GeoGraphical and SeaSonal diStributionS of the SeedeaterS Sporophila bouvreuil and Sporophila pileata ( aveS : emberizidae )

Many species in the genus Sporophila are migratory. Migration patterns, while poorly studied, may be influenced by seed production which can be very seasonal in some regions. The distribution of S. bouvreuil extends from the Amazon and Suriname south through a large part of the open regions of Brazil. Sporophila pileata, on the other hand, is found in southeastern and southern Brazil as well as Argentina and Paraguay. Both of these species migrate, but their movement patterns are poorly known. To better understand the geographical and the seasonal distributions of S. bouvreuil and S. pileata, we grouped the records into two categories: the breeding season (September to March) and the putative migration season (April to August). We found two areas of sympatry between S. bouvreuil and S. pileata in the Brazilian states of Minas Gerais and São Paulo. For S. bouvreuil we suggest that populations that breed in the Amazon migrate to the Cerrado or Caatinga, where they will encounter resident populations of the same species. These resident populations may take part in short distance migrations. Sporophila pileata, on the other hand, occur in the Cerrado and open areas within the Atlantic Forest and it is not yet possible to determine migratory tendencies or destinations in the non-breeding season.

Few studies address seasonal movement patterns and migration in the genus Sporophila (Silva, 1995(Silva, , 1999;;D'Angelo-Neto & Vasconcelos, 2007;Ortiz & Capllonch, 2007).Data gathering to examine migration patterns is very difficult, due to the mixture of non-migratory with migratory individuals (Willis, 1986).On the other hand, an effective bird banding program with these birds will be indispensable as a tool for uncovering migration patterns (Sick, 1983), such as that used in the study of migration in S. caerulescens in Argentina (Ortiz & Capllonch, 2007).
In an analysis of the seasonal distribution and migration in two populations of S. lineola, two populations, one from the Caatinga and another from southern Brazil, could be described (Silva, 1995).Subsequently, three types of movement patterns were discovered in the Sporophila: local (five species), regional (four species) and long distance (14 species, including Sporophila bouvreuil).Also, wintering grounds (non-breeding) were identified for 11 of the 14 long distance migrants (Silva, 1999).
Recent study of migration in S. c. caerulescens in northern Argentina determined that males arrived on the breeding grounds in mid November, followed approximately two weeks later by females.Captures increased through December when all individuals showed signs of reproductive activity.By early January reproduction was at its peak and young birds were observed in February, while in April migration began again.However, the wintering ground is still unknown (Ortiz & Capllonch, 2007).Field observations with S. lineola in Minas Gerais suggest that birds from the south migrate from the southern and eastern part of the state in towards the western central region of the state (D' Angelo-Neto & Vasconcelos, 2007).
Other species of seedeaters have been found in Minas Gerais during September on the São Francisco River (S. castaneiventris and S. cinnamomea) and in Pirapora (S. palustris and S. melanogaster; Sick, 1997).In grasslands on an island in the upper São Francisco River hundreds of seedeaters were seen flocking together, including S. hypoxantha, S. melanogaster, S. ruficollis, S. palustris, S. bouvreuil, S. nigricollis, S. cinnamomea and S. castaneiventris (Sick, 1997).Sporophila melanogaster is known to breed in the state of Santa Catarina in October and November and populations begin to disappear by late February (Piacentini, pers. com.).In March in Rio Grande do Sul this species was observed foraging, while also in February and March it was seen in the Distrito Federal (Sick, 1997), suggesting that it also has sedentary and migratory populations.Sporophila hypoxantha is found in large flocks between November and December in the Pantanal of Mato Grosso and in the high plains of São Joaquim in the state of Santa Catarina where it was nesting at the same time (Sick, 1997).This species has also been seen along with S. hypochroma and S. palustris at the Parque Nacional das Emas, in Goiás, in October 1989 in a large, mixed species flock (Sick, 1997).Despite all these sightings and observations, movement patterns, migration and sedentary populations of Sporophila are still unknown.
Similarly, migrational or sedentary tendencies in S. bouvreuil and S. pileata are also very poorly understood, with few anecdotal observations.Sporophila bouvreuil disappears in the central plains during the driest months of the year and returns in higher concentrations along the Lago Paranoá in Brasília during October and November with the occasional rare appearance in August (Antas, 1988).This species is common during February and March in Poconé in the Pantanal in the state of Mato Grosso (Cintra & Yamashita, 1990).Sporophila pileata is apparently resident and common in the summer in the central part of the state of Rio Grande do Sul (Belton, 1994).Little else is known about the movement patterns of these two species.Here, we attempt to define summer (reproductive) and winter regions for these two species of seedeaters in Brazil.

