Alternative Ecosystem States Research Articles

Widespread forest-savanna coexistence but limited bistability at a landscape scale in Central Africa

Abstract Tropical forest and savanna frequently coexist under the same climatic conditions, which has led to the hypothesis that they could represent alternative ecosystem states, stabilized by internal feedbacks. An implication of this hypothesis is that forest and savanna may be bistable and exhibit tipping behavior in response to changing conditions. However, we pose that the local presence of forest and savanna within coexistence landscapes is not sufficient evidence that these are alternative stable states at larger ecosystem scales. Therefore, we explore forest-savanna coexistence and bistability at landscape scale in Central Africa. Using remote sensing data on tree cover, we classify 0.1° × 0.1° (approx. 10 × 10 km) landscapes as homogeneous forest, homogeneous savanna, or coexistence, and analyze the roles of climate, topography and soil sand content in driving their distributions. We find that local coexistence of forest and savanna within landscapes is common and occurs for the whole range of mean annual precipitation in our study area. At low precipitation, however, coexistence increases with topographic roughness and is therefore likely driven by local redistribution of resources rather than internal feedbacks. Coexistence within intermediate and high precipitation landscapes remains unexplained by the studied variables, and may be caused either by heterogeneity in unmeasured drivers or by feedback-driven bistability. At landscape scale, the precipitation ranges for which homogeneous forest and savanna occur have only limited overlap, and this overlap can be largely explained by other external drivers, such as seasonality, soil sand content, and topography. This lack of evidence that homogeneous forest and savanna in Central Africa are alternative ecosystem states at this landscape scale means that transitions between them may be mostly local, resulting in coexistence states. Therefore, we conclude that the likelihood of large-scale tipping between homogeneous forest and savanna ecosystems may be lower than previously thought.

Interactions between soil and other environmental variables modulate forest expansion and ecotone dynamics in humid savannas of Central Africa.

Forest expansion into savanna is a pervasive phenomenon in West and Central Africa, warranting comparative studies under diverse environmental conditions. We collected vegetation data from the woody and grassy components within 73 plots of 0.16 ha distributed along a successional gradient from humid savanna to forest in Central Africa. We associated spatially collocated edaphic parameters and fire frequency derived from remote sensing to investigate their combined influence on the vegetation. Soil texture was more influential in shaping savanna structure and species distribution than soil fertility, with clay-rich soils promoting higher grass productivity and fire frequency. Savanna featuring woody aboveground biomass surpassing 40 Mg ha-1 could escape the grass-fire feedback loop, by depressing grass biomass below 4 Mg ha-1. This thicker woody layer also favoured the establishment of fire-tolerant forest pioneers, which synergically contributed to the expansion of forests. Conversely, savannas below this fire suppression threshold sustained a balance between trees and grasses through the grass-fire feedback mechanism. This hysteresis loop, particularly pronounced on clayey soils, suggests that the contrast between grassy savanna and young forests might represent alternative ecosystem states, although savannas with low woody biomass remained vulnerable to forest edge encroachment.

Reassessing the alternative ecosystem states proposition in the African savanna-forest domain.

Ecologists are being challenged to predict how ecosystems will respond to climate changes. According to the Multi-Colored World (MCW) hypothesis, climate impacts may not manifest because consumers such as fire and herbivory can override the influence of climate on ecosystem state. One MCW interpretation is that climate determinism fails because alternative ecosystem states (AES) are possible at some locations in climate space. We evaluated theoretical and empirical evidence for the proposition that forest and savanna are AES in Africa. We found that maps which infer where AES zones are located were contradictory. Moreover, data from longitudinal and experimental studies provide inconclusive evidence for AES. That is, although the forest-savanna AES proposition is theoretically sound, the existing evidence is not yet convincing. We conclude by making the case that the AES proposition has such fundamental consequences for designing management actions to mitigate and adapt to climate change in the savanna-forest domain that it needs a more robust evidence base before it is used to prescribe management actions.

