Predicting ecosystem carbon balance in a warming Arctic: the importance of long‐term thermal acclimation potential and inhibitory effects of light on respiration

The carbon balance of Arctic ecosystems is particularly sensitive to global environmental change. Leaf respiration (R), a temperature‐dependent key process in determining the carbon balance, is not well‐understood in Arctic plants. The potential for plants to acclimate to warmer conditions could strongly impact future global carbon balance. Two key unanswered questions are (1) whether short‐term temperature responses can predict long‐term respiratory responses to growth in elevated temperatures and (2) to what extent the constant daylight conditions of the Arctic growing season inhibit leaf respiration. In two dominant Arctic species E riophorum vaginatum (tussock grass) and B etula nana (woody shrub), we assessed the extent of respiratory inhibition in the light (R _L/R _D), respiratory response to short‐term temperature change, and respiratory acclimation to long‐term warming treatments. We found that R of both species is strongly inhibited by light (averaging 35% across all measurement temperatures). In E . vaginatum both R _L and R _D acclimated to the long‐term warming treatment, reducing the magnitude of respiratory response relative to the short‐term response to temperature increase. In B . nana, both R _L and R _D responded to short‐term temperature increase but showed no acclimation to the long‐term warming. The ability to predict plant respiratory response to global warming with short‐term temperature responses will depend on species‐specific acclimation potential and the differential response of R _L and R _D to temperature. With projected woody shrub encroachment in Arctic tundra and continued warming, changing species dominance between these two functional groups, may impact ecosystem respiratory response and carbon balance.     

全球气候变化对陆地植物碳氮磷生态化学计量学特征的影响

全球气候变化背景下生物地球化学循环的响应规律和陆地植物适应对策已受到广泛关注.本文在分析气候变暖和降水变化对不同生态系统植物C∶N∶P的影响、CO₂浓度升高对不同光合途径物种元素的影响,以及氮沉降对土壤 植物元素影响的短期和长期效应等基础上,从植物生理特性和土壤有效营养元素变化等方面揭示了其可能存在的内在机理,以期为研究C、N、P化学元素在土壤 植物之间传递与调节机制、陆地生态系统结构和功能,以及生物地球化学元素循环对气候变化的响应提供理论依据.最后提出了该领域研究中存在的问题及对今后研究的展望.     

Contrasting mechanisms underlie short‐ and longer‐term soil respiration responses to experimental warming in a dryland ecosystem

Soil carbon losses to the atmosphere through soil respiration are expected to rise with ongoing temperature increases, but available evidence from mesic biomes suggests that such response disappears after a few years of experimental warming. However, there is lack of empirical basis for these temporal dynamics in soil respiration responses, and for the mechanisms underlying them, in drylands, which collectively form the largest biome on Earth and store 32% of the global soil organic carbon pool. We coupled data from a 10 year warming experiment in a biocrust‐dominated dryland ecosystem with laboratory incubations to confront 0–2 years (short‐term hereafter) versus 8–10 years (longer‐term hereafter) soil respiration responses to warming. Our results showed that increased soil respiration rates with short‐term warming observed in areas with high biocrust cover returned to control levels in the longer‐term. Warming‐induced increases in soil temperature were the main drivers of the short‐term soil respiration responses, whereas longer‐term soil respiration responses to warming were primarily driven by thermal acclimation and warming‐induced reductions in biocrust cover. Our results highlight the importance of evaluating short‐ and longer‐term soil respiration responses to warming as a mean to reduce the uncertainty in predicting the soil carbon–climate feedback in drylands.     

Decadal warming causes a consistent and persistent shift from heterotrophic to autotrophic respiration in contrasting permafrost ecosystems

