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INTRODUCTION
Widespread concern exists over the health of marine ecosystems given the diversity and impact of human activities affecting these systems worldwide (Halpern et al. 2008), with overfishing consistently identified as a key threat to all parts of the world's oceans (Pauly et al. 1998). Overexploitation has resulted in dire predictions about the ability of oceans to continue supporting fisheries (Worm et al. 2006), in turn leading to calls for fishery management reform (Costello et al. 2008) and increased protection to maintain some locations in a more 'pristine' ecosystem state (Lubchenco et al. 2003). No-take marine reserves, which prohibit all exploitative activities, have become widely recognized as an effective conservation tool for protecting marine resources, resulting in increases in the abundance, biomass and diversity of many species within their boundaries (Halpern 2003; Micheli et al. 2004; Lester et al. 2009). However, the ability for reserves to provide conservation or fisheries benefits to adjacent waters remains highly controversial (Roberts et al. 2001; Tupper 2002; Gell & Roberts 2003; Hilborn et al. 2004).
Growing evidence from theoretical models and empirical studies suggests that higher abundances inside reserves can lead to spillover of adults to nearby fished areas (Rowley 1994; Roberts et al. 2001; Goni et al. 2008; Kellner et al. 2008; Perez-Ruzafa et al. 2008). Spillover is typically observed as declining patterns of abundance or catch across reserve boundaries, although there may be no detectable pattern of decline in abundance if fishing effort is intense surrounding the reserve. Additional evidence comes from ecosystem-level models evaluating the effect of large closures on fish stocks and landings (for example see Guenette et al. 2000; Pitcher et al. 2002; Zeller & Reinert 2004), measures of changing fisheries yield or profit adjacent to reserves (McClanahan & Mangi 2001; Alcala et al. 2005; Abesamis et al. 2006a) and from studies of small-scale adult movement patterns across reserve boundaries (Samoilys 1997; Zeller et al. 2003; Topping et al. 2005). However, these studies cannot address the extent to which small-scale reserves may benefit local fishers (in the case of the ecosystem-level models) or determine if reserves can...