Fowler, 1983; Farji-Brener & Ruggiero, 1994; Gusmão & Loeck, 1999; Farji-Brener et al., 2016), among others. According to Brandão et al. (2011), the genus Atta includes 19 species, of which nine occur in Brazil, Atta bisphaerica Forel, 1908, Atta capiguara Gonçalves, 1944, 1 - Vegetal Protection Department, Laboratory of Social Insects-Pests, State University São Paulo, School of Agricultural Sciences, Botucatu, Brazil 2 - Vegetal Production Department, State University of Northern Paraná, Campus Luiz Meneghel, Bandeirantes, Brazil 3 - Departament Crop Science and Animal Science, State University Southwestern Bahia, Vitória da Conquista, Brazil RESEARCH ARTICLE - ANTS Occurrence of Leaf-Cutting and Grass-Cutting Ants of the Genus Atta (Hymenoptera: Formicidae) in Geographic Regions of Brazil Sociobiology 67(4): 514-525 (December, 2020) 515 Atta cephalotes (Linnaeus, 1758), Atta goiana, Gonçalves, 1942, Atta laevigata (Smith, F. 1858), Atta opaciceps Borgmeier, 1939, Atta robusta Borgmeier, 1939, Atta sexdens Linnaeus, 1758 and Atta wollenweideri Forel, 1893 (Bolton et al., 2006; Delabie et al., 2011). The species A. sexdens comprises three subspecies, A. sexdens sexdens, A. sexdens piriventris and A. sexdens rubropilosa (Mariconi, 1970; Della Lucia et al., 1993; Bacci et al., 2009). The available information on the distribution of leaf- cutting ants in Brazil is found in a dispersed and unevenly updated form (Delabie et al., 2011). For the Brazilian regions, few studies on the subject have been conducted, highlighting the pioneering works of Gonçalves (1942, 1945, 1951, 1955, 1960, 1961, 1967, 1971) and Kempf (1972), who generated a broader knowledge on the distribution of species in the different regions of Brazil, and Paula (1956) for the State of Paraná and Mariconi (1966, 1970) for the State of São Paulo. More recent are the studies developed by Loeck and Grützmacher (2001) and Grürzmacher et al. (2002), for leaf- cutting ants in Rio Grande do Sul and Corrêa et al. (2005) and Brito et al. (2012) for the Northeastern region, States of Alagoas and Bahia, respectively. The contributions on timely reports of the occurrence of leaf-cutting ants, such as Carvalho and Tarragô (1982) for Rio Grande do Sul, Delabie (1989) for Bahia and Souza et al. (2009) for Alagoas are also noteworthy. Some authors have compiled the existing literature on the occurrence and/or geographic distribution of leaf-cutting ants in Brazil, such as the work of Della Lucia et al. (1993), Forti and Boaretto (1997) and Delabie et al. (2011). In general, A. sexdens presents the broadest geographical distribution, occurring in all regions of Brazil, followed by A. laevigata, while the other species are restricted to certain Brazilian regions or states, such as A. robusta, which only occurs in Rio de Janeiro (Mariconi, 1970; Della Lucia et al., 1993) and Espírito Santo (Teixeira et al., 2003). It is observed that studies on the distribution of leaf-cutting ants of the genus Atta in Brazil date from at least five decades. The objective of this study was to describe the current occurrence of leaf- cutting and grass-cutting ants of the genus Atta in geographic regions of Brazil. Material and Methods For the survey of leaf-cutting and grass-cutting ants of the genus Atta, five routes comprising some of the main highways in Brazil were sampled, beginning with the city of Botucatu, São Paulo (22°50’46”S and 48°26’02”W). The sampling was performed in 218 municipalities of the five itineraries established: 1) Botucatu (São Paulo State) to Iepê (São Paulo State), covering 48 municipalities; 2) Ibiporã (Paraná State) to Quintana (São Paulo State), with 30 municipalities; 3) Oiapoque (Amapa State) to Santa Isabel do Pará (Para State), covering four municipalities, 4) Pirajuí (São Paulo State) to Avaré (São Paulo State), with 83 municipalities; 5) Aparecida do Taboado (Mato Grosso Sul State) to Ivinhema (Mato Grosso Sul State), comprising 53 municipalities (Table 1). Every 100 km, in a 500 m long by 50 m wide strip, marked at random, specimens of soldier ants were collected from the colonies found. The colonies were visually searched for and selected because of the huge size of their nests (large amount of loose soil removed). We selected the municipalities nearest to the main roads. At each sampling site, at least five exemplars of the ants found on the nests or on the foraging trails were collected. All material collected at each sampling site was stored in glass flasks containing 70% alcohol and labeled with the data obtained for subsequent analysis. Latitude and longitude