Soil carbon in permafrost ecosystems has the potential to become a major positive feedback to climate change if permafrost thaw increases heterotrophic decomposition. However, warming can also stimulate autotrophic production leading to increased ecosystem carbon storage—a negative climate change feedback. Few studies partitioning ecosystem respiration examine decadal warming effects or compare responses among ecosystems. Here, we first examined how 11 years of warming during different seasons affected autotrophic and heterotrophic respiration in a bryophyte‐dominated peatland in Abisko, Sweden. We used natural abundance radiocarbon to partition ecosystem respiration into autotrophic respiration, associated with production, and heterotrophic decomposition. Summertime warming decreased the age of carbon respired by the ecosystem due to increased proportional contributions from autotrophic and young soil respiration and decreased proportional contributions from old soil. Summertime warming's large effect was due to not only warmer air temperatures during the growing season, but also to warmer deep soils year‐round. Second, we compared ecosystem respiration responses between two contrasting ecosystems, the Abisko peatland and a tussock‐dominated tundra in Healy, Alaska. Each ecosystem had two different timescales of warming (<5 years and over a decade). Despite the Abisko peatland having greater ecosystem respiration and larger contributions from heterotrophic respiration than the Healy tundra, both systems responded consistently to short‐ and long‐term warming with increased respiration, increased autotrophic contributions to ecosystem respiration, and increased ratios of autotrophic to heterotrophic respiration. We did not detect an increase in old soil carbon losses with warming at either site. If increased autotrophic respiration is balanced by increased primary production, as is the case in the Healy tundra, warming will not cause these ecosystems to become growing season carbon sources. Warming instead causes a persistent shift from heterotrophic to more autotrophic control of the growing season carbon cycle in these carbon‐rich permafrost ecosystems.     

Thermal acclimation modulates the impacts of temperature and enrichment on trophic interaction strengths and population dynamics

Global change affects individual phenotypes and biotic interactions, which can have cascading effects up to the ecosystem level. However, the role of environmentally induced phenotypic plasticity in species interactions is poorly understood, leaving a substantial gap in our knowledge of the impacts of global change on ecosystems. Using a cladoceran–dragonfly system, we experimentally investigated the effects of thermal acclimation, acute temperature change and enrichment on predator functional response and metabolic rate. Using our experimental data, we next parameterized a population dynamics model to determine the consequences of these effects on trophic interaction strength and food‐chain stability. We found that (1) predation and metabolic rates of the dragonfly larvae increase with acute warming, (2) warm‐acclimated larvae have a higher maximum predation rate than cold‐acclimated ones, and (3) long‐term interaction strength increases with enrichment but decreases with both acclimation and acute temperatures. Overall, our experimental results show that thermal acclimation can buffer negative impacts of environmental change on predators and increase food‐web stability and persistence. We conclude that the effect of acclimation and, more generally, phenotypic plasticity on trophic interactions should not be overlooked if we aim to understand the effects of climate change and enrichment on species interaction strength and food‐web stability.     

Assessing ecosystem threats from global and regional change: hierarchical modeling of risk to sagebrush ecosystems from climate change, land use and invasive species in Nevada, USA

Global change poses significant challenges for ecosystem conservation. At regional scales, climate change may lead to extensive shifts in species distributions and widespread extirpations or extinctions. At landscape scales, land use and invasive species disrupt ecosystem function and reduce species richness. However, a lack of spatially explicit models of risk to ecosystems makes it difficult for science to inform conservation planning and land management. Here, I model risk to sagebrush ( Artemisia spp.) ecosystems in the state of Nevada, USA from climate change, land use/land cover change, and species invasion. Risk from climate change is based on an ensemble of 10 atmosphere-ocean general circulation model (AOGCM) projections applied to two bioclimatic envelope models (Mahalanobis distance and Maxent). Risk from land use is based on the distribution of roads, agriculture, and powerlines, and on the spatial relationships between land use and probability of cheatgrass Bromus tectorum invasion in Nevada. Risk from land cover change is based on probability and extent of pinyon-juniper ( Pinus monophylla; Juniperus spp.) woodland expansion. Climate change is most likely to negatively impact sagebrush ecosystems at the edges of its current range, particularly in southern Nevada, southern Utah, and eastern Washington. Risk from land use and woodland expansion is pervasive throughout Nevada, while cheatgrass invasion is most problematic in the northern part of the state. Cumulatively, these changes pose major challenges for conservation of sagebrush and sagebrush obligate species. This type of comprehensive assessment of ecosystem risk provides managers with spatially explicit tools important for conservation planning.     

