Extreme climatic events and vegetation: the role of stabilizing processes

Current climatic trends involve both increasing temperatures and climatic variability, with extreme events becoming more frequent. Increasing concern on extreme climatic events has triggered research on vegetation shifts. However, evidences of vegetation shifts resulting from these events are still relatively rare. Empirical evidence supports the existence of stabilizing processes minimizing and counteracting the effects of these events, reinforcing community resilience. We propose a demographic framework to understand this inertia to change based on the balance between adult mortality induced by the event and enhanced recruitment or adult survival after the event. The stabilizing processes potentially contributing to this compensation include attenuation of the adult mortality caused by the event, due to site quality variability, to tolerance, phenotypic variability, and plasticity at population level, and to facilitative interactions. Mortality compensation may also occur by increasing future survival due to beneficial effect on growth and survival of the new conditions derived from global warming and increased climatic variability, to lowered competition resulting from reduced density in affected stands, or to antagonistic release when pathogens or predators are vulnerable to the event or the ongoing climatic conditions. Finally, mortality compensation may appear by enhanced recruitment due to release of competition with established vegetation, for instance as a consequence of gap openings after event‐caused mortality, or to the new conditions, which may be more favorable for seedling establishment, or to enhanced mutualistic interactions (pollination, dispersal). There are important challenges imposed by the need of long‐term studies, but a research agenda focused on potentially stabilizing processes is well suited to understand the variety of responses, including lack of sudden changes and community inertia that are frequently observed in vegetation under extreme events. This understanding is crucial for the establishment of sound management strategies and actions addressed to improve ecosystem resilience under climate change scenarios.     

Climate variability and vulnerability to climate change: a review

The focus of the great majority of climate change impact studies is on changes in mean climate. In terms of climate model output, these changes are more robust than changes in climate variability. By concentrating on changes in climate means, the full impacts of climate change on biological and human systems are probably being seriously underestimated. Here, we briefly review the possible impacts of changes in climate variability and the frequency of extreme events on biological and food systems, with a focus on the developing world. We present new analysis that tentatively links increases in climate variability with increasing food insecurity in the future. We consider the ways in which people deal with climate variability and extremes and how they may adapt in the future. Key knowledge and data gaps are highlighted. These include the timing and interactions of different climatic stresses on plant growth and development, particularly at higher temperatures, and the impacts on crops, livestock and farming systems of changes in climate variability and extreme events on pest‐weed‐disease complexes. We highlight the need to reframe research questions in such a way that they can provide decision makers throughout the food system with actionable answers, and the need for investment in climate and environmental monitoring. Improved understanding of the full range of impacts of climate change on biological and food systems is a critical step in being able to address effectively the effects of climate variability and extreme events on human vulnerability and food security, particularly in agriculturally based developing countries facing the challenge of having to feed rapidly growing populations in the coming decades.     

Ecosystem feedbacks constrain the effect of day-to-day weather variability on land–atmosphere carbon exchange

Whole-ecosystem interactions and feedbacks constrain ecosystem responses to environmental change. The effects of these constraints on responses to climate trends and extreme weather events have been well studied. Here we examine how these constraints respond to changes in day-to-day weather variability without changing the long-term mean weather. Although environmental variability is recognized as a critical factor affecting ecological function, the effects of climate change on day-to-day weather variability and the resultant impacts on ecosystem function are still poorly understood. Changes in weather variability can alter the mean rates of individual ecological processes because many processes respond non-linearly to environmental drivers. We assessed how these individual-process responses to changes in day-to-day weather variability interact with one another at an ecosystem level. We examine responses of arctic tundra to changes in weather variability using stochastic simulations of daily temperature, precipitation, and light to drive a biogeochemical model. Changes in weather variability altered ecosystem carbon, nitrogen, and phosphorus stocks and cycling rates in our model. However, responses of some processes (e.g., respiration) were inconsistent with expectations because ecosystem feedbacks can moderate, or even reverse, direct process responses to weather variability. More weather variability led to greater carbon losses from land to atmosphere; less variability led to higher carbon sequestration on land. The magnitude of modeled ecosystem response to weather variability was comparable to that predicted for the effects of climate mean trends by the end of the century.     

