Geographical and seasonal distributions of the seedeaters Sporophila bouvreuil and Sporophila pileata (aves: Emberizidae)

Geographical and seasonal distributions of the seedeaters Sporophila bouvreuil and Sporophila pileata (aves: Emberizidae)

Érika Machado^I,II; Luís Fábio Silveira^I,III

^IDepartamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11.461, CEP 05422-970, São Paulo, SP, Brasil and Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42.494, CEP 04218-970, São Paulo, SP, Brasil

^IIE-mail: [e-mail]

^IIIE-Mail: [e-mail]

ABSTRACT

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.

Keywords: Aves; Sporophila pileata; Sporophila bouvreuil; Migration; Seasonality.

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 algumas regiões. A distribuição de Sporophila bouvreuil se estende desde a região amazônica e Suriname passando pelas áreas abertas de boa parte do Brasil, enquanto que S. pileata distribui-se pelo sul e sudeste do Brasil, além da Argentina e Paraguai. Os movimentos migratórios destas duas espécies são pouco conhecidos e há evidências de que algumas populações podem ser residentes durante todo o ano. Com o objetivo de definir a distribuição geográfica e sazonal de S. bouvreuil e S. pileata, os registros foram agrupados em dois tipos, um durante o período reprodutivo (setembro a março) e outro durante os meses de abril a agosto, quando supostamente estas espécies podem migrar. Foram identificadas duas áreas de simpatria entre S. bouvreuil e S. pileata, nos estados de Minas Gerais e São Paulo. Para S. bouvreuil sugere-se que as populações que reproduzem na Amazônia se deslocam para o Cerrado ou a Caatinga, onde se encontram com as populações residentes nestes biomas. As populações desta diagonal seca se reproduzem nestes domínios e provavelmente realizam apenas migrações de curta distância. Na Mata Atlântica parte das populações são residentes, permanecendo neste bioma durante todo o ano. As populações de S. pileata que ocorrem no Cerrado e nas áreas abertas da Mata Atlântica se reproduzem nos seus respectivos biomas, mas não foi possível identificar as possíveis áreas de invernagem para esta espécie.

Palavras-Chave: Aves; Caboclinho; Sporophila; Migração; Sazonalidade.

INTRODUCTION

The genus Sporophila is widespread in the Americas with species found from Texas in the United States (S. torqueola) to Argentina (e.g., S. caerulescens, S. ruficollis). They are typically found in open habitats, while some species may be found in the forest interior (Schauensee, 1952; Austin, 1962; Paynter, 1970; Ridgely & Tudor, 1989; Sibley & Monroe, 1990). Four subspecies were traditionally recognized within the species Sporophila bouvreuil. A recent review (Machado, 2008; Machado & Silveira, submitted) shows that only two of those taxa are valid, but as valid species: S. bouvreuil (Statius Müller, 1776) and S. pileata (Sclater, 1864). Sporophila bouvreuil is found from the Brazilian state of Amapá, through the state of Amazonas, the islands of Mexiana and Marajó, in the state of Pará, to central Brazil, in the states of Goiás and Mato Grosso, to Maranhão, Pernambuco, Bahia, Espírito Santo, Minas Gerais, Rio de Janeiro and northeastern São Paulo. There is also an isolated population in the savannas of southern Suriname (Ridgely & Tudor, 1989; Silva et al., 1997; O'Shea, 2005; Aleixo & Poletto, 2007). Sporophila pileata, in contrast, is restricted more to the southwest of Brazil, in the states of São Paulo, Minas Gerais, Mato Grosso, southern Goiás and Mato Grosso do Sul, and reaching in the south the states of Parana and Rio Grande do Sul and barely into Argentina (Corrientes and Missiones) and Paraguay (Schauensee, 1952, 1966; Sick, 1967, 1997; Paynter, 1970; Ridgely & Tudor, 1989; Sibley & Monroe, 1990; Belton, 1994; Howard & Moore, 1994; Mikich & Bérnils, 2004).

