Abstract. Woody encroachment is a widespread and acute phenomenon affecting grasslands and savannas worldwide. We performed a meta-analysis of 29 studies from 13 different grassland/savanna communities in North America to determine the consequences of woody encroachment on plant species richness. In all 13 communities, species richness declined with woody plant encroachment (average decline ¼ 45%). Species richness declined more in communities with higher precipitation (r 2 ¼ 0.81) and where encroachment was associated with a greater change in annual net primary productivity (ANPP; r 2 ¼ 0.69). Based on the strong positive correlation between precipitation and ANPP following encroachment (r 2 ¼ 0.87), we hypothesize that these relationships occur because water-limited woody plants experience a greater physiological and demographic release as precipitation increases. The observed relationship between species richness and ANPP provides support for the theoretical expectation that a trade-off occurs between richness and productivity in herbaceous communities. We conclude that woody plant encroachment leads to significant declines in species richness in North American grassland/savanna communities.

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Abstract. Over the last century, many grasslands worldwide have transitioned from a graminoid to a tree/shrub-dominated state in a short period of time, a phenomenon referred to as woody encroachment. Positive feedbacks and bi-stability are thought to be important drivers of woody encroachment, but there is little empirical evidence to suggest that positive feedbacks accelerate the woody encroachment of mesic grasslands. In mesic tallgrass prairie, shrub establishment does not directly facilitate seedling establishment. Yet, shrub establishment may facilitate the clonal spread of existing shrubs into nearby patches, because clonal reproduction might circumvent barriers that typically limit woody seedlings. Our results show that when Cornus drummondii (the predominate encroacher of mesic tallgrass prairie) extends rhizomatous stems into open grasslands, these stems use the same deep soil water sources as mature stems-thereby avoiding competition with grasses and gaining access to a reliable water source. In addition, herbaceous fuel concentrations are lower at the shrub/grass interface than in open grasslands, reducing the potential impacts of subsequent grassland fires. We propose that the release from resource and fire limitation results in a positive feedback loop as clonal stems are able to extend into surrounding patches, circumvent demographic barriers, mature, and spread by developing their own clonal stems. Long-term data on site (26 years) corroborates this interpretation: the size of deep-rooted clonal shrub species has increased 16-fold and their cover has increased from 0 to 27%, whereas the cover of shallowrooted species (both clonal and non-clonal) has only increased marginally. Together, these results suggest that (1) positive feedbacks can facilitate mesic woody encroachment and (2) bi-stability exists in mesic tallgrass prairie.

Summary1. This review synthesizes evidence that altered fire frequency drives discontinuous ecosystem transitions from mesic grasslands to shrublands or woodlands in the Central Great Plains, USA. 2. Long-term fire manipulations reveal that grassland to shrubland transitions are triggered when fire-free intervals increase from 1-3 years to ≥ 3-8 years, and longer fire returns (~10 years or more) result in transitions to woodlands. Grazing and soil properties alter these fire thresholds. 3. Grassland to shrubland transitions are abrupt and exhibit nonlinear relationships between driver and state variables. Transitions to shrublands and woodlands exhibit hysteresis, where reintroducing frequent fires does not reverse transitions in management-relevant time-scales (decades). 4. Nonlinear transitions and hysteresis emerge because grasses generate positive feedbacks with fire that create strong demographic barriers for shrub and tree establishment. Fire-free intervals allow shrubs and trees to reach a size sufficient to survive fire, reproduce and disrupt the fire feedback loop through competition. 5. Synthesis. Mesic grasslands, shrublands and woodlands constitute self-reinforcing states (alternative attractors) separated by critical fire frequency thresholds. Even without major shifts in climate, altered fire frequency can produce dramatic state changes, highlighting the importance of fire for predicting future ecosystem states. Local management should focus on prevention of unwanted transitions rather than post hoc restoration.

Ecologists have long studied patterns, directions and tempos of change, but there is a pressing need to extend current understanding to empirical observations of abrupt changes as climate warming accelerates. Abrupt changes in ecological systems (ACES)—changes that are fast in time or fast relative to their drivers—are ubiquitous and increasing in frequency. Powerful theoretical frameworks exist, yet applications in real-world landscapes to detect, explain and anticipate ACES have lagged. We highlight five insights emerging from empirical studies of ACES across diverse ecosystems: (i) ecological systems show ACES in some dimensions but not others; (ii) climate extremes may be more important than mean climate in generating ACES; (iii) interactions among multiple drivers often produce ACES; (iv) contingencies, such as ecological memory, frequency and sequence of disturbances, and spatial context are important; and (v) tipping points are often (but not always) associated with ACES. We suggest research priorities to advance understanding of ACES in the face of climate change. Progress in understanding ACES requires strong integration of scientific approaches (theory, observations, experiments and process-based models) and high-quality empirical data drawn from a diverse array of ecosystems. This article is part of the theme issue ‘Climate change and ecosystems: threats, opportunities and solutions’

Ecosystems with alternative attractors are susceptible to abrupt regime shifts that are often difficult to predict and reverse. In this study, we quantify multiple system dynamics to determine whether the transition of mesic grassland to shrubland, a widespread phenomenon, represents a linear reversible process, a nonlinear but reversible threshold process, or a transition between alternative attractors that is nonlinear and prone to hysteresis. Using a 28‐yr data set with annual resolution and extensive spatial replication, we found that shrub cover is correlated with distinct thresholds of fire and C4 grass cover, resulting in temporal bimodality of shrub cover and abrupt shifts of shrub cover despite gradual changes in grass cover. These abrupt increases in shrub cover are the most rapid ever reported in grasslands, and illustrate internal thresholds that separate grasslands and shrublands. Nonlinear transitions from low to high shrub cover were also closely associated with positive feedback mechanisms that alter fire and competition (r2 = 0.65), suggesting that grasslands and shrublands could show hysteresis, and by definition exist as alternative attractors. Thus, the response of this ecosystem to anthropogenic activity should tend to be rapid, nonlinear, and perhaps difficult to reverse. Regime shifts in this mesic grassland were predictable: we found that grassland and shrubland attractors were differentiated by critical thresholds of ∼50–70% grass cover, 5–10% shrub cover, and a fire return interval of ∼3 yr. These thresholds may provide adaptive potential for managing nonlinear behavior in socio‐ecological systems in a changing environment.

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While a growing proportion of global food consumption is obtained through international trade, there is an ongoing debate on whether this increased reliance on trade benefits or hinders food security, and specifically, the ability of global food systems to absorb shocks due to local or regional losses of production. This paper introduces a model that simulates the short-term response to a food supply shock originating in a single country, which is partly absorbed through decreases in domestic reserves and consumption, and partly transmitted through the adjustment of trade flows. By applying the model to publicly-available data for the cereals commodity group over a 17 year period, we find that differential outcomes of supply shocks simulated through this time period are driven not only by the intensification of trade, but as importantly by changes in the distribution of reserves. Our analysis also identifies countries where trade dependency may accentuate the risk of food shortages from foreign production shocks; such risk could be reduced by increasing domestic reserves or importing food from a diversity of suppliers that possess their own reserves. This simulation-based model provides a framework to study the short-term, nonlinear and out-of-equilibrium response of trade networks to supply shocks, and could be applied to specific scenarios of environmental or economic perturbations.