Toward a conceptual synthesis for climate change responses

Aim To identify hypotheses for how climate change affects long‐term population persistence that can be used as a framework for future syntheses of ecological responses to climate change. Location Global. Methods We surveyed ecological and evolutionary concepts related to how a changing climate might alter population persistence. We organized established concepts into a two‐stage framework that relates abiotic change to population persistence via changes in the rates or outcomes of ecological and evolutionary processes. We surveyed reviews of climate change responses, and evaluated patterns in light of our conceptual framework. Results We classified hypotheses for population responses to climate change as one of two types: (1) hypotheses that relate rates of ecological and evolutionary processes (plasticity, dispersal, population growth and evolution) to abiotic change, and (2) hypotheses that relate changes in these processes to four fundamental population‐level responses (colonization, acclimatization, adaptation or extinction). We found that a disproportionate emphasis on response in the climate change literature is difficult to reconcile with ecological and evolutionary theories that emphasize processes. We discuss a set of 24 hypotheses that represent gaps in the literature that limit our ability determine whether observed climate change responses are sufficient to facilitate persistence through future climate change. Main conclusions Though theory relates environmental change to fundamental ecological and evolutionary processes and population‐level responses, clear hypotheses based on theory have not been systematically formulated and tested in the context of climate change. Stronger links between basic theory and observed impacts of climate change are required to assess which responses are common, likely or able to facilitate population persistence despite ongoing environmental change. We anticipate that a hypothesis‐testing framework will reveal that indirect effects of climate change responses are more pervasive than previously thought and related to a few general processes, even though the patterns they create are incredibly diverse.     

森林土壤酶对环境变化的响应研究进展

全球气候变化已是不争的事实,对陆地生态系统特别是森林生态系统物质循环将产生显著的影响。土壤酶是森林土壤物质循环的主要限制因素之一,对气候变化的响应近年来受到广泛关注。由于森林土壤酶对全球气候变化的响应研究是预测未来环境变化对森林生态系统过程影响的关键,因此,着重综述了森林土壤酶对环境变化尤其是全球变暖和氮沉降响应方面的研究,并分析了未来研究的主要方向。环境变化会引起土壤p H、水分及其营养成分的变化,而这些变化会反作用于土壤酶的活性和稳定性。森林土壤酶对增温的响应,不仅与酶的种类以及增温的温度范围和持续时间有关,还与土壤类型有关,是多种因子综合作用的结果。森林土壤酶对氮添加的响应与林分类型和土层类型有关,受复合氮的影响更大。建议未来的研究应加强酶的基本性质对环境变化的响应研究,注重林分类型、土层类型导致的差异,强化多因素的交互作用,并进行长期、综合的观测。     

Climate change and evolution: disentangling environmental and genetic responses

Rapid climate change is likely to impose strong selection pressures on traits important for fitness, and therefore, microevolution in response to climate-mediated selection is potentially an important mechanism mitigating negative consequences of climate change. We reviewed the empirical evidence for recent microevolutionary responses to climate change in longitudinal studies emphasizing the following three perspectives emerging from the published data. First, although signatures of climate change are clearly visible in many ecological processes, similar examples of microevolutionary responses in literature are in fact very rare. Second, the quality of evidence for microevolutionary responses to climate change is far from satisfactory as the documented responses are often - if not typically - based on nongenetic data. We reinforce the view that it is as important to make the distinction between genetic (evolutionary) and phenotypic (includes a nongenetic, plastic component) responses clear, as it is to understand the relative roles of plasticity and genetics in adaptation to climate change. Third, in order to illustrate the difficulties and their potential ubiquity in detection of microevolution in response to natural selection, we reviewed the quantitative genetic studies on microevolutionary responses to natural selection in the context of long-term studies of vertebrates. The available evidence points to the overall conclusion that many responses perceived as adaptations to changing environmental conditions could be environmentally induced plastic responses rather than microevolutionary adaptations. Hence, clear-cut evidence indicating a significant role for evolutionary adaptation to ongoing climate warming is conspicuously scarce.     

