Coupling microscale vegetation–soil water and macroscale vegetation–precipitation feedbacks in semiarid ecosystems

At macroscale, land–atmosphere exchange of energy and water in semiarid zones such as the Sahel constitutes a strong positive feedback between vegetation density and precipitation. At microscale, however, additional positive feedbacks between hydrology and vegetation such as increase of infiltration due to increase of vegetation, have been reported and have a large impact on vegetation distribution and spatial pattern formation. If both macroscale and microscale positive feedbacks are present in the same region, it is reasonable to assume that these feedback mechanisms are connected. In this study, we develop and analyse a soil‐vegetation‐atmosphere model coupling large‐scale evapotranspiration–precipitation feedback with a model of microscale vegetation–hydrology feedback to study the integration of these nonlinearities at disparate scales. From our results, two important conclusions can be drawn: (1) it is important to account for spatially explicit vegetation dynamics at the microscale in climate models (the strength of the precipitation feedback increased up to 35% by accounting for these microscale dynamics); (2) studies on resilience of ecosystems to climate change should always be cast within a framework of possible large‐scale atmospheric feedback mechanism (substantial changes in vegetation resilience resulted from incorporating macroscale precipitation feedback). Analysis of full‐coupled modelling shows that both type of feedbacks markedly influence each other and that they should both be accounted for in climate change models.     

Feedbacks between nutrient cycling and vegetation predict plant species coexistence and invasion

To investigate the role of species‐specific litter decomposability in determining plant community structure, we constructed a theoretical model of the codevelopmental dynamics of soil and vegetation. This model incorporates feedback between vegetation and soil. Vegetation changes the nutrient conditions of soil by affecting mineralization processes; soil, in turn, has an impact on plant community structure. The model shows that species‐level traits (decomposability, reproductive and competitive abilities) determine whether litter feedback effects are positive or negative. The feedback determines community‐level properties, such as species composition and community stability against invasion. The model predicts that positive feedback may generate multiple alternative steady states of the plant community, which differ in species richness or community composition. In such cases, the realized state is determined by initial abundance of co‐occurring species. Further, the model shows that the importance of species‐level traits depends on environmental conditions such as system fertility.     

Combined statistical and dynamical assessment of simulated vegetation–rainfall interactions in North Africa during the mid‐Holocene1

A negative feedback of vegetation cover on subsequent annual precipitation is simulated for the mid‐Holocene over North Africa using a fully coupled general circulation model with dynamic vegetation, FOAM‐LPJ (Fast Ocean Atmosphere Model‐Lund Potsdam Jena Model). By computing a vegetation feedback parameter based on lagged auto‐covariances, the simulated impact of North African vegetation on precipitation is statistically quantified. The feedback is also dynamically assessed through initial value ensemble experiments, in which North African grass cover is initially reduced and the climatic response analyzed. The statistical and dynamical assessments of the negative vegetation feedback agree in sign and relative magnitude for FOAM‐LPJ. The negative feedback on annual precipitation largely results from a competition between bare soil evaporation and plant transpiration, with increases in the former outweighing reductions in the latter given reduced grass cover. This negative feedback weakens and eventually reverses sign over time during a transient simulation from the mid‐Holocene to present. A similar, but weaker, negative feedback is identified in Community Climate System Model Version 2 (CCSM2) over North Africa for the mid‐Holocene.     

Moisture‐induced greening of the South Asia over the past three decades

South Asia experienced a weakening of summer monsoon circulation in the past several decades, resulting in rainfall decline in wet regions. In comparison with other tropical ecosystems, quantitative assessments of the extent and triggers of vegetation change are lacking in assessing climate‐change impacts over South Asia dominated by crops. Here, we use satellite‐based Normalized Difference Vegetation Index (NDVI) to quantify spatial–temporal changes in vegetation greenness, and find a widespread annual greening trend that stands in contrast to the weakening of summer monsoon circulation particularly over the last decade. We further show that moisture supply is the primary factor limiting vegetation activity during dry season or in dry region, and cloud cover or temperature would become increasingly important in wet region. Enhanced moisture conditions over dry region, coinciding with the decline in monsoon, are mainly responsible for the widespread greening trend. This result thereby cautions the use of a unified monsoon index to predict South Asia's vegetation dynamics. Current climate–carbon models in general correctly reproduce the dominant control of moisture in the temporal characteristics of vegetation productivity. But the model ensemble cannot exactly reproduce the spatial pattern of satellite‐based vegetation change mainly because of biases in climate simulations. The moisture‐induced greening over South Asia, which is likely to persist into the wetter future, has significant implications for regional carbon cycling and maintaining food security.     

