Abstract
The effects of grazing by stream invertebrates on algal biomass and spatial heterogeneity were tested experimentally in flow-through microcosms with natural substrates (rocks). One experiment tested the effects of fixed densities of three species of grazers (the caddisfly Allomyia sp. and two mayflies, Epeorus deceptivus and Baetis bicaudatus) on periphyton. Baetis was tested with and without chemical cues from fish predators, which reduced grazer foraging activity to levels similar to the less mobile mayfly (Epeorus). Mean algal biomass (chlorophyll a; chl a) was reduced in grazer treatments compared to ungrazed controls, but there were no differences among grazer treatments. Algal heterogeneity (Morisita index) increased with grazer mobility, with the highest heterogeneity occurring in the Baetis-no fish treatment (most mobile grazer) and the lowest in the caddisfly treatment (most sedentary grazer). A second experiment used a three factorial design, and tested whether initial resource distribution (homogeneous vs. heterogeneous), Baetis density (high vs. low) and fish odor (present vs. absent) affected grazer impact on algal resources. Abundances of Baetis and chl a on individual rocks were recorded to explore the mechanisms responsible for the observed distributions of algae. Initial resource heterogeneity was maintained despite being subjected to grazing. Mean chl a was highest in controls, as in experiment I, and effects of Baetis on algal biomass increased with grazer density. There were no fish effects on algal biomass and no effects of grazer density or fish on algal heterogeneity. At the scale of individual rocks Baetis was unselective when food was homogeneously distributed, but chose high-food rocks when it was heterogeneously distributed. Results of these mechanistic experiments showed that Baetis can track resources at the scale of single rocks; and at moderate densities mobile grazers could potentially maintain periphyton distributions observed in natural streams.
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Acknowledgements
We are especially grateful to Wendy Brown, Andrea Encalada, Matt Harper, Bryan Horn, Ben Koch, Brad Taylor and Mark Wallin (“the Benthettes”), for their field and laboratory assistance at the Rocky Mountain Biological Laboratory. Special thanks to Barbara Downes for invaluable statistical advice. Comments by Angus McIntosh, Brad Taylor, Bernhard Statzner and an anonymous reviewer greatly improved the manuscript. Discussion with some members of the Entomology and Evolution Department from Cornell University also provided insight on data interpretation. We also thank LeeAnne Martinez for building the floating raft. Fellowships from the Spanish Education Council (MCYT), Xunta de Galicia and Caixanova supported Maruxa Álvarez during the summers of 2002 and 2003 at RMBL, and NSF Award no. DEB 00-89863 to B. L. P. provided funds for equipment, supplies and field assistants. The authors of this manuscript declare that the experiments conducted throughout this study comply with the current USA laws.
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Álvarez, M., Peckarsky, B.L. How do grazers affect periphyton heterogeneity in streams?. Oecologia 142, 576–587 (2005). https://doi.org/10.1007/s00442-004-1759-0
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DOI: https://doi.org/10.1007/s00442-004-1759-0