Demographic processes underpinning post-fire resilience in California closed-cone pine forests: the importance of fire interval, stand structure, and climate

The resilience of serotinous obligate-seeding plants to fire may be compromised if increasing fire frequency curtails time available for canopy seed bank accumulation (i.e., immaturity risk), but how various drivers affect seed availability at the time of fire is poorly understood. Using field data from California closed-cone pine (Pinus attenuata and P. muricata) stands, we assess two critical demographic processes during the inter-fire period-reproductive capacity and mortality. At tree- and stand-levels, we test how these processes are affected by stand age and are mediated by biotic and abiotic factors. We found that stand age was the key driver of reproductive capacity; older stands had a greater proportion of reproductively mature individuals and greater closed cone density. Stand density mediated the effect of age; greater stand density resulted in greater closed cone density and a lower proportion of reproductively mature individuals, but reproductive capacity in low- and high-density stands converged over time. Increased moisture stress reduced the stand-level proportion reproductively mature trees but had no effect on closed cone density. Mortality was strongly associated with density-dependent thinning and increased in stands with high moisture stress. Reproductive capacity began to increase sharply 10 years post-fire and by 20 years immaturity risk was low. However, prior to 20 years, low-density stands with high moisture stress may be more susceptible to immaturity risk. Understanding these relationships is critical to predicting serotinous population persistence under changing climate and disturbance conditions.