of the sampling municipalities were obtained with a global positioning system (Sony GPS – 360), whenever possible. After each trip, all material collected was sent to the Laboratory of Social Insect Pests (LISP) at FCA/UNESP, Botucatu, SP. In addition to the collections made in routes, biological material from other 82 municipalities in different regions of Brazil were duly collected and sent for identification, making up the total of 300 sampled municipalities (Table 2). After screening, the specimens were mounted and identified under a stereo microscope based on the data published by Gonçalves (1961) and using the identification keys of Borgmeier (1959) and Mariconi (1970). For Atta sexdens, the division into subspecies according to Bacci et al. (2009) was considered. All material obtained was compared with specimens stored in the Museum of Zoology, University of São Paulo, and in the Ângelo Moreira da Costa Lima Entomology Collection of Universidade Federal Rural do Rio de Janeiro. The material of this study was stored in Museum of Zoology, University of São Paulo. Results Considering the 300 sampled municipalities (Table 3), frequency of leaf-cutting ant species are: Atta laevigata (32.6%), A. capiguara (20.0%), A. sexdens rubropilosa (10.0%), A. sexdens piriventris (9.7%), A. sexdens sexdens (2.7%), A. cephalotes (1.6%), A. opaciceps (0.3%), A. bisphaerica (0.3%), and A. vollenweideri (0.3%). Our results showed that Atta laevigata and A. capiguara were the species present in the largest number of the sampled municipalities (Fig 1); Atta sexdens piriventris was only recorded in the South region of Brazil, with predominance in the states of Santa Catarina and Rio Grande do Sul; Atta bisphaerica presented restricted occurrence, only at the state of São Paulo. Atta cephalotes and A. opaciceps, were collected in the expected regions, North and Northeast, respectively. The Southeast region was characterized by the occurrence of A. capiguara and A. laevigata, in this order, with A. laevigata being the most frequent species in Minas Gerais and A. capiguara in Paraná. The occurrence of A. laevigata was recorded for the first time in Amapá and Rio Grande do Sul. LC Forti, JS Rando, RS Camargo, AA Moreira, MA Castellani, SA Leite, KKA Sousa, N Caldato – Leaf-cutting in geographic regions of Brazil516 State County Longitude Latitude Alagoas Maceió -09o66’58” -35o73’52” Amapá Porto Grande -00o71’33” -51o41’33” Bahia Barra do Cacau -11o08’94” -43o14’16” Bahia Barra do Rocha -14o21’05” -39o60’19” Bahia Mangue Seco -11o13’42” -36o29’14” Bahia Maraú -14o10’30” -39o01’47” Bahia Salvador -12o97’11” -38o51’08” Distrito Federal Brasília -15o77’97” -47o93’00” Goiás Caldas Novas -17°74’16” -48o63’05” Maranhão São Luís -02o52’97” -44o30’27” Minas Gerais Araxá -19o59’33” -46o04’05” Minas Gerais Itaguara -20o30’22” -44o48’75” Minas Gerais Jaíba -15o33’83” -43o67’44” Minas Gerais Piumhi -20o46’52” -45o95’80” Minas Gerais São Gonçalo do Abaeté -18°33’83” -45°83’33” Mato Grosso Campo Novo dos Parecis -13°67’52” -57°89’19” Mato Grosso Sapezal -12o98’94” -58o76’41” Pernambuco Petrolina -09o39’86” -40o50’08” Paraná Arapoti -24°15’07” -49°82’66” Paraná Bandeirantes -23o11’00” -50o36’75” Paraná Cornélio Procópio -23o18’11” -50o64’66” Paraná Curitiba -25o42’77” -49o27’30” Paraná Doutor Camargo -23o55’58” -52o21’80” Paraná Guarapuava -25o39’52” -51o45’80” Paraná Matelândia -25o24’08” -53o99’63” Paraná Palotina -24o28’38” -53o84’00” Paraná Paranaguá -25°31’00” -48°31’00” Paraná Paula Freitas -26o20’83” -50o93’80” Paraná Porto Vitória -26°16’11” -51°23’16” Paraná Telêmaco Borba -24°32’38” -50°61’55” Rio de Janeiro Angra dos Reis -23o06’61” -44o31’80” Rio de Janeiro Parati -23o21’77” -44o71’30” Rio Grande do Sul Passo Fundo -28o26’27” -52o40’66” Rio Grande do Sul Arroio dos Ratos -30o07’72” -51o72’91” Rio Grande do Sul Butiá -30o11’97” -51o96’22” Rio Grande do Sul Charqueadas -30o73’27” -51o64’53” Rio Grande do Sul Gravataí -29o94’44” -50o99’19” Rio Grande do Sul Porto Alegre -30o03’30” -51o23’00” Rio Grande do Sul Tapes -30o67’33” -51o39’58” Santa Catarina Abelardo Luz -26°56’47” -52°32’83” Santa Catarina Caibí -27°07’16” -53°24’77” Santa Catarina Cordilheira Alta -26°95’88” -52°76’11” Table 2. Geographical location of the collect sites of material received in the laboratory of Social Insects-Prague for identification. Santa Catarina Coronel Freitas -26°90’86” -52°70’30” Santa Catarina Cunha Porã -26°89’36” -53°16’80” Santa Catarina Descanso -26°82’61” -53°50’16” Santa Catarina Dionisio Cerqueira -26°25’05” -53°63’97” Santa Catarina Galvão -26°45’05” -52°68’58” Santa Catarina Irani -27°02’47” -51°90’16” Santa Catarina Ita -27°29’05” -52°32’30” Santa Catarina Itapiranga -27°16’94” -53°73’22” Santa Catarina Jupiá -26°39’83” -52°72’77” Santa Catarina Maravilha -26°76’08” -53°17’25” Santa Catarina Nova Erexim -26°90’25” -52°90’58” Santa Catarina Palmitos -27°06’75” -53°16’11” Santa Catarina Pinhalzinho -26°84’80” -52°99’19” Santa Catarina Piratuba -27°41’97” -51°77’19” Santa Catarina Quilombo -26°72’61” -52°72’05” Santa Catarina São Carlos -27°07’75” -53°00’38” Santa Catarina São José Cedro -26°45’05” -53°49’41” Santa Catarina São Lourenço do Oeste -26°35’91” -52°85’11” Santa Catarina São Miguel do Oeste -26°72’52” -53°51’80” Santa Catarina Saudades -26°92’41” -53°00’30” Santa Catarina Vargeão -26°86’36” -52°15’05” Santa Catarina Xaxim -26°96’16” -52°53’47” São Paulo Agudos -22o46’91” -48o98’75” São Paulo Altinópolis -21°02’55” -47°37’38” São Paulo Assis -22°66’16” -50°41’22” São Paulo Cananéia -25°01’47” -47°92’66” São Paulo Eldorado -24o52’00” -48o10’80” São Paulo Franca -20o53’86” -47o43’03” São Paulo Igarapava -20o03’83” -47o74’69” São Paulo Itapetininga -23°59’16” -48°05’30” São Paulo Itatinga -23o10’16” -48o61’58” São Paulo Jaboticabal -21°25’47” -48°32’22” São Paulo Juquiá -24o32’08” -47o63’47” São Paulo Lençóis Paulista -22o59’86” -48o80’02” São Paulo Mongaguá -24°09’16” -46°61’77” São Paulo Nazaré Paulista -23o18’11” -46o39’05” São Paulo Peruíbe -24°32’00” -46°99’83” São Paulo Piracicaba -22°72’52” -47°64’91” São Paulo Santa Rosa de Viterbo -21°47’27” -47°36’30” São Paulo Ubatuba -23o43’38” -45o07’11” State County Longitude Latitude Sociobiology 67(4): 514-525 (December, 2020) 517 Species North Northeast Midwest Southeast South Brazil A. laevigata 6.0 0.3 20.3 5.0 1.0 32.6 A. capiguara -* - 8.7 7.0 4.3 20.0 A. sexdens rubropilosa - 0.3 2.7 2.0 5.0 10.0 A. sexdens piriventris - - - - 9.7 9.7 A. sexdens sexdens 2.7 - - - - 2.7 A. cephalotes 1.3 0.3 - - - 1.6 A. bisphaerica - - - 0.3 - 0.3 A. opaciceps - 0.3 - - - 0.3 A. vollenweideri - - - - 0.3 0.3 *No collection Discussion Atta laevigata Atta laevigata was the only species found in all regions of Brazil (Fig 1). The results are consistent with previous studies that already indicated the occurrence of this species in the five Brazilian geographic regions (Castro et al., 1961; Della Lucia et al., 1993). Gonçalves (1967) reported the presence of A. laevigata in Manaus, Óbidos and São Gabriel (Northern region), where this species was found on roadsides and in crops, including under the shade of trees in covered areas. The Cerrado region is an adequate habitat for A. laevigata, as observed by high-density nests recorded in Cerrado areas also highlight the suitability of this vegetation type (Costa & Vieira-Neto, 2016). Although in the present study the occurrence of A. laevigata was not recorded in the Northeast of Brazil, this species was found by Gonçalves (1951) in Ceará, Pernambuco and Bahia in dicotyledons, wild grasses, eucalyptus and cassava. Subsequently, Brito et al. (2012), recorded the occurrence of A. laevigata in four municipalities in southwestern Bahia, in areas of pasture, eucalyptus and forest. These municipalities are inserted in environments of Caatinga, Cerrado and deciduous seasonal forest. The Southeastern region of Brazil showed the third highest frequency of A. laevigata According to Antunes (1996), part of the southeastern region is occupied by Cerrado and a large area of Minas Gerais is covered with this type of vegetation. For the two states of the Southeastern Region (Minas Gerais and São Paulo), the occurrence of A. laevigata had already been recorded (Della Lucia et al., 1993). The Cerrado biome (Brazilian Savanna) covers 2 million km2 representing 23% of the area of the country. It is an ancient biome with rich biodiversity, estimated at 160,000 species of plants, fungi and animals (MMA, 2020). The predominant vegetation of the woods is composed of small trees with twisted trunks with irregular ramifications; with shrubs and sub-shrubs that may have xylopodios, underground organs, which allow regrowth after burning or cutting, this being denominate cerrado senso stricto, one of the various components of the physiognomy mosaic of the Cerrado biome. In the Southern region, A. laevigata was rarely frequent (Table 3). However, the record of its occurrence in Rio Grande do Sul (Southern Region, municipality of Tapes). In general, A. laevigata partially present in many municipalities (Fig 1), mainly due to the opening of highways and expansion of livestock, which serve as means for the dispersion of