Beyond a warming fingerprint: individualistic biogeographic responses to heterogeneous climate change in California

Understanding recent biogeographic responses to climate change is fundamental for improving our predictions of likely future responses and guiding conservation planning at both local and global scales. Studies of observed biogeographic responses to 20th century climate change have principally examined effects related to ubiquitous increases in temperature – collectively termed a warming fingerprint. Although the importance of changes in other aspects of climate – particularly precipitation and water availability – is widely acknowledged from a theoretical standpoint and supported by paleontological evidence, we lack a practical understanding of how these changes interact with temperature to drive biogeographic responses. Further complicating matters, differences in life history and ecological attributes may lead species to respond differently to the same changes in climate. Here, we examine whether recent biogeographic patterns across California are consistent with a warming fingerprint. We describe how various components of climate have changed regionally in California during the 20th century and review empirical evidence of biogeographic responses to these changes, particularly elevational range shifts. Many responses to climate change do not appear to be consistent with a warming fingerprint, with downslope shifts in elevation being as common as upslope shifts across a number of taxa and many demographic and community responses being inconsistent with upslope shifts. We identify a number of potential direct and indirect mechanisms for these responses, including the influence of aspects of climate change other than temperature (e.g., the shifting seasonal balance of energy and water availability), differences in each taxon's sensitivity to climate change, trophic interactions, and land‐use change. Finally, we highlight the need to move beyond a warming fingerprint in studies of biogeographic responses by considering a more multifaceted view of climate, emphasizing local‐scale effects, and including a priori knowledge of relevant natural history for the taxa and regions under study.     

Ecological and methodological drivers of species’ distribution and phenology responses to climate change

Climate change is shifting species’ distribution and phenology. Ecological traits, such as mobility or reproductive mode, explain variation in observed rates of shift for some taxa. However, estimates of relationships between traits and climate responses could be influenced by how responses are measured. We compiled a global data set of 651 published marine species’ responses to climate change, from 47 papers on distribution shifts and 32 papers on phenology change. We assessed the relative importance of two classes of predictors of the rate of change, ecological traits of the responding taxa and methodological approaches for quantifying biological responses. Methodological differences explained 22% of the variation in range shifts, more than the 7.8% of the variation explained by ecological traits. For phenology change, methodological approaches accounted for 4% of the variation in measurements, whereas 8% of the variation was explained by ecological traits. Our ability to predict responses from traits was hindered by poor representation of species from the tropics, where temperature isotherms are moving most rapidly. Thus, the mean rate of distribution change may be underestimated by this and other global syntheses. Our analyses indicate that methodological approaches should be explicitly considered when designing, analysing and comparing results among studies. To improve climate impact studies, we recommend that (1) reanalyses of existing time series state how the existing data sets may limit the inferences about possible climate responses; (2) qualitative comparisons of species’ responses across different studies be limited to studies with similar methodological approaches; (3) meta‐analyses of climate responses include methodological attributes as covariates; and (4) that new time series be designed to include the detection of early warnings of change or ecologically relevant change. Greater consideration of methodological attributes will improve the accuracy of analyses that seek to quantify the role of climate change in species’ distribution and phenology changes.     

Bat responses to climate change: a systematic review

Understanding how species respond to climate change is key to informing vulnerability assessments and designing effective conservation strategies, yet research efforts on wildlife responses to climate change fail to deliver a representative overview due to inherent biases. Bats are a species-rich, globally distributed group of organisms that are thought to be particularly sensitive to the effects of climate change because of their high surface-to-volume ratios and low reproductive rates. We systematically reviewed the literature on bat responses to climate change to provide an overview of the current state of knowledge, identify research gaps and biases and highlight future research needs. We found that studies are geographically biased towards Europe, North America and Australia, and temperate and Mediterranean biomes, thus missing a substantial proportion of bat diversity and thermal responses. Less than half of the published studies provide concrete evidence for bat responses to climate change. For over a third of studied bat species, response evidence is only based on predictive species distribution models. Consequently, the most frequently reported responses involve range shifts (57% of species) and changes in patterns of species diversity (26%). Bats showed a variety of responses, including both positive (e.g. range expansion and population increase) and negative responses (range contraction and population decrease), although responses to extreme events were always negative or neutral. Spatial responses varied in their outcome and across families, with almost all taxonomic groups featuring both range expansions and contractions, while demographic responses were strongly biased towards negative outcomes, particularly among Pteropodidae and Molossidae. The commonly used correlative modelling approaches can be applied to many species, but do not provide mechanistic insight into behavioural, physiological, phenological or genetic responses. There was a paucity of experimental studies (26%), and only a small proportion of the 396 bat species covered in the examined studies were studied using long-term and/or experimental approaches (11%), even though they are more informative about the effects of climate change. We emphasise the need for more empirical studies to unravel the multifaceted nature of bats' responses to climate change and the need for standardised study designs that will enable synthesis and meta-analysis of the literature. Finally, we stress the importance of overcoming geographic and taxonomic disparities through strengthening research capacity in the Global South to provide a more comprehensive view of terrestrial biodiversity responses to climate change.     