Modelling climate‐change‐induced nonlinear thresholds in cephalopod population dynamics

A significant global challenge lies in our current inability to anticipate, and therefore prepare for, critical ecological thresholds (i.e. tipping points in ecosystems). This deficit stems largely from an inadequate understanding of the many complex interactions between species and the environment at the ecosystem level, and the paucity of mechanistic models relating environment to population dynamics at the species level. In marine ecosystems, abundant, short‐lived and fast‐growing species such as anchovies or squids, consistently function as ‘keystone’ groups whose population dynamics affect entire ecosystems. Increasing exploitation coupled with climate change impacts has the potential to affect these ecological groups and consequently, the entire marine ecosystem. There are currently very few models that predict the impact of climate change on these keystone groups. Here we use a combination of individual‐based bioenergetics and stage‐structured population models to characterize the fundamental capacity of cephalopods to respond to climate change. We demonstrate the potential for, and mechanisms behind, two unfavourable climate‐change‐induced thresholds in future population dynamics. Although one threshold was the direct consequence of a decrease in incubation time caused by ocean warming, the other threshold was linked to survivorship, implying the possibility of management through a modification of fishing mortality. Additional substantive changes in phenology were also predicted, with a possible loss in population resilience. Our results demonstrate the feasibility of predicting complex nonlinear dynamics with a reasonably simplistic mechanistic model, and highlight the necessity of developing such approaches for other species if attempts to moderate the impact of climate change on natural resources are to be effective.     

Effects of climate warming on carbon fluxes in grasslands— A global meta‐analysis

Climate warming will affect terrestrial ecosystems in many ways, and warming‐induced changes in terrestrial carbon (C) cycling could accelerate or slow future warming. So far, warming experiments have shown a wide range of C flux responses, across and within biome types. However, past meta‐analyses of C flux responses have lacked sufficient sample size to discern relative responses for a given biome type. For instance grasslands contribute greatly to global terrestrial C fluxes, and to date grassland warming experiments provide the opportunity to evaluate concurrent responses of both plant and soil C fluxes. Here, we compiled data from 70 sites (in total 622 observations) to evaluate the response of C fluxes to experimental warming across three grassland types (cold, temperate, and semi‐arid), warming methods, and short (≤3 years) and longer‐term (>3 years) experiment lengths. Overall, our meta‐analysis revealed that experimental warming stimulated C fluxes in grassland ecosystems with regard to both plant production (e.g., net primary productivity (NPP) 15.4%; aboveground NPP (ANPP) by 7.6%, belowground NPP (BNPP) by 11.6%) and soil respiration (Rs) (9.5%). However, the magnitude of C flux stimulation varied significantly across cold, temperate and semi‐arid grasslands, in that responses for most C fluxes were larger in cold than temperate or semi‐arid ecosystems. In semi‐arid and temperate grasslands, ecosystem respiration (Reco) was more sensitive to warming than gross primary productivity (GPP), while the opposite was observed for cold grasslands, where warming produced a net increase in whole‐ecosystem C storage. However, the stimulatory effect of warming on ANPP and Rs observed in short‐term studies (≤3 years) in both cold and temperate grasslands disappeared in longer‐term experiments (>3 years). These results highlight the importance of conducting long‐term warming experiments, and in examining responses across a wide range of climate.     

Climate change impacts on wildlife in a High Arctic archipelago – Svalbard,Norway

The Arctic is warming more rapidly than other region on the planet, and the northern Barents Sea, including the Svalbard Archipelago, is experiencing the fastest temperature increases within the circumpolar Arctic, along with the highest rate of sea ice loss. These physical changes are affecting a broad array of resident Arctic organisms as well as some migrants that occupy the region seasonally. Herein, evidence of climate change impacts on terrestrial and marine wildlife in Svalbard is reviewed, with a focus on bird and mammal species. In the terrestrial ecosystem, increased winter air temperatures and concomitant increases in the frequency of ‘rain‐on‐snow’ events are one of the most important facets of climate change with respect to impacts on flora and fauna. Winter rain creates ice that blocks access to food for herbivores and synchronizes the population dynamics of the herbivore–predator guild. In the marine ecosystem, increases in sea temperature and reductions in sea ice are influencing the entire food web. These changes are affecting the foraging and breeding ecology of most marine birds and mammals and are associated with an increase in abundance of several temperate fish, seabird and marine mammal species. Our review indicates that even though a few species are benefiting from a warming climate, most Arctic endemic species in Svalbard are experiencing negative consequences induced by the warming environment. Our review emphasizes the tight relationships between the marine and terrestrial ecosystems in this High Arctic archipelago. Detecting changes in trophic relationships within and between these ecosystems requires long‐term (multidecadal) demographic, population‐ and ecosystem‐based monitoring, the results of which are necessary to set appropriate conservation priorities in relation to climate warming.     

