Functional diversity affects tree vigor, growth, and mortality in mixed-conifer/hardwood forests in California, U.S.A, in the absence of fire

Abstract

Mixed tree-species forest management can increase forest resilience by reducing the impacts of disturbances that disproportionately affect a single tree species or closely related groups of tree species. Beyond disturbance-risk reduction, tree-species diversification may foster functional-diversity effects (e.g., complementarity or facilitation) that alter the performance of a given tree species in mixed versus pure stands, potentially benefitting carbon sequestration and wildlife habitat. Tree species-mixture effects have been explored to only a limited degree in western US forests and, particularly, in California. Establishing whether vigor, growth, and mortality of common tree species vary with stand composition would help inform restoration and modeling of these forests under climate change. Using data from USDA Forest Service Forest Inventory and Analysis (FIA) plots from California, we examined how individual-tree vigor, as indicated by live crown ratio (LCR), periodic basal area increment (BAI), and mortality odds varied with functional dissimilarity (FDis). We quantified FDis using an index based on 11 traits related to resource acquisition, competition, environmental tolerances, and fire ecology. We classified major tree species into ponderosa pine (Pinus ponderosa), Jeffrey pine (Pinus jeffreyi), incense-cedar (Calocedrus decurrens), true firs (Abies spp.), Douglas-fir (Pseudotsuga menziesii), live oaks (Quercus spp.), and deciduous oaks (Quercus spp.) response groups. We tested for the main effects of FDis on tree responses, as well as for interactions with tree, site, stand, and climate factors. We found that initial tree height modulated the effects of FDis on ponderosa pine, Jeffrey pine, incense-cedar, and true fir LCR, whereas FDis interacted with climate to alter live oak and deciduous oak LCR. FDis decreased BAI in ponderosa pine and increased BAI in live oaks. FDis interacted with tree size to influence BAI for Jeffrey pine, Douglas-fir, and true firs. We found no evidence that climate or site quality modulated FDis effects on BAI for any species group. Tree mortality was not responsive to FDis, except for the true firs, where both initial tree height and competition interacted with FDis to increase and decrease mortality odds, respectively. FDis effects commonly shifted from positive to negative along gradients of stand structure and site quality, indicating that these effects vary with site and stand conditions. Our results have implications for balancing the ecosystem benefits of mixed stands, such as disturbance risk, carbon sequestration, and habitat during forest restoration projects in the region, as well as for more accurate modeling of complex stands.