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A number of ecosystems can exhibit abrupt shifts between alternative stable states. Because of their important ecological and economic consequences, recent research has focused on devising early warning signals for anticipating such abrupt ecological transitions. In particular, theoretical studies show that changes in spatial characteristics of the system could provide early warnings of approaching transitions. However, the empirical validation of these indicators lag behind their theoretical developments. Here, we summarize a range of currently available spatial early warning signals, suggest potential null models to interpret their trends, and apply them to three simulated spatial data sets of systems undergoing an abrupt transition. In addition to providing a step-by-step methodology for applying these signals to spatial data sets, we propose a statistical toolbox that may be used to help detect approaching transitions in a wide range of spatial data. We hope that our methodology together with the computer codes will stimulate the application and testing of spatial early warning signals on real spatial data.

CONTEXT: More than a century of forest and fire management of Inland Pacific landscapes has transformed their successional and disturbance dynamics. Regional connectivity of many terrestrial and aquatic habitats is fragmented, flows of some ecological and physical processes have been altered in space and time, and the frequency, size and intensity of many disturbances that configure these habitats have been altered. Current efforts to address these impacts yield a small footprint in comparison to wildfires and insect outbreaks. Moreover, many current projects emphasize thinning and fuels reduction within individual forest stands, while overlooking large-scale habitat connectivity and disturbance flow issues. METHODS: We provide a framework for landscape restoration, offering seven principles. We discuss their implication for management, and illustrate their application with examples. RESULTS: Historical forests were spatially heterogeneous at multiple scales. Heterogeneity was the result of variability and interactions among native ecological patterns and processes, including successional and disturbance processes regulated by climatic and topographic drivers. Native flora and fauna were adapted to these conditions, which conferred a measure of resilience to variability in climate and recurrent contagious disturbances. CONCLUSIONS: To restore key characteristics of this resilience to current landscapes, planning and management are needed at ecoregion, local landscape, successional patch, and tree neighborhood scales. Restoration that works effectively across ownerships and allocations will require active thinking about landscapes as socio-ecological systems that provide services to people within the finite capacities of ecosystems. We focus attention on landscape-level prescriptions as foundational to restoration planning and execution.

Humans and climate affect ecosystems and their services, which may involve continuous and discontinuous transitions from one stable state to another. Discontinuous transitions are abrupt, irreversible and among the most catastrophic changes of ecosystems identified. For terrestrial ecosystems, it has been hypothesized that vegetation patchiness could be used as a signature of imminent transitions. Here, we analyse how vegetation patchiness changes in arid ecosystems with different grazing pressures, using both field data and a modelling approach. In the modelling approach, we extrapolated our analysis to even higher grazing pressures to investigate the vegetation patchiness when desertification is imminent. In three arid Mediterranean ecosystems in Spain, Greece and Morocco, we found that the patch-size distribution of the vegetation follows a power law. Using a stochastic cellular automaton model, we show that local positive interactions among plants can explain such power-law distributions. Furthermore, with increasing grazing pressure, the field data revealed consistent deviations from power laws. Increased grazing pressure leads to similar deviations in the model. When grazing was further increased in the model, we found that these deviations always and only occurred close to transition to desert, independent of the type of transition, and regardless of the vegetation cover. Therefore, we propose that patch-size distributions may be a warning signal for the onset of desertification.

Fire is an important disturbance in many forest landscapes, but there is heightened concern regarding recent wildfire activity in western North America. Several regional‐scale studies focus on high‐severity fire, but a comprehensive examination at all levels of burn severity (i.e., low, moderate, and high) is needed to inform our understanding of the ecological effects of contemporary fires and how they vary among vegetation zones at sub‐regional scales. We integrate Landsat time series data with field measurements of tree mortality to map burn severity in forests of the Pacific Northwest, USA, from 1985 to 2010. We then examine temporal trends in fire extent and spatial patterns of burn severity in relation to drought and annual fire extent. Finally, we compare results among vegetation zones and with expectations based on studies of historical landscape dynamics and fire regimes. Small increases in fire extent over time were associated with drought in all vegetation zones, but fire cumulatively affected <3% of wet vegetation zones, and most dry vegetation zones experienced less fire than expectations from fire history studies. Although the proportion of fire at any level of severity did not increase over time, temporal trends toward larger patches of high‐severity fire were related to drought and annual fire extent, depending on vegetation zone. In vegetation zones with historically high‐severity regimes, high‐severity fire accounted for a large proportion of recent fire extent (43–48%) and occurred primarily in patches ≥100 ha. In vegetation zones with historically low‐ and mixed‐severity regimes, low (45–54%)‐ and moderate‐severity (24–36%) fires were prevalent, but proportions of high‐severity fire (23–26%), almost half of which occurred in patches ≥100 ha, were much greater than expectations from most fire history studies. Our results support concerns about large patches of high‐severity fire in some dry forests but also suggest that spatial patterns of burn severity across much of the extent burned are generally consistent with current understanding of historical landscape dynamics in the region. This study highlights the importance of considering the ecological effects of fire at all levels of severity in management and policy initiatives intended to promote forest biodiversity and resilience to future fire activity.

