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Toco Toucan Feeding Ecology and Local Abundance in a Habitat Mosaic in the Brazilian Cerrado

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Toco Toucan Feeding Ecology and Local Abundance in a Habitat Mosaic in the Brazilian Cerrado ORNITOLOGIA NEOTROPICAL 19: 345–359, 2008 © The Neotropical Ornithological Society TOCO TOUCAN FEEDING ECOLOGY AND LOCAL ABUNDANCE IN A HABITAT MOSAIC IN THE BRAZILIAN CERRADO José Ragusa-Netto Departamento de Ciências Naturais, Campus Três Lagoas, Universidade Federal do Mato Grosso do Sul, C.P. 210, 79620-080, Três Lagoas, MS, Brazil. E-mail: [email protected] Resumo. – Ecologia alimentar e abundância local do Tucano Toco, em um mosaico de habitats do cerrado. – Os tucanos são frugívoros do dossel que exploram áreas amplas e heterogêneas. O Tucano Toco (Ramphastos toco) é comum no Brasil Central, principalmente no cerrado. Nesse estudo avaliei a pro- dução de frutos, a abundância do Tucano Toco e seus hábitos alimentares em um mosaico de habitats do cerrado. Tanto as variações espaciais quanto temporais de abundância dos tucanos coincidiram com o perí- odo de frutificação das espécies consumidas extensivamente. Essas espécies, principalmente Virola sebifera na mata ciliar e Schefflera macrocarpa no cerrado, exibiram períodos prolongados de frutificação, além de serem conhecidas por produzirem diásporos com elevado teor de lipídeos. Por outro lado, com exceção de Eugenia punicifolia e Miconia albicans, os tucanos consumiram moderadamente muitos tipos de frutos ricos em açúcares, que estiveram disponíveis por breves períodos. Portanto, em razão do Tucano Toco explorar, durante a maior parte do tempo, proporções elevadas de poucas espécies de frutos e oportunamente alter- nar para uma dieta mais variada, ele exibiu variações acentuadas de amplitude de nicho alimentar. As pro- fundas variações espaciais e temporais de abundância, ao longo do ano, sugerem que os tucanos exploram áreas amplas e heterogênas em resposta à disponibilidade de frutos que são importantes em sua dieta. Potencialmente, toda essa flexibilidade, pelo menos em parte, está relacionada com a abundância do tucano toco no altamente heterogêneo e fortemente sazonal cerrado, onde nenhuma outra espécie de Ram- phastos é comum. Abstract. – Toucans are canopy frugivores which forage over large and diverse areas. Toco Toucan (Ramphastos toco) is common in the dry interior of Brazil, mainly in the ‘cerrado’ areas, dominated by savanna like habitats. In this study, I evaluated fruit production, Toco Toucan abundance and their feeding activity within a habitat mosaic in the Brazilian cerrado. Toco Toucan exhibited substantial temporal and spatial variations of abundance, which coincided with the availability of specific fruits. These food resources, mainly Virola sebifera in gallery forests and Schefflera macrocarpa in the cerrado, presented such traits as prolonged availability and lipid-rich diaspores. Conversely, with the exception of both Eugenia punicifolia and Miconia albicans fruits, toucans foraged moderately on a variety of briefly available sugar-rich fruits. As a result, due to an extensive use of few fruit types alternated with periods of an opportunistic broad diet, Toco Toucan exhibited substantial variations of niche breadth. The enhanced spatial and tem- poral variations of Toco Toucan abundance suggest a year-round use of wide and diverse areas mainly in response to fruiting species, which comprise the bulk of their diet. Potentially, all those flexibility, at least partly, may explain Toco Toucan abundance in the highly diverse and marked seasonal cerrado, in which no other Ramphastos species is common. Accepted 15 April 2008. Key words: Ramphastidae, Ramphastos toco, Toco Toucan, toucans, cerrado, abundance, frugivory, feeding ecology, canopy phenology, central Brazil. 345 RAGUSA-NETTO INTRODUCTION 2002). The typical habitat types of cerrado differ both in soil quality and moisture Toucans (Ramphastidae) inhabit a wide range (Oliveira-Filho et al. 1990), besides tree spe- of Neotropical areas occurring from dry to cies composition (Ribeiro & Walter 1998). tall rain forests (Short & Horne 2002). These Consequently, variable fruiting patterns are prominent birds use large home ranges in expected within this habitat mosaic (van which they forage mostly for canopy fruits Schaik et al. 1993, Oliveira 1998), which can (Terborgh et al. 1990). In this respect, toucans potentially affect the feeding ecology and recurrently may experience food shortage local patterns of abundance of Toco Toucans. due to the seasonality of fruit production in Detailed knowledge on the relationships this vegetation layer (Frankie et al. 1974, between Toco Toucans and food resources Bullock & Solis-Magallanes 1990). Therefore, may allow the identification of important fruit toucans often move from one to another species, habitats, and critical areas for conser- habitat type in response to fruit availability vation. This issue is of special concern (Graham 2001, Ragusa-Netto 2006). In because, despite Toco Toucans persisted in fact, food resources are among the major fragmented areas (Short & Horne 