Estimating forest net primary production under changing climate: adding pests into the equation.

The current approach to modelling pest impacts on forest net primary production (NPP) is to apply a constant modifier. This does not capture the large spatial and temporal variability in pest abundance and activity that can occur, meaning that overestimates or underestimates of pest impacts on forest NPP are likely. Taking a more mechanistic approach that incorporates an understanding of how physiology is influenced by pest attack, enables us to better capture system feedbacks and dynamics, thereby improving the capacity to predict into novel situations such as changing climate, and to account for both changes in pest activity and host responses to the growing environment now and into the future. We reviewed the effects of pests on forest NPP and found a range of responses and physiological mechanisms underlying those responses. Pest outbreaks can clearly be a major perturbation to forest NPP, and it seems likely that the frequency and intensity of pest outbreaks, and the ways in which host species respond to pest damage, will change in the future. We summarized these impacts in the form of a conceptual model at leaf, tree and stand scales, and compared the physiological processes embedded within that framework with the capacity of a representative range of NPP models to capture those processes. We found that some models can encapsulate some of the processes, but no model can comprehensively account for the range of physiological responses to pest attack experienced by trees. This is not surprising, given the paucity of empirical data for most of the world's forests, and that the models were developed primarily for other purposes. We conclude with a list of the key physiological processes and pathways that need to be included in forest growth models in order to adequately capture pest impacts on forest NPP under current and future climate scenarios, the equations that might enable this and the empirical data required to support them.