Community and ecosystem responses to recent climate change

There is ample evidence for ecological responses to recent climate change. Most studies to date have concentrated on the effects of climate change on individuals and species, with particular emphasis on the effects on phenology and physiology of organisms as well as changes in the distribution and range shifts of species. However, responses by individual species to climate change are not isolated; they are connected through interactions with others at the same or adjacent trophic levels. Also from this more complex perspective, recent case studies have emphasized evidence on the effects of climate change on biotic interactions and ecosystem services. This review highlights the ‘knowns’ but also ‘unknowns’ resulting from recent climate impact studies and reveals limitations of (linear) extrapolations from recent climate-induced responses of species to expected trends and magnitudes of future climate change. Hence, there is need not only to continue to focus on the impacts of climate change on the actors in ecological networks but also and more intensively to focus on the linkages between them, and to acknowledge that biotic interactions and feedback processes lead to highly complex, nonlinear and sometimes abrupt responses.     

Game theory sheds new light on ecological responses to current climate change when phenology is historically mismatched

Phenological changes are well documented biological effects of current climate change but their adaptive value and demographic consequences are poorly known. Game theoretical models have shown that deviating from the fitness-maximising phenology can be evolutionary stable under frequency-dependent selection. We study eco-evolutionary responses to climate change when the historical phenology is mismatched in this way. For illustration we model adaptation of arrival dates in migratory birds that compete for territories at their breeding grounds. We simulate climate change by shifting the timing and the length of the favourable season for breeding. We show that initial trends in changes of population densities can be either reinforced or counteracted during the ensuing evolutionary adaptation. We find in total seven qualitatively different population trajectories during the transition to a new evolutionary equilibrium. This surprising diversity of eco-evolutionary responses provides adaptive explanations to the observed variation in phenological responses to recent climate change.     

The influence of historical climate changes on Southern Ocean marine predator populations: a comparative analysis

The Southern Ocean ecosystem is undergoing rapid physical and biological changes that are likely to have profound implications for higher‐order predators. Here, we compare the long‐term, historical responses of Southern Ocean predators to climate change. We examine palaeoecological evidence for changes in the abundance and distribution of seabirds and marine mammals, and place these into context with palaeoclimate records in order to identify key environmental drivers associated with population changes. Our synthesis revealed two key factors underlying Southern Ocean predator population changes; (i) the availability of ice‐free ground for breeding and (ii) access to productive foraging grounds. The processes of glaciation and sea ice fluctuation were key; the distributions and abundances of elephant seals, snow petrels, gentoo, chinstrap and Adélie penguins all responded strongly to the emergence of new breeding habitat coincident with deglaciation and reductions in sea ice. Access to productive foraging grounds was another limiting factor, with snow petrels, king and emperor penguins all affected by reduced prey availability in the past. Several species were isolated in glacial refugia and there is evidence that refuge populations were supported by polynyas. While the underlying drivers of population change were similar across most Southern Ocean predators, the individual responses of species to environmental change varied because of species specific factors such as dispersal ability and environmental sensitivity. Such interspecific differences are likely to affect the future climate change responses of Southern Ocean marine predators and should be considered in conservation plans. Comparative palaeoecological studies are a valuable source of long‐term data on species’ responses to environmental change that can provide important insights into future climate change responses. This synthesis highlights the importance of protecting productive foraging grounds proximate to breeding locations, as well as the potential role of polynyas as future Southern Ocean refugia.     

