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

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In order for similar species to avoid competitive exclusion they must be capable<br />

<strong>of</strong> partition<strong>in</strong>g some aspect <strong>of</strong> the trophic, spatial, or temporal variables <strong>of</strong> habitat usage<br />

(MacArthur <strong>and</strong> Lev<strong>in</strong>s 1964, Pianka 1974). Effects <strong>of</strong> competition, such as decreased<br />

survivorship <strong>and</strong> fecundity, are reduced through natural selection <strong>in</strong> lieu <strong>of</strong> more<br />

favorable traits (Murray 1986). Differentiation <strong>of</strong> breed<strong>in</strong>g strategies, behavior,<br />

susceptibility to biochemical hazards, or resistance to the effects <strong>of</strong> environmental<br />

stochasticity is needed for <strong>in</strong>terspecific traits to rema<strong>in</strong> dist<strong>in</strong>ct (Harper et al. 1961). Such<br />

evolutionary specializations are more pronounced <strong>in</strong> complex, heterogeneous<br />

environments (Murray 1986). Increased niche dimensionality allows greater ecological<br />

pack<strong>in</strong>g, thus allow<strong>in</strong>g an environment to accommodate a greater number <strong>of</strong> species<br />

(MacArthur <strong>and</strong> Lev<strong>in</strong>s 1967). Similarly, it is believed that the number <strong>of</strong> related species<br />

coexist<strong>in</strong>g <strong>in</strong> a given ecosystem is strongly dependent on the width <strong>and</strong> overlap <strong>of</strong> those<br />

species’ niches (Pielou 1972). Complete, non-<strong>in</strong>terbreed<strong>in</strong>g competitors cannot coexist<br />

without evolutionary divergence, readjusted niche widths, or competitive elim<strong>in</strong>ation <strong>of</strong><br />

one <strong>of</strong> the species (Hard<strong>in</strong> 1960, Pianka 1966, Pianka 1974). Problems occur when<br />

ecological processes set <strong>in</strong>to place through disturbances disrupt stable evolutionary<br />

processes (Pianka 1966). The <strong>in</strong>troduction <strong>of</strong> a non-<strong>in</strong>digenous species that acquires<br />

already utilized niche space <strong>in</strong> a new region would be an example <strong>of</strong> such a disturbance.<br />

The chance exists for competitive exclusion to occur if two similar, geographically<br />

separated species are mixed (Hard<strong>in</strong> 1960). To make matters worse, established<br />

<strong>in</strong>troduced species <strong>of</strong>ten utilize a realized niche that more closely resembles its<br />

fundamental niche <strong>in</strong> the host community (Li <strong>and</strong> Moyle 1981). Thus, species<br />

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