the species (Vieira-Neto et al., 2016), as well as the opening of agricultural frontiers in Cerrado areas, which involve the deforestation of natural forests and the plantation of monocultures and pastures. The first reference of this species in Espírito Santo was confirmed in the work of Delabie (1998), whose author, after morphological studies of several samples of A. laevigata from different regions of Brazil, concluded that Atta silvai Gonçalves, registered in that state, is a junior synonym of A. laevigata. These species occurred in a greater percentage on sampled municipalities in relation to A. sexdens. This result is probably related to the collection sites that covered areas close to the highways and that favor the nesting of A. laevigata (Forti et al., 2011). A. laevigata is aggressive in the selection of nesting areas, since nests are built in both sunny and shady (Mariconi, 1970; Pereira da Silva, 1975; Moreira et al., 2004) and the forager workers select mono and dicotyledonous plants as growth substrate of the symbiont fungus (Della Lucia et al., 1993; Forti et al., 2011). Atta capiguara The species A. capiguara, commonly known as brown leaf-cutting ant, exploits monocotyledon plants, basically grasses, and is economically important in sugarcane fields and pastures (Amante, 1967; Mariconi, 1970; Forti, 1985). Atta capiguara was found in the Southeast, Mid-west and South regions in 7.0%, 8.7% and 4.3% of sampled municipalities more restricted in its Brazilian distribution range. Its occurrence had already been reported for the states of São Paulo, Mato Grosso and Minas Gerais (Della Lucia et al., 1993) and also for Paraná, Mato Grosso do Sul and Goiás (Forti & Boaretto, 1997). The highest concentration of A. capiguara was observed in the State of Paraná (Southern Region) (Fig 1). The common Table 3. Percentage of municipalities (n = 300) with the occurrence of species and subspecies of Atta North, Northeast, Midwest, Southeast and South regions of Brazil. LC Forti, JS Rando, RS Camargo, AA Moreira, MA Castellani, SA Leite, KKA Sousa, N Caldato – Leaf-cutting in geographic regions of Brazil518 Fig 1. Atta species recorded in the municipalities of the North, Northeast, Midwest, Southeast and Southern regions of Brazil. characteristics of municipalities where A. capiguara was found are sandy soil and plant nutrient deficiency, as well as the native vegetation classified as Tropical Forest (BRAZIL, 1998). Smaller size forests resembling secondary forests occur in these soils were naturally fertility is poor and used as pasture lands (Wons, 1985). The characteristics of the Cerrado are typical in most parts of this ecoregion, but gradual changes are observed due to climatic influences of neighboring regions (Adámoli et al., 1986). Two parameters should be highlighted since they define the characteristics of the seasonal regime of this region: the average annual rainfall of 1,200 to 1,800 mm and the duration of the dry season that ranges from 5 to 6 months. The average annual temperature is 22 °C in the southern part of this ecoregion and 27 °C in the northern part. Its vegetation consists of isolated or groups of winding small trees on a continuous grass rug (BRAZIL, 1998). The importance of A. capiguara has been known since the 1940s, with observations of nests in some localities with predominance of grasses in municipalities of the State of São Paulo (Gonçalves, 1945). Surveys conducted in other municipalities increased the occurrence of this species to 17 municipalities (Mariconi et al., 1961). Mariconi (1966b) also recorded the occurrence of A. capiguara in 104 municipalities in the western region of the State of São Paulo. Amante (1967) cited the occurrence of A. capiguara in Mato Grosso do Sul and Minas Gerais. Although not mentioning the municipalities, Thomas (1990) reported that A. capiguara had become a serious problem in two municipalities of the northwestern region of Paraná after 1975 and, by 1990, this species had damaged pastures in more than 32 municipalities of that state. Many factors could have contributed to the population explosion of this species, such as disturbances caused by humans when they replaced the natural vegetation with grass monocultures (Forti & Boaretto, 1997). According to Thomas (1990), the reduction of areas with natural vegetation and the substitution of crops such as coffee, peanuts and other crops for pastures facilitated the rapid growth of A. capiguara infestation in Paraná. The increased availability of monocotyledonous plants, especially exotic grasses, allows grass-cutting