Respiration, phytoplankton size and the metabolic balance in the Atlantic gyres

The balance between plankton photosynthesis (GPP) and community respiration (CR) in the euphotic zone (net community production, NCP) is an essential driver of the biological carbon pump. Deficient datasets and a lack of knowledge of the mechanisms regulating CR cause poor empirical models and oversimplified parameterisations that maintain NCP as one of the most important unknowns for projections of the carbon pump. One important unresolved issue is the unexpected lack of empirical relationships between CR and the biomass or size-structure of the phytoplankton, which undermines the use of remotely sensed observations to predict net community metabolism. Here we analyse the spatial variation of plankton metabolism, chlorophyll a concentration (Chla), pico- and nanophytoplankton abundance and size-fractionated primary production (14CPP) along a latitudinal (49°N–46°S) transect of 73 stations across the Atlantic Ocean (AMT-22 cruise). The use of depth-weighted rates (rates integrated to the depth of 0.1% PAR, divided by the regionally varying depth of integration) markedly improved the depiction of latitudinal patterns and the significance of relationships, over volumetric or integrated rates. Depth-weighted CR showed clear and consistent latitudinal patterns with relevance for the distribution of NCP. Depth-weighted Chla and CR exhibited a significant relationship (CRZ=1.42ChlaZ-0.21, r2 = 0.69, N=37, p<0.001) with potential for the difficult prediction of CR. A general ratio of 1.42 mmolO2 mgChla-1 d-1 and a threshold Chla for net heterotrophy of ca. 0.25 mgChla m-3 can be tentatively proposed for the Atlantic, although further analyses of spatial and seasonal variation are necessary. We observed unusually positive NCP rates in the central part of the N gyre, due to a marked decrease of CR in a patch of high Synechococcus spp. abundance and high 14CPP by large phytoplankton. However, no relationship was observed between size-fractionated 14CPP and CR or the GPP : CR ratio during the cruise, contradicting the hypothesis that food web functioning is determined by the phytoplankton size structure. Such independence, together with the persistence of distinct GPP : CR and 14CPP : NCP relationships in distinct biogeographic provinces suggest a resilience of trophic dynamics and the existence of alternative ecosystem states, whose implications for projections of the metabolic state of the ocean are discussed.

Forest resilience and regeneration dynamics following wildfire disturbance

AbstractSpecies that are adapted to specific disturbance regimes may be sensitive to changes such as severity, frequency, and extent of ecosystem perturbation. Along range margins, these sensitivities may be exacerbated by concurrent changes in climate resulting in loss of resilience and shifts to alternative ecosystem states. Pinus banksiana (jack pine) is a serotinous species dependent on fire disturbance to open cones, release seed, reduce understory competition, and expose mineral soil. The Altona Flat Rock in northern New York is a globally rare sandstone pine barrens, at the southern edge of jack pine's range. In the summer of 2018, a wildfire burned approximately 200 ha of the Flat Rock barrens, providing the opportunity to study post‐disturbance jack pine and understory resilience. We aimed to establish if jack pine regeneration would occur, and if so, would it be influenced by fire severity, pre‐fire stand characteristics, and competition. We predicted that higher seedling densities would be associated with higher burn severity, higher pre‐fire jack pine stand density, and lesser understory competition. We collected seedling density and understory composition data annually from 2019 to 2022 across 45 plots spanning the disturbed and adjacent undisturbed barrens. In 2021 and 2022, seedling heights were also collected. Seedling densities following the disturbance ranged from 275,385 to 390,513 ha−1; = 357,821 ha−1. Jack pine seedling density was positively associated with fire severity and moss abundance for all years. Pre‐fire stand density along with ericaceous shrub, grass, and duff abundance were also positively associated with seedling density in some years. Seedling height was positively associated with fire severity along with grass and blueberry abundance. These factors, for both establishment/survival and growth, point to the importance of seed supply, microsite conditions, and facilitation in jack pine regeneration success. At this point, it does not appear that the interaction of disturbance with changing climate has shifted enough to prevent the recovery of this ecosystem to its pre‐disturbance jack pine‐dominated state. However, ongoing climate change may still have an impact on seedlings/saplings as this forest continues to mature.

Limited climatic space for alternative ecosystem states in Africa.

