Regeneration strategies and forest resilience to changing fire regimes: Insights from a Goldilocks model.

Disturbances are ubiquitous in ecological systems, and species have evolved a range of strategies to resist or rebound following disturbance. Understanding how the presence and complementarity of regeneration traits will affect community responses to disturbance is increasingly urgent as disturbance regimes shift beyond their historical ranges of variability. We define "disturbance niche" as a species' fitness across a range of disturbance sizes and frequencies that can reflect the fundamental or realized niche, that is, whether the species occurs alone or with other species. We developed a model of intermediate complexity (i.e., a Goldilocks model) to infer the disturbance niche. We parameterized the model for subalpine forests in Yellowstone National Park (USA) adapted to infrequent stand-replacing fires and included the three major tree-regeneration strategies: (1) obligate seeders that rely on ex situ seeding into burned areas (non-serotinous lodgepole pine, Pinus contorta var. latifola), (2) obligate seeders that depend on in situ seedbanks (serotinous lodgepole pine, Pinus contorta var. latifola), and (3) species that can resprout from surviving roots following fire (quaking aspen, Populus tremuloides). Our results showed which regeneration strategies increase or decrease in prevalence as fire rotation declines. Non-serotinous pines were extirpated when fire rotation was below 50 years in a monoculture and 100 years in a mixed forest; serotinous pines were extirpated when fire rotation was below 20 years; and aspen was extirpated when fire rotation fell below 6 years. The fundamental and realized disturbance niches pinpointed the key mechanisms limiting regeneration for each strategy, namely, increasing fire size for non-serotinous pine (ex situ seeders), decreasing fire frequency for serotinous pine (in situ seeders), and interspecific competition for aspen (resprouters). In a mixed forest, the three regeneration strategies were complementary and each dominated at different combinations of fire size and frequency. Consequently, diversity of regeneration strategies enhanced forest resilience to declining fire rotations. Despite its simplicity, our Goldilocks model produced realistic dynamics and could be readily adapted to other disturbance-prone ecosystems to explore the generality of these results. The disturbance niche is a key concept for anticipating community resilience to changing disturbance regimes.