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TITLE PAGE - acumen - The University of Alabama

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are potentially capable <strong>of</strong> supporting relatively large populations <strong>of</strong> obligate cave species.<br />

Additionally, the energetic budgets provide the first quantitative explanation <strong>of</strong> why K-selected<br />

life-history characteristics and highly efficient physiologies are an evolutionary advantage to<br />

obligate cave species.<br />

Introduction<br />

<strong>The</strong> movement <strong>of</strong> resources among ecosystems, commonly referred to as resource<br />

subsidies (sensu Polis & Strong, 1996), is an ever-present feature within the ecological<br />

landscape. <strong>The</strong> roles that such subsidies play in structuring recipient ecosystems can vary widely<br />

as a function <strong>of</strong> resource quantity, quality (e.g., C:N:P ratios), and type (e.g., nutrient vs. organic<br />

matter vs. prey; see Cebrian, 1999; Moore et al., 2004; Marczak et al., 2007). Cave ecosystems,<br />

however, appear to represent an extreme endpoint along the resource subsidy spectrum because<br />

the lack <strong>of</strong> light prevents primary production, which causes nearly all caves (except those<br />

systems based on chemolithoautotrophy; see Engel et al., 2004) to be reliant on inputs <strong>of</strong> organic<br />

matter from surface environments to support biological productivity (Poulson & Lavoie, 2001;<br />

Simon et al. 2007). However, reduced surface connectivity typically limits allochthonous inputs,<br />

which has caused caves to be characterized as energy-limited ecosystems (Hüppop, 2001 &<br />

2005).<br />

One line <strong>of</strong> evidence that indicates cave ecosystems are energy-limited environments is<br />

the remarkable set <strong>of</strong> traits that are shared among many obligate cave species, such as lower<br />

metabolic rates, increased starvation resistance, and more K-selected life-history characteristics<br />

(e.g. long life span, slow growth rate, and reduced fecundity; see Hüppop, 2001 & 2005;<br />

Venarsky et al., 2012b). Energy limitation also appears to be an important factor structuring<br />

entire cave communities, as several studies have reported community shifts or increased biomass<br />

101

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