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Table S1 also provides covariances and correlations between the three focal traits and fitness.
To meet the strict definition of phenotypic plasticity, a study must demonstrate individual (or genotypic) variation in focal traits (de Jong 1995).
For these analyses, we selected 10 out of the 66 character-pair comparisons that are possible among the 12 focal traits listed in Table 1.
Focal traits were mostly environmental tolerances, which correspond to phenotypic buffering, a plasticity type that maintains a functional phenotype despite external disturbance.
The focal traits were predominantly physiological responses and tolerances, thus 'labile' traits that can be adjusted several times during the life time of an organism.
Because the genetic, environmental and hence phenotypic correlations between the three focal traits were weak, we chose to use single trait models (additional file 1 Table S1).
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Of course, an optimality model, like any evolutionary model, must include background assumptions and constraints that anchor the "local" analysis of the focal trait.
Direct selection on a trait is estimated by its direct effect and effects through intermediate steps on fitness, whereas indirect selection on a focal trait is estimated by its effects via correlations with other traits related to fitness in the model [26].
Note however, that all of these studies have measured the focal trait in a common garden setting.
A fundamental question in evolutionary biology is whether or not genetic variation exists in a focal trait.
If a relationship was found, the implication is that at least some genetic variance is due to neutral or deleterious variants with pleiotropic effects on the focal trait.
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