Abstract. Complex systems science provides a transdisciplinary framework to study systems characterized by (1) heterogeneity, (2) hierarchy, (3) self-organization, (4) openness, (5) adaptation, (6) memory, (7) non-linearity, and (8) uncertainty. Complex systems thinking has inspired both theory and applied strategies for improving ecosystem resilience and adaptability, but applications in forest ecology and management are just beginning to emerge. We review the properties of complex systems using four well-studied forest biomes (temperate, boreal, tropical and Mediterranean) as examples. The lens of complex systems science yields insights into facets of forest structure and dynamics that facilitate comparisons among ecosystems. These biomes share the main properties of complex systems but differ in specific ecological properties, disturbance regimes, and human uses. We show how this approach can help forest scientists and managers to conceptualize forests as integrated social-ecological systems and provide concrete examples of how to manage forests as complex adaptive systems.

Predators impact prey populations not only by consuming individuals, but also by altering their behaviours. These nonlethal effects can influence food web properties as much as lethal effects. The mechanisms of nonlethal effects include chronic and temporary anti-predator behaviours, the nature of which depends on the spatial dynamics of predators and the range over which prey perceive risk. The relation between chronic and ephemeral responses to risk determines predator-prey interactions, with consequences that can ripple across the food web. Nonetheless, few studies have quantified the spatio-temporal scales over which prey respond to predation threat, and how this response varies with habitat features. We evaluated the reaction of radio-collared caribou and moose to the passage of radio-collared wolves, by considering changes in movement characteristics during winter and summer. We used an optimization algorithm to identify the rate at which the impact of prior passage of wolves decreases over time and with the predator's distance. The spatial and temporal scales of anti-predator responses varied with prey species and season. Caribou and moose displayed four types of behaviour following the passage of wolves: lack of response, increased selection of safe land cover types, decreased selection of risky cover types and increased selection of food-rich forest stands. For example, moose increased their avoidance of open conifer stands with lichen in summer, which are selected by wolves in this season. Also in winter, caribou increased their selection of conifer stands with lichen for nearly 10 days following a wolf's passage. This stronger selection for food-rich patches could indicate that the recent passage of wolves informs caribou on the current predator distribution and reveals the rate at which this information become less reliable over time. Caribou and moose used anti-predator responses that combine both long- and short-term behavioural adjustments. The spatial game between wolves and their prey involves complex and nonlinear mechanisms that vary between species and seasons. A comprehensive assessment of risk effects on ecosystem dynamics thus requires the characterization of chronic and temporary anti-predator behaviours.

The world's forests and forestry sector are facing unprecedented biological, political, social, and climatic challenges. The development of appropriate, novel forest management and restoration approaches that adequately consider uncertainty and adaptability are hampered by a continuing focus on production of a few goods or objectives, strong control of forest structure and composition, and most importantly the absence of a global scientific framework and long-term vision. Ecosystem-based approaches represent a step in the right direction, but are limited in their ability to deal with the rapid pace of social, climatic, and environmental changes. We argue here that viewing forest ecosystems as complex adaptive system provides a better alternative for both production-and conservation-oriented forests and forestry. We propose a set of broad principles and changes to increase the adaptive capacity of forests in the face of future uncertainties. These span from expanding the sustainedyield, single-good paradigm to developing policy incentives and interventions that promote self-organization and integrated social-ecological adaptation.

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