Ecological and evolutionary impacts of changing climatic variability

While average temperature is likely to increase in most locations on Earth, many places will simultaneously experience higher variability in temperature, precipitation, and other climate variables. Although ecologists and evolutionary biologists widely recognize the potential impacts of changes in average climatic conditions, relatively little attention has been paid to the potential impacts of changes in climatic variability and extremes. We review the evidence on the impacts of increased climatic variability and extremes on physiological, ecological and evolutionary processes at multiple levels of biological organization, from individuals to populations and communities. Our review indicates that climatic variability can have profound influences on biological processes at multiple scales of organization. Responses to increased climatic variability and extremes are likely to be complex and cannot always be generalized, although our conceptual and methodological toolboxes allow us to make informed predictions about the likely consequences of such climatic changes. We conclude that climatic variability represents an important component of climate that deserves further attention.     

Effects of climate extremes on the terrestrial carbon cycle: concepts,processes and potential future impacts

Extreme droughts, heat waves, frosts, precipitation, wind storms and other climate extremes may impact the structure, composition and functioning of terrestrial ecosystems, and thus carbon cycling and its feedbacks to the climate system. Yet, the interconnected avenues through which climate extremes drive ecological and physiological processes and alter the carbon balance are poorly understood. Here, we review the literature on carbon cycle relevant responses of ecosystems to extreme climatic events. Given that impacts of climate extremes are considered disturbances, we assume the respective general disturbance‐induced mechanisms and processes to also operate in an extreme context. The paucity of well‐defined studies currently renders a quantitative meta‐analysis impossible, but permits us to develop a deductive framework for identifying the main mechanisms (and coupling thereof) through which climate extremes may act on the carbon cycle. We find that ecosystem responses can exceed the duration of the climate impacts via lagged effects on the carbon cycle. The expected regional impacts of future climate extremes will depend on changes in the probability and severity of their occurrence, on the compound effects and timing of different climate extremes, and on the vulnerability of each land‐cover type modulated by management. Although processes and sensitivities differ among biomes, based on expert opinion, we expect forests to exhibit the largest net effect of extremes due to their large carbon pools and fluxes, potentially large indirect and lagged impacts, and long recovery time to regain previous stocks. At the global scale, we presume that droughts have the strongest and most widespread effects on terrestrial carbon cycling. Comparing impacts of climate extremes identified via remote sensing vs. ground‐based observational case studies reveals that many regions in the (sub‐)tropics are understudied. Hence, regional investigations are needed to allow a global upscaling of the impacts of climate extremes on global carbon–climate feedbacks.     

Moving forward in global‐change ecology: capitalizing on natural variability

Natural resources managers are being asked to follow practices that accommodate for the impact of climate change on the ecosystems they manage, while global‐ecosystems modelers aim to forecast future responses under different climate scenarios. However, the lack of scientific knowledge about short‐term ecosystem responses to climate change has made it difficult to define set conservation practices or to realistically inform ecosystem models. Until recently, the main goal for ecologists was to study the composition and structure of communities and their implications for ecosystem function, but due to the probable magnitude and irreversibility of climate‐change effects (species extinctions and loss of ecosystem function), a shorter term focus on responses of ecosystems to climate change is needed. We highlight several underutilized approaches for studying the ecological consequences of climate change that capitalize on the natural variability of the climate system at different temporal and spatial scales. For example, studying organismal responses to extreme climatic events can inform about the resilience of populations to global warming and contribute to the assessment of local extinctions. Translocation experiments and gene expression are particular useful to quantitate a species' acclimation potential to global warming. And studies along environmental gradients can guide habitat restoration and protection programs by identifying vulnerable species and sites. These approaches identify the processes and mechanisms underlying species acclimation to changing conditions, combine different analytical approaches, and can be used to improve forecasts of the short‐term impacts of climate change and thus inform conservation practices and ecosystem models in a meaningful way.     

