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Changing climate and disturbance regimes are increasingly challenging the resilience of forest ecosystems around the globe. A powerful indicator for the loss of resilience is regeneration failure, that is, the inability of the prevailing tree species to regenerate after disturbance. Regeneration failure can result from the interplay among disturbance changes (e.g., larger and more frequent fires), altered climate conditions (e.g., increased drought), and functional traits (e.g., method of seed dispersal). This complexity makes projections of regeneration failure challenging. Here we applied a novel simulation approach assimilating data-driven fire projections with vegetation responses from process modeling by means of deep neural networks. We (i) quantified the future probability of regeneration failure; (ii) identified spatial hotspots of regeneration failure; and (iii) assessed how current forest types differ in their ability to regenerate under future climate and fire. We focused on the Greater Yellowstone Ecosystem (2.9 × 106  ha of forest) in the Rocky Mountains of the USA, which has experienced large wildfires in the past and is expected to undergo drastic changes in climate and fire in the future. We simulated four climate scenarios until 2100 at a fine spatial grain (100 m). Both wildfire activity and unstocked forest area increased substantially throughout the 21st century in all simulated scenarios. By 2100, between 28% and 59% of the forested area failed to regenerate, indicating considerable loss of resilience. Areas disproportionally at risk occurred where fires are not constrained by topography and in valleys aligned with predominant winds. High-elevation forest types not adapted to fire (i.e., Picea engelmannii-Abies lasiocarpa as well as non-serotinous Pinus contorta var. latifolia forests) were especially vulnerable to regeneration failure. We conclude that changing climate and fire could exceed the resilience of forests in a substantial portion of Greater Yellowstone, with profound implications for carbon, biodiversity, and recreation.

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'.

Changes in climate and land management over the last half‐century have favoured woody plants native to grasslands and led to the rapid expansion of woody species. Despite this being a global phenomenon, it is unclear why some woody species have rapidly expanded while others have not. We assessed whether the most abundant woody encroaching species in tallgrass prairie have common growth forms and physiology or unique traits that differentiate their resource‐use strategies. We characterized the abundance, above‐ground carbon allocation, and leaf‐level physiological and structural traits of seven woody encroaching species in tallgrass prairie that span an order of magnitude in abundance. To identify species‐specific increases in abundance, we used a 34‐year species composition dataset at Konza Prairie Biological Station (Central Great Plains, USA). We then compared biomass allocation and leaf‐level traits to determine differences in carbon and water use strategies among species. While all focal species increased in abundance over time, encroachment in this system is primarily driven by three species: Cornus drummondii, Prunus americana and Rhus glabra. The most dominant species, Cornus drummondii, had the most extreme values for several traits, including the lowest leaf:stem mass ratios, lowest photosynthetic capacity and highest turgor loss point. Two of the most abundant species, Cornus drummondii and Rhus glabra, had opposing growth forms and resource‐use strategies. These species had significantly different above‐ground carbon allocation, leaf‐level drought tolerance and photosynthetic capacity. There were surprisingly few interspecific differences in specific leaf area and leaf dry matter content, suggesting these traits were poor predictors of species‐level encroachment. Synthesis. Woody encroaching species in tallgrass prairie encompass a spectrum of growth forms and leaf physiology. Two of the most abundant woody species fell at opposite ends of this spectrum. Our results suggest niche differences among a community of woody species facilitate the rapid encroachment by a few species. This study shows that woody encroaching species do not conform to a ‘one‐size‐fits‐all’ strategy, and a diversity of growth forms and physiological strategies may make it more challenging to reach management goals that aim to conserve or restore grassland communities. [ABSTRACT FROM AUTHOR]

