1. Growing recognition of the importance of climate extremes as drivers of contemporary and future ecological dynamics has led to increasing interest in studying these locally and globally important ...phenomena. 2. Many ecological studies examining the impacts of what are deemed climate extremes, such as heat waves and severe drought, do not provide a definition of extremity, either from a statistical context based on the long-term climatic record or from the perspective of the response of the system - are the effects extreme (unusual or profound) in comparison to normal variability? 3. A synthetic definition of an extreme climatic event (ECE) is proposed that includes ‘extremeness' in both the driver and the response: an ECE is as an episode or occurrence in which a statistically rare or unusual climatic period alters ecosystem structure and/or function well outside the bounds of what is considered typical or normal variability. This definition is accompanied by a mechanistic framework based on the concept that extreme response thresholds associated with significant community change and altered ecosystem function must be crossed in order for an ECE to occur. 4. Synthesis. A definition and mechanistic framework for ECEs is used to identify priorities for future research that will enable ecologists to more fully assess the ecological consequences of climate extremes for ecosystem structure and function today and in a future world where their frequency and intensity are expected to increase.
1. Climate extremes, such as severe drought, heat waves and periods of heavy rainfall, can have profound consequences for ecological systems and for human welfare. Global climate change is expected ...to increase both the frequency and the intensity of climate extremes and there is an urgent need to understand their ecological consequences. 2. Major challenges for advancing our understanding of the ecological consequences of climate extremes include setting a climatic baseline to facilitate the statistical determination of when climate conditions are extreme, having sufficient knowledge of ecological systems so that extreme ecological responses can be identified, and finally, being able to attribute a climate extreme as the driver of an extreme ecological response, defined as an extreme climatic event (ECE). 3. The papers in this issue represent a cross-section of the emerging field of climate extremes research, including an examination of the palaeo-ecological record to assess patterns and drivers of extreme ecological responses in the late Quaternary, experiments in grasslands assessing a range of ecological responses and the role of ecotypic variation in determining responses to climate extremes, and the quantification of the ecological consequences of a recent ECE in the desert Southwest of the USA. 4. Synthesis. The papers in this Special Feature suggest that although the occurrence of ECEs may be common in palaeo-ecological and observational studies, studies in which climate extremes have been experimentally imposed often do not result in ecological responses outside the bounds of normal variability of a system. Thus, ECEs occur much less frequently than their potential drivers and even less frequently than observational studies suggest. Future research is needed to identify the types and time-scales of climate extremes that result in ECEs, the potential for interactions among different types of climate changes and extremes, and the role of genetic, species and trait diversity in determining ecological responses and their evolutionary consequences. These research priorities require the development of alternative research approaches to impose realistic climate extremes on a broad range of organisms and ecosystems.
Precipitation (PPT) is a primary climatic determinant of plant growth and aboveground net primary production (ANPP) over much of the globe. Thus, PPT–ANPP relationships are important both ...ecologically and to land–atmosphere models that couple terrestrial vegetation to the global carbon cycle. Empirical PPT–ANPP relationships derived from long-term site-based data are almost always portrayed as linear, but recent evidence has accumulated that is inconsistent with an underlying linear relationship. We review, and then reconcile, these inconsistencies with a nonlinear model that incorporates observed asymmetries in PPT–ANPP relationships. Although data are currently lacking for parameterization, this new model highlights research needs that, when met, will improve our understanding of carbon cycle dynamics, as well as forecasts of ecosystem responses to climate change.
Climate change forecasts of more frequent climate extremes suggest that such events will become increasingly important drivers of future ecosystem dynamics and function. Because the rarity and ...unpredictability of naturally occurring climate extremes limits assessment of their ecological impacts, we experimentally imposed extreme drought and a mid-summer heat wave over two years in a central U.S. grassland. While the ecosystem was resistant to heat waves, it was not resistant to extreme drought, which reduced aboveground net primary productivity (ANPP) below the lowest level measured in this grassland in almost 30 years. This extreme reduction in ecosystem function was a consequence of reduced productivity of both C
4
grasses and C
3
forbs. However, the dominant forb was negatively impacted by the drought more than the dominant grass, and this led to a reordering of species abundances within the plant community. Although this change in community composition persisted post-drought, ANPP recovered completely the year after drought due to rapid demographic responses by the dominant grass, compensating for loss of the dominant forb. Overall, these results show that an extreme reduction in ecosystem function attributable to climate extremes (e.g., low resistance) does not preclude rapid ecosystem recovery. Given that dominance by a few species is characteristic of most ecosystems, knowledge of the traits of these species and their responses to climate extremes will be key for predicting future ecosystem dynamics and function.