Means and extremes: building variability into community‐level climate change experiments

Experimental studies assessing climatic effects on ecological communities have typically applied static warming treatments. Although these studies have been informative, they have usually failed to incorporate either current or predicted future, patterns of variability. Future climates are likely to include extreme events which have greater impacts on ecological systems than changes in means alone. Here, we review the studies which have used experiments to assess impacts of temperature on marine, freshwater and terrestrial communities, and classify them into a set of ‘generations’ based on how they incorporate variability. The majority of studies have failed to incorporate extreme events. In terrestrial ecosystems in particular, experimental treatments have reduced temperature variability, when most climate models predict increased variability. Marine studies have tended to not concentrate on changes in variability, likely in part because the thermal mass of oceans will moderate variation. In freshwaters, climate change experiments have a much shorter history than in the other ecosystems, and have tended to take a relatively simple approach. We propose a new ‘generation’ of climate change experiments using down‐scaled climate models which incorporate predicted changes in climatic variability, and describe a process for generating data which can be applied as experimental climate change treatments.     

森林衰退与全球气候变化

工业革命以来的人类活动,改变着大气的化学组成,从而改变着全球的气候,气候变率影响着森林的生长,甚至能导致森林衰退,全球气候变化引起森林衰退的机近几年制可能有几种,全球变暖,降水模型的改变以及蒸散作用的提高,将给一些森林带来高温胁迫和水分胁迫,危害植物的生理过程,ＣＯ２的施肥作用及其与气温升高的协同效应,将促进植物的新陈代谢,加速树木的成熟和衰老,在全球气候变化条件下,植物种竞争和分异将加强,气候变     

Phenotypic plasticity increases exposure to extreme climatic events that reduce individual fitness

Climate models, and empirical observations, suggest that anthropogenic climate change is leading to changes in the occurrence and severity of extreme climatic events (ECEs). Effects of changes in mean climate on phenology, movement, and demography in animal and plant populations are well documented. In contrast, work exploring the impacts of ECEs on natural populations is less common, at least partially due to the challenges of obtaining sufficient data to study such rare events. Here, we assess the effect of changes in ECE patterns in a long-term study of great tits, near Oxford, over a 56-year period between 1965 and 2020. We document marked changes in the frequency of temperature ECEs, with cold ECEs being twice as frequent in the 1960s than at present, and hot ECEs being ~three times more frequent between 2010 and 2020 than in the 1960s. While the effect of single ECEs was generally quite small, we show that increased exposure to ECEs often reduces reproductive output, and that in some cases the effect of different types of ECE is synergistic. We further show that long-term temporal changes in phenology, resulting from phenotypic plasticity, lead to an elevated risk of exposure to low temperature ECEs early in reproduction, and hence suggest that changes in ECE exposure may act as a cost of plasticity. Overall, our analyses reveal a complex set of risks of exposure and effects as ECE patterns change and highlight the importance of considering responses to changes in both mean climate and extreme events. Patterns in exposure and effects of ECEs on natural populations remain underexplored and continued work will be vital to establish the impacts of ECEs on populations in a changing climate.     

Riparian Ecosystems in the 21st Century: Hotspots for Climate Change Adaptation?

Riparian ecosystems in the 21st century are likely to play a critical role in determining the vulnerability of natural and human systems to climate change, and in influencing the capacity of these systems to adapt. Some authors have suggested that riparian ecosystems are particularly vulnerable to climate change impacts due to their high levels of exposure and sensitivity to climatic stimuli, and their history of degradation. Others have highlighted the probable resilience of riparian ecosystems to climate change as a result of their evolution under high levels of climatic and environmental variability. We synthesize current knowledge of the vulnerability of riparian ecosystems to climate change by assessing the potential exposure, sensitivity, and adaptive capacity of their key components and processes, as well as ecosystem functions, goods and services, to projected global climatic changes. We review key pathways for ecological and human adaptation for the maintenance, restoration and enhancement of riparian ecosystem functions, goods and services and present emerging principles for planned adaptation. Our synthesis suggests that, in the absence of adaptation, riparian ecosystems are likely to be highly vulnerable to climate change impacts. However, given the critical role of riparian ecosystem functions in landscapes, as well as the strong links between riparian ecosystems and human well-being, considerable means, motives and opportunities for strategically planned adaptation to climate change also exist. The need for planned adaptation of and for riparian ecosystems is likely to be strengthened as the importance of many riparian ecosystem functions, goods and services will grow under a changing climate. Consequently, riparian ecosystems are likely to become adaptation ‘hotspots’ as the century unfolds.     