Thermal acclimation in widespread heterotrophic soil microbes

Respiration by plants and microorganisms is primarily responsible for mediating carbon exchanges between the biosphere and atmosphere. Climate warming has the potential to influence the activity of these organisms, regulating exchanges between carbon pools. Physiological ‘down‐regulation’ of warm‐adapted species (acclimation) could ameliorate the predicted respiratory losses of soil carbon under climate change scenarios, but unlike plants and symbiotic microbes, the existence of this phenomenon in heterotrophic soil microbes remains controversial. Previous studies using complex soil microbial communities are unable to distinguish physiological acclimation from other community‐scale adjustments. We explored the temperature‐sensitivity of individual saprotrophic basidiomycete fungi growing in agar, showing definitively that these widespread heterotrophic fungi can acclimate to temperature. In almost all cases, the warm‐acclimated individuals had lower growth and respiration rates at intermediate temperatures than cold‐acclimated isolates. Inclusion of such microbial physiological responses to warming is essential to enhance the robustness of global climate‐ecosystem carbon models.     

Convergent ecosystem responses to 23‐year ambient and manipulated warming link advancing snowmelt and shrub encroachment to transient and long‐term climate–soil carbon feedback

Ecosystem responses to climate change can exert positive or negative feedbacks on climate, mediated in part by slow‐moving factors such as shifts in vegetation community composition. Long‐term experimental manipulations can be used to examine such ecosystem responses, but they also present another opportunity: inferring the extent to which contemporary climate change is responsible for slow changes in ecosystems under ambient conditions. Here, using 23 years of data, we document a shift from nonwoody to woody vegetation and a loss of soil carbon in ambient plots and show that these changes track previously shown similar but faster changes under experimental warming. This allows us to infer that climate change is the cause of the observed shifts in ambient vegetation and soil carbon and that the vegetation responses mediate the observed changes in soil carbon. Our findings demonstrate the realism of an experimental manipulation, allow attribution of a climate cause to observed ambient ecosystem changes, and demonstrate how a combination of long‐term study of ambient and experimental responses to warming can identify mechanistic drivers needed for realistic predictions of the conditions under which ecosystems are likely to become carbon sources or sinks over varying timescales.     

Projecting climate change impacts on species distributions in megadiverse South African Cape and Southwest Australian Floristic Regions: Opportunities and challenges

Increasing evidence shows that anthropogenic climate change is affecting biodiversity. Reducing or stabilizing greenhouse gas emissions may slow global warming, but past emissions will continue to contribute to further unavoidable warming for more than a century. With obvious signs of difficulties in achieving effective mitigation worldwide in the short term at least, sound scientific predictions of future impacts on biodiversity will be required to guide conservation planning and adaptation. This is especially true in Mediterranean type ecosystems that are projected to be among the most significantly affected by anthropogenic climate change, and show the highest levels of confidence in rainfall projections. Multiple methods are available for projecting the consequences of climate change on the main unit of interest – the species – with each method having strengths and weaknesses. Species distribution models (SDMs) are increasingly applied for forecasting climate change impacts on species geographic ranges. Aggregation of models for different species allows inferences of impacts on biodiversity, though excluding the effects of species interactions. The modelling approach is based on several further assumptions and projections and should be treated cautiously. In the absence of comparable approaches that address large numbers of species, SDMs remain valuable in estimating the vulnerability of species. In this review we discuss the application of SDMs in predicting the impacts of climate change on biodiversity with special reference to the species‐rich South West Australian Floristic Region and South African Cape Floristic Region. We discuss the advantages and challenges in applying SDMs in biodiverse regions with high levels of endemicity, and how a similar biogeographical history in both regions may assist us in understanding their vulnerability to climate change. We suggest how the process of predicting the impacts of climate change on biodiversity with SDMs can be improved and emphasize the role of field monitoring and experiments in validating the predictions of SDMs.     