Summary
Drought is intensifying globally with climate change, creating an urgency to understand ecosystem response to drought both during and after these events end to limit loss of ecosystem ...functioning. The literature is replete with studies of how ecosystems respond during drought, yet there are far fewer studies focused on ecosystem dynamics after drought ends. Furthermore, while the terms used to describe drought can be variable and inconsistent, so can those that describe ecosystem responses following drought. With this review, we sought to evaluate and create clear definitions of the terms that ecologists use to describe post‐drought responses. We found that legacy effects, resilience and recovery were used most commonly with respect to post‐drought ecosystem responses, but the definitions used to describe these terms were variable. Based on our review of the literature, we propose a framework for generalizing ecosystem responses after drought ends, which we refer to as ‘the post‐drought period’. We suggest that future papers need to clearly describe characteristics of the imposed drought, and we encourage authors to use the term post‐drought period as a general term that encompasses responses after drought ends and use other terms as more specific descriptors of responses during the post‐drought period.
•We review current understanding of plant responses to climate extremes.•Various mechanisms underlying plant responses to climate extremes are summarized.•We propose future research effort in extreme ...event research.
Ongoing climate change has caused extreme climatic events to happen more frequently, which can fundamentally threaten plant growth and survivorship. In this review paper, we found that extreme climatic events, such as heat waves, frost, drought and flooding, usually reduces plant production and induces mortality. The magnitude of impacts on production and mortality are exceedingly variable, which likely result from different severities of the climate extremes, sensitivities of various processes, vegetation types, and inherent regulatory mechanisms of plants and ecosystems. Climatologically severe events may not necessarily trigger plant responses. Different processes respond to the same extreme events differently. Such different responses also vary with species. Moreover, plants likely activate a variety of physiological and molecular mechanisms regulate their responses to extremes. Documenting those variable responses and identifying their causes are critical to advancing our understanding. Nevertheless, our research has to move beyond the documentation of phenomenon to reveal fundamental mechanisms underlying plant responses to climate extremes. Toward that goal, we need to define extreme climatic events under a plant perspective and evaluate different response patterns of various processes to climate extremes. In this review, we also propose to focus our future research on manipulative field experiments and coordinated networks of experiments at multiple sites over different regions to understand the real-world responses of plants and ecosystems.
In contrast to pulses in resource availability following disturbance events, many of the most pressing global changes, such as elevated atmospheric carbon dioxide concentrations and nitrogen ...deposition, lead to chronic and often cumulative alterations in available resources. Therefore, predicting ecological responses to these chronic resource alterations will require the modification of existing disturbance-based frameworks. Here, we present a conceptual framework for assessing the nature and pace of ecological change under chronic resource alterations. The "hierarchical-response framework" (HRF) links well-documented, ecological mechanisms of change to provide a theoretical basis for testing hypotheses to explain the dynamics and differential sensitivity of ecosystems to chronic resource alterations. The HRF is based on a temporal hierarchy of mechanisms and responses beginning with individual (physiological/metabolic) responses, followed by species reordering within communities, and finally species loss and immigration. Each mechanism is hypothesized to differ in the magnitude and rate of its effects on ecosystem structure and function, with this variation depending on ecosystem attributes, such as longevity of dominant species, rates of biogeochemical cycling, levels of biodiversity, and trophic complexity. Overall, the HRF predicts nonlinear changes in ecosystem dynamics, with the expectation that interactions with natural disturbances and other global-change drivers will further alter the nature and pace of change. The HRF is explicitly comparative to better understand differential sensitivities of ecosystems, and it can be used to guide the design of coordinated, cross-site experiments to enable more robust forecasts of contemporary and future ecosystem dynamics.
