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australis are comparable to other estimates for a modest assemblage of both cave and surface species of crayfish for which credible age estimate exist. Regardless, these shorter longevity and time-to-maturity estimates for O. australis are still relatively great compared with surface species in the same genus, indicating that this species has evolved K-selected life history traits and has a high degree of specialization to cave habitats. Support for the energy-limitation hypothesis in cave ecosystems has historically come from either laboratory-based studies focused on individual physiological characteristics (e.g. metabolic rates) of obligate cave species or field-based population- or community-level studies examining for correlations between resource availability and species biomass or productivity. While each approach has its strengths, neither places its results within the context of energy dynamics of actual cave ecosystems (e.g. resource supply rates vs. consumption and growth). In Chapter Five, the mark-recapture data set for O. australis from Chapter Four was combined with both the trophic basis of production approach (sensu Benke & Wallace, 1980) and estimates of resource supply rates (e.g. organic matter and macroinvertebrate prey) to place the energetic demands of O. australis within the context of cave energy dynamics. Similar to the results of Chapter 3, macroinvertebrate biomass increased with organic matter standing stock among the three cave streams. Both the biomass and secondary production of O. australis were positively related to resource standing stocks. The energy budgets showed no indication of resource deficits. The energetic models, however, indicated that nearly all prey production is necessary to support the populations of O. australis, which suggests that inter- and intra-specific competition for resources within these caves is likely high. Thus, the energetic budgets constructed for O. australis in this study provide the first quantitative explanation of why K-selected life history 135

characteristics, highly efficient physiologies, and enhanced sensory systems for food acquisition are an evolutionary advantage to obligate cave species. Collectively, this dissertation represents the most robust examination to date of how energy availability shapes the structure and function of cave communities. On evolutionary time scales the low quantities of energy inputs appear to have influenced the evolution of K-selected life history characteristics (see Chapters 4 and 5). These adaptations likely allow obligate cave species to respond (e.g. increased population size) to long-term (see Chapter 5) rather than shortterm (see Chapter 3) increases in energy availability. Surface-adapted taxa dominated the biomass of cave communities (see Chapters 2 and 3), suggesting that their effects on cave ecosystem processes may be greater than those of cave-adapted taxa, which have been the traditional focus of cave studies. Lastly, this dissertation has demonstrated that some cave and surface (e.g., forested headwater streams) ecosystems are fundamentally similar and appear to be linked to one another along a detrital subsidy spectrum that includes both high (e.g. surface) and low (e.g. cave) rates of detrital inputs (Chapter 3). References Benke A.C. & Wallace J.B. (1980) Trophic basis of production among net-spinning caddisflies in a southern Appalachian stream. Ecology, 61, 108-118. Cooney T.J. & Simon K.S. (2009) Influence of dissolved organic matter and invertebrates on the function of microbial films in groundwater. Microbial ecology, 58, 599-610. Cooper J. (1975) Ecological and behavorial studies in Shelta Cave, Alabama, with emphasis on decapod crustaceans. Ph. D., University of Kentucky. Culver D.C., Kane T.C. & Fong D.W. (1995) Adaptation and natural selection in caves: the evolution of Gammarus minus, Harvard University Press. Graening G.O. & Brown A.V. (2003) Ecosystem dynamics and pollution effects in an Ozark cave stream. Journal of the American Water Resources Association, 39, 1497-1507. Huntsman B.M., Venarsky M.P. & Benstead J.P. (2011a) Relating carrion breakdown rates to 136

characteristics, highly efficient physiologies, and enhanced sensory systems for food acquisition<br />

are an evolutionary advantage to obligate cave species.<br />

Collectively, this dissertation represents the most robust examination to date <strong>of</strong> how<br />

energy availability shapes the structure and function <strong>of</strong> cave communities. On evolutionary time<br />

scales the low quantities <strong>of</strong> energy inputs appear to have influenced the evolution <strong>of</strong> K-selected<br />

life history characteristics (see Chapters 4 and 5). <strong>The</strong>se adaptations likely allow obligate cave<br />

species to respond (e.g. increased population size) to long-term (see Chapter 5) rather than shortterm<br />

(see Chapter 3) increases in energy availability. Surface-adapted taxa dominated the<br />

biomass <strong>of</strong> cave communities (see Chapters 2 and 3), suggesting that their effects on cave<br />

ecosystem processes may be greater than those <strong>of</strong> cave-adapted taxa, which have been the<br />

traditional focus <strong>of</strong> cave studies. Lastly, this dissertation has demonstrated that some cave and<br />

surface (e.g., forested headwater streams) ecosystems are fundamentally similar and appear to be<br />

linked to one another along a detrital subsidy spectrum that includes both high (e.g. surface) and<br />

low (e.g. cave) rates <strong>of</strong> detrital inputs (Chapter 3).<br />

References<br />

Benke A.C. & Wallace J.B. (1980) Trophic basis <strong>of</strong> production among net-spinning caddisflies<br />

in a southern Appalachian stream. Ecology, 61, 108-118.<br />

Cooney T.J. & Simon K.S. (2009) Influence <strong>of</strong> dissolved organic matter and invertebrates on the<br />

function <strong>of</strong> microbial films in groundwater. Microbial ecology, 58, 599-610.<br />

Cooper J. (1975) Ecological and behavorial studies in Shelta Cave, <strong>Alabama</strong>, with emphasis on<br />

decapod crustaceans. Ph. D., <strong>University</strong> <strong>of</strong> Kentucky.<br />

Culver D.C., Kane T.C. & Fong D.W. (1995) Adaptation and natural selection in caves: the<br />

evolution <strong>of</strong> Gammarus minus, Harvard <strong>University</strong> Press.<br />

Graening G.O. & Brown A.V. (2003) Ecosystem dynamics and pollution effects in an Ozark<br />

cave stream. Journal <strong>of</strong> the American Water Resources Association, 39, 1497-1507.<br />

Huntsman B.M., Venarsky M.P. & Benstead J.P. (2011a) Relating carrion breakdown rates to<br />

136

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