Persistent versus transient tree encroachment of temperate peat bogs: effects of climate warming and drought events

Peatlands store approximately 30% of global soil carbon, most in moss‐dominated bogs. Future climatic changes, such as changes in precipitation patterns and warming, are expected to affect peat bog vegetation composition and thereby its long‐term carbon sequestration capacity. Theoretical work suggests that an episode of rapid environmental change is more likely to trigger transitions to alternative ecosystem states than a gradual, but equally large, change in conditions. We used a dynamic vegetation model to explore the impacts of drought events and increased temperature on vegetation composition of temperate peat bogs. We analyzed the consequences of six patterns of summer drought events combined with five temperature scenarios to test whether an open peat bog dominated by moss (Sphagnum) could shift to a tree‐dominated state. Unexpectedly, neither a gradual decrease in the amount of summer precipitation nor the occurrence of a number of extremely dry summers in a row could shift the moss‐dominated peat bog permanently into a tree‐dominated peat bog. The increase in tree biomass during drought events was unable to trigger positive feedbacks that keep the ecosystem in a tree‐dominated state after a return to previous ‘normal’ rainfall conditions. In contrast, temperature increases from 1 °C onward already shifted peat bogs into tree‐dominated ecosystems. In our simulations, drought events facilitated tree establishment, but temperature determined how much tree biomass could develop. Our results suggest that under current climatic conditions, peat bog vegetation is rather resilient to drought events, but very sensitive to temperature increases, indicating that future warming is likely to trigger persistent vegetation shifts.     

Dispersal strategies and spatial organization of vegetation in arid ecosystems

Seed dispersal and establishment are critical stages for plants in arid environments, where vegetation is spatially organized in patches with suitable and unsuitable sites for establishment. Theoretical studies suggest that the ability of vegetation to self‐organize in patchy spatial patterns is a critical property for plant survival in arid environments, and is a consequence of a scale‐dependent feedback between plants and resource availability. Field observations show that plants of arid environments evolved towards short dispersal distance (proxichory) and that the investment in reproduction increases along an aridity gradient. Here, we investigated how plant dispersal strategies affect spatial organization and associated scale‐dependent feedback in arid ecosystems. We addressed this research question using a model where the spatio‐temporal vegetation patterns were driven by scale‐dependent feedbacks between plants and soil water availability. In the model, water availability limited vegetation growth, seed production and establishment ability. Seed dispersal was modelled with an integrodifferential equation that mimicked important plant dispersal characteristics (i.e. fecundity, mean dispersal distance and establishment ability). Results showed that, when the investment in fecundity was relatively high, short seed dispersal helped maintaining higher mean biomass in the system, improving the vegetation efficiency in water use. However, higher fecundity induced a large cost, and high mean biomass could be sustained only with high establishment ability. Considering low establishment ability, intermediate fecundity was more efficient than low fecundity in maintaining high plant biomass under the most arid conditions. Consistently, plant dispersal strategies that maintained more biomass were related to a vegetation spatial organization that allowed the most efficient soil water redistribution, through the strengthening of the scale‐dependent feedback. The efficient dispersal strategies and spatial patterns in the model are commonly observed in plants of arid environments. Thus, dispersal strategies in arid environments might contribute to a favourable spatial organization and associated scale‐dependent feedback.     