Thermal plasticity of photosynthesis: the role of acclimation in forest responses to a warming climate

The increasing air temperatures central to climate change predictions have the potential to alter forest ecosystem function and structure by exceeding temperatures optimal for carbon gain. Such changes are projected to threaten survival of sensitive species, leading to local extinctions, range migrations, and altered forest composition. This study investigated photosynthetic sensitivity to temperature and the potential for acclimation in relation to the climatic provenance of five species of deciduous trees, Liquidambar styraciflua, Quercus rubra, Quercus falcata, Betula alleghaniensis, and Populus grandidentata. Open‐top chambers supplied three levels of warming (+0, +2, and +4 °C above ambient) over 3 years, tracking natural temperature variability. Optimal temperature for CO₂ assimilation was strongly correlated with daytime temperature in all treatments, but assimilation rates at those optima were comparable. Adjustment of thermal optima was confirmed in all species, whether temperatures varied with season or treatment, and regardless of climate in the species' range or provenance of the plant material. Temperature optima from 17° to 34° were observed. Across species, acclimation potentials varied from 0.55 °C to 1.07 °C per degree change in daytime temperature. Responses to the temperature manipulation were not different from the seasonal acclimation observed in mature indigenous trees, suggesting that photosynthetic responses should not be modeled using static temperature functions, but should incorporate an adjustment to account for acclimation. The high degree of homeostasis observed indicates that direct impacts of climatic warming on forest productivity, species survival, and range limits may be less than predicted by existing models.     

Long‐term monitoring of an amphibian community after a climate change‐ and infectious disease‐driven species extirpation

Infectious disease and climate change are considered major threats to biodiversity and act as drivers behind the global amphibian decline. This is, to a large extent, based on short‐term studies that are designed to detect the immediate and strongest biodiversity responses to a threatening process. What few long‐term studies are available, although typically focused on single species, report outcomes that often diverge significantly from the short‐term species responses. Here, we report the results of an 18‐year survey of an amphibian community exposed to both climate warming and the emergence of lethal chytridiomycosis. Our study shows that the impacts of infectious disease are ongoing but restricted to two out of nine species that form the community, despite the fact all species can become infected with the fungus. Climate warming appears to be affecting four out of the nine species, but the response of three of these is an increase in abundance. Our study supports a decreasing role of infectious disease on the community, and an increasing and currently positive effect of climate warming. We caution that if the warming trends continue, the net positive effect will turn negative as amphibian breeding habitat becomes unavailable as water bodies dry, a pattern that already may be underway.     

Population trends of European common birds are predicted by characteristics of their climatic niche

Temperate species are projected to experience the greatest temperature increases across a range of modelled climate change scenarios, and climate warming has been linked to geographical range and population changes of individual species at such latitudes. However, beyond the multiple modelling approaches, we lack empirical evidence of contemporary climate change impacts on populations in broad taxonomic groups and at continental scales. Identifying reliable predictors of species resilience or susceptibility to climate warming is of critical importance in assessing potential risks to species, ecosystems and ecosystem services. Here we analysed long‐term trends of 110 common breeding birds across Europe (20 countries), to identify climate niche characteristics, adjusted to other environmental and life history traits, that predict large‐scale population changes accounting for phylogenetic relatedness among species. Beyond the now well‐documented decline of farmland specialists, we found that species with the lowest thermal maxima (as the mean spring and summer temperature of the hottest part of the breeding distribution in Europe) showed the sharpest declines between 1980 and 2005. Thermal maximum predicted the recent trends independently of other potential predictors. This study emphasizes the need to account for both land‐use and climate changes to assess the fate of species. Moreover, we highlight that thermal maximum appears as a reliable and simple predictor of the long‐term trends of such endothermic species facing climate change.     

'As Earth's testimonies tell': wilderness conservation in a changing world

Too often, wilderness conservation ignores a temporal perspective greater than the past 50 years, yet a long‐term perspective (centuries to millennia) reveals the dynamic nature of many ecosystems. Analysis of fossil pollen, charcoal and stable isotopes, combined with historical analyses and archaeology can reveal how ongoing interactions between climatic change, human activities and other disturbances have shaped today's landscapes over thousands of years. This interdisciplinary approach can inform wilderness conservation and also contribute to interpreting current trends and predicting how ecosystems might respond to future climate change. In this paper, we review literature that reveals how increasing collaboration among palaeoecologists, archaeologists, historians, anthropologists and ecologists is improving understanding of ecological complexity. Drawing on case studies from forested and non‐forested ecosystems in Europe, the Americas, Africa and Australia, we discuss how this integrated approach can inform wilderness conservation and ecosystem management.     