ants to expand their distribution and increase population density (Cherrett, 1981). Atta sexdens The subspecies A. sexdens rubropilosa was registered in all Brazilian regions (Fig 1), except in the North, while A. sexdens piriventris and A. sexdens sexdens were of more restricted occurrence, only in the South and North regions, respectively. This result reflects the adaptability de A. sexdens rubropilosa to the different environmental conditions of the country (Farji-Brener & Ghermandi, 2008). A large number of studies on this leaf-cutting ant subspecies are available and its occurrence has been reported in several states, including Minas Gerais, Espírito Santo, Goiás, Mato Grosso, São Paulo, Paraná, Rio de Janeiro and Distrito Federal (Gonçalves, 1945; Della Lucia et al., 1993). Atta sexdens piriventris was observed in the South region. The occurrence of this subspecies was Sociobiology 67(4): 514-525 (December, 2020) 519 predominant in Santa Catarina. In the state of Rio Grande do Sul, A. sexdens piriventris occurred in most of the sampled municipalities, corroborating the studies by Loeck and Grützmacher (2001) and Grürzmacher et al. (2002). The main environmental characteristics that distinguish this region from other areas are lower average temperatures (16° C and 20° C) and uniform rainfall throughout the year, which are typical of subtropical climates (Antunes, 1996). Associated with this climate, the vegetation ranges from Atlantic Rainforest, Tropical Forest and Araucaria Forest to fields and coastal vegetation (Carraro, 1994). Supporting the hypothesis that this species tends to occupy climate environments such as those described above, in the State of Paraná this species was only found in the municipality of Paula Freitas, located at the southern end of the state (Fig 1). The north of Paraná, where a significant number of the municipalities visited are located, is found in an intertropical transition zone characterized by high average temperatures and a rainy period concentrated in the summer months (Antunes, 1996). This distribution in Paraná resembles that observed in the Southeast and, within this region, in the State of São Paulo (Fig 1). This fact is possibly associated with the similar environmental conditions in these border states, since the municipalities sampled in Paraná are concentrated in the northern and northwestern regions of the state as mentioned above. The subspecie Atta sexdens sexdens was exclusively recorded in the northern region (Fig 1). However, the occurrence of this subspecies has already been reported by Della Lucia et al. (1993) for the Mid-west, Northeast and Southeast regions. According to Delabie (1989), this leaf-cutting ant occurs in the State of Bahia (Northeast Region of Brazil) in more open areas and is more generalist in its foraging, attacking several dicotyledonous plants of economic importance (cacao, cassava and citrus) and also forage grasses, being more abundant in pastures. The occurrence of this ant in the state of Alagoas (Northeast Region of Brazil) was recorded by Souza et al. (2009). Atta cephalotes Atta cephalotes was found in the North region, in 1.3% of sampled municipalities. The State of Amazonas (North Region) was represented by only one locality (Guajará) where A. cephalotes was collected. The occurrence of A. cephalotes was recorded during the travels in Oiapoque (Amapá) and in Cruzeiro do Sul and Mâncio Lima (Acre) (Fig 1), as well as in material sent for identification from Maraú in Bahia (North region) (Fig 1), sites of occurrence of this species already cited in the literature. This result was expected, because A. cephalotes is a species easily found in the forests of Amazonia and is the most demanding leaf-cutting ant in terms of soil moisture (Gonçalves, 1960). In the Northeast region, this ant had already been reported to the States of Maranhão, Pernambuco, Bahia (Mariconi, 1970; Kempf, 1972) and Alagoas (Corrêa et al., 2005). Atta bisphaerica In this work, A. bisphaerica occurred only in the Southeast region (Fig 1), in the municipality of Botucatu, São Paulo, which was not expected, considering that the possibilities of expansion of this ant, as well as in A. capiguara, are associated with the substitution of natural vegetation and agricultural crops by pasture and sugarcane, since this species preferentially cuts monocotyledonous plants, it was expected an increase of its occurrence. For example, in the state of São Paulo, A. capiguara (Amante, 1972) and A. bisphaerica (Precetti et al., 1988) are of great economic importance in sugarcane and