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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]

Riparian zones and the streams they border provide vital habitat for organisms, water quality protection, and other important ecosystem services. These areas are under pressure from local (land use/land cover change) to global (climate change) processes. Woody vegetation is expanding in grassland riparian zones worldwide. Here we report on a decade‐long watershed‐scale mechanical removal of woody riparian vegetation along 4.5 km of stream channel in a before–after control impact experiment. Prior to this removal, woody plants had expanded into grassy riparian areas, associated with a decline in streamflow, loss of grassy plant species, and other ecosystem‐scale impacts. We confirmed some expected responses, including rapid increases in stream nutrients and sediments, disappearance of stream mosses, and decreased organic inputs to streams via riparian leaves. We were surprised that nutrient and sediment increases were transient for 3 years, that there was no recovery of stream discharge, and that areas with woody removal did not shift back to a grassland state, even when reseeded with grassland species. Rapid expansion of shrubs (Cornus drummondii, Prunus americana) in the areas where trees were removed allowed woody vegetation to remain dominant despite repeating the cutting every 2 years. Our results suggest woody expansion can fundamentally alter terrestrial and aquatic habitat connections in grasslands, resulting in inexorable movement toward a new ecosystem state. Human pressures, such as climate change, atmospheric CO2 increases, and elevated atmospheric nitrogen deposition, could continue to push the ecosystem along a trajectory that is difficult to change. Our results suggest that predicting relationships between riparian zones and the streams they border could be difficult in the face of global change in all biomes, even in well‐studied sites. [ABSTRACT FROM AUTHOR]

As temperatures continue rising, the direction, magnitude, and tempo of change in disturbance‐prone forests remain unresolved. Even forests long resilient to stand‐replacing fire face uncertain futures, and efforts to project changes in forest structure and composition are sorely needed to anticipate future forest trajectories. We simulated fire (incorporating fuels feedbacks) and forest dynamics on five landscapes spanning the Greater Yellowstone Ecosystem (GYE) to ask the following questions: (1) How and where are forest landscapes likely to change with 21st‐century warming and fire activity? (2) Are future forest changes gradual or abrupt, and do forest attributes change synchronously or sequentially? (3) Can forest declines be averted by mid‐21st‐century stabilization of atmospheric greenhouse gas (GHG) concentrations? We used the spatially explicit individual‐based forest model iLand to track multiple attributes (forest extent, stand age, tree density, basal area, aboveground carbon stocks, dominant forest types, species occupancy) through 2100 for six climate scenarios. Hot‐dry climate scenarios led to more fire, but stand‐replacing fire peaked in mid‐century and then declined even as annual area burned continued to rise. Where forest cover persisted, previously dense forests were converted to sparse young woodlands. Increased aridity and fire drove a ratchet of successive abrupt declines (i.e., multiple annual landscape‐level changes ≥20%) in tree density, basal area, and extent of older (>150 yr) forests, whereas declines in carbon stocks and mean stand age were always gradual. Forest changes were asynchronous across landscapes, but declines in stand structure always preceded reductions in forest extent and carbon stocks. Forest decline was most likely in less topographically complex landscapes dominated by fire‐sensitive tree species (Picea engelmannii, Abies lasiocarpa, Pinus contorta var. latifolia) and where fire resisters (Pseudotsuga menziesii var. glauca) were not already prevalent. If current GHG emissions continue unabated (RCP 8.5) and aridity increases, a suite of forest changes would transform the GYE, with cascading effects on biodiversity and myriad ecosystem services. However, stabilizing GHG concentrations by mid‐century (RCP 4.5) would slow the ratchet, moderating fire activity and dampening the magnitude and rate of forest change. Monitoring changes in forest structure may serve as an operational early warning indicator of impending forest decline. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Hysteresis is a fundamental characteristic of alternative stable state theory, yet evidence of hysteresis is rare. In mesic grasslands, fire frequency regulates transition from grass‐ to shrub‐dominated system states. It is uncertain, however, if increasing fire frequency can reverse shrub expansion, or if grass‐shrub dynamics exhibit hysteresis. We implemented annual burning in two infrequently burned grasslands and ceased burning in two grasslands burned annually. With annual fires, grassland composition converged on that of long‐term annually burned vegetation due to rapid recovery of grass cover, although shrubs persisted. When annual burning ceased, shrub cover increased, but community composition did not converge with a long‐term infrequently burned reference site because of stochastic and lagged dispersal by shrubs, reflecting hysteresis. Our results demonstrated that annual burning can slow, but not reverse, shrub encroachment. In addition, reversing fire frequencies resulted in hysteresis because vegetation trajectories from grassland to shrubland differed from those of shrubland to grassland. [ABSTRACT FROM AUTHOR]

