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

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

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native species pairs were partially segregated, species <strong>in</strong> syntopy <strong>in</strong>teracted as a mixedgroup<br />

without <strong>in</strong>terspecific antagonistic behavior. Based on the concept <strong>of</strong> competitive<br />

displacement <strong>and</strong> depend<strong>in</strong>g on a species’ generality <strong>of</strong> habitat use, one would expect a<br />

reduced range <strong>of</strong> water-column distribution <strong>of</strong> a native species follow<strong>in</strong>g the <strong>in</strong>troduction<br />

<strong>of</strong> a closely-related species (Pielou 1972). However, competitive displacement is not a<br />

general rule <strong>in</strong> the ecology <strong>of</strong> fishes, <strong>and</strong> many studies have demonstrated coexistence <strong>of</strong><br />

closely-related species, as well as benefits <strong>of</strong> mixed-species shoals (Magurran <strong>and</strong> Pitcher<br />

1983, 1987).<br />

Water depths dur<strong>in</strong>g normal flows with<strong>in</strong> the upper Greenbrier River system are<br />

greater than those available with<strong>in</strong> the study aquaria, <strong>and</strong> therefore allow additional space<br />

for segregation between the study species. Species <strong>in</strong>tegration, however, likely results<br />

dur<strong>in</strong>g periods <strong>of</strong> reduced water column depths dur<strong>in</strong>g dry summers (Gorman 1988).<br />

Thus, vertical position patterns may change dur<strong>in</strong>g the low flows <strong>of</strong> summer, dur<strong>in</strong>g a<br />

time <strong>of</strong> competitive crunch (Wiens 1977). Our results suggest that with<strong>in</strong> a water column<br />

space <strong>of</strong> 51cm, N. scabriceps <strong>and</strong> N. telescopus rarely used space with<strong>in</strong> 17 cm <strong>of</strong> the<br />

surface. Therefore, these two species may be <strong>in</strong> close association dur<strong>in</strong>g low <strong>and</strong> high<br />

flows, although direct observational studies (such as snorkel<strong>in</strong>g) are needed to quantify<br />

<strong>in</strong>teractions between the study species with<strong>in</strong> the Greenbrier River dra<strong>in</strong>age. Both C.<br />

spiloptera <strong>and</strong> C. galactura used the upper water column position <strong>in</strong> the study aquaria,<br />

<strong>and</strong> separation between these two species may <strong>in</strong>crease with<strong>in</strong> deeper habitats <strong>of</strong> the<br />

Greenbrier River.<br />

<strong>Vertical</strong> segregation between species <strong>in</strong> this experiment, consistent with a large<br />

body <strong>of</strong> literature (Baker <strong>and</strong> Ross 1981, Gorman 1988, Chipps et al. 1993a), supports a<br />

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