Species and community responses to short‐term climate manipulation: Microarthropods in the sub‐Antarctic

Abstract: Both species and community‐level investigations are important for understanding the biotic impacts of climate change, because current evidence suggests that individual species responses are idiosyncratic. However, few studies of climate change impacts have been conducted on entire terrestrial arthropod communities living in the same habitat in the southern Hemisphere, and the effects of precipitation changes on them are particularly poorly understood. Here we investigate the species‐ and community‐level responses of microarthropods inhabiting a keystone plant species, on sub‐Antarctic Marion Island, to experimental reduction in precipitation, warming and shading. These climate manipulations were chosen based on observed climate trends and predicted indirect climate change impacts on this system. The dry‐warm and shade inducing treatments that were imposed effected significant species‐ and community‐level responses after a single year. Although the strongest community‐level trends included a dramatic decline in springtail abundance and total biomass under the dry‐warm and shade treatments, species responses were generally individualistic, that is, springtails responded differently to mites, and particular mite and springtail species responded differently to each other. Our results therefore provide additional support for the dynamic rather than static model for community responses to climate change, in the first such experiment in the sub‐Antarctic. In conclusion, these results show that an ongoing decline in precipitation and increase in temperature is likely to have dramatic direct and indirect effects on this microarthropod community. Moreover, they indicate that while at a broad scale it may be possible to make generalizations regarding species responses to climate change, these generalizations are unlikely to translate into predictable effects at the community level.     

Tracking unstable states: ecosystem dynamics in a changing world

Ecological systems can show complex and sometimes abrupt responses to environmental change, with important implications for their resilience. Theories of alternate stable states have been used to predict regime shifts of ecosystems as equilibrium responses to sufficiently slow environmental change. The actual rate of environmental change is a key factor affecting the response, yet we are still lacking a non-equilibrium theory that explicitly considers the influence of this rate of environmental change. We present a metacommunity model of predator–prey interactions displaying multiple stable states, and we impose an explicit rate of environmental change in habitat quality (carrying capacity) and connectivity (dispersal rate). We study how regime shifts depend on the rate of environmental change and compare the outcome with a stability analysis in the corresponding constant environment. Our results reveal that in a changing environment, the community can track states that are unstable in the constant environment. This tracking can lead to regime shifts, including local extinctions, that are not predicted by alternative stable state theory. In our metacommunity, tracking unstable states also controls the maintenance of spatial heterogeneity and spatial synchrony. Tracking unstable states can also lead to regime shifts that may be reversible or irreversible. Our study extends current regime shift theories to integrate rate-dependent responses to environmental change. It reveals the key role of unstable states for predicting transient dynamics and long-term resilience of ecological systems to climate change.     

Evolutionary responses to global change: lessons from invasive species

Biologists have recently devoted increasing attention to the role of rapid evolution in species' responses to environmental change. However, it is still unclear what evolutionary responses should be expected, at what rates, and whether evolution will save populations at risk of extinction. The potential of biological invasions to provide useful insights has barely been realised, despite the close analogies to species responding to global change, particularly climate change; in both cases, populations encounter novel climatic and biotic selection pressures, with expected evolutionary responses occurring over similar timescales. However, the analogy is not perfect, and invasive species are perhaps best used as an upper bound on expected change. In this article, we review what invasive species can and cannot teach us about likely evolutionary responses to global change and the constraints on those responses. We also discuss the limitations of invasive species as a model and outline directions for future research.     

Breathing some air into the single-species vacuum: multi-species responses to environmental change

Studies of ecological responses to climate change have often analysed species independently of each other, yet interactions between species are fundamental aspects of ecology. Mutshinda, O'Hara & Woiwod (2011) used light-trapping data for Lepidoptera (moths) to examine population responses to intraspecific effects and effects of winter rainfall and temperature. They show how Bayesian hierarchical models can analyse residual correlations among species' responses, illustrating an approach to account for and measure dependencies that are not fully explained by the candidate explanatory variables. A key result is that the responses of the different moth species did not appear to have strong residual correlation (Mutshinda, O'Hara & Woiwod 2011). These analyses provide an approach for synthesising across species and can better inform ecological responses to environmental change.     

