Crowding, climate, and the case for social distancing among trees.

In an emerging era of megadisturbance, bolstering forest resilience to wildfire, insects, and drought has become a central objective in many western forests. Climate has received considerable attention as a driver of these disturbances, but few studies have examined the complexities of climate-vegetation-disturbance interactions. Current strategies for creating resilient forests often rely on retrospective approaches, seeking to impart resilience by restoring historical conditions to contemporary landscapes, but historical conditions are becoming increasingly unattainable amidst modern bioclimatic conditions. What becomes an appropriate benchmark for resilience when we have novel forests, rapidly changing climate, and unprecedented disturbance regimes? We combined two longitudinal datasets-each representing some of the most comprehensive spatially explicit, annual tree mortality data in existence-in a post-hoc factorial design to examine the nonlinear relationships between fire, climate, forest spatial structure, and bark beetles. We found that while prefire drought elevated mortality risk, advantageous local neighborhoods could offset these effects. Surprisingly, mortality risk (Pm ) was higher in crowded local neighborhoods that burned in wet years (Pm  = 42%) compared with sparse neighborhoods that burned during drought (Pm  = 30%). Risk of beetle attack was also increased by drought, but lower conspecific crowding impeded the otherwise positive interaction between fire and beetle attack. Antecedent fire increased drought-related mortality over short timespans (<7 years) but reduced mortality over longer intervals. These results clarify interacting disturbance dynamics and provide a mechanistic underpinning for forest restoration strategies. Importantly, they demonstrate the potential for managed fire and silvicultural strategies to offset climate effects and bolster resilience to fire, beetles, and drought.

Spatially nonrandom tree mortality and ingrowth maintain equilibrium pattern in an old-growth Pseudotsuga-Tsuga forest.

Mortality processes in old-growth forests are generally assumed to be driven by gap-scale disturbance, with only a limited role ascribed to density-dependent mortality, but these assumptions are rarely tested with data sets incorporating repeated measurements. Using a 12-ha spatially explicit plot censused 13 years apart in an approximately 500-year-old Pseudotsuga-Tsuga forest, we demonstrate significant density-dependent mortality and spatially aggregated tree recruitment. However, the combined effect of these strongly nonrandom demographic processes was to maintain tree patterns in a state of dynamic equilibrium. Density-dependent mortality was most pronounced for the dominant late-successional species, Tsuga heterophylla. The long-lived, early-seral Pseudotsuga menziesii experienced an annual stem mortality rate of 0.84% and no new recruitment. Late-seral species Tsuga and Abies amabilis had nearly balanced demographic rates of ingrowth and mortality. The 2.34% mortality rate for Taxus brevifolia was higher than expected, notably less than ingrowth, and strongly affected by proximity to Tsuga. Large-diameter Tsuga structured both the regenerating conspecific and heterospecific cohorts with recruitment of Tsuga and Abies unlikely in neighborhoods crowded with large-diameter competitors (P < 0.001). Density-dependent competitive interactions strongly shape forest communities even five centuries after stand initiation, underscoring the dynamic nature of even equilibrial old-growth forests.

Response to Comment on "Plant diversity increases with the strength of negative density dependence at the global scale".

Hülsmann and Hartig suggest that ecological mechanisms other than specialized natural enemies or intraspecific competition contribute to our estimates of conspecific negative density dependence (CNDD). To address their concern, we show that our results are not the result of a methodological artifact and present a null-model analysis that demonstrates that our original findings-(i) stronger CNDD at tropical relative to temperate latitudes and (ii) a latitudinal shift in the relationship between CNDD and species abundance-persist even after controlling for other processes that might influence spatial relationships between adults and recruits.

Response to Comment on "Plant diversity increases with the strength of negative density dependence at the global scale".

Chisholm and Fung claim that our method of estimating conspecific negative density dependence (CNDD) in recruitment is systematically biased, and present an alternative method that shows no latitudinal pattern in CNDD. We demonstrate that their approach produces strongly biased estimates of CNDD, explaining why they do not detect a latitudinal pattern. We also address their methodological concerns using an alternative distance-weighted approach, which supports our original findings of a latitudinal gradient in CNDD and a latitudinal shift in the relationship between CNDD and species abundance.

Plant diversity increases with the strength of negative density dependence at the global scale.

Theory predicts that higher biodiversity in the tropics is maintained by specialized interactions among plants and their natural enemies that result in conspecific negative density dependence (CNDD). By using more than 3000 species and nearly 2.4 million trees across 24 forest plots worldwide, we show that global patterns in tree species diversity reflect not only stronger CNDD at tropical versus temperate latitudes but also a latitudinal shift in the relationship between CNDD and species abundance. CNDD was stronger for rare species at tropical versus temperate latitudes, potentially causing the persistence of greater numbers of rare species in the tropics. Our study reveals fundamental differences in the nature of local-scale biotic interactions that contribute to the maintenance of species diversity across temperate and tropical communities.