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An Experimental Study of Vertical Habitat Use and Habitat Shifts in ...

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Introduction<br />

Non-native fishes negatively impact native populations through multiple<br />

pathways (Meffe 1985, Ross 1991, Mooney <strong>and</strong> Clel<strong>and</strong> 2001), <strong>in</strong>clud<strong>in</strong>g competition for<br />

food or space (Hocutt <strong>and</strong> Hambrick 1973, Mathur 1975, Walser et al. 2000). Follow<strong>in</strong>g<br />

ecological theories <strong>of</strong> competitive exclusion (Hard<strong>in</strong> 1960), limit<strong>in</strong>g similarity<br />

(MacArthur <strong>and</strong> Lev<strong>in</strong>s 1967) <strong>and</strong> niche (Pielou 1972, Pianka 1974), competitive<br />

<strong>in</strong>teractions are expected between non-native species <strong>and</strong> closely-related native species.<br />

Aquarium-based studies <strong>of</strong> vertical segregation <strong>in</strong> m<strong>in</strong>nows have emphasized expected<br />

similarities with<strong>in</strong> behaviors <strong>and</strong> habitat use <strong>of</strong> closely-related species (i.e., an historical<br />

ecology emphasis on phylogenetic constra<strong>in</strong>ts on habitat use, Gorman 1988, 1992). Also,<br />

researchers have emphasized morphological correlates on use <strong>and</strong> segregation <strong>of</strong> habitats<br />

among closely-related species, such as mouth position <strong>and</strong> its determ<strong>in</strong>istic <strong>in</strong>fluence on<br />

benthic, pelagic, or surface forag<strong>in</strong>g (Mendelson 1975, Surat et al. 1982, Page <strong>and</strong><br />

Sw<strong>of</strong>ford 1984). Alternatively, some closely-related species <strong>in</strong>tegrate as mixed-species<br />

shoals, such as m<strong>in</strong>nows (Cypr<strong>in</strong>idae), <strong>and</strong> this strategy may improve forag<strong>in</strong>g success or<br />

predator avoidance (Mendelson 1975, Magurran <strong>and</strong> Pitcher 1983, 1987). Multispecific<br />

aggregations suggest a historic legacy <strong>of</strong> strong behavioral relationships amongst<br />

cypr<strong>in</strong>id guilds. Both mutual responsiveness <strong>and</strong> selective forces, required to reduce<br />

competition <strong>and</strong> ma<strong>in</strong>ta<strong>in</strong> species identity, are required for mixed-species shoals to<br />

function (Mendelson 1975). However, little is known <strong>of</strong> the occurrence <strong>and</strong><br />

consequences <strong>of</strong> mixed-species shoals <strong>of</strong> native <strong>and</strong> non-native m<strong>in</strong>nows. Given recent<br />

conservation concerns <strong>of</strong> fish <strong>in</strong>troductions <strong>and</strong> impacts <strong>of</strong> non-natives (Rahel 2000),<br />

additional research needs focused on native/non-native species <strong>in</strong>teractions.<br />

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