Global Change Biology

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Online ISSN: 1365-2486
Print ISSN: 1354-1013
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Mytilus edulis shell and somatic growth during the experiment. Weekly measured shell length of mussels from the four pCO2 and three feeding treatments (a–c). Shell length growth (SLG) could best be described with the following two factorial linear regression: SLG (μm 7 weeks−1) = 1312.2 (± 150.6) − 0.148 (± 0.06) pCO2 (μatm) + 46.83 (± 3.81) energy supply (J), R2 = 0.66, F(2,81) = 78.4, P < 0.01, means ± SE (a–c). Final shell CaCO3 growth (filled bars) and total organic growth (SFDM + OSC, striped bars, CaCO3: Two-way anova: factor energy: F(2,72) = 40.6, P < 0.01, factor pCO2: F(3,72) = 3.27, P < 0.05, interaction: F(6,72) = 1.07, P < 0.5, total organic: two-way anova: factor energy: F(2,72) = 51.84, P < 0.01, factor pCO2: F(3,72) = 1.19, P > 0.05, interaction: F(6,72) = 1.02, P < 0.5 (d). n = 7 for all treatments. Bars represent means ± SD.
Box-whisker-plot of Kiel Fjord pCO2 in 2009/2010 measured at stations 1–4 in surface (light grey, S) and bottom (dark grey, B) water samples. Circles depict individual measurements (a); Time course of surface pHNBS in inner Kiel Fjord (station IF, closed circles) between 2008 and 2010 (data from 2008–2009 adopted from Thomsen et al., 2010) and at station OF during the field experiment (open circles), (b).
Results of the field study (outer Fjord: white, inner Fjord: black): Final mean CaCO3 mass of Mytilus edulis per settlement panel (a), Final shell mass and length of individual M. edulis grown at the outer and inner Fjord (b), POC concentrations (c), Mean values for abiotic conditions during the experimental period at both sites (d–f), Ωaragonite (d), pHtotal (e), pCO2 in μatm (f). Bars represent means ± SD.
Relationship of haemolymph pH (pHHL) of Mytilus edulis and ambient seawater pHNBS (pHSW), pHHL = 0.413 (± 0.08) × pHSW + 4.14 (± 0.64), F(1,60) = 26.5 P < 0.01, means ± SE; each data point represents mean ± SD of n = 5 mussels from one sampling date.
Ocean acidification is expected to decrease calcification rates of bivalves. Nevertheless, in many coastal areas high pCO2 variability is encountered already today. Kiel Fjord (Western Baltic Sea) is a brackish (12-20 g kg(-1) ) and CO2 enriched habitat, but the blue mussel Mytilus edulis dominates the benthic community. In a coupled field and laboratory study we examined the annual pCO2 variability in this habitat and the combined effects of elevated pCO2 and food availability on juvenile M. edulis growth and calcification. In the laboratory experiment, mussel growth and calcification were found to chiefly depend on food supply, with only minor impacts of pCO2 up to 3350 μatm. Kiel Fjord was characterized by strong seasonal pCO2 variability. During summer, maximal pCO2 values of 2500 μatm were observed at the surface and >3000 μatm at the bottom. However, the field growth experiment revealed seven times higher growth and calcification rates of M. edulis at a high pCO2 inner fjord field station (mean pCO2 ca. 1000 μatm) in comparison to a low pCO2 outer fjord station (ca. 600 μatm). In addition, mussels were able to out-compete the barnacle Amphibalanus improvisus at the high pCO2 site. High mussel productivity at the inner fjord site was enabled by higher particulate organic carbon concentrations. Kiel Fjord is highly impacted by eutrophication, which causes bottom water hypoxia and consequently high seawater pCO2 . At the same time, elevated nutrient concentrations increase the energy availability for filter feeding organisms such as mussels. Thus, M. edulis can dominate over a seemingly more acidification resistant species such as A. improvisus. We conclude that benthic stages of M. edulis tolerate high ambient pCO2 when food supply is abundant and that important habitat characteristics such as species interactions and energy availability need to be considered to predict species vulnerability to ocean acidification.
 
Ecosystem models play a crucial role in understanding and evaluating the combined impacts of rising atmospheric CO2 concentration and changing climate on terrestrial ecosystems. However, we are not aware of any studies where the capacity of models to simulate intra- and inter-annual variation in responses to elevated CO2 has been tested against long-term experimental data. Here we tested how well the ecosystem model APSIM/AgPasture was able to simulate the results from a Free Air Carbon Dioxide Enrichment (FACE) experiment on grazed pasture. At this FACE site, during 11 years of CO2 enrichment, a wide range in annual plant production response to CO2 (-6 to +28%) was observed. As well as running the full model, which includes three plant CO2 response functions (plant photosynthesis, nitrogen (N) demand and stomatal conductance), we also tested the influence of these three functions on model predictions. Model/data comparisons showed that: 1) overall the model over-predicted the mean annual plant production response to CO2 (18.5% cf 13.1%) largely because years with small or negative responses to CO2 were not well simulated, 2) in general seasonal and inter-annual variation in plant production responses to elevated CO2 were well represented by the model, 3) the observed CO2 enhancement in overall mean legume content was well simulated but year-to-year variation in legume content was poorly captured by the model, 4) the best fit of the model to the data required all three CO2 response functions to be invoked, 5) using actual legume content and reduced N fixation rate under elevated CO2 in the model provided the best fit to the experimental data. We conclude that in temperate grasslands the N dynamics (particularly the legume content and N fixation activity) play a critical role in pasture production responses to elevated CO2 , and are processes for model improvement. This article is protected by copyright. All rights reserved.
 
Dot density graph of global methane sources. Horizontal lines are the median for each category. Anthropogenic sources include rice fields and natural sources include freshwater aquatic ecosystems and wetlands, but they are also presented separately. Data for figure are in Tables S1 and S2, and when ranges are given, the mean value was used.
Methanogenic community structure among different ecosystems or root-associated niches within an ecosystem. Y-axis represents the fraction (%) of each group in 16S rRNA gene-derived clone libraries. Methanogenic groups were classified at the family level plus uncultured clusters. Data were summarized from previous reports taken as representative samples of: acidic bog (Cadillo-Quiroz et al., 2006), minerotrophic fen (Cadillo-Quiroz et al., 2008), rice paddy soil (Lueders & Friedrich, 2000), temperate wetland (Castro et al., 2004), lakes (Borrel et al., 2011), root surface (rhizoplane) in acidic bog (Cadillo-Quiroz et al., 2010), and root-associated soil (rhizosphere) in rice paddy (Chin et al., 2004).
Understanding the dynamics of methane (CH4) emissions is of paramount importance because CH4 has 25 times the global warming potential of carbon dioxide (CO2) and is currently the second most important anthropogenic greenhouse gas. Wetlands are the single largest natural CH4 source with median emissions from published studies of 164 Tgyr1, which is about a third of total global emissions. We provide a perspective on important new frontiers in obtaining a better understanding of CH4 dynamics in natural systems, with a focus on wetlands. One of the most exciting recent developments in this field is the attempt to integrate the different methodologies and spatial scales of biogeochemistry, molecular microbiology, and modeling, and thus this is a major focus of this review. Our specific objectives are to provide an up-to-date synthesis of estimates of global CH4 emissions from wetlands and other freshwater aquatic ecosystems, briefly summarize major biogeophysical controls over CH4 emissions from wetlands, suggest new frontiers in CH4 biogeochemistry, examine relationships between methanogen community structure and CH4 dynamics in situ, and to review the current generation of CH4 models. We highlight throughout some of the most pressing issues concerning global change and feedbacks on CH4 emissions from natural ecosystems. Major uncertainties in estimating current and future CH4 emissions from natural ecosystems include the following: (i) A number of important controls over CH4 production, consumption, and transport have not been, or are inadequately, incorporated into existing CH4 biogeochemistry models. (ii) Significant errors in regional and global emission estimates are derived from large spatial-scale extrapolations from highly heterogeneous and often poorly mapped wetland complexes. (iii) The limited number of observations of CH4 fluxes and their associated environmental variables loosely constrains the parameterization of process-based biogeochemistry models.
 