Climate extremes will elicit responses from the individual to the ecosystem level. However, only recently have ecologists begun to synthetically assess responses to climate extremes across multiple ...levels of ecological organization. We review the literature to examine how plant responses vary and interact across levels of organization, focusing on how individual, population and community responses may inform ecosystem-level responses in herbaceous and forest plant communities. We report a high degree of variability at the individual level, and a consequential inconsistency in the translation of individual or population responses to directional changes in community- or ecosystem-level processes. The scaling of individual or population responses to community or ecosystem responses is often predicated upon the functional identity of the species in the community, in particular, the dominant species. Furthermore, the reported stability in plant community composition and functioning with respect to extremes is often driven by processes that operate at the community level, such as species niche partitioning and compensatory responses during or after the event. Future research efforts would benefit from assessing ecological responses across multiple levels of organization, as this will provide both a holistic and mechanistic understanding of ecosystem responses to increasing climatic variability.
This article is part of the themed issue ‘Behavioural, ecological and evolutionary responses to extreme climatic events’.
In terrestrial ecosystems, climate change forecasts of increased frequencies and magnitudes of wet and dry precipitation anomalies are expected to shift precipitation–net primary productivity ...(PPT–NPP) relationships from linear to nonlinear. Less understood, however, is how future changes in the duration of PPT anomalies will alter PPT–NPP relationships. A review of the literature shows strong potential for the duration of wet and dry PPT anomalies to impact NPP and to interact with the magnitude of anomalies. Within semi‐arid and mesic grassland ecosystems, PPT gradient experiments indicate that short‐duration (1 year) PPT anomalies are often insufficient to drive nonlinear aboveground NPP responses. But long‐term studies, within desert to forest ecosystems, demonstrate how multi‐year PPT anomalies may result in increasing impacts on NPP through time, and thus alter PPT–NPP relationships. We present a conceptual model detailing how NPP responses to PPT anomalies may amplify with the duration of an event, how responses may vary in xeric vs. mesic ecosystems, and how these differences are most likely due to demographic mechanisms. Experiments that can unravel the independent and interactive impacts of the magnitude and duration of wet and dry PPT anomalies are needed, with multi‐site long‐term PPT gradient experiments particularly well‐suited for this task.
The cumulative impacts to terrestrial ecosystems resulting from longer durations of droughts and wet periods may differentially alter temporal precipitation–productivity relationships in xeric vs. mesic ecosystems, thus producing different types of nonlinear ecosystem response dynamics. However, such divergence in ecosystem responses is proposed to result from a shared mechanisms in the form of shifting vegetation demographics.
Despite asymmetric competition and a wide array of functional similarities, two ecologically important C4 perennial grasses, Andropogon gerardii and Sorghastrum nutans, frequently codominate areas of ...the mesic tallgrass prairie of the US Great Plains. A subtle difference in their vegetative reproduction strategies may play a role in preventing the exclusion of S. nutans, the presumed weaker competitor in such regions.
While A. gerardii vegetative tiller densities peak in the early growing season and decline thereafter (determinate recruitment), those of S. nutans may continue to increase throughout the growing season (indeterminate recruitment), providing a potential avenue for recovery from more intensive early season competition. However, until now these patterns have only been informally observed in the field.
We examined the year‐to‐year consistency of growing season vegetative tiller dynamics (measured as seasonal change in tiller densities) of each grass species from an intact tallgrass prairie in Kansas – a site within the core of both species' distributions – over a period of 8 years. Then, to investigate environmental effects on these dynamics, we examined whether they differ across a Kansas landscape varying in topography, fire management regimes, and the abundances of the study species. Finally, we expanded the investigation of environmental effects on growing season tiller dynamics by observing them at the periphery of the species' distributions in central Colorado, where climatic conditions are dryer and the study species' abundances are reduced.
Synthesis. We found that the tiller densities of A. gerardii decline within seasons with striking consistency regardless of spatio‐temporal scale or environmental factors (topography and fire regimes). In contrast, we found the seasonal dynamics of S. nutans tiller densities were dependent on environmental factors, with seasonal tiller density increases occurring only within the Kansas populations but not consistent between years. These observations lay the groundwork for establishing differences in tiller recruitment determinacy as a potentially important yet underappreciated mechanism for promoting coexistence and codominance among perennial plant species.
Differences in recruitment determinacy – whether populations can continue to grow throughout the growing season or have genetically predetermined, annual time limits – may underly the cryptic mechanism keeping big bluestem and Indiangrass (pictured) codominant in mesic tallgrass prairies. We found that these reproductive characteristics are associated with consistently differential intra‐seasonal population dynamics despite many functional similarities between the two grasses.
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