One of the foundational premises of ecology is that climate determines ecosystems. This has been challenged by alternative ecosystem state models, which illustrate that internal ecosystem dynamics acting on the initial ecosystem state can overwhelm the influence of climate, and by observations suggesting that climate cannot reliably discriminate forest and savanna ecosystem types. Using a novel phytoclimatic transform, which estimates the ability of climate to support different types of plants, we show that climatic suitability for evergreen trees and C4 grasses are sufficient to discriminate between forest and savanna in Africa. Our findings reassert the dominant influence of climate on ecosystems and suggest that the role of feedbacks causing alternative ecosystem states is less prevalent than has been suggested.

Can remotely sensed vegetation patterns signal dryland restoration success?

Active restoration is frequently implemented to restore degraded drylands globally, yet predicting the overall success of these restoration projects remains challenging. Here, we aim to explore if vegetation spatial patterns can be used to monitor ecosystem recovery and anticipate the success of restoration practices. We combined field surveys and high‐resolution drone images to analyze how vegetation spatial patterns change after the large‐scale straw checkerboards restoration projects in the sand dune systems of the Tengger Desert in northern China. We found that vegetation cover, plant species richness, and diversity increased rapidly, approaching the level of the naturally vegetated sand dunes, after 5 years of restoration. Soil fertility remained low despite the positive vegetation change. Along with the recovery process, we found a larger diversity of patch sizes (i.e. spatial variance) with an increasing proportion of large‐size patches. These patterns, combined with a constant positive recovery rate in vegetation cover, are consistent with theoretical predictions used to anticipate transitions to alternative ecosystem states. Although some indicators may be masked by the artificial planting scheme and community succession process during recovery, our results show that vegetation spatial patterns can be used to forecast and monitor the possible recovery of drylands.

Adaptive Evolution Can Both Prevent Ecosystem Collapse and Delay Ecosystem Recovery.

AbstractThere is growing concern about the dire socioecological consequences of abrupt transitions between alternative ecosystem states in response to environmental changes. At the same time, environmental change can trigger evolutionary responses that could stabilize or destabilize ecosystem dynamics. However, we know little about how coupled ecological and evolutionary processes affect the risk of transition between alternative ecosystem states. Using shallow lakes as a model ecosystem, we investigate how trait evolution of a key species affects ecosystem resilience under environmental stress. We find that adaptive evolution of macrophytes can increase ecosystem resilience by shifting the critical threshold, which marks the transition from a clear-water state to a turbid-water state to a higher level of environmental stress. However, following the transition, adaptation to the turbid-water state can delay the ecosystem recovery back to the clear-water state. This implies that restoration could be more effective when implemented early enough after a transition occurs and before organisms adapt to the alternative state. Our findings provide new insights into how to prevent and mitigate the occurrence of regime shifts in ecosystems and highlight the need to understand ecosystem responses to environmental change in the context of coupled ecological and evolutionary processes.

Transition from Fire-Dependent Open Forests: Alternative Ecosystem States in the Southeastern United States

Land use and fire exclusion have influenced ecosystems worldwide, resulting in alternative ecosystem states. Here, I provide two examples from the southeastern United States of fire-dependent open pine and pine-oak forest loss and examine dynamics of the replacement forests, given continued long-term declines in foundation longleaf (Pinus palustris) and shortleaf (Pinus echinata) pines and recent increases in commercial loblolly (Pinus taeda) and slash (Pinus elliottii var. elliottii) pines. Shortleaf pine-oak forest historically may have been dominant on about 32 to 38 million ha, a provisional estimate based on historical composition of 75% of all trees, and has decreased to about 2.5 million ha currently; shortleaf pine now is 3% of all trees in the northern province. Longleaf pine forest decreased from about 30 million ha, totaling 75% of all trees, to 1.3 million ha and 3% of all trees in contemporary forests of the southern province. The initial transition from open pine ecosystems to closed forests, primarily comprised of broadleaf species, was countered by conversion to loblolly and slash pine plantations. Loblolly pine now accounts for 37% of all trees. Loss of fire-dependent ecosystems and their foundation tree species affect associated biodiversity, or the species that succeed under fire disturbance.

