Abstract
We tested the hypothesis that brook trout growth rates are controlled by a complex interaction of food availability, water temperature, and competitor density. We quantified trout diet, growth, and consumption in small headwater tributaries characterized as cold with low food and high trout density, larger tributaries characterized as cold with moderate food and moderate trout density, and large main stems characterized as warm with high food and low trout density. Brook trout consumption was highest in the main stem where diets shifted from insects in headwaters to fishes and crayfish in larger streams. Despite high water temperatures, trout growth rates also were consistently highest in the main stem, likely due to competitively dominant trout monopolizing thermal refugia. Temporal changes in trout density had a direct negative effect on brook trout growth rates. Our results suggest that competition for food constrains brook trout growth in small streams, but access to thermal refugia in productive main stem habitats enables dominant trout to supplement growth at a watershed scale. Brook trout conservation in this region should seek to relieve the “temperature–productivity squeeze,” whereby brook trout productivity is constrained by access to habitats that provide both suitable water temperature and sufficient prey.






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Amundsen, P. A. & H. M. Gabler, 2008. Food consumption and growth of Atlantic salmon Salmo salar parr in sub-Arctic rivers: empirical support for food limitation and competition. Journal of Fish Biology 73: 250–261.
Anderson, R. O. & R. M. Neumann, 1996. Length, weight, and associated structural indices. In Murphy, B. R. & D. W. Willis (eds), Fisheries Techniques, 2nd ed. American Fisheries Society, Bethesda, MD: 447–482.
Bates, B. C., Z. Q. Kundzewicz, S. Wu & J. P. Palutikof (eds), 2008. Climate Change and Water. Technical Paper of the Intergovernmental Panel on Climate Change. IPCC Secretariat, Geneva.
Benke, A. C., A. D. Huryn, L. A. Smock & J. B. Wallace, 1999. Length–mass relationships for freshwater macroinvertebrates in North America with particular reference to the southeastern United States. Journal of the North American Benthological Society 18: 308–343.
Bopp, J., 2002. The Combined Effects of Water Chemistry, Canopy Cover, and Basin Area on Benthic Macroinvertebrates along a Central Appalachian Stream Continuum. West Virginia University, Morgantown.
Borror, D. J., C. A. Triplehorn & N. F. Johnson, 1989. An Introduction to the Study of Insects, 6th ed. Saunders College Publishers, Ft. Worth, TX.
Breau, C., R. A. Cunjak & G. Bremset, 2007. Age-specific aggregation of wild juvenile Atlantic salmon Salmo salar at cool water sources during high temperature events. Journal of Fish Biology 71: 1179–1191.
Carlson, S. M., A. P. Hendry & B. H. Letcher, 2007. Growth rate differences between resident native brook trout and non-native brown trout. Journal of Fish Biology 71: 1430–1447.
Chaloner, D. T., M. S. Wipfli & J. P. Caouette, 2002. Mass loss and macroinvertebrate colonisation of Pacific salmon carcasses in southeastern Alaskan streams. Freshwater Biology 47: 263–273.
Cummins, K. W. & J. C. Wuycheck, 1971. Caloric equivalents for investigations in ecological energetics. International Association of Theoretical and Applied Limnology 18: 1–158.
Daufresne, M. & P. Boet, 2007. Climate change impacts on structure and diversity of fish communities in rivers. Global Change Biology 13: 2467–2478.
Dunham, J. B. & G. L. Vinyard, 1997. Relationships between body mass, population density, and the self-thinning rule in stream-living salmonids. Canadian Journal of Fisheries and Aquatic Sciences 54: 1025–1030.
Ensign, W. E., R. J. Strange & S. E. Moore, 1990. Summer food limitation reduced brook and rainbow trout biomass in a southern Appalachian stream. Transactions of the American Fisheries Society 119: 894–901.
Fausch, K. D., C. E. Torgersen, C. V. Baxter & H. W. Li, 2002. Landscape to riverscapes: bridging the gap between research and conservation of stream fishes. Bioscience 52: 483–498.
Flebbe, P. A., L. D. Roghair & J. L. Bruggink, 2006. Spatial modeling to project Southern Appalachian trout distribution in a warmer climate. Transactions of the American Fisheries Society 135: 1371–1382.
Fretwell, S. D. & H. L. Lucas Jr, 1970. On territorial behavior and other factors influencing habitat distribution in birds. 1. Theoretical development. Acta Biotheoretica 1970: 16–36.