In wind erosion models, previous parameters related to vegetation morphology and density are limited in describing the spatial distribution of vegetation that influences surface heterogeneity. Thus, it is not fully understood how spatial vegetation patterns affect wind erosion on a field-scale. Based on an investigation of 36 plots of vegetation in Alxa Plateau, northwestern China, we established a multivariate linear model for temporally and spatially averaged aerodynamic roughness length ( Z 0 ) incorporating the height, roughness density, regularity of vegetation patches (curvature) and spacing between patches (connectivity). The curvature positively interacted with the connectivity in affecting the mean Z 0 , while it was the most important factor affecting the standard deviation of Z 0 . The connectivity modulated the roughness density in affecting the standard deviation of Z 0 . The spatial-related terms contributed 37% and 62% to the model variance of the mean and standard deviation of Z 0 , respectively. Our results validate the importance of spatial vegetation structure in the vegetation-airflow interactions, with a suggestion of estimating the heterogeneity of surface erodibility by intuitive spatial parameters. Based on that spatial vegetation patterns reflect the ecosystem states, a strengthened linkage between wind erosion and vegetation stability may be useful in erosion regulation in drylands.

AimsInteractions among plants are the main biotic driver for the development of spatial vegetation patterns in dryland ecosystems. The modification of soil properties by perennial plants is an important mechanism. We aimed to examine how spatial vegetation patterns were affected by the local changes in soil physiochemical properties beneath shrubs in different communities.MethodsWe investigated 36 plots containing shrub vegetation on the Alxa Plateau, northwestern China. Patch sizes of all shrubs were measured. The median and curvature of patch size distribution (obtained by quadratic regression) were selected as indicators for spatial patterns. We determined soil physiochemical properties beneath the canopy of dominant and other shrub species and in adjacent intershrub spaces. A multi-model inference procedure was employed to analyze the standardized effect of the changes in soil properties and their interactions on the spatial vegetation patterns.ResultsLocal changes in soil physiochemical properties associated with shrub-soil feedbacks affected spatial vegetation patterns in different aspects. Salt aggregation was positively correlated with deviations from the power-law distribution, while nutrient concentration had no significant effect. Interaction between nutrient and salinity concentration positively affected the median patch size, maintaining a stable Turing-like pattern. Heterogeneity analysis showed that the dominant shrub species are representative of plant-soil interactions for the entire community.ConclusionsOur study suggests that soil salinity accumulated under the canopy of shrubs is the key soil property driving the spatial pattern of vegetation patches on the Alta Plateau.

The spatial configuration of vascular vegetation has been linked to variations in land degradation and ecosystem functioning in drylands. However, most studies on spatial patterns conducted to date have focused on a single or a few study sites within a particular region, specific vegetation types, or in landscapes characterized by a certain type of spatial patterns. Therefore, little is known on the general typology and distribution of plant spatial patterns in drylands worldwide, and on the relative importance of biotic and abiotic factors as predictors of their variations across geographical regions and habitat types. We analyzed 115 dryland plant communities from all continents except Antarctica to: 1) investigate the general typology of spatial patterns, and 2) assess the relative importance of biotic (plant cover, frequency of facilitation, soil amelioration, height of the dominant species) and abiotic (aridity, rainfall seasonality and sand content) factors as predictors of spatial patterns (median patch size, shape of patch‐size distribution and regularity) across contrasting habitat types (shrublands and grasslands). Precipitation during the warmest period and sand content were particularly strong predictors of plant spatial patterns in grasslands and shrublands, respectively. Facilitation associated with power‐law like and irregular spatial patterns in both shrublands and grasslands, although it was mediated by different mechanisms (respectively soil ammelioration and percentage of facilitated species). The importance of biotic attributes as predictors of the shape of patch‐size distributions declined with aridity in both habitats, leading to the emergence of more regular patterns under the most arid conditions. Our results expand our knowledge about patch formation in drylands and the habitat‐dependency of their drivers. They also highlight different ways in which facilitation affects ecosystem structure through the formation of plant spatial patterns.