pasture, but only A. capiguara has high occurrence. Nevertheless, A. bisphaerica has been more associated to sugarcane and A. capiguara to pasture areas. In sugarcane crops, the chemical control of leaf-cutting ants is quite intense, with the systematic use of thermonebulization, contributing to a significant reduction of nest density, which usually does not occur in pastures, which could explain the lower expansion of A. bisphaerica in relation to A. capiguara. The occurrence of A. bisphaerica had already been reported in other states of Southeastern Brazil (Minas Gerais and Rio de Janeiro), as well as in the Mid-western region (Mato Grosso) (Della Lucia et al., 1993). Atta opaciceps The occurrence of A. opaciceps was observed in the Northeast region, state of Alagoas (municipality of Maceió) (Fig 1), a finding also reported by Souza et al. (2009), but was distributed in the Caatinga region. The occurrence of this species was also reported for the states of Sergipe (Delabie et al., 1997) and Bahia (Delabie et al., 1997; Brito et al., 2012) for the North and Southweste regions of Brazil. The hypothesis raised by Fowler et al. (1990) that this leaf-cutting ant could be in extinction has not been confirmed. Atta vollenweideri Atta vollenweideri, which usually cuts the leaves of grasses and dicotyledons, was recorded only in the South region, in the municipality of Arroio dos Ratos, state of Rio Grande do Sul (Fig 1). This species is of very restricted occurrence, being recorded only for two Brazilian states, Rio Grande do Sul (Gonçalves, 1960; 1971; Jonkman, 1978; Della Lucia et al., 1993) and Mato Grosso (Della Lucia et al., 1993). In general, the results expand the knowledge about leaf- cutting ants of the Atta genus in Brazilian regions, since the work encompassed many municipalities until then not sampled. The first record of the occurrence of A. laevigata in Amapá and Rio Grande do Sul, as well as the presence of A. opaciceps in municipalities of Alagoas, a quite what positive aspect once it were already raice possibility treat of being a species extinction. Atta laevigata, A. sexdens (considering its subspecies) and A. capiguara are species of wide occurrence in the Brazilian territory. Atta cephalotes, A. bisphaerica, A. opaciceps and A. vollenweideri are species of very restricted occurrence. LC Forti, JS Rando, RS Camargo, AA Moreira, MA Castellani, SA Leite, KKA Sousa, N Caldato – Leaf-cutting in geographic regions of Brazil520 We consider that the data presented in this work can be integrated into the databases on leaf-cutting ants, the example of created and used by Delabie et al. (2011), to expand the studies of regional clusters with identification of the main associated vectors and that allow the prediction of the occurrence of Atta species. Acknowledgements LCF was the recipient of a fellowship from Conselho Nacional de Desenvolvimento Cientifico e Tecnológico (Grant 301718/2013-0). State Route County Longitude Latitude São Paulo I Botucatu -22°50’46” -48°26’02” São Paulo I Bauru -22o31’47” -49o06’05” São Paulo I Marília -22o21’38” -49o94’58” São Paulo I Rancharia -22°25’50.8” -50°59’33.7” São Paulo I Presidente Prudente -22°21’42.4” -51°08’09.4” São Paulo I Ourinhos -23°00’16.2” -49°51’31.2” São Paulo I Santa Cruz do Rio Pardo -22°89’88” -49°63’25” São Paulo I Iepê -22°66’05” -51°07’61” Paraná I Santo Inácio -22°42’59.7” -51°46’09,8” Paraná I Colorado -22o83’75” -51o97’30” Paraná I Paranacity -22°55’00” -52°09’09.4” Paraná I Inajá -22o74’91” -52o19’80” Paraná I São João do Caiuá -22°56’25.5” -52°22’32.4” Paraná I Santo Antonio do Caiuá -22o73’47” -52o 34’22” Paraná I Tamboara -23°13’29.2” -52°35’19.9” Paraná I Santa Helena -24o86’02” -54o33’27” Paraná I Umuarama -23o76’63” -53°32’05” Paraná I Assis Chateaubriand -24o42’00” -53°52’13” Paraná I Toledo -24o71’36” -53°74’30” Paraná I Vera Cruz do Oeste -25o05’77” -53°87’69” Paraná I Missal -25o09’19” -54o24’75” Paraná I São Miguel do Iguaçú -25o34’80” -54o23’77” Paraná I Foz do Iguaçú -25°54’77” -54°58’80” Paraná I Cascavel -24o95’58” -53o45’52” Paraná I Ampere -25o91’05” -53o47’27” Paraná I Capitão Leônidas Marques -25o47’91” -53o61’41” Paraná I Rio Negro -26°06’00” -49°48’00” Paraná I Campo do Tenente -25°97’80” -49°68’27” Paraná I Ponta Grossa -25°06’00” -50°10’00” Paraná I Ortigueira -24o20’83” -50o94’94” Paraná I Imbaú -24°44’05” -50°76’08” Paraná I Astorga -23o23’25” -51°66’55” Paraná I Maringá -23o42’52” -51°93’86” Paraná I Londrina -23°31’02” -51°16’27” Paraná I Assaí -23°37’33” -50°84’13” Table 1. Geographical location of the collect sites visited by routes (I