Behavioral responses across a mosaic of ecosystem states restructure a sea otter–urchin trophic cascade

Consumer and predator foraging behavior can impart profound trait-mediated constraints on community regulation that scale up to influence the structure and stability of ecosystems. Here, we demonstrate how the behavioral response of an apex predator to changes in prey behavior and condition can dramatically alter the role and relative contribution of top-down forcing, depending on the spatial organization of ecosystem states. In 2014, a rapid and dramatic decline in the abundance of a mesopredator (Pycnopodia helianthoides) and primary producer (Macrocystis pyrifera) coincided with a fundamental change in purple sea urchin (Strongylocentrotus purpuratus) foraging behavior and condition, resulting in a spatial mosaic of kelp forests interspersed with patches of sea urchin barrens. We show that this mosaic of adjacent alternative ecosystem states led to an increase in the number of sea otters (Enhydra lutris nereis) specializing on urchin prey, a population-level increase in urchin consumption, and an increase in sea otter survivorship. We further show that the spatial distribution of sea otter foraging efforts for urchin prey was not directly linked to high prey density but rather was predicted by the distribution of energetically profitable prey. Therefore, we infer that spatially explicit sea otter foraging enhances the resistance of remnant forests to overgrazing but does not directly contribute to the resilience (recovery) of forests. These results highlight the role of consumer and predator trait-mediated responses to resource mosaics that are common throughout natural ecosystems and enhance understanding of reciprocal feedbacks between top-down and bottom-up forcing on the regional stability of ecosystems.

生态系统多稳态研究进展

在多稳态的生态系统中，外力可能导致生态系统状态突然之间发生不可逆转的转变，从而达到一个新的平衡状态。但目前对多稳态理论的系统研究很少，如何使用预警信号来预测生态系统的状态转变依旧是个难题。通过多稳态理论的梳理提出了一个更加综合的多稳态定义，并以放牧模型为例，系统总结了多稳态理论的相关概念，将多稳态理论应用在生态系统演替和扰沌理论的解释中；通过对生态系统稳态转换预警信号的原理、优缺点和应用条件的分析，对不同尺度下多稳态的研究方法进行了归纳；最后提出了目前多稳态领域的研究问题和未来的研究重点。结果表明：（1）将时间和空间预警信号结合在一起，并量化正确预警信号的概率，对错误预警信号的比例进行加权，可能会提供更准确的稳态转换的预报。（2）定量观测试验适用于小尺度的研究，而较大尺度的研究则采用简化的模型来模拟研究，选择正确的尺度极有可能改变预警信号的可靠性。（3）结合多稳态理论研究生态系统临界转换和反馈控制机制，并将基于性状的特征指标和进化动力学纳入其中，是生态系统修复实践的重要研究方向。（4）将多稳态相关理论和生态保护管理政策的实践相结合，是多稳态理论未来应用的前景。本研究为多稳态理论和实践的深入研究提供科学支撑。;In an alternative stable state (ASS) ecosystem, tipping points lead to abrupt and possibly irreversible shifts between alternative ecosystem states. However, there are only a few systematic studies on the ASS theory at present, and how to use a variety of early warning signals to predict the regime shift in the ecosystem at different scales is still a difficult problem. Besides, critical transitions are difficult to predict because the state of the system usually shows little change before the transition. In this review, firstly, a more integrated definition of ASS has been put forward through reorganizing of ASS theory from different previous literatures. Alternative stable states are defined:under the same external force or internal force driving, the structure, material composition and energy flow of the ecosystem will change into two or more different stable states, which are maintained by negative feedback loops. Then, the relevant concepts of ASS theory were systematically summarized by the model of the grazing animals on pasture land. Furthermore, ASS theory has been originally applied to interpret succession and panarchy theory in the ecosystem. Secondly, the principles, advantages, disadvantages, as well as application conditions of the early warning signal of regime shift in an ecosystem were analyzed, and the research methods of ASS theory at different ecological scales were summarized. The mathematical reason for critical slowing down leads to an increase in the autocorrelation and variance of the fluctuations in a stochastically forced system approaching a tipping point is illustrated by the autoregression model. Finally, the current research problems in the field of ASS and the focus of future research were put forward. The review results show that:(1) combining different early warning signals could provide a more accurate prediction of the tipping point by quantifying the probability of correct early warning signals and weighting the proportion of wrong early warning signals. (2) The quantitative experimental observation experiment is suitable for small-scale research, while larger-scale research usually uses a simplified model to simulate the results. Moreover, choosing the correct scale is very likely to improve the reliability of the early warning signal in predicting the regime shift. (3) The application of ASS theory would be important for ecosystem restoration by studying the critical transition and feedback control mechanism combining the trait-based characteristic indicators and evolutionary dynamics. (4) Applying ASS theory into the practice of ecological protection and management policy is very promising. This study provides scientific support for the further research of ASS theory and practice.