Freund, J. G. & J. T. Petty, 2007. Response of fish and macroinvertebrate bioassessment indices to water chemistry in a mined Appalachian watershed. Environmental Management 39: 707–720.
Goniea, T. M., M. L. Keefer, T. C. Bjornn, C. A. Peery, D. H. Bennet & L. C. Stuehrenberg, 2006. Behavioral thermoregulations and slowed migration by adult fall Chinook salmon in response to high Columbia River water temperatures. Transactions of the American Fisheries Society 135: 408–419.
Gowan, C. & K. D. Fausch, 2002. Why do foraging stream salmonids move during summer? Environmental Biology of Fishes 64: 139–153.
Gross, M. R., R. M. Coleman & R. M. McDowall, 1988. Aquatic productivity and the evolution of diadromous fish migration. Science 239: 1291–1293.
Grossman, G. D., R. E. Ratajczak, C. M. Wagner & J. T. Petty, 2010. Dynamics and regulation of the southern brook trout (Salvelinus fontinalis) population in an Appalachian stream. Freshwater Biology 55: 1494–1508.
Grossman, G. D., A. Nuhfer, T. Zorn, G. Sundin & G. Alexander, 2012. Population regulation of brook trout (Salvelinus fontinalis) in Hunt Creek, Michigan: a 50-year study. Freshwater Biology 57: 1434–1448.
Haak, A. L., J. E. Williams, H. M. Neville, D. C. Dauwalter & W. T. Colyer, 2010. Conserving peripheral trout populations: the values and risks of life on the edge. Fisheries 35: 530–549.
Hartman, K. J. & K. M. Cox, 2008. Refinement and testing of a brook trout bioenergetics model. Transactions of the American Fisheries Society 137: 357–363.
Hartman, K. J. & J. A. Sweka, 2001. Development of a bioenergetics model for Appalachian brook trout. Proceedings of the Southeastern Association of Fish and Wildlife Agencies 55: 38–51.
Hense, Z., R. W. Martin & J. T. Petty, 2010. Electrofishing capture efficiencies for common stream fish species to support watershed-scale studies in the central Appalachians. North American Journal of Fisheries Management 30: 1041–1050.
Hilderbrand, R. H. & J. L. Kershner, 2004. Are there differences in growth and condition between mobile and resident cutthroat trout? Transactions of the American Fisheries Society 133: 1042–1046.
Hill, D. K. & J. J. Magnuson, 1990. Potential effects of global climate warming on the growth and prey consumption of great lakes fish. Transactions of the American Fisheries Society 119: 265–275.
Hill, J. & G. D. Grossman, 1993. An energetic model of microhabitat use for rainbow trout and rosyside dace. Ecology 74: 685–698.
Hudy, M., T. M. Theiling, N. Gillespie & E. P. Smith, 2008. Distribution, status, and land use characteristics of subwatersheds within the native range of brook trout in the eastern United States. North American Journal of Fisheries Management 28: 1069–1085.
Hughes, N. F., 1998. A model of habitat selection by drift-feeding stream salmonids at different scales. Ecology 79: 281–294.
Hughes, N. F. & L. M. Dill, 1990. Position choice by drift-feeding salmonids: model and test for arctic grayling (Thymallus arcticus) in subarctic mountain streams, interior Alaska. Canadian Journal of Fisheries and Aquatic Sciences 47: 2039–2047.
Isaak, D. J., C. H. Luce, B. E. Rieman, D. E. Nagel, E. E. Peterson, D. L. Horan, S. Parkes & G. L. Chandler, 2010. Effect of climate change and wildfire on stream temperatures and salmonid thermal habitat in a mountain river network. Ecological Applications 20: 1350–1371.
Jensen, A. J., 1990. Growth of young migratory brown trout Salmo trutta correlated with water temperature in Norwegian rivers. Journal of Animal Ecology 59: 603–614.
Keefer, M. L., C. A. Peery & B. High, 2009. Behavioral thermoregulation and associated mortality trade-offs in migrating adult steelhead (Oncorhynchus mykiss): variability among sympatric populations. Canadian Journal of Fisheries and Aquatic Sciences 66: 1734–1747.
Letcher, B. H., K. H. Nislow, J. A. Coombs, M. J. O’Donnell & T. L. Dubreuil, 2007. Population response to habitat fragmentation in a stream-dwelling brook trout population. PLoS One 2(11): e1139.