to V), during the trips. Paraná I Ibaiti -23°84’86” -50°18’77” Paraná I Santo Antônio da Platina -23°29’05” -50°07’72” Santa Catarina I Chapecó -27°07’00” -52°37’00” Santa Catarina I Cunhaporã -26°89’36” -53°16’80” Santa Catarina I Xanxerê -26°56’02.2” -52°29’55.4” Santa Catarina I Catanduvas -27°07’05” -51°66’16” Santa Catarina I Campos Novos -27°40’16” -51°22’05” Santa Catarina I Curitibanos -27°18’10.7” -50°42’24.4” Santa Catarina I Blumenau -26°55’00” -49°03’00” Santa Catarina I Lages -27°81’61” -50°32’61” Santa Catarina I Pouso Redondo -27°15’29” -49°46’02” Santa Catarina I Ibirama -27°05’69” -49°51’77” Santa Catarina I Corupá -26°42’52” -49°24’30” Paraná II Ibiporã -23o26’91” -51°04’80” Paraná II Sertanópolis -23°05’86” -51°03’63” Paraná II Bela Vista do Paraíso -22°99’66” -51°19’05” Paraná II Florestópolis -22°86’33” -51°38’72” Paraná II Jaguapitã -23°11’27” -51°53’19” Paraná II Guaraci -22°97’30” -51°64’97” Paraná II Santa Fé -23°03’75” -51°80’52” Paraná II Flórida -23°08’72” -51°95’36” Paraná II Atalaia -23°10’50.6” -52°05’50.7” Paraná II Nova Esperança -23°12’41” -52°11’31” Paraná II Paranavaí -23°07’30” -52°46’52” Paraná II Presidente Castelo Branco -23°27’80” -52°15’16” Paraná II São Jorge do Ivaí -23°43’27” -52°29’30” Paraná II Cianorte -23°66’33” -52°60’05” Paraná II Campo Mourão -24°04’55” -52°38’30” Paraná II Jaguariaíva -24°79’11” -50°01’19” Paraná II Guaíra -24°08’00” -54°25’58” State Route County Longitude Latitude Sociobiology 67(4): 514-525 (December, 2020) 521 Table 1. Geographical location of the collect sites visited by routes (I to V), during the trips. (Continuation) Paraná II Pérola -23°80’47” -53°68’36” Paraná II Cafezal do Sul -23°90’22” -53°51’25” Paraná II Santa Isabel do Ivaí -23°00’27” -53°19’66” São Paulo II Porto Primavera -22°29’08” -52°35’43” São Paulo II Teodoro Sampaio -22°53’25” -52°16’75” São Paulo II Mirante do Paranapanema -22°29’19” -51°90’63” São Paulo II Presidente Epitácio -21°76’33” -52°11’55” São Paulo II Presidente Venceslau -21°54’24” -51°48’08” São Paulo II Dracena -21°48’25” -51°53’27” São Paulo II Flórida Paulista -21°61’47” -51°17’36” São Paulo II Osvaldo Cruz -21°79’66” -50°87’86” São Paulo II Universo -22°16’27” -50°68’15” São Paulo II Quintana -22°07’25” -50°87’86” Amapá III Oiapoque -03°84’30” -51°83’05” Amapá III Clevelândia do Norte -03°92’21” -51°84’10” Amapá III Macapá -00°03’88” -51°06’64” Pará III Santa Isabel do Pará -01°29’86” -48°16’05” São Paulo IV Pirajuí -21°99’86” -49°45’72” São Paulo IV Penápolis -21°41’97” -50°07’05” São Paulo IV Andradina -20°89’61” -51°37’94” Mato Grosso do Sul IV Três Lagoas -20°75’11” -51°67’83” Mato Grosso do Sul IV Água Clara -20°31’30” -52°35’43” Mato Grosso do Sul IV Mutum -20°17’23” -52°39’51” Mato Grosso do Sul IV Ribas do Rio Pardo -20°44’30” -53°75’91” Mato Grosso do Sul IV Campo Grande -20°29’55” -54°36’39” Mato Grosso do Sul IV Anhanduí -20°37’65” -54°13’49” Mato Grosso do Sul IV Nova Alvorada do Sul -21°46’58” -54°38’38” Mato Grosso do Sul IV Jaraguari -19°46’27” -54°21’42” Mato Grosso do Sul IV Bandeirantes -19°36’35” -54°23’17” Mato Grosso do Sul IV São Gabriel do Oeste -19°03’03” -54°47’58” Mato Grosso do Sul IV Rio Verde de Mato Grosso -18°91’80” -54°84’41” Mato Grosso do Sul IV Coxim -18°50’66” -54°76’00” Mato Grosso do Sul IV Piúva -18°27’33” -54°15’02” Mato Grosso do Sul IV Sonora -17°57’69” -54°75’77” Mato Grosso IV Anhumas -17°23’47” -54°45’27” Mato Grosso IV Pedra Preta -16°62’30” -54°47’38” Mato Grosso IV Jaciara -16°42’13” -54°72’57” Mato Grosso IV Juscimeira -16°05’05” -54°88’44” Mato Grosso IV São Pedro da Cipa -16°00’05” -54°92’13” Mato Grosso IV São Vicente -15°49’20” -55°25’00” Mato Grosso IV Cuiaba -15°23’32” -55°59’11” Mato Grosso IV Jangada -15°23’55” -56°48’91” Mato Grosso IV Rosário Oeste -14°17’25” -55°33’68” Mato Grosso IV Nobres -14°36’10” -55°14’36” Mato Grosso IV Posto Gil -14°26’42” -55°11’36” Mato Grosso IV Nova Mutum -13°37’57” -52°02’04” Mato Grosso IV Piúva -13°18’45” -56°24’21” Mato Grosso IV Lucas do Rio Verde -13°05’02” -55°91’11” Mato Grosso IV Sorriso -12°54’52” -55°71’13” Mato Grosso IV Sinop -11°86’41” -55°50’25” Mato Grosso IV Chapada dos Guimarães -15°26’04” -55°46’49” Mato Grosso IV Campo Verde -15°39’27” -55°12’29” Mato Grosso IV Coronel Ponce -15°18’45” -53°44’07” Mato Grosso IV Primavera do Leste -16°10’21” -56°28’54” Mato Grosso IV Poxoréo -16°10’20” -56°28’54” Mato Grosso IV Presidente Murtinho -15°37’59” -53°57’49” Mato Grosso IV Paredão Grande -15°48’73” -53°22’81” Mato Grosso IV Coronel Meruri -15°44’23” -53°19’67” Mato Grosso IV Alto Araguaia -17°31’47” -53°21’52” Mato