The long history of reciprocal transplant studies testing the hypothesis of local adaptation has shown that populations are often adapted to their local environments. Yet many studies have not demonstrated local adaptation, suggesting that sometimes native populations are no better adapted than are genotypes from foreign environments. Local adaptation may also lead to trade-offs, in which adaptation to one environment comes at a cost of adaptation to another environment. I conducted a survey of published studies of local adaptation to quantify its frequency and magnitude and the costs associated with local adaptation. I also quantified the relationship between local adaptation and environmental differences and the relationship between local adaptation and phenotypic divergence. The overall frequency of local adaptation was 0.71, and the magnitude of the native population advantage in relative fitness was 45%. Divergence between home site environments was positively associated with the magnitude of local adaptation, but phenotypic divergence was not. I found a small negative correlation between a population's relative fitness in its native environment and its fitness in a foreign environment, indicating weak trade-offs associated with local adaptation. These results suggest that populations are often locally adapted but stochastic processes such as genetic drift may limit the efficacy of divergent selection.
The fundamental equation in evolutionary quantitative genetics, the Lande equation, describes the response to directional selection as a product of the additive genetic variance and the selection gradient of trait value on relative fitness. Comparisons of both genetic variances and selection gradients across traits or populations require standardization, as both are scale dependent. The Lande equation can be standardized in two ways. Standardizing by the variance of the selected trait yields the response in units of standard deviation as the product of the heritability and the variance-standardized selection gradient. This standardization conflates selection and variation because the phenotypic variance is a function of the genetic variance. Alternatively, one can standardize the Lande equation using the trait mean, yielding the proportional response to selection as the product of the squared coefficient of additive genetic variance and the mean-standardized selection gradient. Mean-standardized selection gradients are particularly useful for summarizing the strength of selection because the mean-standardized gradient for fitness itself is one, a convenient benchmark for strong selection. We review published estimates of directional selection in natural populations using mean-standardized selection gradients. Only 38 published studies provided all the necessary information for calculation of mean-standardized gradients. The median absolute value of multivariate mean-standardized gradients shows that selection is on average 54% as strong as selection on fitness. Correcting for the upward bias introduced by taking absolute values lowers the median to 31%, still very strong selection. Such large estimates clearly cannot be representative of selection on all traits. Some possible sources of overestimation of the strength of selection include confounding environmental and genotypic effects on fitness, the use of fitness components as proxies for fitness, and biases in publication or choice of traits to study.
Climate change has the potential to affect the ecology and evolution of every species on Earth. Although the ecological consequences of climate change are increasingly well documented, the effects of climate on the key evolutionary process driving adaptationnatural selection-are largely unknown. We report that aspects of precipitation and potential evapotranspiration, along with the North Atlantic Oscillation, predicted variation in selection across plant and animal populations throughout many terrestrial biomes, whereas temperature explained little variation. By showing that selection was influenced by climate variation, our results indicate that climate change may cause widespread alterations in selection regimes, potentially shifting evolutionary trajectories at a global scale.C limate affects organisms in ways that ultimately shape patterns of biodiversity (1). Consequently, the rapid changes in Earth's recent climate impose challenges for many organisms, often reducing population fitness (2-4). Although some species may migrate and undergo range shifts to avoid climate-induced declines and potential extinction (5), an alternative outcome is adaptive evolution in response to selection imposed by climate (6). However, we lack a general understanding of whether local and global climatic factors such as temperature, precipitation, and water availability influence selection (2, 7). Understanding these effects is critical for predicting the consequences of increasing droughts, heat waves, and extreme precipitation events that are expected in many regions (8, 9).To quantify how climate variation influences selection, we assembled a large database of standardized directional selection gradients and differentials from spatially [mean = 4.6 ± 5.4 (SD) populations, range = 2 to 59 populations] and temporally [mean = 5.2 ± 6.8 (SD) years, range = 2 to 45 years] replicated selection studies (N = 168) in plant and animal populations (Table 1 and database S1). We focused on directional selection that can generate increases or decreases in trait values because it is well characterized and is likely to drive rapid evolution (10) in response to variation in climatic factors. However, selection acting on trait combinations and trait variance may also be affected by climate (7). Selection gradients estimate the strength and direction of selection acting directly on a trait, whereas differentials estimate "total selection" on a trait via both direct and indirect selection because of trait correlations (11). These standardized selection coefficients describe selection in terms of the relationship between relative fitness and quantitative traits measured in standard deviations, thus facilitating cross-study comparisons (11,12).Geographically, the database contains many estimates of selection from temperate, mid-latitude regions centered at 40°N (Fig. 1A). The populations in this database span many terrestrial biomes on Earth, with the exception of tundra and tropical rainforests where selection has rarely been quantified (Fig. 1B...
A pervasive hypothesis at the interface of ecology and evolution is that biotic interactions contribute to regional biodiversity by accelerating adaptation and speciation. We investigated this question in the context of closely related, bumble bee‐pollinated plants (Pedicularis spp.) in the Hengduan Mountains of south‐central China, where they exhibit spectacular levels of richness, endemism, and floral diversity. Because these species co‐occur frequently, flower synchronously, and share pollinators during the brief reproductive season, we predict that pollinator‐mediated interactions may influence their community assembly and evolutionary diversification. If disparity in floral traits reduces competitive interactions between species, as would happen if floral isolation mitigates reproductive interference caused by heterospecific pollen flow, then species with dissimilar flowers should co‐occur more often, yielding greater floral diversity at local scales than expected by chance. Moreover, if such interactions have repeatedly driven character displacement, then floral traits should exhibit homoplasy, the phylogenetic signature of labile evolution. We present evidence supporting these predictions, and find that local species richness is best explained by a model including both floral diversity and phylogenetic distance. Our results suggest that a dynamic mosaic of pollinator‐mediated interactions among Pedicularis in the Hengduan region promotes ecological sorting through recurrent selection against reproductive interference, causing rapid species turnover at local scales, and accelerating the rate of floral divergence among species. Together these processes may have contributed to the remarkable accumulation of florally diverse species of Pedicularis endemic to the Hengduan Mountains biodiversity hotspot.
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