Location of the six study sites along the Transverse Volcanic Axis of Mexico. Sites were distributed from the side of the Ocean Pacific to the side of Gulf of Mexico, along the parallel ~19°N, with similar conditions of soils, topography, and vegetation cover.
Trends in tree growth for the periods: 1910–1950 (green circles), 1950–1980 (red circles), and 1980–2010 (black circles). Circles represent average Basal Area Increments for both species at each study site. Lines represent significant relationships between growth rates and calendar year. Recent growth decline is statistically significant for all sites and species, except for pine at the NC site. Only trees with age ranging between 50 and 100 years (n = 108) were used. Trends in growth for older and younger trees are presented in Appendix S1.
Average tree ring δ13C (a and b), δ18O (c and d), and estimated source to product fractionation – ∆13C (e and f) – and intrinsic water use efficiency – iWUE (g and h) – at each study site. Dotted line in (b) shows changes in atmospheric δ13C values over time. Estimates of ∆13C and iWUE were corrected for changes atmospheric CO2 concentration and 13C composition following Eqns (3)-(5). Asterisks indicated significant differences between species at each site according to the t-test (P < 0.05).
Average tree ring d 13 C (a and b), d 18 O (c and d), and estimated source to product fractionation-Δ 13 C (e and f)-and intrinsic water use efficiency-iWUE (g and h)-at each study site. Dotted line in (b) shows changes in atmospheric d 13 C values over time. Estimates of Δ 13 C and iWUE were corrected for changes atmospheric CO 2 concentration and 13 C composition following Eqns (3-5). Asterisks indicated significant differences between species at each site according to the t-test (P < 0.05). 
Changes in intrinsic water use efficiency (iWUE) at all study sites. Black circles (a, c, e, g, i and k) correspond to Abies religiosa along the six mountains in the study transect from the Pacific to the Gulf of Mexico. White circles (b, d, f, h, j and l) are for Pinus hartwegii, in the same direction of the transect. In all cases significant increases in iWUE were observed over time in response to rising atmospheric CO2 levels.
Human-induced changes in atmospheric composition are expected to affect primary productivity across terrestrial biomes. Recent changes in productivity have been observed in many forest ecosystems, but low-latitude upper tree line forests remain to be investigated. Here we use dendrochronological methods and isotopic analysis to examine changes in productivity, and their physiological basis, in Abies religiosa (Ar) and Pinus hartwegii (Ph) trees growing in high-elevation forests of central Mexico. Six sites were selected across a longitudinal transect (Transverse Volcanic Axis), from the Pacific Ocean toward the Gulf of Mexico, where mature dominant trees were sampled at altitudes ranging from 3200 to 4000m asl. A total of 60 Ar and 84 Ph trees were analyzed to describe changes in growth (annual-resolution) and isotopic composition (decadal-resolutions) since the early 1900s. Our results show an initial widespread increase in basal area increment (BAI) during the first half of the past century. However, BAI has decreased significantly since the 1950s with accentuated decline after the 1980s in both species and across sites. We found a consistent reduction in atmosphere to wood (13) C discrimination, resulting from increasing water use efficiency (20-60%), coinciding with rising atmospheric CO2 . Changes in (13) C discrimination were not followed, however, by shifts in tree ring δ(18) O, indicating site- and species-specific differences in water source or uptake strategy. Our results indicate that CO2 stimulation has not been enough to counteract warming induced drought stress, but other stressors, such as progressive nutrient limitation, could also have contributed to growth decline. Future studies should explore the distinct role of resource limitation (water vs. nutrients) in modulating the response of high-elevation ecosystems to atmospheric change. © 2013 Blackwell Publishing Ltd.
 
Ecosystem respiration (Reco ) is one of the largest terrestrial carbon (C) fluxes. The effect of climate change on Reco depends on the responses of its autotrophic and heterotrophic components. How autotrophic and heterotrophic respiration sources respond to climate change is especially important in ecosystems underlain by permafrost. Permafrost ecosystems contain vast stores of soil C (1672 Pg) and are located in northern latitudes where climate change is accelerated. Warming will cause a positive feedback to climate change if heterotrophic respiration increases without corresponding increases in primary production. We quantified the response of autotrophic and heterotrophic respiration to permafrost thaw across the 2008 and 2009 growing seasons. We partitioned Reco using Δ(14) C and δ(13) C into four sources-two autotrophic (above - and belowground plant structures) and two heterotrophic (young and old soil). We sampled the Δ(14) C and δ(13) C of sources using incubations and the Δ(14) C and δ(13) C of Reco using field measurements. We then used a Bayesian mixing model to solve for the most likely contributions of each source to Reco . Autotrophic respiration ranged from 40 to 70% of Reco and was greatest at the height of the growing season. Old soil heterotrophic respiration ranged from 6 to 18% of Reco and was greatest where permafrost thaw was deepest. Overall, growing season fluxes of autotrophic and old soil heterotrophic respiration increased as permafrost thaw deepened. Areas with greater thaw also had the greatest primary production. Warming in permafrost ecosystems therefore leads to increased plant and old soil respiration that is initially compensated by increased net primary productivity. However, barring large shifts in plant community composition, future increases in old soil respiration will likely outpace productivity, resulting in a positive feedback to climate change.
 
Location of all study sites. Size of the dot represents the original number of species in each community.
Shifting size distributions over time for Plethodon cheoah (a), P. cinerus (b), P. cylindraceus (c), P. jordani (d), P. ventralis (e), and P. yonahlossee (f). The number of sites sampled in each decade is represented by the number in parentheses above the sample size of animals measured for that time period.
Grand mean changes in standardized difference in mean body size per generation, relative to within-population standard deviation for populations in areas that have become colder and wetter (blue), warmer and drier (red), and either colder and drier or warmer and wetter (khaki). To compare among species and populations with different generation times, we converted body size change into Haldane Ratios. The greatest body size reductions, as indicated by Haldane Ratios, were found in populations that experienced both an increase in temperature and a decrease in precipitation (95% CRI = 0.732, 6.837). (b) Spatial distribution of actual climate trends during the study period; areas with the darkest reds experienced the greatest amount of both warming and drying, blue colors are areas that have become both colder and wetter (data from Girvetz et al., 2009).
Results of modeling annual activity (upper row) and annual energy expenditure (bottom row) for a 10 g Plethodon at Catoctin Mountain N.P., MD (FDR), Mountain Lake area, VA (ATW), and Mt. Rogers National Recreation Area, VA (LIM).
Reduction in body size is a major response to climate change, yet evidence in globally imperiled amphibians is lacking. Shifts in average population body size could indicate either plasticity in the growth response to changing climates through changes in allocation and energetics, or through selection for decreased size where energy is limiting. We compared historic and contemporary size measurements in 15 Plethodon species from 102 populations (9450 individuals) and found that six species exhibited significant reductions in body size over 55 years. Biophysical models, accounting for actual changes in moisture and air temperature over that period, showed a 7.1-7.9% increase in metabolic expenditure at three latitudes but showed no change in annual duration of activity. Reduced size was greatest at southern latitudes in regions experiencing the greatest drying and warming. Our results are consistent with a plastic response of body size to climate change through reductions in body size as mediated through increased metabolism. These rapid reductions in body size over the past few decades have significance for the susceptibility of amphibians to environmental change, and relevance for whether adaptation can keep pace with climate change in the future.
 
Model predictions of extinction risks from anthropogenic climate change are dire, but still overly simplistic. To reliably predict at-risk species we need to know which species are currently responding, which are not, and what traits are mediating the responses. For mammals, we have yet to identify overarching physiological, behavioral, or biogeographic traits determining species' responses to climate change, but they must exist. To date, 73 mammal species in North America and eight additional species worldwide have been assessed for responses to climate change, including local extirpations, range contractions and shifts, decreased abundance, phenological shifts, morphological or genetic changes. Only 52% of those species have responded as expected, 7% responded opposite to expectations, and the remaining 41% have not responded. Which mammals are and are not responding to climate change is mediated predominantly by body size and activity times (phylogenetic multivariate logistic regressions, P < 0.0001). Large mammals respond more, for example, an elk is 27 times more likely to respond to climate change than a shrew. Obligate diurnal and nocturnal mammals are more than twice as likely to respond as mammals with flexible activity times (P < 0.0001). Among the other traits examined, species with higher latitudinal and elevational ranges were more likely to respond to climate change in some analyses, whereas hibernation, heterothermy, burrowing, nesting, and study location did not influence responses. These results indicate that some mammal species can behaviorally escape climate change whereas others cannot, analogous to paleontology's climate sheltering hypothesis. Including body size and activity flexibility traits into future extinction risk forecasts should substantially improve their predictive utility for conservation and management.
 