Tailoring restoration interventions to the grassland‐savanna‐forest complex in central Brazil

Defining the reference system for restoration projects in regions characterized by complex vegetation mosaics is challenging. Here we use the Cerrado region of Brazil as an example of the importance of clearly defining multiple natural and anthropogenically altered states in grassland‐savanna‐forest mosaics. We define three main, natural vegetation types–grassland, savanna, and scleromorphic (cerradão) forest–to (1) distinguish between original and degraded states and (2) set appropriate targets for and guide restoration. We contend that the differences in Cerrado vegetation composition originally were driven by soil conditions and secondarily by fire frequency and precipitation patterns that differ from the core to the edge of the Cerrado region. Grasslands are found on the shallowest, least fertile soils and/or in waterlogged soils; scleromorphic forests are generally located on deeper, more fertile soils; and savannas occupy an intermediate position. In recent decades, this biophysical template has been overlain by a range of human land‐use intensities that strongly affect resilience, resulting in alternative anthropogenic states. For example, areas that were originally scleromorphic forest are likely to regenerate naturally following low‐ or medium‐intensity land use due to extensive resprouting of woody plants, whereas grassland restoration requires reintroduction of grass and forb species that do not tolerate soil disturbance and exotic grass competition. Planting trees into historic grasslands results in inappropriate restoration targets and often restoration failure. Correctly identifying original vegetation types is critical to most effectively allocate scarce restoration funding.     

Synergy between small- and large-scale feedbacks of vegetation on the water cycle

Predictions of the effects of climate change on the extent of forests, savannas and deserts are usually based on simple response models derived from actual vegetation distributions. In this review, we show two major problems with the implicitly assumed straightforward cause–effect relationship. Firstly, several studies suggest that vegetation itself may have considerable effects on regional climate implying a positive feedback, which can potentially lead to large‐scale hysteresis. Secondly, vegetation ecologists have found that effects of plants on microclimate and soils can cause a microscale positive feedback, implying that critical precipitation conditions for colonization of a site may differ from those for disappearance from that site. We argue that it is important to integrate these nonlinearities at disparate scales in models to produce more realistic predictions of potential effects of climate change and deforestation.     

Adopting a spatially explicit perspective to study the mysterious fairy circles of Namibia

The mysterious ‘fairy circles’ are vegetation‐free discs that cover vast areas along the pro‐Namib Desert. Despite 30 yr of research their origin remains unknown. Here we adopt a novel approach that focuses on analysis of the spatial patterns of fairy circles obtained from representative 25‐ha aerial images of north‐west Namibia. We use spatial point pattern analysis to quantify different features of their spatial structures and then critically inspect existing hypotheses with respect to their ability to generate the observed circle patterns. Our working hypothesis is that fairy circles are a self‐organized vegetation pattern. Finally, we test if an existing partial‐differential‐equation model, that was designed to describe vegetation pattern formation, is able to reproduce the characteristic features of the observed fairy circle patterns. The model is based on key‐processes in arid areas such as plant competition for water and local resource‐biomass feedbacks. The fairy circles showed at all three study areas the same regular spatial distribution patterns, characterized by Voronoi cells with mostly six corners, negative correlations in their size up to a distance of 13 m, and remarkable homogeneity over large spatial scales. These results cast doubts on abiotic gas‐leakage along geological lines or social insects as causal agents of their origin. However, our mathematical model was able to generate spatial patterns that agreed quantitatively in all of these features with the observed patterns. This supports the hypothesis that fairy circles are self‐organized vegetation patterns that emerge from positive biomass‐water feedbacks involving water transport by extended root systems and soil‐water diffusion. Future research should search for mechanisms that explain how the different hypotheses can generate the patterns observed here and test the ability of self‐organization to match the birth‐ and death dynamics of fairy circles and their regional patterns in the density and size with respect to environmental gradients.     

The global biogeography of semi‐arid periodic vegetation patterns

Aim Vegetation exhibiting landscape‐scale regular spatial patterns has been reported for arid and semi‐arid areas world‐wide. Recent theories state that such structures are bound to low‐productivity environments and result from a self‐organization process. Our objective was to test this relationship between periodic pattern occurrence and environmental factors at a global scale and to parametrize a predictive distribution model. Location Arid and semi‐arid areas world‐wide. Methods We trained an empirical predictive model (Maxent) for the occurrence of periodic vegetation patterns, based on environmental predictors and known occurrences verified on Landsat satellite images. Results This model allowed us to discover previously unreported pattern locations, and to report the first ever examples of spotted patterns in natural systems. Relationships to the main environmental drivers are discussed. Main conclusions These results confirm that periodic patterned vegetations are ubiquitous at the interface between arid and semi‐arid regions. Self‐organized patterning appears therefore to be a biome‐scale response to environmental conditions, including soil and topography. The set of correlations between vegetation patterns and their environmental conditions presented in this study will need to be reproduced in future modelling attempts.     