Condition thresholds in Australia’s threatened ecological community listings hinder conservation of dynamic ecosystems

Environmental degradation is threatening biodiversity and ecosystem function globally. Mandating ecosystem-level protection in policy and legislative frameworks is essential to prevent biodiversity loss. Australia’s Environment Protection and Biodiversity Conservation Act 1999 is the key legislative mechanism for supporting biodiversity at the national level, but has so far been ineffective at protecting habitat and ecological communities. Here we identify a major flaw in the current approach to listing threatened ecological communities (TECs): restrictive condition thresholds that threaten ecosystem function in dynamic ecosystems. Using two wetland TECs as a case study (Upland Wetlands and Coolibah-Black Box Woodlands), we argue that Australia’s environmental legislation should adopt a landscape-scale approach to TEC protection that acknowledges ecosystem function, accounts for different states in temporally dynamic systems, and sustains landscape connectivity of TEC distribution. We present a state-and-transition model for each TEC to show how human activities affect the reference-state continuum of wet and dry phases. We also show that the current listed condition thresholds do not acknowledge alternative ecosystem states and exclude areas that may be important for restoration and conservation of the TEC at the landscape-scale. Description of alternative and transitional states for dynamic systems, including how, when and why ecological communities shift between different states, should be formally integrated into the TEC listing process to protect Australia’s vulnerable ecosystems from further degradation and loss.

Vegetation Trajectories and Shortwave Radiative Forcing Following Boreal Forest Disturbance in Eastern Siberia

AbstractMajor boreal forest disturbance and associated carbon emissions have been reported in the coldest region of the Northern Hemisphere. Related biophysical feedbacks to climate remain highly uncertain but might reduce warming effects expected from carbon emissions. This study quantifies albedo change after disturbance, primarily fires, in larch‐dominated forests around Yakutsk as compared to undisturbed areas with natural albedo variability, using satellite‐based time series. The related annual mean shortwave radiative forcing was −6.015 W/m2 for the 13 years following forest disturbance. It was highly negative during snow‐covered months (−3.738 to −13.638 W/m2), but positive (+5.441 W/m2) for the summer months in the first year after disturbance, decreasing afterward and also turning into a negative forcing after 5 years. Forcing by surface shortwave radiation must be considered to assess the impact of boreal forest disturbance on climate and additional feedbacks, such as increased permafrost thaw or transition to alternative ecosystem states.

Planktonic stages of the ecologically important sea urchin, Diadema africanum: larval performance under near future ocean conditions

Abstract Diadema africanum is a recently described sea urchin from the Eastern Atlantic archipelagos, and adults play a major ecological role mediating the transition between two alternative ecosystem states: macroalgal beds and urchin barrens. The aim of this study was to describe for the first time the egg characteristics, fertilization and larval development. To determine basic life-history characteristics for this species, we reared larvae through to metamorphic competence under an energy shortage experiment and temperature–pH experiments to characterize the morphological plasticity of larval responses to actual and future oceanic conditions. D. africanum produces eggs that are larger both in diameter (82.7 μm) and volume (0.30 nl) than the eggs of both Diadema antillarum (70.0 μm, 0.18 nl) and Diadema mexicanum (68.0 μm, 0.16 nl). Larval development is similar to other species within the Family Diadematidae, with a Echinopluteus transversus larval type morphology. The combined effects of the climate change-related environmental factors resulted in a reduction in fitness of D. africanum at the warmer limit of its thermal range when combined with low pH. Results suggest that the egg and larval life-history characteristics of D. africanum may have evolved to facilitate long-distance oceanic transport; however, near-future oceanic conditions may compromise larval survival.