Enhanced soil respiration in response to global warming may substantially increase atmospheric CO2 concentrations above the anthropogenic contribution, depending on the mechanisms underlying the temperature sensitivity of soil respiration. Here, we compared short-term and seasonal responses of soil respiration to a shifting thermal environment and variable substrate availability via laboratory incubations. To analyze the data from incubations, we implemented a novel process-based model of soil respiration in a hierarchical Bayesian framework. Our process model combined a Michaelis-Menten-type equation of substrate availability and microbial biomass with an Arrhenius-type nonlinear temperature response function. We tested the competing hypotheses that apparent thermal acclimation of soil respiration can be explained by depletion of labile substrates in warmed soils, or that physiological acclimation reduces respiration rates. We demonstrated that short-term apparent acclimation can be induced by substrate depletion, but that decreasing microbial biomass carbon (MBC) is also important, and lower MBC at warmer temperatures is likely due to decreased carbon-use efficiency (CUE). Observed seasonal acclimation of soil respiration was associated with higher CUE and lower basal respiration for summer- vs. winter-collected soils. Whether the observed short-term decrease in CUE or the seasonal acclimation of CUE with increased temperatures dominates the response to long-term warming will have important consequences for soil organic carbon storage.
 
We combine satellite and ground observations during 1950-2011 to study the long-term links between multiple climate (air temperature and cryospheric dynamics) and vegetation (greenness and atmospheric CO2 concentrations) indicators of the growing season of northern ecosystems (>45 degrees N) and their connection with the carbon cycle. During the last three decades, the thermal potential growing season has lengthened by about 10.5days (P<0.01, 1982-2011), which is unprecedented in the context of the past 60years. The overall lengthening has been stronger and more significant in Eurasia (12.6days, P<0.01) than North America (6.2days, P>0.05). The photosynthetic growing season has closely tracked the pace of warming and extension of the potential growing season in spring, but not in autumn when factors such as light and moisture limitation may constrain photosynthesis. The autumnal extension of the photosynthetic growing season since 1982 appears to be about half that of the thermal potential growing season, yielding a smaller lengthening of the photosynthetic growing season (6.7days at the circumpolar scale, P<0.01). Nevertheless, when integrated over the growing season, photosynthetic activity has closely followed the interannual variations and warming trend in cumulative growing season temperatures. This lengthening and intensification of the photosynthetic growing season, manifested principally over Eurasia rather than North America, is associated with a long-term increase (22.2% since 1972, P<0.01) in the amplitude of the CO2 annual cycle at northern latitudes. The springtime extension of the photosynthetic and potential growing seasons has apparently stimulated earlier and stronger net CO2 uptake by northern ecosystems, while the autumnal extension is associated with an earlier net release of CO2 to the atmosphere. These contrasting responses may be critical in determining the impact of continued warming on northern terrestrial ecosystems and the carbon cycle.
 
Location of phenological stations with the four tree species observations in different climate regions.
Parameters of regional unified models and assessment of the models in spatial extrapolation over 1981 to 2005 for the four tree species in the four climate regions
Comparison between observed and predicted leaf unfolding dates over 2006–2009 based on regional unified species-specific models in the humid/subhumid middle temperate region (HSMT), humid/subhumid warm temperate region (HSWT), semiarid middle/warm temperate region (SMWT), and arid middle/warm temperate region (AMWT). (a–d) Salix matsudana, (e–h) Populus simonii, (i–l) Ulmus pumila, (m–p) Prunus armeniaca.
Spatial pattern of linear trends in leaf unfolding dates in northern China over 1960–2009. (a) Salix matsudana, (b) Populus simonii, (c) Ulmus pumila, (d) Prunus armeniaca.
Linear trends in leaf unfolding dates of the four tree species in the humid/subhumid middle temperate region (HSMT), humid/subhumid warm temperate region (HSWT), semiarid middle/warm temperate region (SMWT), and arid middle/warm temperate region (AMWT) over 1960–2009. (a–d) Salix matsudana, (e–h) Populus simonii, (i–l) Ulmus pumila, (m–p) Prunus armeniaca.
Using first leaf unfolding data of Salix matsudana, Populus simonii, Ulmus pumila, and Prunus armeniaca, and daily mean temperature data during the 1981-2005 period at 136 stations in northern China, we fitted unified forcing and chilling phenology models and selected optimum models for each species at each station. Then, we examined performances of each optimum local species-specific model in predicting leaf unfolding dates at all external stations within the corresponding climate region and selected 16 local species-specific models with maximum effective predictions as the regional unified models in different climate regions. Furthermore, we validated the regional unified models using leaf unfolding and daily mean temperature data beyond the time period of model fitting. Finally, we substituted gridded daily mean temperature data into the regional unified models, and reconstructed spatial patterns of leaf unfolding dates of the four tree species across northern China during 1960-2009. At local scales, the unified forcing model shows higher simulation efficiency at 83% of data sets, whereas the unified chilling model indicates higher simulation efficiency at 17% of data sets. Thus, winter temperature increase so far has not yet significantly influenced dormancy and consequent leaf development of deciduous trees in most parts of northern China. Spatial and temporal validation confirmed capability and reliability of regional unified species-specific models in predicting leaf unfolding dates in northern China. Reconstructed leaf unfolding dates of the four tree species show significant advancements by 1.4-1.6 days per decade during 1960-2009 across northern China, which are stronger for the earlier than the later leaf unfolding species. Our findings suggest that the principal characteristics of plant phenology and phenological responses to climate change at regional scales can be captured by phenological and climatic data sets at a few representative locations.
 
Improved crop yield forecasts could enable more effective adaptation to climate variability and change. Here, we explore how to combine historical observations of crop yields and weather with climate model simulations to produce crop yield projections for decision relevant timescales. Firstly, the effects on historical crop yields of improved technology, precipitation and daily maximum temperatures are modelled empirically, accounting for a nonlinear technology trend and interactions between temperature and precipitation, and applied specifically for a case study of maize in France. The relative importance of precipitation variability for maize yields in France has decreased significantly since the 1960s, likely due to increased irrigation. In addition, heat stress is found to be as important for yield as precipitation since around 2000. A significant reduction in maize yield is found for each day with a maximum temperature above 32 °C, in broad agreement with previous estimates. The recent increase in such hot days has likely contributed to the observed yield stagnation. Furthermore, a general method for producing near-term crop yield projections, based on climate model simulations, is developed and utilized. We use projections of future daily maximum temperatures to assess the likely change in yields due to variations in climate. Importantly, we calibrate the climate model projections using observed data to ensure both reliable temperature mean and daily variability characteristics, and demonstrate that these methods work using retrospective predictions. We conclude that, to offset the projected increased daily maximum temperatures over France, improved technology will need to increase base level yields by 12% to be confident about maintaining current levels of yield for the period 2016-2035; the current rate of yield technology increase is not sufficient to meet this target.
 
Circumpolar expansion of tall shrubs and trees into Arctic tundra is widely thought to be occurring as a result of recent climate warming, but little quantitative evidence exists for northern Siberia, which encompasses the world's largest forest-tundra ecotonal belt. We quantified changes in tall shrub and tree canopy cover in eleven, widely-distributed Siberian ecotonal landscapes by comparing very-high-resolution photography from the Cold War-era "Gambit" and "Corona" satellite surveillance systems (1965-1969) with modern imagery. We also analyzed within-landscape patterns of vegetation change to evaluate the susceptibility of different landscape components to tall shrub and tree increase. The total cover of tall shrubs and trees increased in nine of eleven ecotones. In northwest Siberia, alder (Alnus) shrubland cover increased 5.3 - 25.9% in five ecotones. In Taymyr and Yakutia, larch (Larix) cover increased 3.0 - 6.7% within three ecotones, but declined 16.8% at a fourth ecotone due to thaw of ice-rich permafrost. In Chukotka, the total cover of alder and dwarf pine (Pinus) increased 6.1% within one ecotone and was little-changed at a second ecotone. Within most landscapes, shrub and tree increase was linked to specific geomorphic settings, especially those with active disturbance regimes such as permafrost patterned-ground, floodplains, and colluvial hillslopes. Mean summer temperatures increased at most ecotones since the mid-1960s, but rates of shrub and tree canopy cover expansion were not strongly correlated with temperature trends and were better correlated with mean annual precipitation. We conclude that shrub and tree cover is increasing in tundra ecotones across most of northern Siberia, but rates of increase vary widely regionally and at the landscape-scale. Our results indicate that extensive changes can occur within decades in moist, shrub-dominated ecotones, as in northwest Siberia, while changes are likely to occur much more slowly in the highly continental, larch-dominated ecotones of central and eastern Siberia. This article is protected by copyright. All rights reserved.
 