Evidence suggests that many species in the genus Sporophila are migratory (Schauensee, 1952; Ridgely & Tudor, 1989; Sick, 1997; Areta & Almirón, 2009; Areta et al., 2009), yet very little is known of their migratory patterns and movements (Silva, 1995). In Brazil, migration in Sporophila seems to be determined by rainfall, temperature and availability of food, such as seed production, that may be very seasonal in some regions (Silva, 1995; Sick, 1997; Capllonch, pers. com.). These seedeaters are social and often are found in small groups or large flocks, except during reproduction (Schauensee, 1952; Austin, 1962; Ridgely & Tudor, 1989).

Few studies address seasonal movement patterns and migration in the genus Sporophila (Silva, 1995, 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: Departamento de Zoologia, Universidade Federal de Minas Gerais (DZUFMG), in Belo Horizonte, Minas Gerais; Museu de Biologia Melo Leitão (MBML) in Santa Teresa, Espírito Santo; Museu de Ciências Naturais da Fundação Zoobotânica (MCN) in Porto Alegre, Rio Grande do Sul; Museu de História Natural de Taubaté (MHNT), Taubaté, São Paulo; Museu Paraense Emílio Goeldi (MPEG) in Belém, Pará; Museu de Zoologia da Universidade de São Paulo (MZUSP) in São Paulo; Museu Nacional do Rio de Janeiro (MNRJ) in Rio de Janeiro; and Coleção Rolf Grantsau (RG), São Bernardo do Campo, São Paulo. Additionally other international institutions were consulted: American Museum of Natural History (AMNH), New York; The Academy of Natural Sciences (ANSP), Philadelphia; Field Museum of Natural History (FMNH), Chicago; Museum of Comparative Zoology (MCZ), Cambridge; Natural History Museum of Los Angeles County (LACM), Los Angeles; Naturhistorisches Museum Wien (NMW), Vienna; United States National Museum, Smithsonian Institution (USMN), Washington; Muséum National d'Histoire Naturelle (MNHN), Paris; British Museum of Natural History (BMNH), Tring; Zoologisches Staatssammlung München (ZSM), Munich; Rijksmuseum van Natuurlijke Histoire (RMNH), Leiden. Records from international institutions provided 26 specimens of S. bouvreuil and 30 of S. pileata. Finally, additional data from published records and field observations (ours as well as those of our colleagues) supplemented the above sources. 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.

RESULTS AND DISCUSSION

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 and 2005, 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.

ACKNOWLEDGMENTS

We thank the Programa de Coordenação e Aperfeiçoamento de Pessoal do Ensino Superior (Capes) and the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP Process number 06/53819-3) and the Programa de Apoio à Pós-Graduação (PROAP) for their financial support. We greatly appreciate the help and cooperation of all the curators at the collections we visited in Brazil, especially M. Rodrigues (DZUFMG), H. Fernandes (MBML), H. Alvarenga (MHNT), A. Aleixo (MPEG), M. Raposo (MNRJ), G.A. Bencke (MCN) and R. Grantsau (RG), as well as the curators of international museums and collections: Paul Sweet (American Museum of Natural History – AMNH); Natham H. Rice (The Academy of Natural Sciences – ANSP); Kimball Garret (Natural History Museum of Los Angeles County – LACM); Claudia Angle (United States National Museum, Smithsonian Institution – USMN); Gerald Mayr (Natural History Museum of Frankfurt), R. Prys-Jones and Mark Adams (British Museum of Natral History – BMNH), C. Voisin and E. Pasquet (Muséum National d'Histoire Naturelle – MNHN), J. Reichholf and R. Diesner (Zoologisches Staatssammlung München – ZSM), and R. Dekker and C. Peppermans (Rijksmuseum van Natuurlijke Histoire Leiden – RMNH). We also appreciate the help, records and photos of many ornithologists and friends who contributed information about the distribution patterns of these seedeaters. Herculano Alvarenga, Mônica Toledo-Piza and Marcelo Vasconcelos commented the first draft of the manuscript. James J. Roper translated this text from the original Portuguese. LFS receives a grant from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) and is an Associate Researcher of the World Pheasant Association.

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Recebido em: 20.07.2009

Impresso em: 20.09.2010

Aceito em: 10.12.2010

Appendix 1

Publication Dates

• Publication in this collection
  13 Dec 2010
• Date of issue
  2010

History

• Received
  20 July 2009
• Accepted
  10 Dec 2010