Responses of High Arctic wet sedge tundra to climate warming since 1980

The global climate is changing rapidly and Arctic regions are showing responses to recent warming. Responses of tundra ecosystems to climate change have been examined primarily through short‐term experimental manipulations, with few studies of long‐term ambient change. We investigated changes in above‐ and belowground biomass of wet sedge tundra to the warming climate of the Canadian High Arctic over the past 25 years. Aboveground standing crop was harvested from five sedge meadow sites and belowground biomass was sampled from one of the sites in the early 1980s and in 2005 using the same methods. Aboveground biomass was on average 158% greater in 2005 than in the early 1980s. The belowground biomass was also much greater in 2005: root biomass increased by 67% and rhizome biomass by 139% since the early 1980s. Dominant species from each functional group (graminoids, shrubs and forbs) showed significant increases in aboveground biomass. Responsive species included the dominant sedge species Carex aquatilis stans, C. membranacea, and Eriophorum angustifolium, as well as the dwarf shrub Salix arctica and the forb Polygonum viviparum. However, diversity measures were not different between the sample years. The greater biomass correlated strongly with increased annual and summer temperatures over the same time period, and was significantly greater than the annual variation in biomass measured in 1980–1983. Increased decomposition and mineralization rates, stimulated by warmer soils, were likely a major cause of the elevated productivity, as no differences in the mass of litter were found between sample periods. Our results are corroborated by published short‐term experimental studies, conducted in other wet sedge tundra communities which link warming and fertilization with elevated decomposition, mineralization and tundra productivity. We believe that this is the first study to show responses in High Arctic wet sedge tundra to recent climate change.     

The effects of climate charge on land-atmosphere feedbacks in arctic tundra regions

Recently reported high-latitude warming has the potential to affect arctic ecosystem structure and function in the short and long term. Arctic ecosystems are known sources of atmospheric CH(4), and recent CO(2) flux measurements indicate that these ecosystems are now, at least regionally, net sources of atmospheric Co(2). It appears that over the short term (decades to centuries), arctic ecosystems may represent a positive feedback on global atmospheric CO(2) concentrations and associated greenhouse gas-Induced climate change. In addition, short-term feedbacks may be large enough to affect both local and global surface temperatures. Over the long term, changes in the structure, function and composition of arctic ecosystems may increase C accumulation relatively more than the amount lost, thus restoring the sink status of arctic ecosystems.     

Impacts of anthropogenic climate change on tropical montane forests: an appraisal of the evidence

In spite of their small global area and restricted distributions, tropical montane forests (TMFs) are biodiversity hotspots and important ecosystem services providers, but are also highly vulnerable to climate change. To protect and preserve these ecosystems better, it is crucial to inform the design and implementation of conservation policies with the best available scientific evidence, and to identify knowledge gaps and future research needs. We conducted a systematic review and an appraisal of evidence quality to assess the impacts of climate change on TMFs. We identified several skews and shortcomings. Experimental study designs with controls and long-term (≥10 years) data sets provide the most reliable evidence, but were rare and gave an incomplete understanding of climate change impacts on TMFs. Most studies were based on predictive modelling approaches, short-term (<10 years) and cross-sectional study designs. Although these methods provide moderate to circumstantial evidence, they can advance our understanding on climate change effects. Current evidence suggests that increasing temperatures and rising cloud levels have caused distributional shifts (mainly upslope) of montane biota, leading to alterations in biodiversity and ecological functions. Neotropical TMFs were the best studied, thus the knowledge derived there can serve as a proxy for climate change responses in under-studied regions elsewhere. Most studies focused on vascular plants, birds, amphibians and insects, with other taxonomic groups poorly represented. Most ecological studies were conducted at species or community levels, with a marked paucity of genetic studies, limiting understanding of the adaptive capacity of TMF biota. We thus highlight the long-term need to widen the methodological, thematic and geographical scope of studies on TMFs under climate change to address these uncertainties. In the short term, however, in-depth research in well-studied regions and advances in computer modelling approaches offer the most reliable sources of information for expeditious conservation action for these threatened forests.