Soil organic matter not only affects soil properties and productivity but also has an essential role in global carbon (C) cycle. We studied changes in the topsoil C content of Finnish croplands using a dataset produced in nationwide soil monitoring. The monitoring network consisting of fields on both mineral and organic soils was established in 1974 and resampled in 1987, 1998, and 2009. Over the monitoring period from 1974 to 2009, cultivated soils showed a continuous decline in C concentration (g kg(-1) ). In organic soils, C concentration decreased at a mean rate of 0.2-0.3% yr(-1) relative to the existing C concentration. In mineral soils, the relative decrease was 0.4% yr(-1) corresponding to a C stock (kg m(-2) ) loss of 220 kg ha(-1) yr(-1) . The change in management practices in last decades toward increasing cultivation of annual crops has contributed to soil C losses noted in this study. The results, however, suggest that the C losses result partly from other processes affecting cultivated soils such as climatic change or the continuing long-term effect of forest clearance. We estimated that Finnish cropland soils store 161 Tg carbon nationwide in the topmost 15 cm of which 117 Tg is in mineral soils. C losses from mineral soils can therefore total up to 0.5 Tg yearly.
 
Does agricultural intensification reduce the area used for agricultural production in Brazil? Census and other data for time periods 1975-1996 and 1996-2006 were processed and analyzed using GIS and statistical tools to investigate whether and if so, how, changes in yield and stocking rate coincide with changes in cropland and pasture area. Complementary medium-resolution data on total farmland area changes were used in a spatially explicit assessment of the land use transitions that occurred in Brazil during 1960-2006. The analyses show that in agriculturally consolidated areas (mainly southern and southeastern Brazil), land use intensification (both on cropland and pastures) coincided with either contraction of both cropland and pasture areas, or cropland expansion at the expense of pastures, both cases resulting in farmland stability or contraction. In contrast, in agricultural frontier areas (i.e. the deforestation zones in central and northern Brazil), land use intensification coincided with expansion of agricultural lands. These observations provide support for the thesis that (i) technological improvements create incentives for expansion in agricultural frontier areas and (ii) farmers are likely to reduce their managed acreage only if land becomes a scarce resource. The spatially explicit examination of land use transitions since 1960 reveals an expansion and gradual movement of the agricultural frontier toward the interior (center-western Cerrado) of Brazil. It also indicates a possible initiation of a reversed trend in line with the forest transition theory, i.e., agricultural contraction and recurring forests in marginally suitable areas in southeastern Brazil, mainly within the Atlantic Forest biome. The significant reduction in deforestation that has taken place recent years, despite rising food commodity prices, indicates that policies put in place to curb conversion of native vegetation to agriculture land might be effective. This can improve the prospects for protecting native vegetation by investing in agricultural intensification. © 2013 Blackwell Publishing Ltd.
 
In this paper we used recent (2010) and herbarium material (1980) of six bryophyte species to assess long-term atmospheric deposition in natural forested areas in northern Spain. For this purpose, tissue nitrogen and carbon content, as well as δ(13) C and δ(15) N signatures of samples of Hypnum cupressiforme, Polytrichastrum formosum, Leucobryum juniperoideum, Rhytidiadelphus loreus, Homalothecium lutescens and Diplophyllum albicans were analysed and comparisons made between years and species. In addition, the usefulness of each of the six species was evaluated. The range of values observed was similar to that in other studies carried out in rural areas. Significantly lower values were found in 2010 for N (H. cupressiforme), δ(15) N (R. loreus and D. albicans), C (R. loreus) and δ(13) C (all except L. juniperoideum). Our natural areas are thus now less influenced by atmospheric pollutants than they were, most probably due to changes in some traditional local activities. Differences were observed between species for all the four parameters studied, so different species must not be analysed together. Finally, R. loreus and H. lutescens seem to be good bioindicators, sensitive even with a few samples, although further studies are needed to corroborate their usefulness. © 2013 Blackwell Publishing Ltd.
 
Non-native, invasive grasses have been linked to altered grass-fire cycles worldwide. Although a few studies have quantified resulting changes in fire activity at local scales, and many have speculated about larger scales, regional alterations to fire regimes remain poorly documented. We assessed the influence of large-scale Bromus tectorum (hereafter cheatgrass) invasion on fire size, duration, spread rate, and interannual variability in comparison to other prominent land cover classes across the Great Basin, USA. We compared regional land cover maps to burned area measured using the Moderate Resolution Imaging Spectroradiometer (MODIS) for 20002009 and to fire extents recorded by the USGS registry of fires from 1980 to 2009. Cheatgrass dominates at least 6% of the central Great Basin (650 000 km(2)). MODIS records show that 13% of these cheatgrass-dominated lands burned, resulting in a fire return interval of 78 years for any given location within cheatgrass. This proportion was more than double the amount burned across all other vegetation types (range: 0.56% burned). During the 1990s, this difference was even more extreme, with cheatgrass burning nearly four times more frequently than any native vegetation type (16% of cheatgrass burned compared to 15% of native vegetation). Cheatgrass was also disproportionately represented in the largest fires, comprising 24% of the land area of the 50 largest fires recorded by MODIS during the 2000s. Furthermore, multi-date fires that burned across multiple vegetation types were significantly more likely to have started in cheatgrass. Finally, cheatgrass fires showed a strong interannual response to wet years, a trend only weakly observed in native vegetation types. These results demonstrate that cheatgrass invasion has substantially altered the regional fire regime. Although this result has been suspected by managers for decades, this study is the first to document recent cheatgrass-driven fire regimes at a regional scale.
 
Based on the crop trial data during 1981-2009 at 57 agricultural experimental stations across the northeastern China Plain (NECP) and the middle and lower reaches of Yangtze River (MLRYR), we investigated how major climate variables had changed and how the climate change had affected crop growth and yield in a setting in which agronomic management practices were taken based on actual weather. We found a significant warming trend during rice growing season, and a general decreasing trend in solar radiation (SRD) in the MLRYR during 1981-2009. Rice transplanting, heading and maturity dates were generally advanced, but the heading and maturity dates of single rice in the MLRYR (YZ_SR) and NECP (NE_SR) were delayed. Climate warming had a negative impact on growth period lengths at about 80% of the investigated stations. Nevertheless, the actual growth period lengths of YZ_SR and NE_SR, as well as the actual length of reproductive growth period (RGP) of early rice in the MLRYR (YZ_ER), were generally prolonged due to adoption of cultivars with longer growth period to obtain higher yield. In contrast, the actual growth period length of late rice in the MLRYR (YZ_LR) was shortened by both climate warming and adoption of early mature cultivars to prevent cold damage and obtain higher yield. During 1981-2009, climate warming and decrease in SRD changed yield of YZ_ER by -0.59% to 2.4%; climate warming during RGP increased yield of YZ_LR by 8.38% to 9.56%; climate warming and decrease in SRD jointly reduced yield of YZ_SR by 7.14% to 9.68%; climate warming and increase in SRD jointly increased yield of NE_SR by 1.01% to 3.29%. Our study suggests rice production in China has been affected by climate change, yet at the same time changes in varieties continue to be the major factor driving yield and growing period trends. This article is protected by copyright. All rights reserved.
 
Land Surface Phenology (LSP) is the most direct representation of intra-annual dynamics of vegetated land surfaces as observed from satellite imagery. LSP plays a key role in characterizing land-surface fluxes, and is central to accurately parameterizing terrestrial biosphere-atmosphere interactions, as well as climate models. In this paper we present an evaluation of Pan-European LSP and its changes over the past 30 years, using the longest continuous record of Normalized Difference Vegetation Index (NDVI) available to date in combination with a landscape-based aggregation scheme. We used indicators of Start-Of-Season, End-Of-Season and Growing Season Length (SOS, EOS and GSL, respectively) for the period 1982-2011 to test for temporal trends in activity of terrestrial vegetation and their spatial distribution. We aggregated pixels into ecologically representative spatial units using the European Landscape Classification (LANMAP) and assessed the relative contribution of spring and autumn phenology. GSL increased significantly by 18-24 days/decade over 18-30% of the land area of Europe, depending on methodology. This trend varied extensively within and between climatic zones and landscape classes. The areas of greatest growing-season lengthening were the Continental and Boreal zones, with hotspots concentrated in southern Fennoscandia, Western Russia and pockets of continental Europe. For the Atlantic and Steppic zones, we found an average shortening of the growing season with hotspots in Western France, the Po valley, and around the Caspian Sea. In many zones, changes in the NDVI-derived end-of-season contributed more to the GSL trend than changes in spring green-up, resulting in asymmetric trends. This underlines the importance of investigating senescence and its underlying processes more closely as a driver of LSP and global change. This article is protected by copyright. All rights reserved.
 