Regime shifts are difficult-to-reverse transitions that occur when an ecosystem reorganizes around a new set of self-reinforcing feedbacks. Regime shifts are predicted to occur when the intensity of some exogenous driver variable, such as temperature, annual harvest rate, or nutrient addition rate, gradually approaches and crosses a threshold value, initiating a transition to an alternative state. However, many driver variables now change rapidly as presses or pulses, not gradually, requiring new conceptual frameworks for understanding and predicting regime shifts. We argue that identifying and controlling regime shifts in response to presses and pulses will require a greater focus on the duration, not just the intensity, of changes in driver variables. In ecosystems with slower dynamics, transitions to an alternative state can take years to decades and as a result, a driver press with an intensity capable of resulting in a regime shift over long time spans may fail to cause a regime shift when applied for shorter durations. We illustrate these ideas using simulations of local-scale alternative stable state models and preliminary evidence from long-term grazing and eutrophication experiments. The simulations also suggest that small changes in the duration of driver presses or pulses can determine whether an ecosystem recovers to its original state. These insights may extend to larger scales. In spatially extended simulations that included patchiness, spatial heterogeneity, and spatial connectivity, all patches recovered to their original state after shorter presses. However, once press duration exceeded a threshold, growing proportions of the landscape shifted to an alternative state as press duration increased. We observed similar patchy transitions in a catchment-scale experiment that reinstated frequent fires approximately halfway through a regime shift from grassland to shrubland, initiated by fire suppression. In both the local- and larger-scale models, the threshold duration needed to elicit regime shifts decreased as press intensity increased or when factors counteracting regime shifts weakened. These multiple lines of evidence suggest that conceptualizing regime shifts as an interactive function of the intensity and duration of driver changes will increase understanding of the varying effects of driver presses, pulses, and cycles on ecosystem dynamics.

State transitions are changes in ecosystem structure and self-reinforcing feedbacks that are initiated when an exogenous driver variable crosses a threshold. Reversing state transitions is difficult and costly. While some state transitions are relatively rapid, many take years to decades. Outside of theoretical models, very little is known about slower state transitions and how they unfold in time and space. We quantified changes in spatial variance as a mesic grassland ecosystem shifts to a shrub-dominated state, using long-term experiments and simulations that maintain grasslands with annual fires or initiate a state transition to shrub dominance by decreasing fire frequency. In the experiments, the susceptibility to state transitions varied substantially in space. In the less frequent fire treatment, some plots became shrub-dominated around year 20 and grass extirpations began in year 25, but a third of the plots were still grass-dominated in year 37. Variable rates of state transition resulted in increasing spatial variance of grass cover over time, whereas shrub cover variance decreased. In the annually burned treatment, grasses remained dominant and the spatial variance of grass cover declined. In a separate experiment, less frequent fires were maintained for 23 years and then switched to annual fires. The switch to annual fires occurred shortly after grass variance started to increase and a majority of these plots quickly returned to a grass dominated state. In simulations, spatial variance remained low and average grass cover was high under frequent fires. If fire frequency decreased below a threshold, the ecosystem transitioned to shrubland, with a transient increase in the spatial variance of grass cover during the transition between states. Synthesis. Spatial variability in the rate and susceptibility to state transitions is indicative of a system with a patchy spatial structure, high spatial heterogeneity and low connectivity between patches. Increases in spatial variance can serve as an indication that some patches have begun a state transition and that management interventions are needed to avoid widespread transitions. This is one of the first empirical examples where altering management after an increase in spatial variance prevented state transitions.

Species interaction effects on populations can vary in both magnitude (i.e. strong vs. weak) and sign (positive, negative, or no effect). Context‐dependent effects of species interactions occur when the sign or strength of the interaction's effect on population growth rate changes across abiotic gradients.We know that species can vary substantially in the degree of context dependence they exhibit, even across similar abiotic gradients. However, few studies have characterised context dependence of co‐occurring species, limiting our ability to understand the implications of context dependence for species interaction effects on community composition.Using over three decades of data collected for 13 tallgrass prairie forbs at the Konza Prairie Biological Station, we parameterise density structured population models that predict population dynamics as functions of abiotic conditions and bison herbivory. We use these models to estimate the degree of context dependence in responses to bison herbivory for 13 species across three abiotic gradients: weather, fire frequency and soil type.All species showed significant context dependence for fire frequency in the same direction, though with variable magnitude, such that herbivory increased cover with more frequent fires. Context dependence with weather and soil type varied dramatically across species in both direction and magnitude. For example, herbivory effects on 3/13 species were stronger in wet conditions, but herbivory effects on 5/13 species were stronger in dry conditions. Thus, context dependence exhibited by individual species, as opposed to effects of abiotic conditions on the relative abundances of species, could generate much of the weather‐dependent effects of herbivory on community composition.Synthesis: Our work suggests that species can vary dramatically in the presence, direction and magnitude of context dependence, even when occurring in the same community and when considering the same species interaction (i.e. response to a herbivore). In addition, we find that context dependence could drive substantial variation in the effect of species interactions on community characteristics (e.g. composition) across multiple abiotic gradients. [ABSTRACT FROM AUTHOR]