McCafferty, W. P. & A. V. Provonsha, 1998. Aquatic Entomology: The Fisherman’s and Ecologist’s Illustrated Guide to Insects and Their Relatives. Jones and Bartlett, Boston.
McClurg, S. E., J. T. Petty, P. M. Mazik & J. L. Clayton, 2007. Stream ecosystem response to limestone treatment in acid impacted watersheds of the Allegheny Plateau. Ecological Applications 17: 1087–1104.
Merritt, R. W. & K. W. Cummins, 1996. An Introduction to the Aquatic Insects of North America, 3rd ed. Kendall and Hunt, Dubuque, IA.
Morinville, G. R. & J. B. Rasmussen, 2003. Early juvenile bioenergetic differences between anadromous and resident brook trout (Salvelinus fontinalis). Canadian Journal of Fisheries and Aquatic Sciences 60: 401–410.
Petty, J. T. & G. D. Grossman, 2010. Giving-up densities and ideal pre-emptive patch use in a predatory benthic stream fish. Freshwater Biology 55: 780–793.
Petty, J. T. & E. P. Merriam, 2012. Brook trout restoration. Nature Education Knowledge 3:17. http://www.nature.com/scitable/knowledge/library/brook-trout-restoration-83031062.
Petty, J. T. & D. Thorne, 2005. An ecologically based approach to identifying restoration priorities in an acid impacted watershed. Restoration Ecology 13: 348–357.
Petty, J. T., J. Freund, P. Lamothe & P. Mazik, 2001. Quantifying instream habitat in the upper Shavers Fork basin at multiple spatial scales. Proceedings of the Southeastern association of Fish and Wildlife Agencies 55: 81–94.
Petty, J. T., P. J. Lamothe & P. M. Mazik, 2005. Spatial and seasonal dynamics of brook trout populations inhabiting a central Appalachian watershed. Transactions of the American Fisheries Society 134: 572–587.
Petty, J. T., J. L. Hansbarger, B. M. Huntsman & P. M. Mazik, 2012. Brook trout movement in response to temperature, flow, and thermal refugia within a complex Appalachian riverscape. Transactions of the American Fisheries Society 141: 1060–1073.
Poff, N. L. & A. D. Huryn, 1998. Multi-scale determinants of secondary production in Atlantic salmon (Salmo salar) streams. Canadian Journal of Fisheries and Aquatic Sciences 55: 201–217.
Poplar-Jeffers, I. O., J. T. Petty, J. T. Anderson, S. J. Kite, M. P. Strager & R. H. Fortney, 2009. Culvert replacement and stream habitat restoration: Implications from brook trout management in an Appalachian watershed, U.S.A. Restoration Ecology 17: 404–413.
Pullium, R. H. & B. J. Danielson, 1991. Sources, sinks, and habitat selection: a landscape perspective on population dynamics. The American Naturalist 137: S50–S66.
Rieman, B. & D. Isaak, 2010. Climate Change, Aquatic Ecosystems and Fishes in the Rocky Mountain West: Implications and Alternatives for Management. USDA Forest Service, Rocky Mountain Research Station, GTR-RMRS-250, Fort Collins, CO.
Rieman, B. E., D. Isaak, S. Adams, D. Horan, D. Nagel, C. Luce & D. Myers, 2007. Anticipated climate warming effects on bull trout habitats and populations across the interior Columbia River basin. Transactions of the American Fisheries Society 136: 1552–1565.
Ries, R. D. & S. A. Perry, 1995. Potential effects of global climate warming on brook trout growth and prey consumption in central Appalachian streams, USA. Climate Research 5: 197–206.
Robillard, M. M., R. L. McLaughlin & R. W. Mackereth, 2011. Diversity in habitat use and trophic ecology of brook trout in Lake Superior and tributary streams revealed through stable isotopes. Transactions of the American Fisheries Society 140: 943–953.
Sabo, J. L., J. L. Bastow & M. E. Power, 2002. Length-mass relationships for adult aquatic and terrestrial invertebrates in a California watershed. Journal of the North American Benthological Society 21: 336–343.
Sample, B. E., R. J. Cooper, R. D. Greer & R. C. Whitmore, 1993. Estimation of insect biomass by length and width. American Midland Naturalist 129: 234–240.
Smock, L. A., 1980. Relationships between body size and biomass of aquatic insects. Freshwater Biology 10: 375–383.
Sotiropoulos, J. C., K. H. Nislow & M. R. Ross, 2006. Brook trout, Salvelinus fontinalis, microhabitat selection and diet under low summer stream flows. Fisheries Management and Ecology 13: 149–155.
