How to improve scaling from traits to ecosystem processes.

Scaling approaches in ecology assume that traits are the main attributes by which organisms influence ecosystem functioning. However, several recent empirical papers have found only weak links between traits and ecosystem functioning, questioning the usefulness of trait-based ecology (TBE). We argue that these studies often suffer from one or more widespread misconceptions. Specifically, these studies often (i) conflict with the conceptual foundations of TBE, (ii) lack theory- or hypothesis-driven selection and use of traits, (iii) tend to ignore intraspecific variation, and (iv) use experimental or study designs that are not well suited to make strong tests of TBE assumptions. Addressing these aspects could significantly improve our ability to scale from traits to ecosystem functioning.

Tree resistance and recovery from drought mediated by multiple abiotic and biotic processes across a large geographic gradient.

Worldwide, increasing severity of droughts threatens to change forest ecosystem functioning and community structure. Understanding how forest resilience is determined by its two underlying components, resistance and recovery, will help elucidate the mechanisms of drought responses and help inform management strategies. However, drought responses are shaped by complex processes across different scales, including species-specific drought strategies, tree size, competition, local environmental conditions, and the intensity of the drought event. Here, we quantified the reduction in tree growth during drought (an inverse measure of drought resistance) and post-drought recovery for three montane conifers (Abies concolor, Pinus jeffreyi, and Pinus lambertiana) in California. We used tree ring analysis to quantify responses to drought events of varying intensity between 1895 and 2018 across a geographic climatic gradient, to examine the roles of tree size (DBH) and competition (tree density) in mediating drought responses. We found that years of more intense drought corresponded with larger growth reductions and recovery rates were lower following drought years where trees suffered larger reductions. We found little variation among species in their growth reductions during drought events, but significant differences among species in their recovery post-drought. Across the geographic gradient, trees in the driest locations were susceptible to large growth reductions, signaling either strong sensitivity to drought intensity or exposure to the most extreme drought conditions. These growth reductions were not always compensated for by higher recovery rates. We also found that larger trees were more susceptible to drought due to a steeper negative relationship between recovery rates and the intensity of growth reduction during the drought. Contrary to expectations, recovery rates following the most detrimental drought years were higher in denser forests. Our results demonstrate the importance of considering how factors at various spatial and temporal scales affect the different components of drought responses.

Frequency-dependent tree growth depends on climate.

Climate and competition interact to affect species' performance, such as growth and survival, and help determine species distributions and coexistence. However, it is unclear how climatic conditions modulate frequency-dependent performance, that is, how performance changes as a species becomes locally rare or common. This is critical because declines in performance as a species becomes more common (negative frequency dependence) is a signature of niche differences among species that stabilize coexistence, whereas positive frequency dependence leads to priority effects and hampers species coexistence. Here, we used dendrochronology and hierarchical models to test whether frequency-dependent growth of sugar pine (Pinus lambertiana) depends on climatic conditions. We found that growth rates were strongly dependent on annual precipitation, but no frequency dependence was evident across all years. However, there was a strong interaction between precipitation and frequency dependence, revealing stabilizing niche differences in dry years but positive frequency dependence in wet years. These differences emerged because of precipitation-driven changes in the direction and strength of both con- and heterospecific competition. Overall, these results show how stabilizing and destabilizing effects can be temporally dynamic for long-lived species and interact with climate variation.

Extensive mismatches between species distributions and performance and their relationship to functional traits.

Mismatches between species distributions and their optimal habitat are predicted by ecological theory and will affect species responses to changing climate. However, empirical tests lack consensus on the prevalence of such mismatches and their underlying mechanisms. Here we present a conceptual framework to quantify the mismatch between optimal conditions for species occurrence and multiple measures of population and individual performance (density, adult growth and survival, and recruitment) and the associated performance reduction, or cost. We quantified these mismatches for 59 tree species in the western US along a soil water balance gradient and found high variability in mismatches among species and among performance measures, often resulting in high costs. We used functional traits to explore how dispersal limitation, migration lags, and competitive exclusion may cause mismatches. Overall, the large variability in mismatches, their costs and the relationship with functional traits highlight the nuanced relationships between species' performance and their distributions.

Testing the roles of vertical transmission and drought stress in the prevalence of heritable fungal endophytes in annual grass populations.

Beneficial inherited symbionts are expected to reach high prevalence in host populations, yet many are observed at intermediate prevalence. Theory predicts that a balance of fitness benefits and efficiency of vertical transmission may interact to stabilize intermediate prevalence. We established populations of grass hosts (Lolium multiflorum) that varied in prevalence of a heritable fungal endophyte (Epichloё occultans), allowing us to infer long-term equilibria by tracking change in prevalence over one generation. We manipulated an environmental stressor (elevated precipitation), which we hypothesized would reduce the fitness benefits of symbiosis, and altered the efficiency of vertical transmission by replacing endophyte-positive seeds with endophyte-free seeds. Endophytes and elevated precipitation both increased host fitness, but symbiont effects were not stronger in the drier treatment, suggesting that benefits of symbiosis were unrelated to drought tolerance. Reduced transmission suppressed the inferred equilibrium prevalence from 42.6% to 11.7%. However, elevated precipitation did not modify prevalence, consistent with the result that it did not modify fitness benefits. Our results demonstrate that failed transmission can influence the prevalence of heritable microbes and that intermediate prevalence can be a stable equilibrium due to forces that allow symbionts to increase (fitness benefits) but prevent them from reaching fixation (failed transmission).