Modelling global terrestrial vegetation-climate interaction

By coupling an atmospheric general circulation model asynchronously with an equilibrium vegetation model, manifold equilibrium solutions of the atmosphere-biosphere system have been explored. It is found that under present-day conditions of the Earth''s orbital parameters and sea-surface temperatures, two stable equilibria of vegetation patterns are possible: one corresponding to present-day sparse vegetation in the Sahel, the second solution yielding savannah which extends far into the south-western part of the Sahara. A similar picture is obtained for conditions during the last glacial maximum (21 000 years before present (BP)). For the mid-Holocene (6000 years BP), however, the model finds only one solution: the green Sahara. We suggest that this intransitive behaviour of the atmosphere-biosphere is related to a westward shift of the Hadley-Walker circulation. A conceptual model of atmosphere-vegetation dynamics is used to interpret the bifurcation as well as its change in terms of stability theory.     

Tipping points in tropical tree cover: linking theory to data

It has recently been found that the frequency distribution of remotely sensed tree cover in the tropics has three distinct modes, which seem to correspond to forest, savanna, and treeless states. This pattern has been suggested to imply that these states represent alternative attractors, and that the response of these systems to climate change would be characterized by critical transitions and hysteresis. Here, we show how this inference is contingent upon mechanisms at play. We present a simple dynamical model that can generate three alternative tree cover states (forest, savanna, and a treeless state), based on known mechanisms, and use this model to simulate patterns of tree cover under different scenarios. We use these synthetic data to show that the hysteresis inferred from remotely sensed tree cover patterns will be inflated by spatial heterogeneity of environmental conditions. On the other hand, we show that the hysteresis inferred from satellite data may actually underestimate real hysteresis in response to climate change if there exists a positive feedback between regional tree cover and precipitation. Our results also indicate that such positive feedback between vegetation and climate should cause direct shifts between forest and a treeless state (rather than through an intermediate savanna state) to become more likely. Finally, we show how directionality of historical change in conditions may bias the observed relationship between tree cover and environmental conditions.     

Applying ecological site concepts and state‐and‐transition models to a grazed riparian rangeland

Ecological sites and state‐and‐transition models are useful tools for generating and testing hypotheses about drivers of vegetation composition in rangeland systems. These models have been widely implemented in upland rangelands, but comparatively, little attention has been given to developing ecological site concepts for rangeland riparian areas, and additional environmental criteria may be necessary to classify riparian ecological sites. Between 2013 and 2016, fifteen study reaches on five creeks were studied at Tejon Ranch in southern California. Data were collected to describe the relationship between riparian vegetation composition, environmental variables, and livestock management; and to explore the utility of ecological sites and state‐and‐transition models for describing riparian vegetation communities and for creating hypotheses about drivers of vegetation change. Hierarchical cluster analysis was used to classify the environmental and vegetation data (15 stream reaches × 4 years) into two ecological sites and eight community phases that comprised three vegetation states. Classification and regression tree (CART) analysis was used to determine the influence of abiotic site variables, annual precipitation, and cattle activity on vegetation clusters. Channel slope explained the greatest amount of variation in vegetation clusters; however, soil texture, geology, watershed size, and elevation were also selected as important predictors of vegetation composition. The classification tree built with this limited set of abiotic predictor variables explained 90% of the observed vegetation clusters. Cattle grazing and annual precipitation were not linked to qualitative differences in vegetation. Abiotic variables explained almost all of the observed riparian vegetation dynamics—and the divisions in the CART analysis corresponded roughly to the ecological sites—suggesting that ecological sites are well‐suited for understanding and predicting change in this highly variable system. These findings support continued development of riparian ecological site concepts and state‐and‐transition models to aid decision making for conservation and management of rangeland riparian areas.     

