Competition, species interaction and ageing control tree mortality in boreal forests

Summary 1.  Tree mortality has important influences on forest structure and composition, but the mechanisms that cause tree mortality are not well understood. Asymmetric competition is known to be a dominant cause of plant mortality, but this idea has not received much attention in studies of long‐lived trees. 2.  We hypothesised that while tree mortality is dependent on size relative to neighbours as a result of asymmetric competition, tree mortality of shade‐tolerant species varies little with size because of their physiological and morphological adaptations to shaded environments. Furthermore, we hypothesised that tree mortality is higher in more crowded stands because of higher average resource competition, in conspecific stands because of potential negative intra‐specific interactions, and in older stands because of the physiological limitations and susceptibility to minor disturbances of large trees. 3.  Using data from repeatedly measured permanent sampling plots that covered a wide range of tree sizes, stand developmental stages and stand compositions in boreal forests, we simultaneously tested, by boosted regression tree models, the effects of an individual’s relative size, stand crowding, species interaction and ageing on mortality of Pinus banksiana , Populus tremuloides , Betula papyrifera and Picea mariana . 4.  Mortality increased strongly with decreasing relative size for all study species, and the size‐dependent mortality was stronger for shade‐intolerant than for shade‐tolerant species. With increasing stand basal area, mortality increased for Pinus banksiana , Populus tremuloides and Picea mariana but decreased for Betula papyrifera . Mortality was higher in stands with more conspecific neighbours for Populus tremuloides , Betula papyrifera and Picea mariana, but was slightly lower for Pinus banksiana . Mortality also increased with stand age for all species. Furthermore, the size‐dependent mortality was generally stronger in more crowded stands. 5.   Synthesis . Our findings show that tree mortality over a wide range of tree sizes, stand developmental stages and stand compositions in non‐equilibrium boreal forests was strongly controlled by competition, but species interactions and ageing were also important mechanisms. Furthermore, the relative importance of these mechanisms to tree mortality differed with the shade tolerance of species.