MAterIAl And Methods
To describe locations and seasons, we used location information on the labels of 144 museum specimens of S. bouvreuil and 30 of S. pileata, housed in the following ornithological collections in Brazil: From the literature and field observations, we obtained respectively, 26 records of S. bouvreuil and nine of S. pileata, and 73 records of S. bouvreuil and 29 of S. pileata.Each unique location was considered as one coordinate (locations in Appendix 1).When possible the numbers and sexes of individuals were also noted.
Geographical coordinates were mapped with ArcView 3.3.Timing (breeding season -September to March, and wintering season -April to August) information was gathered from the records as much as possible and used to group the information by Brazilian Biomes (Cerrado, Caatinga, Amazon Forest and Atlantic Forest, classified according to Nimer (1989) and Ab'Sáber ( 2003)) to determine migration patterns.

Geographical distribution
Sporophila bouvreuil (sensu Machado, 2008 and Machado & Silveira, submitted) is found in open areas throughout most of northern South America, from Suriname (Ridgely & Tudor, 1989;O'Shea, 2005 and specimens in RMNH) and French Guiana (specimens in ANSP) to Argentina and throughout Brazil in the open habitat (Figure 1).Despite the ample distribution, some states have few records, such as Amazonas (one sighting, Aleixo & Poletto, 2007), Amapá (one, Silva et al., 1997), Tocantins (one specimen and some observations in 2001 and2005, Pacheco, pers. com.) and Rio Grande do Norte (Grupo Ornitológico Potiguar, pers. com.).The record for S. bouvreuil from Santa Teresa, in the state of Espírito Santo, is questioned and believed to be an escaped individual (Willis & Oniki, 2002), although the species had already been seen between the cities of Vitória and Santa Teresa (Forrester, 1993).Thus, the status on this area of Espírito Santo State remains still unresolved.
Sporophila pileata is limited to the southeast of Brazil (in the states of Minas Gerais, São Paulo, Mato Grosso do Sul, Goiás, Paraná, Rio Grande do Sul) and adjacent countries (Paraguay and Argentina).This species is found in the Cerrado and open areas within the Altantic Forest formation and its transition zone with the Cerrado, and is found in the Pantanal of Mato Grosso and the southern grasslands of Rio Grande do Sul.Finally, it is found in the Chaco of Paraguay and Argentina (Figure 1).seasonality Sporophila bouvreuil has been reported from 62 locations in the open areas within the Amazon Forest, found throughout the year except June, and in greatest numbers during September -November (Figure 2).The lack of reports for June may merely be a sampling artifact.September to November, when numbers are greatest, is also the breeding season.This suggests arrival on their breeding territories by September and the drop in abundance in December suggests that is when they migrate away from their breeding grounds, when they move to the Cerrado or possibly the Caatinga.
In the Cerrado and Caatinga, the species has 335 records, in every month of the year but with the greatest numbers (n = 242) during the breeding season between September and December.An increase in numbers of males in December (n = 109) suggests that they arrive after leaving the Amazon or Atlantic Forest and that these individuals will overwinter in Cerrado and Caatinga (Figure 2).And, the populations of Cerrado and Caatinga also reproduce in these regions, perhaps with some short-distance migration (Figure 3).
In the Atlantic Forest, of the total of 132 records, 103 were from the months of September to March.
More records during this period may be due to the increased activity associated with reproduction and territory defense.The few (n = 29) records from April to August suggest that at least some of the individuals do not migrate (Figure 2).Sporophila pileata was recorded 132 times in the Cerrado, Atlantic Forest and southern grasslands together, during the entire year, but more between the months of November to February, during the reproductive season (Figures 4, 5).Where these birds spend their non-breeding time remains unknown.