Impacts of climate change on avian populations

This review focuses on the impacts of climate change on population dynamics. I introduce the MUP (Measuring, Understanding, and Predicting) approach, which provides a general framework where an enhanced understanding of climate‐population processes, along with improved long‐term data, are merged into coherent projections of future population responses to climate change. This approach can be applied to any species, but this review illustrates its benefit using birds as examples. Birds are one of the best‐studied groups and a large number of studies have detected climate impacts on vital rates (i.e., life history traits, such as survival, maturation, or breeding, affecting changes in population size and composition) and population abundance. These studies reveal multifaceted effects of climate with direct, indirect, time‐lagged, and nonlinear effects. However, few studies integrate these effects into a climate‐dependent population model to understand the respective role of climate variables and their components (mean state, variability, extreme) on population dynamics. To quantify how populations cope with climate change impacts, I introduce a new universal variable: the ‘population robustness to climate change.’ The comparison of such robustness, along with prospective and retrospective analysis may help to identify the major climate threats and characteristics of threatened avian species. Finally, studies projecting avian population responses to future climate change predicted by IPCC‐class climate models are rare. Population projections hinge on selecting a multiclimate model ensemble at the appropriate temporal and spatial scales and integrating both radiative forcing and internal variability in climate with fully specified uncertainties in both demographic and climate processes.     

Climate warming and heat waves affect reproductive strategies and interactions between submerged macrophytes

Extreme climatic events, such as heat waves, are predicted to increase in frequency and intensity during the next hundred years, which may accelerate shifts in hydrological regimes and submerged macrophyte composition in freshwater ecosystems. Since macrophytes are profound components of aquatic systems, predicting their response to extreme climatic events is crucial for implementation of climate change adaptation strategies. We therefore performed an experiment in 24 outdoor enclosures (400 L) separating the impact of a 4 °C increase in mean temperature with the same increase, that is the same total amount of energy input, but resembling a climate scenario with extreme variability, oscillating between 0 °C and 8 °C above present conditions. We show that at the moderate nutrient conditions provided in our study, neither an increase in mean temperature nor heat waves lead to a shift from a plant‐dominated to an algal‐dominated system. Instead, we show that species‐specific responses to climate change among submerged macrophytes may critically influence species composition and thereby ecosystem functioning. Our results also imply that more fluctuating temperatures affect the number of flowers produced per plant leading to less sexual reproduction. Our findings therefore suggest that predicted alterations in climate regimes may influence both plant interactions and reproductive strategies, which have the potential to inflict changes in biodiversity, community structure and ecosystem functioning.     

Can life-history traits predict the response of forb populations to changes in climate variability?

1.  Climate change will cause changes in average temperature and precipitation as well as increased fluctuations around the mean, yet few studies have considered the impacts of altered climate variability on plant populations. We tested whether life-history traits (expected life span, generation time and seed size) can predict plant responses to increased environmental variability across similar plant species sharing the same habitat.
2.  We combined long-term demographic data on 10 prairie forb species with stochastic demography techniques to estimate the effects of potential changes in matrix element means and variances on the long-term stochastic population growth rate.
3.  For all 10 species, recruitment had higher contribution and elasticity values than survival, meaning that climate change is more likely to influence population growth through effects on recruitment than on survival for these relatively short-lived forbs. Species with longer generation times had lower elasticities to increases in matrix element variability.
4.   Synthesis. Our analysis of a unique, long-term data set suggests that longer-lived plant species will be less vulnerable to the effects of future increases in climate variability. While this relationship was previously reported for diverse taxa from many locations, our results show that it also applies within a guild of short-lived species from a single community. The generality of the pattern demonstrates the potential for using life-history traits to make predictions about which species may be the most vulnerable to climate change.     