Aim: Simulation models are important tools for quantifying the resilience (i.e., persistence under changed environmental conditions) of forest ecosystems to global change. We synthesized the modelling literature on forest resilience, summarizing common models and applications in resilience research, and scrutinizing the implementation of important resilience mechanisms in these models. Models applied to assess resilience are highly diverse, and our goal was to assess how well they account for important resilience mechanisms identified in experimental and empirical research. Location: Global. Time period: 1994 to 2019. Major taxa studied: Trees. Methods: We reviewed the forest resilience literature using online databases, selecting 119 simulation modelling studies for further analysis. We identified a set of resilience mechanisms from the general resilience literature and analysed models for their representation of these mechanisms. Analyses were grouped by investigated drivers (resilience to what) and responses (resilience of what), as well as by the type of model being used. Results: Models used to study forest resilience varied widely, from analytical approaches to complex landscape simulators. The most commonly addressed questions were associated with resilience of forest cover to fire. Important resilience mechanisms pertaining to regeneration, soil processes, and disturbance legacies were explicitly simulated in only 34 to 46% of the model applications. Main conclusions: We found a large gap between processes identified as underpinning forest resilience in the theoretical and empirical literature, and those represented in models used to assess forest resilience. Contemporary forest models developed for other goals may be poorly suited for studying forest resilience during an era of accelerating change. Our results highlight the need for a new wave of model development to enhance understanding of and management for resilient forests. [ABSTRACT FROM AUTHOR]

Questions: Climate extremes are predicted to become more common in many ecosystems. Climate extremes can promote and interact with disturbances, but the combined effects of climate extremes and disturbances have not been quantified in many ecosystems. In this study, we ask whether the dual impact of a climate extreme and concomitant disturbance (wildfire) has a greater affect than a climate extreme alone. Location: Tallgrass prairie in the Konza Prairie Biological Station, northeastern Kansas, USA. Methods: We quantified the response of a tallgrass prairie plant community to a 2‐year climate extreme of low growing‐season precipitation and high temperatures. In the first year of the climate extreme, a subset of plots was burned by a growing‐season wildfire. This natural experiment allowed us to compare community responses to a climate extreme with and without wildfire. Results: In plots exposed to the climate extreme but not wildfire, community structure, diversity, and composition showed minor to insignificant changes, such as a 20% reduction in grass cover and a slight increase in species diversity. Plots exposed to both the climate extreme and wildfire underwent larger changes, including an 80% reduction in grass cover, 50% increase in forb cover, and increased plant diversity. Two years after the climate extreme, structural shifts in burned plots showed little sign of recovery, indicating a potentially lasting shift in plant community structure. Conclusions: Our results suggest that community responses to climate extremes need to account for climate‐related disturbances — in this case, high temperatures, drought and wildfire. This response diverged from our expectation that heat, drought, and an additional fire would favor grasses. Although growing‐season wildfires in tallgrass prairie have been rare in recent decades, they will likely become more common with climate change, potentially leading to changes in grassland structure. [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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Winter climate is expected to change under future climate scenarios, yet the majority of winter ecology research is focused in cold-climate ecosystems. In many temperate systems, it is unclear how winter climate relates to biotic responses during the growing season. The objective of this study was to examine how winter weather relates to plant and animal communities in a variety of terrestrial ecosystems ranging from warm deserts to alpine tundra. Specifically, we examined the association between winter weather and plant phenology, plant species richness, consumer abundance, and consumer richness in 11 terrestrial ecosystems associated with the U.S. Long-Term Ecological Research ( LTER) Network. To varying degrees, winter precipitation and temperature were correlated with all biotic response variables. Bud break was tightly aligned with end of winter temperatures. For half the sites, winter weather was a better predictor of plant species richness than growing season weather. Warmer winters were correlated with lower consumer abundances in both temperate and alpine systems. Our findings suggest winter weather may have a strong influence on biotic activity during the growing season and should be considered in future studies investigating the effects of climate change on both alpine and temperate systems. [ABSTRACT FROM AUTHOR]

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., 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., 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)., 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., 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Intensified anthropogenic activities have drastically altered many ecosystems, motivating the use of restoration to regain key ecosystem functions and services and to stem biodiversity losses. Restoration is particularly difficult when human activities have pushed an area into a new state with self‐reinforcing feedbacks. This study investigated a long‐term restoration project in a dryland ecosystem of the Tengger Desert in northwestern China, initiated in 1956. We analyzed shrub and grass cover for 49 yr after the installation of restoration infrastructure that altered external conditions (i.e., using packed straw to reduce wind erosion) and system state (by planting shrubs). After 37–40 yr, the re‐vegetation project was successful in restoring the system to a state similar to native vegetation, with high grass cover (30–50%), low shrub cover (8–10%), and a thick biological soil crust (biocrust). However, the shift to high grass cover did not begin until year 37, before which shrub cover was high (15–20%) and grasses were subdominant (usually <10%). The shift from shrub to grass dominance was abrupt, registering significant nonlinear changes over time and relative to a key driver of vegetation dynamics, estimated biocrust thickness. Biocrust thickness increased gradually over time, which reduced rainfall infiltration into deep soil and thus increased soil moisture available for the shallow‐rooted grasses. The shift from the bare soil state to the steppe state exhibited a long time lag, suggesting that it can take decades to determine whether dryland restoration efforts succeed. The results indicate that persistence might be critical to forcing desired state transitions and that dryland restoration can proceed as a series of time lags, punctuated by abrupt changes in ecosystem state. [ABSTRACT FROM AUTHOR]