The change in spring phenology is recognized to exert a major influence on carbon balance dynamics in temperate ecosystems. Over the past several decades, several studies focused on shifts in spring phenology; however, large uncertainties still exist, and one understudied source could be the method implemented in retrieving satellite-derived spring phenology. To account for this potential uncertainty, we conducted a multimethod investigation to quantify changes in vegetation green-up date from 1982 to 2010 over temperate China, and to characterize climatic controls on spring phenology. Over temperate China, the five methods estimated that the vegetation green-up onset date advanced, on average, at a rate of 1.3 ± 0.6 days per decade (ranging from 0.4 to 1.9 days per decade) over the last 29 years. Moreover, the sign of the trends in vegetation green-up date derived from the five methods were broadly consistent spatially and for different vegetation types, but with large differences in the magnitude of the trend. The large intermethod variance was notably observed in arid and semiarid vegetation types. Our results also showed that change in vegetation green-up date is more closely correlated with temperature than with precipitation. However, the temperature sensitivity of spring vegetation green-up date became higher as precipitation increased, implying that precipitation is an important regulator of the response of vegetation spring phenology to change in temperature. This intricate linkage between spring phenology and precipitation must be taken into account in current phenological models which are mostly driven by temperature.
 
Understanding spring phenology changes in response to the rapid climate change at biome-level is crucial for projecting regional ecosystem carbon exchange and climate-biosphere interactions. In this study, we assessed the long-term changes and responses to changing climate of the spring phenology in six temperate biomes of China by analyzing the global inventory monitoring and modeling studies (GIMMS) NOAA/AVHRR Normalized Difference Vegetation Index (NDVI) and concurrent mean temperature and precipitation data for 1982-2006. Results show that the spring phenology trends in the six temperate biomes are not continuous throughout the 25 year period. The spring phenology in most areas of the six biomes showed obvious advancing trends (ranging from -0.09 to -0.65 day/yr) during the 1980s and early 1990s, but has subsequently suffered consistently delaying trends (ranging from 0.22 to 1.22 day/yr). Changes in spring (February-April) temperature are the dominating factor governing the pattern of spring vegetation phenology in the temperate biomes of China. The recently delayed spring phenology in these temperate biomes has been mainly triggered by the stalling or reversal of the warming trend in spring temperatures. Results in this study also reveal that precipitation during November-January can explain 16.1% (P < 0.05), 20.9% (P < 0.05) and 14.2% (P < 0.05) of the variations in temperate deciduous forest (TDF), temperate steppe (TS), temperate desert (TD) respectively, highlighting the important role of winter precipitation in regulating changes in the spring vegetation phenology of water-limited biomes.
 
This research refers to an object-based automatic method combined with a national expert validation to produce regional and national forest cover change statistics over Congo Basin. A total of 547 sampling sites systematically distributed over the whole humid forest domain are required to cover the six Central African countries containing tropical moist forest. High resolution imagery is used to accurately estimate not only deforestation and reforestation but also degradation and regeneration. The overall method consists of four steps: (i) image automatic preprocessing and preinterpretation, (ii) interpretation by national expert, (iii) statistic computation and (iv) accuracy assessment. The annual rate of net deforestation in Congo Basin is estimated to 0.09% between 1990 and 2000 and of net degradation to 0.05%. Between 2000 and 2005, this unique exercise estimates annual net deforestation to 0.17% and annual net degradation to 0.09%. An accuracy assessment reveals that 92.7% of tree cover (TC) classes agree with independent expert interpretation. In the discussion, we underline the direct causes and the drivers of deforestation. Population density, small-scale agriculture, fuelwood collection and forest's accessibility are closely linked to deforestation, whereas timber extraction has no major impact on the reduction in the canopy cover. The analysis also shows the efficiency of protected areas to reduce deforestation. These results are expected to contribute to the discussion on the reduction in CO2 emissions from deforestation and forest degradation (REDD+) and serve as reference for the period.
 
Seasonal pulses of phytoplankton drive seasonal cycles of carbon fixation and particle sedimentation, and might condition recruitment success in many exploited species. Taking advantage of long term series of remotely sensed chlorophyll a (1998-2012), we analyzed changes in phytoplankton seasonality in the North Atlantic Ocean. Phytoplankton phenology was analyzed based on a probabilistic characterization of bloom incidence. This approach allowed us to detect changes in the prevalence of different seasonal cycles and, at the same time, to estimate bloom timing and magnitude taking into account uncertainty in bloom detection. Deviations between different sensors stressed the importance of a prolonged overlap between successive missions to ensure a correct assessment of phenological changes, as well as the advantage of semi-analytical chlorophyll algorithms over empirical ones to reduce biases. Earlier and more intense blooms were detected in the subpolar Atlantic, while advanced blooms of less magnitude were common in the Subtropical gyre. In the temperate North Atlantic, spring blooms advanced their timing and decreased in magnitude, whereas fall blooms delayed and increased their intensity. At the same time, the prevalence of locations with a single autumn/winter bloom or with a bimodal seasonal cycle increased, in consonance with a poleward expansion of subtropical conditions. Changes in bloom timing and magnitude presented a clear signature of environmental factors, especially wind forcing, although changes on incident photosynthetically active radiation and sea surface temperature were also important depending on latitude. Trends in bloom magnitude matched changes in mean chlorophyll a during the study period, suggesting that seasonal peaks drive long term trends in chlorophyll a concentration. Our results link changes in North Atlantic climate with recent trends in the phenology of phytoplankton, suggesting an intensification of these impacts in a near future. This article is protected by copyright. All rights reserved.
 
Drought affects more people than any other natural disaster but there is little understanding of how ecosystems react to droughts. This study jointly analyzed spatio-temporal changes of drought patterns with vegetation phenology and productivity changes between 1999 and 2010 in major European bioclimatic zones. The Standardized Precipitation and Evapotranspiration Index (SPEI) was used as drought indicator whereas changes in growing season length and vegetation productivity were assessed using remote sensing time-series of NDVI (Normalized Difference Vegetation Index). Drought spatio-temporal variability was analyzed using a Principal Component Analysis, leading to the identification of four major drought events between 1999 and 2010 in Europe. Correspondence Analysis showed that at the continental scale the productivity and phenology reacted differently to the identified drought events depending on ecosystem and land cover. Northern and Mediterranean ecosystems proved to be more resilient to droughts in terms of vegetation phenology and productivity developments. Western Atlantic regions and Eastern Europe showed strong agglomerations of decreased productivity and shorter vegetation growing season length, indicating that these ecosystems did not buffer the effects of drought well. In a climate change perspective, increase in drought frequency or intensity may result in larger impacts over these ecosystems, thus management and adaptation strategies should be strengthened in these areas of concerns. This article is protected by copyright. All rights reserved.
 
More than 50% of the world's population feeds on rice. Soils used for rice production are mostly managed under submerged conditions (paddy soils). This management, which favors carbon sequestration, potentially decouples surface from subsurface carbon cycling. The objective of this study was to elucidate the long-term rates of carbon accrual in surface and subsurface soil horizons relative to those of soils under nonpaddy management. We assessed changes in total soil organic as well as of inorganic carbon stocks along a 2000-year chronosequence of soils under paddy and adjacent nonpaddy management in the Yangtze delta, China. The initial organic carbon accumulation phase lasts much longer and is more intensive than previously assumed, e.g., by the Intergovernmental Panel on Climate Change (IPCC). Paddy topsoils accumulated 170-178 kg organic carbon ha(-1) a(-1) in the first 300 years; subsoils lost 29-84 kg organic carbon ha(-1) a(-1) during this period of time. Subsoil carbon losses were largest during the first 50 years after land embankment and again large beyond 700 years of cultivation, due to inorganic carbonate weathering and the lack of organic carbon replenishment. Carbon losses in subsoils may therefore offset soil carbon gains or losses in the surface soils. We strongly recommend including subsoils into global carbon accounting schemes, particularly for paddy fields.
 
