The relative contributions of disease and insects in the decline of a long-lived tree: a stochastic demographic model of whitebark pine (Pinus albicaulis)

Pathogens and insect pests have become increasingly important drivers of tree mortality in forested ecosystems. Unfortunately, understanding the relative contributions of multiple mortality agents to the population decline of trees is difficult, because it requires frequent measures of tree survival, growth, and recruitment, as well as the incidence of mortality agents. We present a population model of whitebark pine (Pinus albicaulis), a high-elevation tree undergoing rapid decline in western North America. The loss of whitebark pine is thought to be primarily due to an invasive pathogen (white pine blister rust; Cronartium ribicola) and a native insect (mountain pine beetle; Dendroctonus ponderosae). We utilized seven plots in Crater Lake National Park (Oregon, USA) where 1220 trees were surveyed for health and the presence of blister rust and beetle activity annually from 2003–2014, except 2008. We constructed size-based projection matrices for nine years and calculated the deterministic growth rate (λ) using an average matrix and the stochastic growth rate (λ_s) by simulation for whitebark pine in our study population. We then assessed the roles of blister rust and beetles by calculating λ and λ_susing matrices in which we removed trees with blister rust and, separately, trees with beetles. We also conducted life-table response experiments (LTRE) to determine which demographic changes contributed most to differences in λ between ambient conditions and the two other scenarios. The model suggests that whitebark pine in our plots are currently declining 1.1% per year (λ = 0.9888, λ_s = 0.9899). Removing blister rust from the models resulted in almost no increase in growth (λ = 0.9916, λ_s = 0.9930), while removing beetles resulted in a larger increase in growth (λ = 1.0028, λ_s = 1.0045). The LTRE demonstrated that reductions in stasis of the three largest size classes due to beetles contributed most to the smaller λ in the ambient condition. Our work demonstrates a method for assessing the relative effects of different mortality agents on declining tree populations, and it shows that the effects of insects and pathogens can be markedly different from one another. In our study, beetle activity significantly reduced tree population growth while a pathogen had minimal effect, thus management actions to stabilize our study population will likely need to include reducing beetle activity.