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.

Resource competition is thought to play a major role in driving evolutionary diversification. For instance, in ecological character displacement, coexisting species evolve to use different resources, reducing the effects of interspecific competition. It is thought that a similar diversifying effect might occur in response to competition among members of a single species. Individuals may mitigate the effects of intraspecific competition by switching to use alternative resources not used by conspecific competitors. This diversification is the driving force in some models of sympatric speciation, but has not been demonstrated in natural populations. Here, we present experimental evidence confirming that competition drives ecological diversification within natural populations. We manipulated population density of three-spine sticklebacks (Gasterosteus aculeatus) in enclosures in a natural lake. Increased population density led to reduced prey availability, causing individuals to add alternative prey types to their diet. Since phenotypically different individuals added different alternative prey, diet variation among individuals increased relative to low-density control enclosures. Competition also increased the diet-morphology correlations, so that the frequency-dependent interactions were stronger in high competition. These results not only confirm that resource competition promotes niche variation within populations, but also show that this increased diversity can arise via behavioural plasticity alone, without the evolutionary changes commonly assumed by theory.

Many apparently generalized species are in fact composed of individual specialists that use a small subset of the population's resource distribution. Niche variation is usually established by testing the null hypothesis that individuals draw from a common resource distribution. This approach encourages a publication bias in which negative results are rarely reported, and obscures variation in the degree of individual specialization, limiting our ability to carry out comparative studies of the causes or consequences of niche variation. To facilitate studies of the degree of individual specialization, this paper outlines four quantitative indices of intrapopulation variation in resource use. Traditionally, such variation has been measured by partitioning the population's total niche width into within-and between-individual, sex, or phenotype components. We suggest two alternative measures that quantify the mean resource overlap between an individual and its population, and we discuss the advantages and disadvantages of all four measures. The utility of all indices depends on the quality of the empirical data. If resources are measured in a coarse-grained manner, individuals may falsely appear generalized. Alternatively, specialization may be overestimated by cross-sectional sampling schemes where diet variation can reflect a patchy environment. Isotope ratios, parasites, or diet-morphology correlations can complement cross-sectional data to establish temporal consistency of individual specialization.

There is extensive evidence that some species of ecological generalists, which use a wide diversity of resources, are in fact heterogeneous collections of relatively specialized individuals. This within-population variation, or ''individual specialization,'' is a key requirement for frequency-dependent interactions that may drive a variety of types of evolutionary diversification and may influence the population dynamics and ecological interactions of species. Consequently, it is important to understand when individual specialization is likely to be strong or weak. The niche variation hypothesis (NVH) suggests that populations tend to become more generalized when they are released from interspecific competition. This niche expansion was proposed to arise via increased variation among individuals rather than increased individual niche breadth. Consequently, we expect ecological generalists to exhibit stronger individual specialization, but this correlation has been repeatedly rejected by empiricists. The drawback with previous empirical tests of the NVH is that they use morphological variation as a proxy for niche variation, ignoring the role of behavior and complex phenotype-function relationships. Here, we used diet data to directly estimate niche variation among individuals. Consistent with the NVH, we show that more generalized populations also exhibit more niche variation. This trend is quite general, appearing in all five case studies examined: three-spine stickleback, Eurasian perch, Anolis lizards, intertidal gastropods, and a community of neotropical frogs. Our results suggest that generalist populations may tend to be more ecologically variable. Whether this translates into greater genetic variation, evolvability, or ecological stability remains to be determined. frequency dependence ͉ individual specialization ͉ niche expansion