Edge effects on nesting success of cavity-nesting birds in fragmented forests
Introduction
Forest fragmentation affects the distribution and abundance of organisms by reducing the amount and proximity of remnant patches of suitable habitat and increasing the amount of edges (Andrén, 1992, Boulinier et al., 1998, Ford et al., 2001, Maina and Jackson, 2003). Edges may affect the organisms by causing changes in the biotic and abiotic conditions some distance into the forest, such as increased amounts of sunlight, high wind speeds, and larger fluctuations in temperature and humidity (Saunders et al., 1991, Murica, 1995). Avian species may respond to one or a combination of these changes in the landscape as a result of different biological mechanisms (Robinson et al., 1995, Donovan et al., 1997). Species that require forest interior may avoid edges due to altered microclimate, vegetation structure, or high density of predators or brood parasites (Yahner and Scott, 1988, Malcolm, 1994, Marini et al., 1995, Pasitschniak-Arts et al., 1998, Stephens et al., 2003). On the other hand, although predators and brood parasites may be more abundant or active in the adjacent habitat and edge, some bird species were concentrated near to edge areas. This paradox of high bird abundance and richness but low nesting success near edges was termed an ecological trap by Gates and Gysel (1978).
Many studies have tested the occurrence of edge effects on nesting success of forest birds and concluded that high predation and brood parasitism rates near edges were main causes of nest failures (Hannon and Cotterill, 1998, Manolis et al., 2002). However, an edge effect has not been found in all studies (Angelstam, 1986, Santos and Telleria, 1992, Hanski et al., 1996, Morrison and Bolger, 2002). Early studies suggested that abrupt or permanent edges generally were thought to be associated with higher rates of predation and parasitism than gradual or regenerating edges (Suarez et al., 1997, Deng et al., 2003), but many contradictory results exist, particularly in forest landscapes (Yahner and Scott, 1988, Rodewald, 2002). These conflicting results may be explained in part by the mediation of edge effects by life history traits of organisms (e.g., open- vs. cavity-nesting bids) and landscape characteristics.
Most early studies were completed on open cup-nesting birds. However, nest predation and brood parasitism are unlikely to provide a general explanation for the edge effects on bird species that nest in more protected sites such as cavities (Matthysen and Adriaensen, 1998). Nesting success of cavity-nesting birds near edges may be influenced by other factors such as changes in competition for nest sites (Deng, 2001). Very few studies have assessed nesting success of cavity-nesting birds near edges in fragmented forests (Nour et al., 1993, Matthysen and Adriaensen, 1998). Many studies of effects on nesting success have used artificial nests (Wilcove, 1985, Picman and Schriml, 1994, Hartley and Hunter, 1998, Githiru et al., 2005). Such studies can identify general patterns of variation in nest predation intensity (Haskell, 1995, Sloan et al., 1998). However, predation and brood parasitism rates on artificial nests may be poorly or inconsistently correlated with predation and parasitism rates on real nests (Ortega et al., 1998, Moore and Robinson, 2004).
Here, we examine edge effects on nesting success for natural nests of four cavity-nesting bird species (two excavators and two nonexcavators) in a forested landscape in which edges were created by farmland reclamation. We also used data from these four species to determine whether life history characteristics (excavators vs. nonexcavators) affect the relationship between nest success and distances to edge.
Section snippets
Study site
Our study site, approximately 200 km2 in size, was located in Zuojia Nature Reserve and included the Tumengling Mountains and Zhujia Mountains ranging from the eastern Changbai Mountains to the western plain (126°1′127°2′N, 44°6′45°9′E). The climate is east monsoon, characterized by hot, dry summers and cold, snowy winters. Mean monthly temperatures ranged from −20.5 °C in January to 23.6 °C in August. Vegetation within the study area was quite diverse, although the forest type was secondary
Nest success by distance to forest edge
A total of 439 cavity nests of four species were located and monitored during 1998–2001 (Table 1). Of this total, 422 nests were entered into our analysis of edge effects. Nests abandoned before egg laying (n = 12) or of uncertain outcome (n = 5) were excluded from analyses. Nest success rates of each cavity-nesting bird did not vary between years (χ2 = 6.13, df = 3, P = 0.11) and plots (χ2 = 9.08, df = 5, P = 0.91). However, nest success rates among species within years were significantly different. Nest
Edge effects on cavity-nesting birds
We found some evidence for an edge effect on nesting success of cavity-nesting species near anthropogenic edges, especially for nonexcavator species. Our results concur with findings from several recent studies on open-cup nesting species (Flaspohler et al., 2001, Manolis et al., 2002). Our results are also consistent with findings from several studies of cavity-nesting birds. Denny and Summers (1996) found that cavity birds were affected by edge-related factors. Huhta and Jokimaki (2001) found
Acknowledgments
We thank Yan-Hui Li, Yang Liu, Zheng-Xin Sun, Ren-Kai Song, Gui-Quan Xiang, Hai-Tao Wang, Jing-Run Cheng, Nan Li, Quan Zhao, and Long Sun for their involvement in the collection of field data. We thank the teachers and students who are in Department of Wildlife, Special Plant and Animal College that they found and monitored the success of the nests. Members of the ecology team, Beijing Normal University, were extremely patient and helpful in providing assistance with data analysis. We thank
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