Long generation times limit species' rapid evolution to changing environments. Trees provide critical global ecosystem services, but are under increasing risk of mortality because of climate changemediated disturbances, such as insect outbreaks. The extent to which disturbance changes the dynamics and strength of selection is unknown, but has important implications on the evolutionary potential of tree populations. Using a 40-y-old Pinus ponderosa genetic experiment, we provide rare evidence of context-dependent fluctuating selection on growth rates over time in a long-lived species. Fast growth was selected at juvenile stages, whereas slow growth was selected at mature stages under strong herbivory caused by a mountain pine beetle (Dendroctonus ponderosae) outbreak. Such opposing forces led to no net evolutionary response over time, thus providing a mechanism for the maintenance of genetic diversity on growth rates. Greater survival to mountain pine beetle attack in slow-growing families reflected, in part, a host-based life-history trade-off. Contrary to expectations, genetic effects on tree survival were greatest at the peak of the outbreak and pointed to complex defense responses. Our results suggest that selection forces in tree populations may be more relevant than previously thought, and have implications for tree population responses to future environments and for tree breeding programs.fluctuating selection | growth-survival trade-offs | selection response | Pinus ponderosa | Dendroctonus ponderosae U nderstanding the dynamics of selection over time is fundamental for understanding life-history evolution (1) and predicting evolutionary change under climate change (2, 3). To date, such understanding is based almost exclusively on data for relatively short-lived species (4, 5), but virtually nonexistent for long-lived organisms, such as trees (ref. 6; but see ref. 7). Trees and forests provide critical ecological and commercial services, including impacts on global carbon cycles, species diversity, water quality, and climate regulation (8). Recent forest mortality (9, 10) highlights the importance of understanding how climate change and climate change-driven disturbances may impact forests (11,12). Trees may live for hundreds of years and experience varying selection pressures associated with fluctuating climate (13), disturbance regimes (14), and biotic interactions (15), all of which may be magnified under climate change. The extent to which these events may change the strength and direction of selection and contribute to the maintenance of genetic diversity and evolutionary potential is unknown. Of special relevance are insect outbreaks, a biotic interaction expected to increase with climate change (16) but unaccounted for in models to predict the evolutionary potential of tree populations (17, 18).Mountain pine beetle (MPB; Dendroctonus ponderosae Hopkins) is a native, irruptive forest insect in western North America that uses numerous Pinus species as hosts. Via pheromone-mediated mass attacks that o...