Facilitation is a major force shaping the structure and diversity of plant communities in terrestrial ecosystems. Detecting positive plant-plant interactions relies on the combination of field experimentation and the demonstration of spatial association between neighboring plants. This has often restricted the study of facilitation to particular sites, limiting the development of systematic assessments of facilitation over regional and global scales. Here we explore whether the frequency of plant spatial associations detected from high-resolution remotely sensed images can be used to infer plant facilitation at the community level in drylands around the globe. We correlated the information from remotely sensed images freely available through Google Earth with detailed field assessments, and used a simple individual-based model to generate patch-size distributions using different assumptions about the type and strength of plant-plant interactions. Most of the patterns found from the remotely sensed images were more right skewed than the patterns from the null model simulating a random distribution. This suggests that the plants in the studied drylands show stronger spatial clustering than expected by chance. We found that positive plant co-occurrence, as measured in the field, was significantly related to the skewness of vegetation patch-size distribution measured using Google Earth images. Our findings suggest that the relative frequency of facilitation may be inferred from spatial pattern signals measured from remotely sensed images, since facilitation often determines positive co-occurrence among neighboring plants. They pave the road for a systematic global assessment of the role of facilitation in terrestrial ecosystems.

Recent studies report that multifunctionality—the simultaneous provision of multiple ecosystem functions—in drylands depends on biodiversity. Others report that specific size distributions of vegetation patches indicate overall ecosystem health and function. Using a biocrust (micro-vegetation of mosses, lichens, and cyanobacteria) model system, and multivariate modeling, we determined the relative importance of biodiversity, patch-size distribution, and total abundance to nutrient cycling and multifunctionality. In most cases we explained at least 20%, and up to 65%, of the variation in ecosystem functions, and 42% of the variation in multifunctionality. Species richness was the most important determinant of C cycling, constituting an uncommonly clear link between diversity and function in a non-experimental field setting. Regarding C cycling in gypsiferous soils, we found that patch size distributions with a greater frequency of small to medium patches, as opposed to very small patches, were more highly functional. Nitrogen cycling was largely a function of biocrust cover in two soil types, whereas in gypsiferous soils, more central-tending patch size distributions were less functional with regards to N cycling. All three community properties were about equally important to multifunctionality. Our results highlight the functional role of biotic attributes other than biodiversity, and indicate that high cover and diversity, together with a particular patch-size distribution, must be attained simultaneously to maximize multifunctionality. The results also agree with trends observed with other terrestrial and aquatic communities that more biodiversity is needed to sustain multifunctionality compared to single functions considered independently.

Background Climate is a main driver of fire regimes, but recurrent fires provide stabilizing feedbacks at several spatial scales that can limit fire spread and severity—potentially contributing to a form of self-regulation. Evaluating the strength of these feedbacks in wildland systems is difficult given the spatial and temporal scales of observation required. Here, we used the REBURN model to directly examine the relative strengths of top-down and bottom-up drivers of fire over a 3000-year simulation period, within a 275,000-ha conifer-dominated landscape in north central Washington State, USA. Results We found strong support for top-down and bottom-up spatial and temporal controls on fire patterns. Fire weather was a main driver of large fire occurrence, but area burned was moderated by ignition frequencies and by areas of limited fuels and fuel contagion (i.e., fire fences). Landscapes comprised of >40% area in fire fences rarely experienced large fire years. When large fires did occur during the simulation period, a recovery time of 100–300 years or more was generally required to recover pre-fire vegetation patterns. Conclusions Simulations showed that interactions between fire weather, fuel contagion, topography, and ignitions manifest variability in fire size and severity patch size distributions. Burned and recovering vegetation mosaics provided functional stabilizing feedbacks, a kind of meta stability, which limited future fire size and severity, even under extreme weather conditions. REBURN can be applied to new geographic and physiographic landscapes to simulate these interactions and to represent natural and culturally influenced fire regimes in historical, current, or future climatic settings.