Evolutionary relationships between drought-related traits and climate shape large hydraulic safety margins in western North American oaks.

Quantitative knowledge of xylem physical tolerance limits to dehydration is essential to understanding plant drought tolerance but is lacking in many long-vessel angiosperms. We examine the hypothesis that a fundamental association between sustained xylem water transport and downstream tissue function should select for xylem that avoids embolism in long-vessel trees by quantifying xylem capacity to withstand air entry of western North American oaks (Quercus spp.). Optical visualization showed that 50% of embolism occurs at water potentials below -2.7 MPa in all 19 species, and -6.6 MPa in the most resistant species. By mapping the evolution of xylem vulnerability to embolism onto a fossil-dated phylogeny of the western North American oaks, we found large differences between clades (sections) while closely related species within each clade vary little in their capacity to withstand air entry. Phylogenetic conservatism in xylem physical tolerance, together with a significant correlation between species distributions along rainfall gradients and their dehydration tolerance, suggests that closely related species occupy similar climatic niches and that species' geographic ranges may have shifted along aridity gradients in accordance with their physical tolerance. Such trends, coupled with evolutionary associations between capacity to withstand xylem embolism and other hydraulic-related traits, yield wide margins of safety against embolism in oaks from diverse habitats. Evolved responses of the vascular system to aridity support the embolism avoidance hypothesis and reveal the importance of quantifying plant capacity to withstand xylem embolism for understanding function and biogeography of some of the Northern Hemisphere's most ecologically and economically important plants.

No local adaptation in leaf or stem xylem vulnerability to embolism, but consistent vulnerability segmentation in a North American oak.

Vulnerability to embolism varies between con-generic species distributed along aridity gradients, yet little is known about intraspecific variation and its drivers. Even less is known about intraspecific variation in tissues other than stems, despite results suggesting that roots, stems and leaves can differ in vulnerability. We hypothesized that intraspecific variation in vulnerability in leaves and stems is adaptive and driven by aridity. We quantified leaf and stem vulnerability of Quercus douglasii using the optical technique. To assess contributions of genetic variation and phenotypic plasticity to within-species variation, we quantified the vulnerability of individuals growing in a common garden, but originating from populations along an aridity gradient, as well as individuals from the same wild populations. Intraspecific variation in water potential at which 50% of total embolism in a tissue is observed (P50 ) was explained mostly by differences between individuals (>66% of total variance) and tissues (16%). There was little between-population variation in leaf/stem P50 in the garden, which was not related to site of origin aridity. Unexpectedly, we observed a positive relationship between wild individual stem P50 and aridity. Although there is no local adaptation and only minor phenotypic plasticity in leaf/stem vulnerability in Q. douglasii, high levels of potentially heritable variation within populations or strong environmental selection could contribute to adaptive responses under future climate change.

Effects of topoclimatic complexity on the composition of woody plant communities.

Topography can create substantial environmental variation at fine spatial scales. Shaped by slope, aspect, hill-position and elevation, topoclimate heterogeneity may increase ecological diversity, and act as a spatial buffer for vegetation responding to climate change. Strong links have been observed between climate heterogeneity and species diversity at broader scales, but the importance of topoclimate for woody vegetation across small spatial extents merits closer examination. We established woody vegetation monitoring plots in mixed evergreen-deciduous woodlands that spanned topoclimate gradients of a topographically heterogeneous landscape in northern California. We investigated the association between the structure of adult and regenerating size classes of woody vegetation and multidimensional topoclimate at a fine scale. We found a significant effect of topoclimate on both single-species distributions and community composition. Effects of topoclimate were evident in the regenerating size class for all dominant species (four Quercus spp., Umbellularia californica and Pseudotsuga menziesii) but only in two dominant species (Quercus agrifolia and Quercus garryana) for the adult size class. Adult abundance was correlated with water balance parameters (e.g. climatic water deficit) and recruit abundance was correlated with an interaction between the topoclimate parameters and conspecific adult abundance (likely reflecting local seed dispersal). However, in all cases, the topoclimate signal was weak. The magnitude of environmental variation across our study site may be small relative to the tolerance of long-lived woody species. Dispersal limitations, management practices and patchy disturbance regimes also may interact with topoclimate, weakening its influence on woody vegetation distributions. Our study supports the biological relevance of multidimensional topoclimate for mixed woodland communities, but highlights that this relationship might be mediated by interacting factors at local scales.