Experimental strategies to assess the biological ramifications of multiple drivers of global ocean change—A review

Marine life is controlled by multiple physical and chemical drivers and by diverse ecological processes. Many of these oceanic properties are being altered by climate change and other anthropogenic pressures. Hence, identifying the influences of multifaceted ocean change, from local to global scales, is a complex task. To guide policy‐making and make projections of the future of the marine biosphere, it is essential to understand biological responses at physiological, evolutionary and ecological levels. Here, we contrast and compare different approaches to multiple driver experiments that aim to elucidate biological responses to a complex matrix of ocean global change. We present the benefits and the challenges of each approach with a focus on marine research, and guidelines to navigate through these different categories to help identify strategies that might best address research questions in fundamental physiology, experimental evolutionary biology and community ecology. Our review reveals that the field of multiple driver research is being pulled in complementary directions: the need for reductionist approaches to obtain process‐oriented, mechanistic understanding and a requirement to quantify responses to projected future scenarios of ocean change. We conclude the review with recommendations on how best to align different experimental approaches to contribute fundamental information needed for science‐based policy formulation.     

植物叶片性状对气候变化的响应研究进展

叶片性状反映了植物对环境的高度适应能力及其在复杂生境下的自我调控能力。叶片性状如何响应和适应气候变化是植物适应性研究的重点内容。该文系统综述了叶片大小、比叶质量、叶片氮含量、碳同位素等指标对气候变化响应的最新研究结果。不同叶片性状对气候变化的响应结果存在差异,所指示的生态学含义也有所不同。单一叶片性状不能全面地反映植物对气候变化的响应;不同尺度的研究(如环境的修饰或筛选作用的研究)还存在很多不确定性。高寒地区的研究工作相对缺乏。该文有助于理解植物与气候之间的相互关系、植物对气候变化的响应与适应对策,对了解植物演化、预测植物在未来气候变化条件下的变化特征具有一定意义。     

Threats to mangroves from climate change and adaptation options: A review

Mangrove ecosystems are threatened by climate change. We review the state of knowledge of mangrove vulnerability and responses to predicted climate change and consider adaptation options. Based on available evidence, of all the climate change outcomes, relative sea-level rise may be the greatest threat to mangroves. Most mangrove sediment surface elevations are not keeping pace with sea-level rise, although longer term studies from a larger number of regions are needed. Rising sea-level will have the greatest impact on mangroves experiencing net lowering in sediment elevation, where there is limited area for landward migration. The Pacific Islands mangroves have been demonstrated to be at high risk of substantial reductions. There is less certainty over other climate change outcomes and mangrove responses. More research is needed on assessment methods and standard indicators of change in response to effects from climate change, while regional monitoring networks are needed to observe these responses to enable educated adaptation. Adaptation measures can offset anticipated mangrove losses and improve resistance and resilience to climate change. Coastal planning can adapt to facilitate mangrove migration with sea-level rise. Management of activities within the catchment that affect long-term trends in the mangrove sediment elevation, better management of other stressors on mangroves, rehabilitation of degraded mangrove areas, and increases in systems of strategically designed protected area networks that include mangroves and functionally linked ecosystems through representation, replication and refugia, are additional adaptation options.     

Mixed effects models with bivariate and univariate association parameters for longitudinal bivariate binary response data

When two binary responses are measured for each study subject across time, it may be of interest to model how the bivariate associations and marginal univariate risks involving the two responses change across time. To achieve such a goal, marginal models with bivariate log odds ratio and univariate logit components are extended to include random effects for all components. Specifically, separate normal random effects are specified on the log odds ratio scale for bivariate responses and on the logit scale for univariate responses. Assuming conditional independence given the random effects facilitates the modeling of bivariate associations across time with missing at random incomplete data. We fit the model to a dataset for which such structures are feasible: a longitudinal randomized trial of a cardiovascular educational program where the responses of interest are change in hypertension and hypercholestemia status. The proposed model is compared to a naive bivariate model that assumes independence between time points and univariate mixed effects logit models.     

Can Boreal and Temperate Forest Management be Adapted to the Uncertainties of 21st Century Climate Change?