Using a high resolution (40 × 40 km) chemical transport model coupled with the Regional Emission inventory in Asia (REAS), we simulated surface ozone concentrations ([O3 ]) and evaluated O3 -induced wheat production loss in China and India for the years 2000 and 2020 using dose-response functions based on AOT40 (accumulated [O3 ] above 40 ppb) and PODY (phytotoxic O3 dose, accumulated stomatal flux of O3 above a threshold of Y nmol m(-2) s(-1) ). Two O3 dose metrics (90-days AOT40 and POD6 ) were derived from European experiments, and the other two (75-days AOT40 and POD12 ) were adapted from Asian studies. Relative yield loss (RYL) of wheat in 2000 was estimated to be 6.4-14.9% for China, and 8.2-22.3% for India. POD6 predicted greater RYL, especially for the warm regions of India, whereas the 90-days AOT40 gave the lowest estimates. For the future projection, all the O3 dose metrics gave comparable estimates of an increase in RYL from 2000 to 2020 in the range of 8.1-9.4% and 5.4-7.7% for China and India, respectively. The lower projected increase in RYL for India may be due to conservative estimation of the emission increase in 2020. Sensitivity tests of the model showed that the PODY -based estimates of RYL are highly sensitive to perturbations in the meteorological inputs, but that the estimated increase of RYL from 2000 to 2020 is much more robust. The projected increase in wheat production loss in China and India in the near future is substantially larger than the uncertainties in the estimation and indicates an urgent need for curbing the rapid increase of surface [O3 ] in these regions. This article is protected by copyright. All rights reserved.
 
Rice straw burning has accompanied paddy management for millennia, introducing black carbon (BC) into soil as the residue of incomplete combustion. This study examined the contribution of BC to soil organic matter and the rate at which it accumulates in paddy soils as a result of prolonged paddy management. Soil depth profiles were sampled along a chronosequence of 0-2000 years of rice-wheat rotation systems and adjacent non-paddy systems (50-700 years) in the Bay of Hangzhou (Zhejiang province, China). The soil BC content and its degree of condensation were assessed using benzene-polycarboxylic acids (BPCAs) as geochemical markers. The results showed that despite regular long term BC input, BC only contributed 7-11% of total soil organic carbon (SOC) in the topsoil horizons. Nevertheless, along with SOC, paddy soils accumulated BC with increasing duration of management until 297 years to reach a steady-state of 13 t BC ha(-1) . This was 1.8 times more than in non-paddy soils. The fate of BC in paddy soils (0-1 m) could be modelled revealing an average annual input of 44 kg ha(-1) yr(-1) , and a mean residence time of 303 years. The subsoils contributed at least 50% to overall BC stocks, which likely derived from periods prior to land embankment and episodic burial of ancient topsoil, as also indicated by BPCA pattern changes. We conclude that there is a limited accumulation of C in charred forms upon prolonged paddy management. The final contribution of BC to total SOC in paddy soils was similar to that in other aerobic ecosystems of the world. This article is protected by copyright. All rights reserved.
 
The rate of vegetation recovery from boreal wildfire influences terrestrial carbon cycle processes and climate feedbacks by affecting the surface energy budget and land-atmosphere carbon exchange. Previous forest recovery assessments using satellite optical-infrared normalized difference vegetation index (NDVI) and tower CO2 eddy covariance techniques indicate rapid vegetation recovery within 5 to 10 years, but these techniques are not directly sensitive to changes in vegetation biomass. Alternatively, the vegetation optical depth (VOD) parameter from satellite passive microwave remote sensing can detect changes in canopy biomass structure and may provide a useful metric of post-fire vegetation response to inform regional recovery assessments. We analyzed a multi-year (2003-2010) satellite VOD record from the NASA AMSR-E (Advanced Microwave Scanning Radiometer for EOS) sensor to estimate forest recovery trajectories for 14 large boreal fires from 2004 in Alaska and Canada. The VOD record indicated initial post-fire canopy biomass recovery within 3 to 7 years, lagging NDVI recovery by 1 to 5 years. The VOD lag was attributed to slower non-photosynthetic (woody) and photosynthetic (foliar) canopy biomass recovery, relative to the faster canopy greenness response indicated from the NDVI. The duration of VOD recovery to pre-burn conditions was also directly proportional (p<0.01) to satellite (MODIS) estimated tree cover loss used as a metric of fire severity. Our results indicate that vegetation biomass recovery from boreal fire disturbance is generally slower than reported from previous assessments based solely on satellite optical-infrared remote sensing, while the VOD parameter enables more comprehensive assessments of boreal forest recovery. This article is protected by copyright. All rights reserved.
 
Total biomass and carbon stocks (dry weight in Pg) by vegetation physiognomy in Brazil's Legal Amazonia
The largest carbon stock in tropical vegetation is in Brazilian Amazonia. In this ~5 million km(2) area, over 750,000 km(2) of forest and ~240,000 km(2) of non-forest vegetation types had been cleared through 2013. We estimate current carbon stocks and cumulative gross carbon loss from clearing of pre-modern vegetation in Brazil's "Legal Amazonia" and "Amazonia biome" regions. Biomass of "pre-modern" vegetation (prior to major increases in disturbance beginning in the 1970s) was estimated by matching vegetation classes mapped at a scale of 1:250,000 and 29 biomass means from 41 published studies for vegetation types classified as forest (2317 1-ha plots) and as either non-forest or contact zones (1830 plots and sub-plots of varied size). Total biomass (above and below-ground, dry weight) underwent a gross reduction of 18.3% in Legal Amazonia (13.1 Pg C) and 16.7% in the Amazonia biome (11.2 Pg C) through 2013, excluding carbon loss from the effects of fragmentation, selective logging, fires, mortality induced by recent droughts and clearing of forest regrowth. In spite of the loss of carbon from clearing, large amounts of carbon were stored in stands of remaining vegetation in 2013, equivalent to 149 Mg C ha(-1) when weighted by the total area covered by each vegetation type in Legal Amazonia. Native vegetation in Legal Amazonia in 2013 originally contained 58.6 Pg C, while that in the Amazonia biome contained 56 Pg C. Emissions per unit area from clearing could potentially be larger in the future because previously cleared areas were mainly covered by vegetation with lower mean biomass than the remaining vegetation. Estimates of original biomass are essential for estimating losses to forest degradation. This study offers estimates of cumulative biomass loss, as well as estimates of pre-modern carbon stocks that have not been represented in recent estimates of deforestation impacts. This article is protected by copyright. All rights reserved.
 
The Pederson et al. (2014) letter deals ostensibly with their opinion that we (Nowacki & Abrams 2014) failed to consider the lessening of drought after the 1930s as a major driver of post-European forest dynamics in the eastern U.S. We are befuddled by this because our paper includes long-term precipitation data for four study regions and we report a slight increase (2-8% change) from 1895 to the present. We report a similar range of increase in temperature for that time period. As such, we accounted for these factors in the context of our board-scale study. This article is protected by copyright. All rights reserved. This article is protected by copyright. All rights reserved.
 
In response to our paper "Pyrogenic organic matter production from wildfires: a missing sink in the global carbon cycle" (Santín et al. 2015), Billings & Schlesinger (2015) argue that pyrogenic organic matter (PyOM) formation is not a missing C sink. This article is protected by copyright. All rights reserved. This article is protected by copyright. All rights reserved.
 
Crop-model specific biases are a key uncertainty affecting our understanding of climate change impacts to agriculture. There is increasing research focus on inter-model variation, but comparisons between mechanistic (MMs) and empirical models (EMs) are rare despite both being used widely in this field. We combined MMs and EMs to project future (2055) changes in the potential distribution (suitability) and productivity of maize and spring wheat in South Africa under 18 downscaled climate scenarios (9 models run under 2 emissions scenarios). EMs projected larger yield losses or smaller gains than MMs. The EMs' median projected maize and wheat yield changes were -3.6% and 6.2%, compared to 6.5% and 15.2% for the MM. The EM projected a 10% reduction in the potential maize growing area, where the MM projected a 9% gain. Both models showed increases in the potential spring wheat production region (EM = 48%, MM = 20%), but these results were more equivocal because both models (particularly the EM) substantially overestimated the extent of current suitability. The substantial water use efficiency gains simulated by the MMs under elevated CO2 accounted for much of the EM-MM difference, but EMs may have more accurately represented crop temperature sensitivities. Our results align with earlier studies showing that EMs may show larger climate change losses than MMs. Crop forecasting efforts should expand to include EM-MM comparisons to provide a fuller picture of crop-climate response uncertainties. This article is protected by copyright. All rights reserved.
 