Interactive effects of foundation species on ecosystem functioning and stability in response to disturbance.

A major challenge in ecology is to understand determinants of ecosystem functioning and stability in the face of disturbance. Some important species can strongly shape community structure and ecosystem functioning, but their impacts and interactions on ecosystem-level responses to disturbance are less well known. Shallow ponds provide a model system in which to study the effects of such species because some taxa mitigate transitions between alternative ecosystem states caused by eutrophication. We performed pond experiments to test how two foundation species (a macrophyte and a mussel) affected the biomass of planktonic primary producers and its stability in response to nutrient additions. Individually, each species reduced phytoplankton biomass and tended to increase rates of recovery from disturbance, but together the species reversed these effects, particularly with larger nutrient additions. This reversal was mediated by high cyanobacterial dominance of the community and a resulting loss of trait evenness. Effects of the foundation species on primary producer biomass were associated with effects on other ecosystem properties, including turbidity and dissolved oxygen. Our work highlights the important role of foundation species and their interactive effects in determining responses of ecosystem functioning to disturbance.

Ecosystem tipping points in an evolving world.

There is growing concern over tipping points arising in ecosystems because of the crossing of environmental thresholds. Tipping points lead to abrupt and possibly irreversible shifts between alternative ecosystem states, potentially incurring high societal costs. Trait variation in populations is central to the biotic feedbacks that maintain alternative ecosystem states, as they govern the responses of populations to environmental change that could stabilize or destabilize ecosystem states. However, we know little about how evolutionary changes in trait distributions over time affect the occurrence of tipping points and even less about how big-scale ecological shifts reciprocally interact with trait dynamics. We argue that interactions between ecological and evolutionary processes should be taken into account in order to understand the balance of feedbacks governing tipping points in nature.

Forests expand as livestock pressure declines in subtropical South America

Bernardi, R. E., M. Buddeberg, M. Arim, and M. Holmgren. 2019. Forests expand as livestock pressure declines in subtropical South America. Ecology and Society 24(2):19. https://doi.org/10.5751/ES-10688-240219

Can novel pest outbreaks drive ecosystem transitions in northern‐boreal birch forest?

Abstract The boreal biome exhibits distinct alternative ecosystem states with high and low levels of tree cover. Insect outbreaks facilitated by climate warming could potentially drive transitions from high to low tree cover states. We investigated whether two key premises for such outbreak‐induced transitions—critical thresholds (tipping points) and positive feedbacks that could maintain alternative states—are present in the northern‐boreal mountain birch forest of Fennoscandia. Here, climate warming has promoted range expansions of defoliating geometrid moths, resulting in novel, severe multispecies outbreaks, most recently during 2002–2010. We conducted regional‐scale field surveys of forest damage and recovery in 280 mountain birch stands in a northeast Norway immediately after the outbreak (2010) and 6 years later (2016). Satellite‐derived time series of the Normalized Difference Vegetation Index (NDVI) provided an index of stand defoliation during the outbreak period. The proportion of dead stems per stand displayed a bimodal distribution, with stands generally being either lightly or severely damaged. This was due to a critical threshold in the relationship between defoliation and stem mortality, with mortality rates increasing abruptly in stands experiencing a mean drop in NDVI of more than 4% during the outbreak. The two key forest regenerative pathways—basal sprouting and sapling production—both displayed positive feedbacks with surviving stems and trees, so that regeneration success declined with increasing damage to the mature tree layer. These feedbacks imply that stands which have been forced across critical defoliation thresholds and suffered collapses of living tree cover may struggle to recover, especially if the loss of positive regenerative feedbacks is compounded by ungulate browsing on birch recruits. Synthesis. The north Fennoscandian mountain birch forest displays critical thresholds and positive feedbacks that conform to theoretical expectations for a system that could be vulnerable to abrupt and persistent changes of state in the face of novel, climatically facilitated insect outbreaks. These findings deepen the understanding of the persistent losses of tree cover that have occasionally been observed after outbreaks in this system in the past, and add to the list of mechanisms that could help explain the bistability of tree cover across the boreal biome.

