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.

Abstract. Whitebark pine (Pinus albicaulis) forests in the western United States have been adversely affected by an exotic pathogen (Cronartium ribicola, causal agent of white pine blister rust), insect outbreaks (Dendroctonus ponderosae, mountain pine beetle), and drought. We monitored individual trees from 2004 to 2013 and characterized stand-level biophysical conditions through a mountain pine beetle epidemic in the Greater Yellowstone Ecosystem. Specifically, we investigated associations between tree-level variables (duration and location of white pine blister rust infection, presence of mountain pine beetle, tree size, and potential interactions) with observations of individual whitebark pine tree mortality. Climate summaries indicated that cumulative growing degree days in years 2006-2008 likely contributed to a regionwide outbreak of mountain pine beetle prior to the observed peak in whitebark mortality in 2009. We show that larger whitebark pine trees were preferentially attacked and killed by mountain pine beetle and resulted in a regionwide shift to smaller size class trees. In addition, we found evidence that smaller size class trees with white pine blister rust infection experienced higher mortality than larger trees. This latter finding suggests that in the coming decades white pine blister rust may become the most probable cause of whitebark pine mortality. Our findings offered no evidence of an interactive effect of mountain pine beetle and white pine blister rust infection on whitebark pine mortality in the Greater Yellowstone Ecosystem. Interestingly, the probability of mortality was lower for larger trees attacked by mountain pine beetle in stands with higher evapotranspiration. Because evapotranspiration varies with climate and topoedaphic conditions across the region, we discuss the potential to use this improved understanding of biophysical influences on mortality to identify microrefugia that might contribute to successful whitebark pine conservation efforts. Using tree-level observations, the National Park Service-led Greater Yellowstone Interagency Whitebark Pine Long-term Monitoring Program provided important ecological insight on the size-dependent effects of white pine blister rust, mountain pine beetle, and water availability on whitebark pine mortality. This ongoing monitoring campaign will continue to offer observations that advance conservation in the Greater Yellowstone Ecosystem.

The study of plant distribution and abundance is a fundamental pursuit in ecology and conservation biology. Measuring plant abundance by visually assessing percent cover and recording a cover class is a common field method that yields ordinal data. Statistical models for ordinal data exist but entail cumbersome interpretations and sometimes restrictive assumptions. We propose a Bayesian hierarchical framework for analysing cover class data that allows for linking ordinal observations to a latent beta distribution and accounts for zero inflation. Harnessing a latent beta distribution supports interpreting changes in abundance in terms of mean percent cover rather than odds ratios of cumulative cover classes as for cumulative link models. The zero augmentation allows for simultaneous inferences on both occurrence (distribution) and abundance. We show how our model can account for true and false zeros, misclassification of cover classes, multiple species and hierarchical sampling designs, using empirical examples and simulations. Simulated observation errors, when ignored, led to models overestimating abundance and underestimating occurrence. Based on simulations, we found no substantial difference between mean percent cover estimates when analyzing ordinal cover classes versus continuous percent cover as the response. Our empirical datasets displayed high probability of detection (>0.85 on average for all species), likely due to the sampling design used and training of observers. Probability of occurrence was slightly underestimated for bare ground, Artemisia tridentata, Elycap medusae, and Poa secunda using a model that ignored imperfect detection. Estimated mean percent cover was not substantially impacted by ignoring measurement error for five plant species and bare ground. Our modelling framework for cover class data allows for an explicit separation of distribution from abundance and, importantly, allows for interpreting species–environment relationships in terms of variation in mean percent cover as compared to cumulative odds ratios. The beta distribution inherently accommodates heteroscedasticity and skewness, statistical properties that are a consequence of spatially aggregated patterns common to plant survey data. Recording cover classes provides a reliable, efficient way to measure plants and our simulations suggest little loss of information compared to assuming continuous percent cover. We provide JAGS and Stan model code for implementation.

Whitebark pine, a foundation species at tree line in the Western U.S. and Canada, has declined due to native mountain pine beetle epidemics, wildfire, and white pine blister rust. These declines are concerning for the multitude of ecosystem and human benefits provided by this species. An understanding of the climatic correlates associated with spread is needed to successfully manage impacts from forest pathogens. Since 2000 mountain pine beetles have killed 75% of the mature cone-bearing trees in the Greater Yellowstone Ecosystem, and 40.9% of monitored trees have been infected with white pine blister rust. We identified models of white pine blister rust infection which indicated that an August and September interaction between relative humidity and temperature are better predictors of white pine blister rust infection in whitebark pine than location and site characteristics in the Greater Yellowstone Ecosystem. The climate conditions conducive to white pine blister rust occur throughout the ecosystem, but larger trees in relatively warm and humid conditions were more likely to be infected between 2000 and 2018. We mapped the infection probability over the past two decades to identify coarse-scale patterns of climate conditions associated with white pine blister rust infection in whitebark pine.

A Correction to this paper has been published: https://doi.org/10.1038/s41467-021-22025-2