Sweka, J. A., 2003. Aquatic-Terrestrial Linkages in Appalachian Streams: Influence of Riparian Inputs on Stream Habitat, Brook Trout Populations, and Trophic Dynamics. West Virginia University, Morgantown.
Sweka, J. A. & K. J. Hartman, 2008. Contribution of terrestrial invertebrates to yearly brook trout prey consumption and growth. Transactions of the American Fisheries Society 137: 224–235.
Thompson, A. R., J. T. Petty & G. D. Grossman, 2001. Multi-scale effects of resource patchiness on foraging behaviour and habitat use by longnose dace, Rhinichthys cataractae. Freshwater Biology 46: 145–160.
Thorp, J. H. & A. P. Covich, 2001. Ecology and Classification of North American Freshwater Invertebrates, 2nd ed. Academic Press, San Diego, CA.
Torgersen, C. E., D. M. Price, H. W. Li & B. A. McIntosh, 1999. Multiscale thermal refugia and stream habitat associations of Chinook salmon in northeastern Oregon. Ecological Applications 9: 301–319.
Utz, R. M. & K. J. Hartman, 2006. Temporal and spatial variation in the energy intake of a brook trout (Salvelinus fontinalis) population in an Appalachian watershed. Canadian Journal of Fisheries and Aquatic Sciences 63: 2675–2686.
Utz, R. M. & K. J. Hartman, 2007. Identification of critical prey items to Appalachian brook trout (Salvelinus fontinalis) with emphasis on terrestrial organisms. Hydrobiologica 575: 259–270.
Utz, R. M. & K. J. Hartman, 2009. Density-dependent individual growth and size dynamics of central Appalachian brook trout (Salvelinus fontinalis). Canadian Journal of Fisheries and Aquatic Sciences 66: 1072–1080.
Warren, D. R., J. M. Robinson, D. C. Josephson, D. R. Sheldon & C. E. Kraft, 2012. Elevated summer temperatures delay spawning and reduce redd construction for resident brook trout (Salvelinus fontinalis). Global Change Biology 18: 1804–1811.
Webster, J. J. & K. J. Hartman, 2005. The role of terrestrial invertebrates in allopatric brook trout headwater streams in the central Appalachian mountains. Journal of Freshwater Ecology 20: 101–107.
Wenger, S. J., D. J. Isaak, C. H. Luce, H. M. Neville, K. D. Fausch, J. B. Dunham, D. C. Dauwalter, M. K. Young, M. M. Elsner, B. E. Rieman, A. F. Hamlet & J. E. Williams, 2011. Flow regime, temperature, and biotic interactions drive differential declines of trout species under climate change PNAS 108: 14175–14180.
Wipfli, M. S., 1997. Terrestrial invertebrates as salmonid prey and nitrogen sources in streams: contrasting old-growth and young-growth riparian forests in southeastern Alaska, U.S.A. Canadian Journal of Fisheries and Aquatic Sciences 54: 1259–1269.
Wipfli, M. S., J. Hudson & J. Caouette, 2003. Marine subsidies in freshwater ecosystems: Salmon carcasses increase the growth rates of stream-resident salmonids. Transactions of the American Fisheries Society 132: 371–381.
Xu, C. L., B. H. Letcher & K. H. Nislow, 2010. Context-specific influence of water temperature on brook trout growth rates in the field. Freshwater Biology 55: 2253–2264.
Young, R. G., J. Wilkinson, J. Hay & J. W. Hayes, 2010. Movement and mortality of adult brown trout in the Motupiko River, New Zealand: effects of water temperature, flow, and flooding. Transactions of the American Fisheries Society 139: 137–146.
Acknowledgments
We thank the U.S. Geological Survey and the U.S. Fish and Wildlife Service for financial support for this study. Comments from Kyle Hartman greatly improved earlier versions of this manuscript. We also thank Mike Shingleton and Steve Brown from the WVDNR for sharing their expertise and ideas. As always, we are grateful to Becky Nestor for her administrative support of this project. Use of trade names does not imply endorsement by the U.S. Government.
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Petty, J.T., Thorne, D., Huntsman, B.M. et al. The temperature–productivity squeeze: constraints on brook trout growth along an Appalachian river continuum. Hydrobiologia 727, 151–166 (2014). https://doi.org/10.1007/s10750-013-1794-0
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DOI: https://doi.org/10.1007/s10750-013-1794-0