Ensuring food security requires food production and distribution systems function throughout disruptions. Understanding the factors that contribute to the global food system's ability to respond and adapt to such disruptions (i.e. resilience) is critical for understanding the long-term sustainability of human populations. Variable impacts of production shocks on food supply between countries indicate a need for national-scale resilience indicators that can provide global comparisons. However, methods for tracking changes in resilience have had limited application to food systems. We developed an indicator-based analysis of food systems resilience for the years 1992-2011. Our approach is based on three dimensions of resilience: socio-economic access to food in terms of income of the poorest quintile relative to food prices, biophysical capacity to intensify or extensify food production, and the magnitude and diversity of current domestic food production. The socio-economic indicator has large variability, but with low values concentrated in Africa and Asia. The biophysical capacity indicator is highest in Africa and Eastern Europe, in part because of high potential for extensification of cropland and for yield gap closure in cultivated areas. However, the biophysical capacity indicator has declined globally in recent years. The production diversity indicator has increased slightly, with a relatively even geographic distribution. Few countries had exclusively high or low values for all indicators. Collectively, these results are the basis for global comparisons of resilience between nations, and provide necessary context for developing generalizations about the resilience in the global food system.

Spatially diverse trends in population growth, climate change, industrialization, urbanization and economic development are expected to change future food supply and demand. These changes may affect the suitability of land for food production, implying elevated risks especially for resourceconstrained, food-importing countries.Wepresent the evolution of biophysical redundancy for agricultural production at country level, from 1992 to 2012. Biophysical redundancy, defined as unused biotic and abiotic environmental resources, is represented by the potential food production of 'spare land', available water resources (i.e., not already used for human activities), as well as production increases through yield gap closure on cultivated areas and potential agricultural areas. In 2012, the biophysical redundancy of 75 (48) countries, mainly in North Africa, Western Europe, the Middle East and Asia, was insufficient to produce the caloric nutritional needs for at least 50% (25%) of their population during a year. Biophysical redundancy has decreased in the last two decades in 102 out of 155 countries, 11 of these went from high to limited redundancy, and nine of these from limited to very low redundancy. Although the variability of the drivers of change across different countries is high, improvements in yield and population growth have a clear impact on the decreases of redundancy towards the very low redundancy category.Wetook a more detailed look at countries classified as 'Low Income Economies (LIEs)' since they are particularly vulnerable to domestic or external food supply changes, due to their limited capacity to offset for food supply decreases with higher purchasing power on the international market. Currently, nine LIEs have limited or very low biophysical redundancy. Many of these showed a decrease in redundancy over the last two decades, which is not always linked with improvements in per capita food availability.

Animals must track resources over relatively fine spatial and temporal scales, particularly in disturbance‐mediated systems like grasslands. Grassland birds respond to habitat heterogeneity by dispersing among sites within and between years, yet we know little about how they make post‐dispersal settlement decisions. Many methods exist to quantify the resource selection of mobile taxa, but the habitat data used in these models are frequently not collected at the same location or time that individuals were present. This spatiotemporal misalignment may lead to incorrect interpretations and adverse conservation outcomes, particularly in dynamic systems. To investigate the extent to which spatially and temporally dynamic vegetation conditions and topography drive grassland bird settlement decisions, we integrated multiple data sources from our study site to predict slope, vegetation height, and multiple metrics of vegetation cover at any point in space and time within the temporal and spatial scope of our study. We paired these predictions with avian mark‐resight data for 8 years at the Konza Prairie Biological Station in NE Kansas to evaluate territory selection for Grasshopper Sparrows (Ammodramus savannarum), Dickcissels (Spiza americana), and Eastern Meadowlarks (Sturnella magna). Each species selected different types and amounts of herbaceous vegetation cover, but all three species preferred relatively flat areas with less than 6% shrub cover and less than 1% tree cover. We evaluated several scenarios of woody vegetation removal and found that, with a targeted approach, the simulated removal of just one isolated tree in the uplands created up to 14 ha of grassland bird habitat. This study supports growing evidence that small amounts of woody encroachment can fragment landscapes, augmenting conservation threats to grassland systems. Conversely, these results demonstrate that drastic increases in bird habitat area could be achieved through relatively efficient management interventions. The results and approaches reported pave the way for more efficient conservation efforts in grasslands and other systems through spatiotemporal alignment of habitat with animal behaviors and simulated impacts of management interventions. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]