Increasing fire severity and warmer, drier postfire conditions are making forests in the western United States (West) vulnerable to ecological transformation. Yet, the relative importance of and interactions between these drivers of forest change remain unresolved, particularly over upcoming decades. Here, we assess how the interactive impacts of changing climate and wildfire activity influenced conifer regeneration after 334 wildfires, using a dataset of postfire conifer regeneration from 10,230 field plots. Our findings highlight declining regeneration capacity across the West over the past four decades for the eight dominant conifer species studied. Postfire regeneration is sensitive to high-severity fire, which limits seed availability, and postfire climate, which influences seedling establishment. In the near-term, projected differences in recruitment probability between low- and high-severity fire scenarios were larger than projected climate change impacts for most species, suggesting that reductions in fire severity, and resultant impacts on seed availability, could partially offset expected climate-driven declines in postfire regeneration. Across 40 to 42% of the study area, we project postfire conifer regeneration to be likely following low-severity but not high-severity fire under future climate scenarios (2031 to 2050). However, increasingly warm, dry climate conditions are projected to eventually outweigh the influence of fire severity and seed availability. The percent of the study area considered unlikely to experience conifer regeneration, regardless of fire severity, increased from 5% in 1981 to 2000 to 26 to 31% by mid-century, highlighting a limited time window over which management actions that reduce fire severity may effectively support postfire conifer regeneration.

Significance: Physiological decline in large, old trees could involve diminished reproductive effort, but evidence is lacking. A global analysis finds that fecundity decline in large trees is pervasive, consistent with physiological decline, thus providing new evidence for tree senescence. For the 20% of species not showing fecundity declines, this lack of evidence was linked not to specific species groups, but rather to lack of large trees in the data. Large trees remain important for their structural attributes, but results highlight the critical role of intermediate trees for their contribution to reproduction.

Indirect climate effects on tree fecundity that come through variation in size and growth (climate-condition interactions) are not currently part of models used to predict future forests. Trends in species abundances predicted from meta-analyses and species distribution models will be misleading if they depend on the conditions of individuals. Here we find from a synthesis of tree species in North America that climate-condition interactions dominate responses through two pathways, i) effects of growth that depend on climate, and ii) effects of climate that depend on tree size. Because tree fecundity first increases and then declines with size, climate change that stimulates growth promotes a shift of small trees to more fecund sizes, but the opposite can be true for large sizes. Change the depresses growth also affects fecundity. We find a biogeographic divide, with these interactions reducing fecundity in the West and increasing it in the East. Continental-scale responses of these forests are thus driven largely by indirect effects, recommending management for climate change that considers multiple demographic rates.

Climate warming is contributing to increases in wildfire activity throughout the western United States, leading to potentially long‐lasting shifts in vegetation. The response of forest ecosystems to wildfire is thus a crucial indicator of future vegetation trajectories, and these responses are contingent upon factors such as seed availability, interannual climate variability, average climate, and other components of the physical environment. To better understand variation in resilience to wildfire across vulnerable dry forests, we surveyed conifer seedling densities in 15 recent (1988–2010) wildfires and characterized temporal variation in seed cone production and seedling establishment. We then predicted postfire seedling densities at a 30‐m resolution within each fire perimeter using downscaled climate data, monthly water balance models, and maps of surviving forest cover. Widespread ponderosa pine (Pinus ponderosa) seed cone production occurred at least twice following each fire surveyed, and pulses of conifer seedling establishment coincided with years of above‐average moisture availability. Ponderosa pine and Douglas‐fir (Pseudotsuga menziesii) seedling densities were higher on more mesic sites and adjacent to surviving trees, though there were also important interspecific differences, likely attributable to drought and shade tolerance. We estimated that postfire seedling densities in 42% (for ponderosa pine) and 69% (for Douglas‐fir) of the total burned area were below the lowest reported historical tree densities in these forests. Spatial models demonstrated that an absence of mature conifers (particularly in the interior of large, high‐severity patches) limited seedling densities in many areas, but 30‐yr average actual evapotranspiration and climatic water deficit limited densities on marginal sites. A better understanding of the limitations to postfire forest recovery will refine models of vegetation dynamics and will help to improve strategies of adaptation to a warming climate and shifting fire activity.

Significance: Suitable habitats for forest trees may be shifting fast with recent climate change. Studies tracking the shift in suitable habitat for forests have been inconclusive, in part because responses in tree fecundity and seedling establishment can diverge. Analysis of both components at a continental scale reveals a poleward migration of northern species that is in progress now. Recruitment and fecundity both contribute to poleward spread in the West, while fecundity limits spread in the East, despite a fecundity hotspot in the Southeast. Fecundity limitation on population spread can confront conservation and management efforts with persistent disequilibrium between forest diversity and rapid climate change.