Grosso IV Barra do Garças -15°46’36” -52°33’44” Goiás IV Bom Jardim de Goiás -16°19’43” -51°56’35” Goiás IV Piranhas -16°26’52” -51°38’31” Goiás IV Arenópolis -16°38’61” -51°56’02” Goiás IV Iporá -16°44’19” -51°11’77” Goiás IV Israelândia -16°31’77” -50°90’80” Goiás IV Jussara -15°85’05” -50°86’80” Goiás IV Fazenda Nova -16°04’58” -50°48’11” Goiás IV Santa Fé de Goiás -15°76’91” -51°10’55” Goiás IV Juscelândia -15°40’67” -51°43’58” Goiás IV Aruanã -15°02’40” -51°05’54” Goiás IV Araguapaz -15°09’08” -50°63’22” Goiás IV Faina -15°44’61” -50°36’05” Goiás IV Goiás -15°09’08” -50°63’22” Goiás IV Itaberaí -16°02’02” -49°81’02” Goiás IV Itauçú -16°11’20” -49°36’29” Goiás IV Inhumas -16°18’26” -49°31’30” State Route County Longitude Latitude State Route County Longitude Latitude LC Forti, JS Rando, RS Camargo, AA Moreira, MA Castellani, SA Leite, KKA Sousa, N Caldato – Leaf-cutting in geographic regions of Brazil522 Goiás IV Anápolis -16°32’66’’ -48°95’27” Goiás IV Leopoldo de Bulhões -16°61’91” -48°74’36” Goiás IV Silvania -16°65’88” -48°60’80” Goiás IV Vianópolis -16°74’19” -48°51’63” Goiás IV Ourizona -17°10’23” -48°18’35” Goiás IV Urutaí -17°46’36” -48°20’16” Goiás IV Ipamerí -17°72’19” -48°15’97” Goiás IV Catalão -18°16’58” -47°94’63” Minas Gerais IV Araguari -18°64’72” -48°18’72” Minas Gerais IV Uberlândia -18°43’54” -48°13’22” Minas Gerais IV Tupassiguara -18°53’36” -48°39’41” Minas Gerais IV Monte Alegre de Minas -18°87’05” -48°88’08” Minas Gerais IV Prata -19°23’42” -48°53’49” Minas Gerais IV Frutal -20°02’47” -48°94’05” São Paulo IV Nova Granada -20°53’38” -49°31’41” São Paulo IV São José do Rio Preto -20°81’97” -49°37’94” São Paulo IV Catanduva -21°13’77” -48°97’27” São Paulo IV Santa Adélia -21°24’27” -48°80’41” São Paulo IV Ururaí -21°39’62” -48°29’34” São Paulo IV Matão -21°60’33” -48°36’58” São Paulo IV Boa Esperança do Sul -21°50’38” -49°34’52” São Paulo IV Barra Bonita -22°49’47” -48°55’80” São Paulo IV Espírito Santo do Turvo -22°69’22” -49°43’02” São Paulo IV Avaré -23°89’06” -48°92’58” Mato Grosso do Sul V Aparecida do Taboado -20°08’66” -51°09’36” Mato Grosso do Sul V Raimundo -19°25’22” -51°23’03” Mato Grosso do Sul V Mâncio Lima -07°34’09” -72°49’42” Mato Grosso do Sul V Guajará -07°03’23” -72°34’24” Mato Grosso do Sul V Dourados -22°15’40” -54°69’18” Mato Grosso do Sul V Caarapó -22°63’41” -54°82’22” Mato Grosso do Sul V Amambai -23°04’58” -55°08’41” Mato Grosso do Sul V Sanga Puitã -22°58’35” -55°87’29” Mato Grosso do Sul V Ponta Porã -22°53’61” -55°72’55” Mato Grosso do Sul V Jateí -22°48’19” -54°30’25” Mato Grosso do Sul V Deodápolis -22°31’52” -54°12’30” Mato Grosso do Sul V Ivinhema -22°30’47” -53°81’52” Goiás V Paranaíba -19°47’06” -52°06’54” Goiás V Itajá -19°05’46” -51°37’50” Goiás V Cassilândia -19°04’58” -51°42’33” Goiás V Aporé -19°02’42” -51°48’51” Goiás V Serranópolis -18°30’61” -51°96’22” Goiás V Jataí -17°88’13” -51°71’44” Goiás V Mineiros -17°56’94” -52°55’11” Goiás V Perolândia -17°52’86” -52°06’41” Mato Grosso V Portelândia -17°35’36” -52°67’86” Mato Grosso V Santa Rita do Araguaia -17°32’55” -53°20’52” Mato Grosso V Alto Garças -16°94’38” -53°52’80” Mato Grosso V Poconé -16°01’11” -56°39’06” Mato Grosso V Cangas -16°23’47” -56°12’65” Mato Grosso V Cáceres -16°07’05” -57°67’88” Mato Grosso V Mirassol D’Oeste -15°49’47” -58°11’03” Mato Grosso V Porto Esperidião -15°85’27” -58°46’02” Mato Grosso V Pontes e Lacerda -15°22’61” -59°33’52” Rondônia V Vila Bela da Santíssima Trindade -15°00’80” -59°95’05” Rondônia V Comodoro -13°66’30” -39°78’58” Rondônia V Padronal -12°55’57” -60°02’35” Rondônia V Vilhena -12°74’05” -60°14’58” Rondônia V Marco Rondon -12°11’32” -60°49’38” Rondônia V Cacoal -11°43’86” -61°44’72” Rondônia V Castanhal -01°29’38” -47°92’63” Rondônia V Ji-Paraná -10°87’55” -61°94’91” Rondônia V Ouro Preto do Oeste -10°71’63” -62°24’77” Rondônia V Jaru -10°43’88” -62°46’63” Rondônia V Nova Vida -10°27’30” -62°15’28” Rondônia V Ariquemes -09°49’47” -58°11’03” Rondônia V Porto Velho -08°44’50” -63°53’08” Rondônia V Jamari -08°45’44” -63°44’41” Rondônia V José Bonifácio -09°43’46” -64°08’37” Rondônia V Jaci Paraná -09°18’05” -64°36’16” Rondônia V Mutum Paraná -09°18’05” -64°36’16” Rondônia V Abunã -09°40’58” -65°05’27” Pará V Pimenta Bueno -11°67’25” -61°19’36” Acre V Vista Alegre do Abunã -09°38’45” -65°09’32” Acre V Extrema de Rondônia -09°44’56” -67°03’42” Acre V Plácido de Castro -10°14’46” -67°40’43” Acre V Rio Branco -10°61’07” -67°45’24” Amazonas V Cruzeiro do Sul -07°36’07” -72°43’23” State Route County Longitude Latitude Table 1. 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