Plants and climate change: complexities and surprises

Background Anthropogenic climate change (ACC) will influence all aspects of plant biology over coming decades. Many changes in wild species have already been well-documented as a result of increased atmospheric CO₂ concentrations, warming climate and changing precipitation regimes. A wealth of available data has allowed the use of meta-analyses to examine plant–climate interactions on more sophisticated levels than before. These analyses have revealed major differences in plant response among groups, e.g. with respect to functional traits, taxonomy, life-history and provenance. Interestingly, these meta-analyses have also exposed unexpected mismatches between theory, experimental, and observational studies.Scope We reviewed the literature on species’ responses to ACC, finding ∼42 % of 4000 species studied globally are plants (primarily terrestrial). We review impacts on phenology, distributions, ecophysiology, regeneration biology, plant–plant and plant–herbivore interactions, and the roles of plasticity and evolution. We focused on apparent deviations from expectation, and highlighted cases where more sophisticated analyses revealed that unexpected changes were, in fact, responses to ACC.Conclusions We found that conventionally expected responses are generally well-understood, and that it is the aberrant responses that are now yielding greater insight into current and possible future impacts of ACC. We argue that inconclusive, unexpected, or counter-intuitive results should be embraced in order to understand apparent disconnects between theory, prediction, and observation. We highlight prime examples from the collection of papers in this Special Issue, as well as general literature. We found use of plant functional groupings/traits had mixed success, but that some underutilized approaches, such as Grime''s C/S/R strategies, when incorporated, have improved understanding of observed responses. Despite inherent difficulties, we highlight the need for ecologists to conduct community-level experiments in systems that replicate multiple aspects of ACC. Specifically, we call for development of coordinating experiments across networks of field sites, both natural and man-made.     

Running to stand still: adaptation and the response of plants to rapid climate change

Climate is a potent selective force in natural populations, yet the importance of adaptation in the response of plant species to past climate change has been questioned. As many species are unlikely to migrate fast enough to track the rapidly changing climate of the future, adaptation must play an increasingly important role in their response. In this paper we review recent work that has documented climate‐related genetic diversity within populations or on the microgeographical scale. We then describe studies that have looked at the potential evolutionary responses of plant populations to future climate change. We argue that in fragmented landscapes, rapid climate change has the potential to overwhelm the capacity for adaptation in many plant populations and dramatically alter their genetic composition. The consequences are likely to include unpredictable changes in the presence and abundance of species within communities and a reduction in their ability to resist and recover from further environmental perturbations, such as pest and disease outbreaks and extreme climatic events. Overall, a range‐wide increase in extinction risk is likely to result. We call for further research into understanding the causes and consequences of the maintenance and loss of climate‐related genetic diversity within populations.     

Climate change accelerates local disease extinction rates in a long‐term wild host–pathogen association

Pathogens are a significant component of all plant communities. In recent years, the potential for existing and emerging pathogens of agricultural crops to cause increased yield losses as a consequence of changing climatic patterns has raised considerable concern. In contrast, the response of naturally occurring, endemic pathogens to a warming climate has received little attention. Here, we report on the impact of a signature variable of global climate change – increasing temperature – on the long‐term epidemiology of a natural host–pathogen association involving the rust pathogen Triphragmium ulmariae and its host plant Filipendula ulmaria. In a host–pathogen metapopulation involving approximately 230 host populations growing on an archipelago of islands in the Gulf of Bothnia we assessed changes in host population size and pathogen epidemiological measures over a 25‐year period. We show how the incidence of disease and its severity declines over that period and most importantly demonstrate a positive association between a long‐term trend of increasing extinction rates in individual pathogen populations of the metapopulation and increasing temperature. Our results are highly suggestive that changing climatic patterns, particularly mean monthly growing season (April‐November) temperature, are markedly influencing the epidemiology of plant disease in this host–pathogen association. Given the important role plant pathogens have in shaping the structure of communities, changes in the epidemiology of pathogens have potentially far‐reaching impacts on ecological and evolutionary processes. For these reasons, it is essential to increase understanding of pathogen epidemiology, its response to warming, and to invoke these responses in forecasts for the future.     

Stochastic resonance in climate reddening increases the risk of cyclic ecosystem extinction via phase-tipping

Human activity is leading to changes in the mean and variability of climatic parameters in most locations around the world. The changing mean has received considerable attention from scientists and climate policy makers. However, recent work indicates that the changing variability, that is, the amplitude and the temporal autocorrelation of deviations from the mean, may have greater and more imminent impact on ecosystems. In this paper, we demonstrate that changes in climate variability alone could drive cyclic predator–prey ecosystems to extinction via so-called phase-tipping (P-tipping), a new type of instability that occurs only from certain phases of the predator–prey cycle. We construct a mathematical model of a variable climate and couple it to two self-oscillating paradigmatic predator–prey models. Most importantly, we combine realistic parameter values for the Canada lynx and snowshoe hare with actual climate data from the boreal forest. In this way, we demonstrate that critically important species in the boreal forest have increased likelihood of P-tipping to extinction under predicted changes in climate variability, and are most vulnerable during stages of the cycle when the predator population is near its maximum. Furthermore, our analysis reveals that stochastic resonance is the underlying mechanism for the increased likelihood of P-tipping to extinction.     