Although the distribution ranges and abundance of many plant species have declined dramatically in recent decades, detailed analysis of these changes and their cause have only become possible following the publication of second- and third-generation national distribution atlases. Decline can now be compared both between species and in different parts of species' ranges. We extracted data from distribution atlases to compare range persistence of 736 plant species common to both the UK and Estonia between survey periods encompassing almost the same years (1969 and 1999 in the UK and 1970 and 2004 in Estonia). We determined which traits were most closely associated with variation in species persistence, whether these were the same in each country, and the extent to which they explained differences in persistence between the countries. Mean range size declined less in Estonia than in the UK (24.3% vs. 30.3%). One-third of species in Estonia (239) maintained >90% of their distribution range compared with one-fifth (141) in the UK. In Estonia, 99 species lost >50% of their range compared with 127 species in the UK. Persistence was very positively related to original range in both countries. Major differences in species persistence between the studied countries were primarily determined by biogeographic (affiliation to floristic element) and ecoevolutionary (plant strategy) factors. In contrast, within-country persistence was most strongly determined by tolerance of anthropogenic activities. Decline of species in the families Orchidaceae and Potamogetonaceae was significantly greater in the UK than in Estonia. Almost all of the 736 common and native European plant species in our study are currently declining in their range due to pressure from anthropogenic activities. Those species with low tolerance of human activity, with biotic pollination vectors and in the families referred to above are the most vulnerable, especially where human population density is high.
 
Rapid climate change has the potential to affect economic, social, and biological systems. A concern for species conservation is whether or not the rate of on-going climate change will exceed the rate at which species can adapt or move to suitable environments. Here we assess the climate velocity (both climate displacement rate and direction) for minimum temperature, actual evapotranspiration, and climatic water deficit (deficit) over the contiguous US during the 20th century (1916-2005). Vectors for these variables demonstrate a complex mosaic of patterns that vary spatially and temporally and are dependent on the spatial resolution of input climate data. Velocities for variables that characterize the climatic water balance were similar in magnitude to that derived from temperature, but frequently differed in direction resulting in the divergence of climate vectors through time. Our results strain expectations of poleward and upslope migration over the past century due to warming. Instead, they suggest that a more full understanding of changes in multiple climatic factors, in addition to temperature, may help explain unexpected or conflicting observational evidence of climate-driven species range shifts during the 20th century.
 
Study area in the Maritime Alps with the main locations and valleys in the region. The surface of the study area covers 1923 km2.
Percentage of current and known occurrences explained by each of the three ensembles 927 of species distri- bution model techniques under the two dispersal scenarios of the two models of persistence
Dynamics of expansion, contraction, and persistence across time intervals, including diffusion and microrefugia (MR); (a) potential distribution at time Tx and formation of MR; (b) persistence in MR during extinction periods [predicted by species distribution models (SDMs)]; (c) expansion / recolonization from active MR to suitable areas (predicted by SDMs). (d) Areas colonized at Tx+2 with previous active and newly forming potential MR as well as suitable areas to be colonized at Tx+3 are displayed to allow better visualization at an intermediate stage as Tx+2*; (e) expansion/recolonization from both predicted suitable areas (macrorefugia) and MR.
Suitable surfaces (expressed in number of pixels ×105) predicted by the three species distribution model (SDM) ensembles raw projections. Suitable surfaces predicted by the microrefugia model under the unlimited and limited dispersal scenarios based on the consensus ensemble of SDMs across the 1 ka time intervals are added for comparison.
Microrefugia (MR) with the three types of supporting functions superimposed on the current potential distribution. (a) MR that have potentially contributed to range expansions and shifts across the 1 ka time intervals (stepping-stones MR), (b) MR that have contributed most recently to forming the current potential distribution (recolonization MR) and (c) MR that have remained in the same location for at least two climatic periods (stable MR).
Ongoing rapid climate change is predicted to cause local extinction of plant species in mountain regions. However, some plant species could have persisted during Quaternary climate oscillations without shifting their range, despite the limited evidence from fossils. Here, we tested two candidate mechanisms of persistence by comparing the macrorefugia and microrefugia (MR) hypotheses. We used the rare and endemic Saxifraga florulenta as a model taxon and combined ensembles of species distribution models (SDMs) with a high-resolution paleoclimatic and topographic dataset to reconstruct its potential current and past distribution since the last glacial maximum. To test the macrorefugia hypothesis, we verified whether the species could have persisted in or shifted to geographic areas defined by its realized niche. We then identified potential MR based on climatic and topographic properties of the landscape and applied refined scenarios of MR dynamics and functions over time. Last, we quantified the number of known occurrences that could be explained by either the macrorefugia or MR model. A consensus of two or three SDM techniques predicted absence between 14-10, 3-4 and 1 ka BP, which did not support the macrorefugia model. In contrast, we showed that S. florulenta could have contracted into MR during periods of absence predicted by the SDMs and later re-colonized suitable areas according to the macrorefugia model. Assuming a limited and realistic seed dispersal distance for our species, we explained a large number of the current occurrences (61-96%). Additionally, we showed that MR could have facilitated range expansions or shifts of S. florulenta. Finally, we found that the most recent and the most stable MR were the ones closest to current occurrences. Hence, we propose a novel paradigm to explain plant persistence by highlighting the importance of supporting functions of MR when forecasting the fate of plant species under climate change.
 
Output from an earth system model is paired with a size-based food web model to investigate the effects of climate change on the abundance of large fish over the 21st century. The earth system model, forced by the Intergovernmental Panel on Climate Change (IPCC) Special report on emission scenario A2, combines a coupled climate model with a biogeochemical model including major nutrients, three phytoplankton functional groups, and zooplankton grazing. The size-based food web model includes linkages between two size-structured pelagic communities: primary producers and consumers. Our investigation focuses on seven sites in the North Pacific, each highlighting a specific aspect of projected climate change, and includes top-down ecosystem depletion through fishing. We project declines in large fish abundance ranging from 0 to 75.8% in the central North Pacific and increases of up to 43.0% in the California Current (CC) region over the 21st century in response to change in phytoplankton size structure and direct physiological effects. We find that fish abundance is especially sensitive to projected changes in large phytoplankton density and our model projects changes in the abundance of large fish being of the same order of magnitude as changes in the abundance of large phytoplankton. Thus, studies that address only climate-induced impacts to primary production without including changes to phytoplankton size structure may not adequately project ecosystem responses.
 
Shoot dark respiration at 20 °C [R, panel (a)], leaf fresh mass per unit leaf area [FA, panel (b)], and leaf and stem dry matter content [DMC, panel (c)] plotted against corresponding shoot DMC values. Individual replicate values are shown. Coefficients of determination (r2) and P values: Shoot R20 – Shoot DMC, r2 = 0.56 (P < 0.001); FA – Shoot DMC, r2 = 0.48 (P < 0.001); (c) Leaf DMC – Shoot DMC, r2 = 0.58 (P < 0.001); Stem DMC – Shoot DMC, r2 = 0.83 (P < 0.001).
Temperature dependence of (a) dark respiration (R, nmol CO2 gDM−1 s−1) and (b) Q10 of B. nana shoots. Measurements were made between the 5–35 °C range. Symbols represent the modeled specific rates of R and Q10 values at 1.0 °C intervals (n = 3–4, ±SE). CT, control treatment; GH, greenhouse; +NP, high N and P treatment; GH+NP, greenhouse grown plants provided with high N and P. In (B), the dashed line shows Q10 values of 1.0 for reference. Leaves were heated at 1.0 °C min−1.
Example R-T curve over the larger range of measurement Ts showing impacts of continuous heating on rates of whole shoot respiration of a single replicate B. nana plant grown under control conditions (i.e. no fertilizer or greenhouse treatments). Data points show measured rates over the 25–70 °C measurement range, with data collected at 15 s intervals; leaves were heated at 1 °C min−1. The maximal rate of leaf R (Rmax) at the corresponding Tmax are noted.
Despite concern about the status of carbon (C) in the Arctic tundra, there is currently little information on how plant respiration varies in response to environmental change in this region. We quantified the impact of long-term nitrogen (N) and phosphorus (P) treatments and greenhouse warming on the short-term temperature (T) response and sensitivity of leaf respiration (R), the high T threshold of R, and associated traits in shoots of the Arctic shrub Betula nana in experimental plots at Toolik Lake, Alaska. Respiration only acclimated to greenhouse warming in plots provided with both N and P (resulting in a ~30% reduction in carbon efflux in shoots measured at 10 and 20 °C), suggesting a nutrient-dependence of metabolic adjustment. Neither greenhouse nor N+P treatments impacted on the respiratory sensitivity to T (Q10 ); overall Q10 values decreased with increasing measuring T, from ~3.0 at 5 °C to ~1.5 at 35 °C. New high-resolution measurements of R across a range of measuring Ts (25-70 °C) yielded insights into the T at which maximal rates of R occurred (Tmax ). Although growth temperature did not affect Tmax , N+P fertilization increased Tmax values ~5 °C, from 53 to 58 °C. N+P fertilized shoots exhibited greater rates of R than non-fertilized shoots, with this effect diminishing under greenhouse warming. Collectively, our results highlight the nutrient-dependence of thermal acclimation of leaf R in B. nana, suggesting that the metabolic efficiency allowed via thermal acclimation may be impaired at current levels of soil nutrient availability. This finding has important implications for predicting carbon fluxes in Arctic ecosystems, particularly if soil N and P become more abundant in the future as the tundra warms. This article is protected by copyright. All rights reserved.
 
