Quantifying foliar trait variation and covariation in sun and shade leaves using leaf spectroscopy in eastern North America

Characterizing foliar trait variation in sun and shade leaves can provide insights into inter- and intra-species resource use strategies and plant response to environmental change. However, datasets with records of multiple foliar traits from the same individual and including shade leaves are sparse, which limits our ability to investigate trait-trait, trait-environment relationships and trait coordination in both sun and shade leaves. We presented a comprehensive dataset of 15 foliar traits from sun and shade leaves sampled with leaf spectroscopy, including 424 individuals of 110 plant species from 19 sites across eastern North America. We investigated trait variation, covariation, scaling relationships with leaf mass, and the effects of environment, canopy position, and taxonomy on trait expression. Generally, sun leaves had higher leaf mass per area, nonstructural carbohydrates and total phenolics, lower mass-based chlorophyll a & thorn; b, carotenoids, phosphorus, and potassium, but exhibited species-specific characteristics. Covariation between sun and shade leaf traits, and trait-environment relationships were overall consistent across species. The main dimensions of foliar trait variation in seed plants were revealed including leaf economics traits, photosynthetic pigments, defense, and structural traits. Taxonomy and canopy position collectively explained most of the foliar trait variation. This study highlights the importance of including intra-individual and intra-specific trait variation to improve our understanding of ecosystem functions. Our findings have implications for efficient field sampling, and trait mapping with remote sensing.