Mean conditions predict salt marsh plant community diversity and stability better than environmental variability

Environmental variability and the frequency of extreme events are predicted to increase in future climate scenarios; however, the role of fluctuations in shaping community composition, diversity and stability is not well understood. Identifying current patterns of association between measures of community stability and climatic means and variability will help elucidate the ways in which altered variability and mean conditions may change communities in the future. Salt marshes provide essential ecosystem services and are increasingly threatened by sea‐level rise, land‐use change, eutrophication and predator loss, yet the effects of temporal environmental variation on salt marshes remain unknown. We synthesized long‐term plant community monitoring data from 11 sites on both coasts of the United States. We used an information‐theoretic approach and linear models to determine the associations among long‐term mean conditions, interannual environmental variability, and plant community stability and diversity. We found that salt marsh community stability and diversity were more strongly related to long‐term means of temperature and precipitation than to interannual variation. Warm and wet environments had fewer species and less turnover among years. Our results suggest that communities in cool, dry environments may be more resilient to climate warming due to greater species richness and turnover. Mean conditions are sufficient to predict contemporary patterns of salt marsh plant community dynamics, but environmental variability may have stronger impacts as it increases with climate change.     

Significant Mean and Extreme Climate Sensitivity of Norway Spruce and Silver Fir at Mid-Elevation Mesic Sites in the Alps

Climate forcing is the major abiotic driver for forest ecosystem functioning and thus significantly affects the role of forests within the global carbon cycle and related ecosystem services. Annual radial increments of trees are probably the most valuable source of information to link tree growth and climate at long-term time scales, and have been used in a wide variety of investigations worldwide. However, especially in mountainous areas, tree-ring studies have focused on extreme environments where the climate sensitivity is perhaps greatest but are necessarily a biased representation of the forests within a region. We used tree-ring analyses to study two of the most important tree species growing in the Alps: Norway spruce (Picea abies) and silver fir (Abies alba). We developed tree-ring chronologies from 13 mesic mid-elevation sites (203 trees) and then compared them to monthly temperature and precipitation data for the period 1846–1995. Correlation functions, principal component analysis and fuzzy C-means clustering were applied to 1) assess the climate/growth relationships and their stationarity and consistency over time, and 2) extract common modes of variability in the species responses to mean and extreme climate variability. Our results highlight a clear, time-stable, and species-specific response to mean climate conditions. However, during the previous-year''s growing season, which shows the strongest correlations, the primary difference between species is in their response to extreme events, not mean conditions. Mesic sites at mid-altitude are commonly underrepresented in tree-ring research; we showed that strong climatic controls of growth may exist even in those areas. Extreme climatic events may play a key role in defining the species-specific responses on climatic sensitivity and, with a global change perspective, specific divergent responses are likely to occur even where current conditions are less limited.     

Integrating mechanistic and empirical model projections to assess climate impacts on tree species distributions in northwestern North America

Empirical and mechanistic models have both been used to assess the potential impacts of climate change on species distributions, and each modeling approach has its strengths and weaknesses. Here, we demonstrate an approach to projecting climate‐driven changes in species distributions that draws on both empirical and mechanistic models. We combined projections from a dynamic global vegetation model (DGVM) that simulates the distributions of biomes based on basic plant functional types with projections from empirical climatic niche models for six tree species in northwestern North America. These integrated model outputs incorporate important biological processes, such as competition, physiological responses of plants to changes in atmospheric CO₂ concentrations, and fire, as well as what are likely to be species‐specific climatic constraints. We compared the integrated projections to projections from the empirical climatic niche models alone. Overall, our integrated model outputs projected a greater climate‐driven loss of potentially suitable environmental space than did the empirical climatic niche model outputs alone for the majority of modeled species. Our results also show that refining species distributions with DGVM outputs had large effects on the geographic locations of suitable habitat. We demonstrate one approach to integrating the outputs of mechanistic and empirical niche models to produce bioclimatic projections. But perhaps more importantly, our study reveals the potential for empirical climatic niche models to over‐predict suitable environmental space under future climatic conditions.