GENETIC RESPONSES TO CLIMATE IN PINUS CONTORTA: NICHE BREADTH, CLIMATE CHANGE, AND REFORESTATION

Fundamental plant–environment relationships were revealed by analyses of 20-yr height and survival of 118 populations representing two subspecies of Pinus contorta growing in common gardens at 60 environmentally disparate test sites in British Columbia. The approach involved (1) preparing models that described the general climate of British Columbia, (2) developing population-specific response functions driven by predicted climate variables, (3) developing general transfer functions that predict performance from the climatic distances over which populations were transferred, and (4) interpreting the results in terms of niche breadth, effects of climate change on adaptedness of populations, and reforestation in a changing environment. Polynomial regression models used physiographic descriptors to predict seven climate variables from normalized records of 513 weather stations. Values of R2 ranged over 0.80–0.97 for thermal variables and 0.54–0.61 for precipitation variables. Validations with independent data from 45 stations were strong and suggested that the models were generally free of bias within the limits of the original data. Response functions describing the height or survival of each population were developed from quadratic regressions using predicted climate variables for each test site. Mean annual temperature and mean temperature in the coldest month were the most effective variables for predicting population height, while the ratio of summer temperature to summer moisture was the best predictor of survival. Validation of the response functions with independent data from two additional test sites produced values of R2 between actual and predicted values that were as high as 0.93 for height and 0.73 for survival. The results demonstrated that natural populations have different climatic optima but tend to occupy suboptimal environments. Nevertheless, the general transfer functions showed that optimal growth and survival of the species as a whole is associated with the null transfer distance. These seemingly anomalous results suggest that the same processes thought to determine the distribution of species control the distribution of genotypes within species: (1) environmental selection to produce a broad fundamental niche, and (2) density-dependent selection to produce a relatively narrow realized niche within which most populations are relegated to suboptimal environments. Consequently, the steep geographic clines typical of P. contorta seem to be driven more by density-dependent selection than by environmental selection. Asymmetric gene flow from the center of distribution toward the periphery is viewed as a primary regulator that provides the fuel for both environmental and density-dependent selection and thereby indirectly perpetuates suboptimality. The response functions predict that small changes in climate will greatly affect growth and survival of forest tree populations and, therefore, that maintaining contemporary forest productivities during global warming will require a wholesale redistribution of genotypes across the landscape. The response functions also provide the climatic bases to current reforestation guidelines and quantify the adjustments necessary for maintaining adaptedness in planted trees during periods of small (∼1°C) temporal temperature shifts.