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      Ecological release from interspecific competition leads to decoupled changes in population and individual niche width

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          Abstract

          A species's niche width reflects a balance between the diversifying effects of intraspecific competition and the constraining effects of interspecific competition. This balance shifts when a species from a competitive environment invades a depauperate habitat where interspecific competition is reduced. The resulting ecological release permits population niche expansion, via increased individual niche widths and/or increased among-individual variation. We report an experimental test of the theory of ecological release in three-spine stickleback ( Gasterosteus aculeatus). We factorially manipulated the presence or absence of two interspecific competitors: juvenile cut-throat trout ( Oncorhynchus clarki) and prickly sculpin ( Cottus asper). Consistent with the classic niche variation hypothesis, release from trout competition increased stickleback population niche width via increased among-individual variation, while individual niche widths remained unchanged. In contrast, release from sculpin competition had no effect on population niche width, because increased individual niche widths were offset by decreased between-individual variation. Our results confirm that ecological release from interspecific competition can lead to increases in niche width, and that these changes can occur on behavioural time scales. Importantly, we find that changes in population niche width are decoupled from changes in the niche widths of individuals within the population.

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          The ecology of individuals: incidence and implications of individual specialization.

          Most empirical and theoretical studies of resource use and population dynamics treat conspecific individuals as ecologically equivalent. This simplification is only justified if interindividual niche variation is rare, weak, or has a trivial effect on ecological processes. This article reviews the incidence, degree, causes, and implications of individual-level niche variation to challenge these simplifications. Evidence for individual specialization is available for 93 species distributed across a broad range of taxonomic groups. Although few studies have quantified the degree to which individuals are specialized relative to their population, between-individual variation can sometimes comprise the majority of the population's niche width. The degree of individual specialization varies widely among species and among populations, reflecting a diverse array of physiological, behavioral, and ecological mechanisms that can generate intrapopulation variation. Finally, individual specialization has potentially important ecological, evolutionary, and conservation implications. Theory suggests that niche variation facilitates frequency-dependent interactions that can profoundly affect the population's stability, the amount of intraspecific competition, fitness-function shapes, and the population's capacity to diversify and speciate rapidly. Our collection of case studies suggests that individual specialization is a widespread but underappreciated phenomenon that poses many important but unanswered questions.
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            Foraging adaptation and the relationship between food-web complexity and stability.

            Ecological theory suggests that complex food webs should not persist because of their inherent instability. "Real" ecosystems often support a large number of interacting species. A mathematical model shows that fluctuating short-term selection on trophic links, arising from a consumer's adaptive food choice, is a key to the long-term stability of complex communities. Without adaptive foragers, food-web complexity destabilizes community composition; whereas in their presence, complexity may enhance community persistence through facilitation of dynamical food-web reconstruction that buffers environmental fluctuations. The model predicts a linkage pattern consistent with field observations.
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              Can intraspecific competition drive disruptive selection? An experimental test in natural populations of sticklebacks.

              Theory suggests that frequency-dependent resource competition will disproportionately impact the most common phenotypes in a population. The resulting disruptive selection forms the driving force behind evolutionary models of niche diversification, character release, ecological sexual dimorphism, resource polymorphism, and sympatric speciation. However, there is little empirical support for the idea that intraspecific competition generates disruptive selection. This paper presents a test of this theory, using natural populations of the three-spine stickleback, Gasterosteus aculeatus. Sticklebacks exhibit substantial individual specialization associated with phenotypic variation and so are likely to experience frequency-dependent competition and hence disruptive selection. Using body size and relative gonad mass as indirect measures of potential fecundity and hence fitness, I show that an important aspect of trophic morphology, gill raker length, is subject to disruptive selection in one of two natural lake populations. To test whether this apparent disruptive selection could have been caused by competition, I manipulated population densities in pairs of large enclosures in each of five lakes. In each lake I removed fish from one enclosure and added them to the other to create paired low- and high-population-density treatments with natural phenotype distributions. Again using indirect measures of fitness, disruptive selection was consistently stronger in high-density than low-density enclosures. These results support long-standing theoretical arguments that intraspecific competition drives disruptive selection and thus may be an important causal agent in the evolution of ecological variation.
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                Author and article information

                Journal
                Proc Biol Sci
                RSPB
                royprsb
                Proceedings of the Royal Society B: Biological Sciences
                The Royal Society
                0962-8452
                1471-2954
                22 June 2010
                17 February 2010
                17 February 2010
                : 277
                : 1689
                : 1789-1797
                Affiliations
                [1 ]simpleSection of Integrative Biology, University of Texas at Austin , Austin, TX 78712, USA
                [2 ]Department of Zoology and Biodiversity Research Center, simpleUniversity of British Columbia , 6270 University Boulevard, Vancouver, British Columbia, CanadaV6T 1Z4
                Author notes
                [* ]Author for correspondence ( danbolnick@ 123456mail.utexas.edu ).
                [†]

                Present address: Department of Plant Biology, North Carolina State University, Raleigh, NC 27695, USA.

                [‡]

                Present address: Department of Biology, University of North Carolina, Chapel Hill, NC 27599, USA.

                Article
                rspb20100018
                10.1098/rspb.2010.0018
                2871882
                20164100
                91b3272e-003c-4469-9b68-e4de80a4571d
                © 2010 The Royal Society

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

                History
                : 6 January 2010
                : 27 January 2010
                Categories
                1001
                14
                60
                Research articles

                Life sciences
                three-spine stickleback,gasterosteus aculeatus,individual specialization,interspecific competition,niche variation hypothesis,ecological release

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