Firescape ecology: how topography determines the contrasting distribution of fire and rain forest in the south‐west of the Tasmanian Wilderness World Heritage Area

Aim To test the hypothesis that ‘islands’ of fire‐sensitive rain forest are restricted to topographic fire refugia and investigate the role of topography–fire interactions in fire‐mediated alternative stable state models. Location A vegetation mosaic of moorland, sclerophyll scrub, wet sclerophyll eucalypt forest and rain forest in the rugged, fire‐prone landscapes of south‐west Tasmania, Australia. Methods We used geospatial statistics to: (1) identify the topographic determinants of rain forest distribution on nutrient‐poor substrates, and (2) identify the vegetation and topographic variables that are important in controlling the spatial pattern of a series of very large fires (> 40,000 ha) that were mapped using Landsat Thematic Mapper (TM) satellite imagery. Results Rain forest was more likely to be found in valleys and on steep south‐facing slopes. Fires typically burned within highly flammable treeless moorland and stopped on boundaries with less flammable surrounding vegetation types such as wet sclerophyll forest and rain forest. Controlling for the effect of vegetation, fires were most likely to burn on flats, ridges and steep north‐facing slopes and least likely to burn in valleys and on steep south‐facing slopes. These results suggest an antagonism between fire and rain forest, in which rain forest preferentially occupies parts of the landscape where fire is least likely to burn. Main conclusions The distribution of rain forest on nutrient‐poor substrates was clearly related to parts of the landscape that are protected from fire (i.e. topographic fire refugia). The relative flammability of vegetation types at the landscape scale offers support to the proposed hierarchy of fire frequencies (moorland > scrub > wet sclerophyll > rain forest) that underpins the ecological models proposed for the region. The interaction between fire occurrence and a range of topographic variables suggests that topography plays an important role in mediating the fire–vegetation feedbacks thought to maintain vegetation mosaics in south‐west Tasmania. We suggest that these fire–topography interactions should be included in models of fire‐mediated alternative stable vegetation states in other fire‐prone landscapes.     

Grazing‐induced changes in plant–soil feedback alter plant biomass allocation

Large vertebrate herbivores, as well as plant–soil feedback interactions are important drivers of plant performance, plant community composition and vegetation dynamics in terrestrial ecosystems. However, it is poorly understood whether and how large vertebrate herbivores and plant–soil feedback effects interact. Here, we study the response of grassland plant species to grazing‐induced legacy effects in the soil and we explore whether these plant responses can help us to understand long‐term vegetation dynamics in the field. In a greenhouse experiment we tested the response of four grassland plant species, Agrostis capillaris, Festuca rubra, Holcus lanatus and Rumex acetosa, to field‐conditioned soils from grazed and ungrazed grassland. We relate these responses to long‐term vegetation data from a grassland exclosure experiment in the field. In the greenhouse experiment, we found that total biomass production and biomass allocation to roots was higher in soils from grazed than from ungrazed plots. There were only few relationships between plant production in the greenhouse and the abundance of conspecifics in the field. Spatiotemporal patterns in plant community composition were more stable in grazed than ungrazed grassland plots, but were not related to plant–soil feedbacks effects and biomass allocation patterns. We conclude that grazing‐induced soil legacy effects mainly influenced plant biomass allocation patterns, but could not explain altered vegetation dynamics in grazed grasslands. Consequently, the direct effects of grazing on plant community composition (e.g. through modifying light competition or differences in grazing tolerance) appear to overrule indirect effects through changes in plant–soil feedback.     

The role of vegetation–microclimate feedback in promoting shrub encroachment in the northern Chihuahuan desert

