Reproducing reproduction: How to simulate mast seeding in forest models
Masting is the highly variable and synchronous production of seeds by plants. Masting can have cascading effects on plant population dynamics and forest properties such as tree growth, carbon stocks, regeneration, nutrient cycling, or future species composition. However, masting has often been missing from forest models. Those few that simulate masting have done so using relatively simple empirical rules, and lack an implementation of process-based mechanisms that control such events. Here we review more than 200 published papers on mechanistic formulations of masting, and summarize how the main processes involved in masting and their related patterns can be incorporated in forest models at different degrees of complexity. Our review showed that, of all proximate causes of masting, resource acquisition, storage and allocation were the processes studied most often. Hormonal and genetic regulation of bud formation, floral induction, and anthesis were less frequently addressed. We outline the building blocks of a general process-based model of masting that can be used to improve the oversimplified functions in different types of forest models, and to implement them where missing. A complete implementation of masting in forest models should include functions for resource allocation and depletion, and for pollination, as regulated by both forest structure and weather in the years prior to seed production. When models operate at spatio-temporal scales mismatched with the main masting processes, or if calibration data are not available, simulation can be based on parameterizing masting patterns (variability, synchrony, or frequency). Also, observed masting patterns have the potential to be used as “reality checks” for more processbased forest models wishing to accurately reproduce masting as an emergent phenomenon.