2002, pers. causal factors influencing both the move- observ.), it is under a severe process of habitat ments and local abundance of canopy frugiv- loss due to the accelerated clearing of cerrado orous birds (Kinnaird et al. 1996, Anggraini et areas (Ratter et al. 1997). Hence, studies on al. 2000, Solorzano et al. 2000). Despite their ecology could contribute to conserva- relying mostly on fleshy fruits, toucans also tion plans, which may be important to pre- prey upon large arthropods and small verte- vent pronounced population declines. To brates and depredate nests (Skutch 1971, Sick improve our knowledge on the dynamics of 1997, Remsen et al. 1993, Short & Horne both Toco Toucan use of foraging areas and 2002). In the rain forests, toucans often for- diet, in this study I evaluated fruit production age on large and oily fruits (Galetti et al. 2000; as well as Toco Toucan abundance in a habitat but see Chaves-Campos 2004), which they mosaic of the cerrado. In addition, I exam- disperse efficiently (Howe 1981, 1993). Nev- ined Toco Toucan feeding activity and ana- ertheless, in dry areas, species such as the lyzed the relationships between this Toco Toucan (Ramphastos toco) tend to exploit parameter and both fruit and Toco Toucan extensively less rewarding diaspores such as abundance in each habitat type. figs and Cecropia catkins (Ragusa-Netto 2002, 2006). METHODS The Toco Toucan, the largest toucan spe- cies (over 500 g), mostly occurs in the dry Study area. This study was carried out in the interior of Brazil (Sick 1997, Short & Horne Emas National Park (hereafter ENP), which 2002). It is particularly common in the Brazil- is in the cerrado core region. The total area is ian “cerrado” (Neotropical savanna), which 133 000 ha, located in the Brazilian Central vegetation ranges from open fields to dense Plateau, in the southwest of the State of deciduous wood (cerradão), besides the ever- Goiás (17°19’–18º28’S and 52º39’–53°10’W, green gallery forests and palm stands (Ribeiro altitude 900–1100 m). The climate is seasonal & Walter 1998). In this respect, the Toco Tou- and marked with wet (October to March) and can is singular in its use of both savanna like prolonged dry (April to September) seasons. and dense habitats, instead of only using con- Annual rainfall is approximately 1500 mm tinuous forests (Sick 1997, Short & Horne (70% in the wet season), and mean annual 346 TOCO TOUCAN AND FRUITS IN THE CERRADO temperature lies around 24.6°C (Batalha & & Walter 1998). Due to the implications of Martins 2002). this heterogeneity in the spatial and temporal The vegetation in the area is a mosaic of patterns of fruit availability, to sample the gallery forest, palm (Mauritia flexuosa) stands, wide cerrado vegetation, as well as both the and the dominant cerrado (93% of the area), slender gallery forest and palm swamp, I which exhibit a gradient from open fields to adopted a stratified sampling design within dense wood vegetation. However, 70% of the which a system of points was positioned. The cerrado is semi-open (savanna like habitat) in number of sample points to assess fruit pro- which trees are interspersed with open grassy duction was defined according to the propor- areas (Batalha & Martins 2002). During the tion of each habitat type in the sampled area dry season, tree species drop their leaves, (rectangle of 9 x 30 km, determined using a mainly in the late dry season (August and Sep- map, scale 1 : 50 000), in the southern part of tember). The richest plant families are Aster- ENP. Also, the distance between points was aceae, Fabaceae, Poaceae, and Myrtaceae (for inversely related to tree density in each habitat details see Batalha & Martins 2002). Data col- type, which crudely was at least twice higher lection was conducted in the southern part of in the gallery forest. In the cerrado vegetation, ENP (18º15’S and 52º53’W, altitude 900 m). I sampled fruit production using 36 points This area is dominated by semi-open cerrado (1000 m apart from each other) along three (hereafter cerrado vegetation) cut by the For- 11–km permanent access trail (12 points/ moso river (west-east direction) and by the trail). In the gallery forest, I positioned 12 Buriti torto stream (north-south direction). points (500 m apart from each other; total 120 The dominant vegetation in the Buriti torto trees) along 6-km permanent access trail (hereafter palm swamp) consists of Mauritia while, in the palm swamp, eight points were flexuosa palms besides scattered trees of spe- used (1000 m from each other). At each point cies such as Xylopia emarginata, and Virola in the cerrado vegetation, the 10 nearest trees sebifera. The soil alongside this stream is wet or with diameters at base equal to or greater than even flooded. The evergreen riparian vegeta- 10 cm were numbered with aluminum tags (n tion of Formoso river (hereafter gallery for- = 360 trees). This procedure assures the est) is very dense and dominated by Xylopia inclusion of mature trees. On the other hand, emarginata. The canopy is around 12–17 m in in the gallery forest, I marked the 10 nearest height, but emergent trees may reach 25 m.
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