Many arid and semi‐arid landscapes around the world are affected by a shift from grassland to shrubland vegetation, presumably induced by climate warming, increasing atmospheric CO₂ concentrations, and/or changing land use. This major change in vegetation cover is likely sustained by positive feedbacks with the physical environment. Recent research has focused on a feedback with microclimate, whereby cold intolerant shrubs increase the minimum nocturnal temperatures in their surroundings. Despite the rich literature on the impact of land cover change on local climate conditions, changes in microclimate resulting from shrub expansion into desert grasslands have remained poorly investigated. It is unclear to what extent such a feedback can affect the maximum extent of shrub expansion and the configuration of a stable encroachment front. Here, we focus on the case of the northern Chihuahuan desert, where creosotebush (Larrea tridentata) has been replacing grasslands over the past 100–150 years. We use a process‐based coupled atmosphere‐vegetation model to investigate the role of this feedback in sustaining shrub encroachment in the region. Simulations indicate that the feedback allows juvenile shrubs to establish in the grassland during average years and, once established, reduce their vulnerability to freeze‐induced mortality by creating a warmer microclimate. Such a feedback is crucial in extreme cold winters as it may reduce shrub mortality. We identify the existence of a critical zone in the surroundings of the encroachment front, in which vegetation dynamics are bistable: in this zone, vegetation can be stable both as grassland and as shrubland. The existence of these alternative stable states explains why in most cases the shift from grass to shrub cover is found to be abrupt and often difficult to revert.     

Scale matters: fire–vegetation feedbacks are needed to explain tropical tree cover at the local scale

At a broad (regional to global) spatial scale, tropical vegetation is controlled by climate; at the local scale, it is believed to be determined by interactions between disturbance, vegetation and local conditions (soil and topography) through feedback processes. It has recently been suggested that strong fire–vegetation feedback processes may not be needed to explain tree‐cover patterns in tropical ecosystems and that climate–fire determinism is an alternative possibility. This conclusion was based on the fact that it is possible to reproduce observed patterns in tropical regions (e.g. a trimodal frequency distribution of tree cover) using a simple model that does not explicitly incorporate fire–vegetation feedback processes. We argue that these two mechanisms (feedbacks versus fire–climate control) operate at different spatial and temporal scales; it is not possible to evaluate the role of a process acting at fine scales (e.g. fire–vegetation feedbacks) using a model designed to reproduce regional‐scale pattern (scale mismatch). While the distributions of forest and savannas are partially determined by climate, many studies are providing evidence that the most parsimonious explanation for their environmental overlaps is the existence of feedback processes. Climate is unlikely to be an alternative to feedback processes; rather, climate and fire–vegetation feedbacks are complementary processes at different spatial and temporal scales.     

Localized states qualitatively change the response of ecosystems to varying conditions and local disturbances

The response of dynamical systems to varying conditions and disturbances is a fundamental aspect of their analysis. In spatially extended systems, particularly in pattern-forming systems, there are many possible responses, including critical transitions, gradual transitions and locally confined responses. Here, we use the context of vegetation dynamics in drylands in order to study the response of pattern-forming ecosystems to oscillating precipitation and local disturbances. We focus on two precipitation ranges, a bistability range of bare soil with a patterned vegetation state, and a bistability range of uniform vegetation with a patterned vegetation state. In these ranges, there are many different stable states, which allow for both abrupt and gradual transitions between the system states to occur. We find that large amplitude oscillations of the precipitation rate can lead to a collapse of the vegetation in one range, while in the other range, they result in the convergence to a patterned state with a preferred wavelength. In addition, we show that a series of local disturbances results in the collapse of the vegetation in one range, while it drives the system toward fluctuations around a finite average biomass in the other range. Moreover, it is shown that under certain conditions, local disturbances can actually increase the overall vegetation density. These significant differences in the system response are attributed to the existence of localized states in one of the bistability ranges.     

Gradual regime shifts in spatially extended ecosystems

Ecosystem regime shifts are regarded as abrupt global transitions from one stable state to an alternative stable state, induced by slow environmental changes or by global disturbances. Spatially extended ecosystems, however, can also respond to local disturbances by the formation of small domains of the alternative state. Such a response can lead to gradual regime shifts involving front propagation and the coalescence of alternative-state domains. When one of the states is spatially patterned, a multitude of intermediate stable states appears, giving rise to step-like gradual shifts with extended pauses at these states. Using a minimal model, we study gradual state transitions and show that they precede abrupt transitions. We propose indicators to probe gradual regime shifts, and suggest that a combination of abrupt-shift indicators and gradual-shift indicators might be needed to unambiguously identify regime shifts. Our results are particularly relevant to desertification in drylands where transitions to bare soil take place from spotted vegetation, and the degradation process appears to involve step-like events of local vegetation mortality caused by repeated droughts.