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Environment and human land use both shape forest composition. Abiotic conditions sift tree species from a regional pool via functional traits that influence species' suitability to the local environment. In addition, human land use can modify species distributions and change functional diversity of forests. However, it is unclear how environment and land use simultaneously shape functional diversity of tree communities. Land-use legacies are especially prominent in temperate forest landscapes that have been extensively modified by humans in the last few centuries. Across a 900-ha temperate deciduous forest in the northeastern United States, comprising a mosaic of different-aged stands due to past human land use, we used four key functional traits-maximum height, rooting depth, wood density, and seed mass-to examine how multiple environmental and land-use variables influenced species distributions and functional diversity. We sampled ~40,000 trees >8 cm DBH within 485 plots totaling 137 ha. Species within plots were more functionally similar than expected by chance when we estimated functional diversity using all traits together (multi-trait), and to a lesser degree, with each trait separately. Multi-trait functional diversity was most strongly correlated with distance from the perennial stream, elevation, slope, and forest age. Environmental and land-use predictors varied in their correlation with functional diversities of the four individual traits. Landscape-wide change in abundances of individual species also correlated with both environment and land-use variables, but magnitudes of trait-environment interactions were generally stronger than trait interactions with land use. These findings can be applied for restoration and assisted regeneration of human-modified temperate forests by using traits to predict which tree species would establish well in relation to land-use history, topography, and soil conditions., (© 2018 by the Ecological Society of America.)

Patterns of diversity and community composition in forests are controlled by a combination of environmental factors, historical events, and stochastic or neutral mechanisms. Each of these processes has been linked to forest community assembly, but their combined contributions to alpha and beta-diversity in forests has not been well explored. Here we use variance partitioning to analyze approximately 40,000 individual trees of 49 species, collected within 137 ha of sampling area spread across a 900-ha temperate deciduous forest reserve in Pennsylvania to ask (1) To what extent is site-to-site variation in species richness and community composition of a temperate forest explained by measured environmental gradients and by spatial descriptors (used here to estimate dispersal-assembly or unmeasured, spatially structured processes)? (2) How does the incorporation of land-use history information increase the importance attributed to deterministic community assembly? and (3) How do the distributions and abundances of individual species within the community correlate with these factors? Environmental variables (i.e., topography, soils, and distance to stream), spatial descriptors (i.e., spatial eigenvectors derived from Cartesian coordinates), and land-use history variables (i.e., land-use type and intensity, forest age, and distance to road), explained about half of the variation in both species richness and community composition. Spatial descriptors explained the most variation, followed by measured environmental variables and then by land- use history. Individual species revealed variable responses to each of these sets of predictor variables. Several species were associated with stream habitats, and others were strictly delimited across opposing north- and south-facing slopes. Several species were also associated with areas that experienced recent (i.e., <100 years) human land-use impacts. These results indicate that deterministic factors, including environmental and land-use history variables, are important drivers of community response. The large amount of "unexplained" variation seen here (about 50%) is commonly observed in other such studies attempting to explain distribution and abundance patterns of plant communities. Determining whether such large fractions of unaccounted for variation are caused by a lack of sufficient data, or are an indication of stochastic features of forest communities globally, will remain an important challenge for ecologists in the future.