Sporophila bouvreuil and S. pileata are sympatric in the western part of the state of Minas Gerais, near Indianópolis and in southwestern and central São Paulo.These two species may both be found in February near Itirapina (São Paulo).In southwestern São Paulo, records are relatively close to one another near Avaré (S. pileata) and Botucatu (S. bouvreuil, the two separated by 52 km) and Aracaçú (S. pileata) and Itapetininga (S. bouvreuil, separated by 32 km).In the southeast S. bouvreuil and S. pileata have been recorded within 40 km of each other near Franco da Rocha and Itatiba, São José dos Campos and Tremembé and finally São José dos Campos and Taubaté (respectively).The two are found even closer together near São Paulo and São Bernardo do Campo (18 km), and Mogi das Cruzes and São Bernardo do Campo (33 km, respectively, Figure 1).For a species that regularly migrates greater distances (see introduction), these must be considered short distances and in fact, suggests a broad contact zone between the two species.
Both species have been recorded in the state of Mato Grosso do Sul, yet not in sympatry.Sporophila pileata has not yet been reported from Santa Catarina, yet, is known from northern Rio Grande do Sul near Santa Catarina (Damiani et al., 2007;Damiani, pers. com.).During field study in the high plains of Santa Catarina, only S. melanogaster and S. hypoxantha were found.Only S. frontalis, S. plumbea, S. collaris, S. caerulescens, S. hypoxantha and S. melanogaster are reported from the state of Santa Catarina (Rosário, 1990).
Most observation of Sporophila pileata is from the breeding season between September and March (Fontana et al., 2003).At this time, birds are very easily seen due to active territory defense including singing and flying to confront other males.The lack of records during the drier, non-reproductive months suggest that they have migrated.
Those S. bouvreuil that reproduce in the Amazon and Atlantic Forest migrate to drier regions, the Cerrado and Caatinga, during the non-breeding period (Figure 3).At that time, the migratory individuals coexist with sedentary individuals already in these regions.In the Amazon as well as the Atlantic Forest, some individuals apparently do not migrate.Other seedeater species are known to have similar partially migratory tendencies, such as S. hypoxantha and S. plumbea, in Paraná and Rio Grande do Sul (Fontana et al., 2003;Straube et al., 2004a,b).Sporophila bouvreuil probably overwinters in the Cerrado (Silva, 1999) and other species in the genus (e.g., S. cinnamomea) also spend their winters in the grasslands (Cerrado) of central Brazil (Fontana et al., 2003).While the wintering grounds of S. pileata are still unknown, it is likely that it shows similar behaviors apparently typical of the genus, such as moving to drier grasslands during the non-breeding season.The south and central regions of Brazil, including the states of santa Catarina, Mato Grosso do Sul and Goiás, is a lacuna in terms of avian sampling and should be studied more intensively.Perhaps this region is the key to elucidate the migration patterns of the Sporophila group, especially S. pileata.resuMo Muitos representantes do gênero Sporophila são migratórios.Estes movimentos são pouco estudados e podem ser determinados por fatores como a produção de gramíneas, que podem ser altamente sazonais em   peer review: All submissions to Papéis Avulsos de Zoologia and Arquivos de Zoologia are subject to review by at least two referees and the Editor-in-Chief.All authors will be notified of submission date.Authors may suggest potential reviewers.Communications regarding acceptance or rejection of manuscripts are made through electronic correspondence with the first or corresponding author only.Once a manuscript is accepted providing changes suggested by the referees, the author is requested to return a revised version incorporating those changes (or a detailed explanation of why reviewer's suggestions were not followed) within fifteen days upon receiving the communication by the editor.
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FIGure 2 :
FIGure 2: Monthly distribution of records of Sporophila bouvreuil by the main Brazilian Biomes.Dark gray: Amazonia; light gray: Cerrado and Caatinga; medium gray: Atlantic Forest and southern grasslands.

FIGure 4 :FIGure 5 :
FIGure 4: Monthly distribution map for the records of Sporophila pileata by Biomes: dark gray: Atlantic Forest; light gray: Cerrado and Caatinga.
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