Considerable uncertainties remain about magnitude and character, if not general direction of anthropogenic climate change. Global mean temperature could increase by 1.5–4.5°C or more over historic levels, and extreme weather events—drought, storms, and flooding—are likely to increase greatly in frequency. Although ecologists and foresters agree that the practice of forestry will be transformed under climate change, these uncertainties compound the challenge of achieving sustainable, adaptive forest management. In this aritcle, we (i) present a multidisciplinary synthesis of current knowledge of responses of temperate and boreal tree species and forest communities to climate change, and (ii) outline silvicultural strategies for adapting temperate and boreal forests to confront climate change. Our knowledge synthesis proceeds through critical appraisals of efforts to model future tree distributions and responses to climate change, and reviews physiological, phenological, acclimation, and epigenetic responses to climate. As is the case of climate change itself, there are numerous uncertainties about tree species and provenance responses to climate change. For example, acclimation of respiration and epigenetic conditioning of seed embryos has the potential to buffer species against limited warming. Provenances within species also display idiosyncratic responses to altered climates, implying that soemm varieties will be more resilient or resistant to climate change than others. Genetically determined limits to climatic tolerance, and the limits of tree community resistance and resilience (speed of recovery from disturbance) in the face of climate-related disturbances are largely unknown. These unknowns require managers to adopt a portfolio of silvicultural strategies, which may range from minor modifications of current practices to design of novel multi-species stands that may have no historical analogue. Forest managers must be prepared to respond nimbly as they develop, incorporate new insights about climate change and species responses to warming into their practices. Marshalling all strategies and sources of knowledge should enable forest managers to mount (at least) a partially successful response to the challenges of climate change.     

Visual processing levels revealed by response latencies to changes in different visual attributes.

Visual latencies, and their variation with stimulus attributes, can provide information about the level in the visual system at which different attributes of the image are analysed, and decisions about them made. A change in the colour, structure or movement of a visual stimulus brings about a highly reproducible transient constriction of the pupil that probably depends on visual cortical mechanisms. We measured this transient response to changes in several attributes of visual stimuli, and also measured manual reaction times to the same stimulus changes. Through analysis of latencies, we hoped to establish whether changes in different stimulus attributes were processed by mechanisms at the same or different levels in the visual pathway. Pupil responses to a change in spatial structure or colour are almost identical, but both are ca. 40 ms slower than those to a change in light flux, which are thought to depend largely on subcortical pathways. Manual reaction times to a change in spatial structure or colour, or to the onset of coherent movement, differ reliably, and all are longer than the reaction time to a change in light flux. On average, observers take 184 ms to detect a change in light flux, 6 ms more to detect the onset of a grating, 30 ms more to detect a change in colour, and 37 ms more to detect the onset of coherent motion. The pattern of latency variation for pupil responses and reaction times suggests that the mechanisms that trigger the responses lie at different levels in cortex. Given our present knowledge of visual cortical organization, the long reaction time to the change in motion is surprising. The range of reaction times across different stimuli is consistent with decisions about the onset of a grating being made in V1 and decisions about the change in colour or change in motion being made in V4.     

Forest Remnants Along Urban-Rural Gradients: Examining Their Potential for Global Change Research

Over the next century, ecosystems throughout the world will be responding to rapid changes in climate and rising levels of carbon dioxide, inorganic N and ozone. Because people depend on biological systems for water, food and other ecosystem services, predicting the range of responses to global change for various ecosystem types in different geographic locations is a high priority. Modeling exercises and manipulative experimentation have been the principle approaches used to place upper and lower bounds on community and ecosystem responses. However, each of these approaches has recognized limitations. Manipulative experiments cannot vary all the relevant factors and are often performed at small spatio-temporal scales. Modeling is limited by data availability and by our knowledge of how current observations translate into future conditions. These weaknesses would improve if we could observe ecosystems that have already responded to global change factors and thus presage shifts in ecosystem structure and function. Here we consider whether urban forest remnants might offer this ability. As urban forests have been exposed to elevated temperature, carbon dioxide, nitrogen deposition and ozone for many decades, they may be ahead of the global change “response curve” for forests in their region. Therefore, not only might forests along urbanization gradients provide us with natural experiments for studying current responses to global change factors, but their legacy of response to past urbanization may also constitute space-for-time substitution experiments for predicting likely regional forest responses to continued environmental change. For this approach to be successful, appropriate criteria must be developed for selecting forest remnants and plots that would optimize our ability to detect incipient forest responses to spatial variation in global change factors along urbanization gradients, while minimizing artifacts associated with remnant size and factors other than those that simulate global change. Studying forests that meet such criteria along urban-to-rural gradients could become an informative part of a mixed strategy of approaches for improving forecasts of forest ecosystem change at the regional scale.