176 Chalker-Scott: Nonnative, Noninvasive Woody Species Can Enhance Urban Landscape Biodiversity Surprisingly, native bird numbers and species richness can increase with the development status of land, perhaps correlating to the diversity of trees and shrubs planted. Suburban, residential areas have greater numbers of birds (Sewell and Catterall 1998; Shwartz et al. 2008; Catterall et al. 2010) and bird species (Sewell and Catterall 1998; Reis et al. 2012; Taylor et al. 2013) compared to other systems, including remnant forests (Sewell and Catterall 1998), rural areas (Leveau and Leveau 2005), and other unmanaged systems (Shwartz et al. 2008). While forests tend to provide mainly seed as a food source, urban gardens provide a wide range of resources, including fruit, nectar, and pollen (Smith et al. 2006; van Heezik et al. 2013). Moreover, an urbanized landscape provides resources throughout the year, which is of particular benefit to native birds during the winter (Atchison and Rodewald 2006). Large trees and large habitats are important to native bird species. In environments that vary from Northern California to tropical Mexico to the Canary Islands, bird species richness is posi- tively related to tree height (Palomino and Carrascal 2005; MacGregor-Fors 2008; Kalinowski et al. 2010; Strubbe et al. 2010; Stagoll et al. 2012). Likewise, native bird species richness increases with increased habitat size, particularly that of insectivores, hollow- nesters, and other forest species (Donnelly and Marzluff 2004; Donnelly and Marzluff 2006; Posa and Sodhi 2006; Lu et al. 2007; Hudson and Bird 2009; Khera et al. 2009; Strubbe et al. 2010; Ikin et al. 2013a; van Heezik et al. 2013; Yu and Guo 2013). Tree provenance appears to influence bird popu- lations more than other animal groups, with several papers associating greater species richness with native tree density (Pennington et al. 2008; Helden et al. 2012; Petrova and Irikov 2012; Reis et al. 2012; Ikin et al. 2013b). This may be more a failure to retain appropriate habitat composition rather than native plant species, however, as an ecologically functional environment is required for improving native bird diversity (Parsons et al. 2004) and may be more important than provenance. Birds, including native forest species, prefer urban sites that retain veg- etative characteristics such as structure (Chace and Walsh 2006; Hodgkison et al. 2007; van Heezik et al. 2008; Khera et al. 2009; Echevarria et al. 2011) and density (Hennings and Edge 2003; Chen et al. 2005; Donnelly and Marzluff 2006; Faggi and Perepelizin ©2015 International Society of Arboriculture 2006; Posa and Sodhi 2006; Lu et al. 2007; Luther et al. 2008; MacGregor-Fors 2008; Pennington et al. 2008; van Heezik et al. 2008; Kalinowski et al. 2010; de Toledo et al. 2012), which could be supplied either by native or carefully selected introduced species. While some native birds are undoubtedly depen- dent on native tree species, others have learned to utilize introduced trees, including invasives, for food [e.g., holly berries (Ilex spp.), figs (Ficus spp.), and honeysuckle (Lonicera spp.) nectar] and habitat for nesting and predator avoidance (e.g., Cytisus scoparius) (Sewell and Catterall 1998; Crooks et al. 2004; Feldman and Krannitz 2004; Hasebe and Franklin 2004; Leveau and Leveau 2005; Atchison and Rodewald 2006; Kath et al. 2009; Gleditsch and Carlo 2011; Caughlin et al. 2012; Helden et al. 2012; Moller et al. 2012). The relationship between native birds, especially fru- givorous species, and introduced plants is com- plex. Not only do many invasive trees and shrubs expand habitat choices for native birds, but these birds contribute to invasive plant spread through fruit consumption and seed dispersal (Reichard et al. 2001; Caughlin et al. 2012; Moller et al. 2012). In summary, native bird species abun- dance and richness are positively influenced by: • habitat connectivity and size (Donnelly and Marzluff 2004; Donnelly and Marzluff 2006; Posa and Sodhi 2006; Hudson and Bird 2009; Khera et al. 2009; Strubbe et al. 2010; van Heezik et al. 2013; Yu and Guo 2013) • abundance and diversity of trees and shrubs (Sewell and Catterall 1998; Hennings and Edge 2003; Crooks et al. 2004; Chen et al. 2005; Palomino and Carrascal 2005; Donnelly and Marzluff 2006; Faggi and Perepelizin 2006; Posa and Sodhi 2006; Hodgkison et al. 2007; Lu et al. 2007; Luther et al. 2008; MacGregor-Fors 2008; Pennington et al. 2008; Shwartz et al. 2008; van Heezik et al. 2008; Hudson and Bird 2009; Kath et al. 2009; Khera et al. 2009; Suarez-Rubio and Thomlinson 2009; Kalinowski et al. 2010; Ortega-Alvarez and MacGregor-Fors 2010; de Toledo et al. 2012) • vertical diversity (Hodgkison et al. 2007; van Heezik et al. 2008; Khera et al. 2009) • moderately disturbed sites, such as suburbs and public greenspaces (Blair 1996; Crooks et
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