Consequences of grazer‐induced vegetation transitions on ecosystem carbon storage in the tundra

Abstract Large herbivores can control plant community composition and, under certain conditions, even induce vegetation shifts to alternative ecosystem states. As different plant assemblages maintain contrasting carbon (C) cycling patterns, herbivores have the potential to alter C sequestration at regional scales. Their influence is of particular interest in the Arctic tundra, where a large share of the world's soil C reservoir is stored. We assessed the influence of grazing mammals on tundra vegetation and C stocks by resampling two sites located along pasture rotation fences in northern Norway. These fences have separated lightly grazed areas from heavily grazed areas (in close proximity to the fences) and moderately grazed areas (further away from the fences) for the past 50 years. Fourteen years earlier, the lightly and moderately grazed areas were dominated by dwarf shrubs, whereas heavy grazing had promoted the establishment of graminoid‐dominated vegetation. Since then, both reindeer densities and temperatures have increased, and more time has passed for transient dynamics to be expressed. We expected that the vegetation and C stocks would have changed under all grazing intensities, but not necessarily in the same way. At the site where relative reindeer numbers and trampling intensity had increased the most, graminoid‐dominated vegetation was now also found in the moderately grazed area. At the other site, the dominant vegetation types under all grazing intensities were the same as 14 years earlier. We show that the heavily grazed, graminoid‐dominated areas stored less C above‐ground than the lightly grazed, shrub‐dominated areas. Yet, the below‐ground consequences of grazing‐induced grassification varied between the sites: Grazing did not alter organic soil C stocks at the site where both evergreen and deciduous shrubs were abundant in the lightly grazed area, whereas heavy grazing increased organic soil C stocks at the site where the deciduous shrub Betula nana was dominant. Our results indicate that, despite the negative impacts of grazers on above‐ground C storage, their impact on below‐ground C may even be positive. We suggest that the site‐specific responses of organic soil C stocks to grazing could be explained by the differences in vegetation under light grazing. This would imply that the replacement of deciduous shrubs by graminoids, as a consequence of grazing could be beneficial for C sequestration in tundra soils. A plain language summary is available for this article.

Ecosystem dynamics and management after forest die‐off: a global synthesis with conceptual state‐and‐transition models

AbstractBroad‐scale forest die‐off associated with drought and heat has now been reported from every forested continent, posing a global‐scale challenge to forest management. Climate‐driven die‐off is frequently compounded with other drivers of tree mortality, such as altered land use, wildfire, and invasive species, making forest management increasingly complex. Facing similar challenges, rangeland managers have widely adopted the approach of developing conceptual models that identify key ecosystem states and major types of transitions between those states, known as “state‐and‐transition models” (S&T models). Using expert opinion and available research, the development of such conceptual S&T models has proven useful in anticipating ecosystem changes and identifying management actions to undertake or to avoid. In cases where detailed data are available, S&T models can be developed into probabilistic predictions, but even where data are insufficient to predict transition probabilities, conceptual S&T models can provide valuable insights for managing a given ecosystem and for comparing and contrasting different ecosystem dynamics. We assembled a synthesis of 14 forest die‐off case studies from around the globe, each with sufficient information to infer impacts on forest dynamics and to inform management options following a forest die‐off event. For each, we developed a conceptual S&T model to identify alternative ecosystem states, pathways of ecosystem change, and points where management interventions have been, or may be, successful in arresting or reversing undesirable changes. We found that our diverse set of mortality case studies fit into three broad classes of ecosystem trajectories: (1) single‐state transition shifts, (2) ecological cascading responses and feedbacks, and (3) complex dynamics where multiple interactions, mortality drivers, and impacts create a range of possible state transition responses. We integrate monitoring and management goals in a framework aimed to facilitate development of conceptual S&T models for other forest die‐off events. Our results highlight that although forest die‐off events across the globe encompass many different underlying drivers and pathways of ecosystem change, there are commonalities in opportunities for successful management intervention.