Climate change has affected plant phenology; increasing temperatures are associated with advancing first flowering dates. The impact on flowering duration, however, has rarely been studied. In this study we analysed first flowering dates and flowering durations from a 27-year dataset of weekly flower observations on 232 plant species from the island of Guernsey in the English Channel. The aim of this study was to explore variation in trends and relationships between first flowering dates, flowering duration and temperature. We specifically looked for evidence that traits, such as life forms and phylogenetic groups, explained variation in sensitivity of first flowering and flowering duration among species. Overall trends revealed significantly earlier flowering over time, by an average of 5.2 days decade(-1) since 1985. A highly significant shortening of flowering duration was observed, by an average of 10 days decade(-1) . Correlations between first flowering, flowering duration and year varied between different species, traits and flowering periods. Significant differences among traits were observed for first flowering and to a lesser degree for flowering duration. Overall, in comparison to first flowering, more species had significant trends in flowering duration. Temperature relationships revealed large differences in strength and direction of response. 55% of the species revealed a significant negative relationship of first flowering dates and temperature. In contrast, only 19% of flowering durations had a significant negative temperature relationship. The advance in first flowering date together with a shortening of flowering duration suggests potentially serious impacts on pollinators, which might pose a major threat to biodiversity, agriculture and horticulture. Human health, in terms of pollen allergies, however, might benefit from a shortening of specific plant pollen seasons. This article is protected by copyright. All rights reserved.
 
The impact of climate change and of other anthropogenic pressures on the structure and composition of phytoplankton communities of large European rivers remains poorly documented. Here we report the findings of a study of the changes in the phytoplankton community of the middle segment of the river Loire over the past 24 years. An attempt is made to distinguish between the impact of changes acting at the local scale and that of those acting more globally. A dramatic reduction in phytoplankton abundance was observed, particularly in the mid -1990s; this was concomitant with an increase in the relative proportion of cyanobacteria. At the same time, the phytoplankton community displayed increasing richness and diversity, and little change in its size structure. All these changes seem to be related to local changes, in particular to the reduction in phosphorus concentrations, as well as to changes in climate, throughout modifications in the river discharge and water temperature. Interestingly, herbicide contamination also appeared to be of particular importance in explaining the unexpected increase in the proportion of cyanobacteria in the phytoplankton community after the 1990s. These findings suggest that combinations of numerous anthropogenic pressures acting at different spatial and temporal scales have led to a mix of predictable and unpredictable changes occurring in the phytoplankton community of the river Loire, with probable consequences for the trophic networks in this river.
 
With their position at the interface between land and ocean and their fragile nature, lagoons are sensitive to environmental change, and it is reasonable to expect these changes would be recorded in well-preserved taxa such as molluscs. To test this, the 4000-year history of molluscs in Great South Bay, a bar-built lagoon, was reconstructed from 24 vibracores. Using x-radiography to identify shell layers, faunal counts, shell condition, organic content, and sediment type were measured in 325 samples. Sample age was estimated by interpolating 40 radiocarbon dates. K-means cluster analysis identified three molluscan assemblages, corresponding to sand-associated and mud-associated groups, and the third associated with inlet areas. Redundancy and regression tree analyses indicated that significant transitions from the sand-associated to mud-associated assemblage occurred over large portions of the bay about 650 and 294years bp. The first date corresponds to the transition from the Medieval Warm Period to the Little Ice Age; this change in climate reduced the frequency of strong storms, likely leading to reduced barrier island breaching, greater bay enclosure, and fine-grained sediment accumulation. The second date marks the initiation of clear cutting by European settlers, an activity that would have increased runoff of fine-grained material. The occurrence of the inlet assemblage in the western and eastern ends of the bay is consistent with a history of inlets in these areas, even though prior to Hurricane Sandy in 2012, no inlet was present in the eastern bay in almost 200years. The mud dominant, Mulinia lateralis, is a bivalve often associated with environmental disturbances. Its increased frequency over the past 300years suggests that disturbances are more common in the bay than in the past. Management activities maintaining the current barrier island state may be contributing to the sand-mud transition and to the bay's susceptibility to disturbances.
 
Many areas of the Arctic are simultaneously affected by rapid climate change and rapid industrial development. These areas are likely to increase in number and size as sea ice melts and abundant Arctic natural resources become more accessible. Documenting the changes that have already occurred is essential to inform management approaches in order to minimize the impacts of future activities. Here we determine the cumulative geoecological effects of 62 years (1949-2011) of infrastructure- and climate-related changes in the Prudhoe Bay Oilfield, the oldest and most extensive industrial complex in the Arctic, and an area with extensive ice-rich permafrost that is extraordinarily sensitive to climate change. We demonstrate that thermokarst has recently affected broad areas of the entire region, and that a sudden increase in the area affected began shortly after 1990 corresponding to a rapid rise in regional summer air temperatures and related permafrost temperatures. We also present a conceptual model that describes how infrastructure-related factors, including road dust and roadside flooding are contributing to more extensive thermokarst in areas adjacent to roads and gravel pads. We mapped the historical infrastructure changes for the Alaska North Slope oilfields for 10 dates from the initial oil discovery in 1968 to 2011. By 2010, over 34% of the intensively mapped area was affected by oil development. In addition, between 1990 and 2001, coincident with strong atmospheric warming during the 1990s, 19% of the remaining natural landscapes (excluding areas covered by infrastructure, lakes and river floodplains) exhibited expansion of thermokarst features resulting in more abundant small ponds, greater microrelief, more active lakeshore erosion and increased landscape and habitat heterogeneity. This transition to a new geoecological regime will have impacts to wildlife habitat, local residents and industry. This article is protected by copyright. All rights reserved.
 
Climate extremes can ultimately reshape grassland services such as forage production and change plant functional type composition. This 3-year field research studied resistance to dehydration and recovery after rehydration of plant community and plant functional types in an upland perennial grassland subjected to climate and cutting frequency (Cut+, Cut-) disturbances by measuring green tissue percentage and above-ground biomass production (ANPP). In year one, a climate disturbance gradient was applied by co-manipulating temperature and precipitation. Four treatments were considered: control and warming-drought climatic treatment, with or without extreme summer event. In year two, control and warming-drought treatments were maintained without extreme. In year three, all treatments received ambient climatic conditions. We found that the grassland community was very sensitive to dehydration during the summer extreme: aerial senescence reached 80% when cumulated climatic water balance fell to -156 mm and biomass declined by 78% at the end of summer. In autumn, canopy greenness and biomass totally recovered in control but not in warming-drought treatment. However ANPP decreased under both climatic treatments, but the effect was stronger on Cut+ (-24%) than Cut- (-15%). This decline was not compensated by the presence of three functional types because they were negatively affected by the climatic treatments, suggesting an absence of buffering effect on grassland production. The following two years, lasting effects of climate disturbance on ANPP were observable. The unexpected stressful conditions of year three induced a decline in grassland production in the Cut+ control treatment. The fact that this treatment cumulated higher (45%) N export over the three years suggests that N plays a key role in ANPP stability. As ANPP in this mesic perennial grassland did not show engineering resilience, long-term experimental manipulation is needed. Infrequent mowing appears more appropriate for sustaining grassland ANPP under future climate extremes. This article is protected by copyright. All rights reserved.
 
Peatlands contain approximately one third of all soil organic carbon (SOC). Warming can alter above- and belowground linkages that regulate soil organic carbon dynamics and C-balance in peatlands. Here we examine the multiyear impact of in situ experimental warming on the microbial food web, vegetation, and their feedbacks with soil chemistry. We provide evidence of both positive and negative impacts of warming on specific microbial functional groups, leading to destabilization of the microbial food web. We observed a strong reduction (70%) in the biomass of top-predators (testate amoebae) in warmed plots. Such a loss caused a shortening of microbial food chains, which in turn stimulated microbial activity, leading to slight increases in levels of nutrients and labile C in water. We further show that warming altered the regulatory role of Sphagnum-polyphenols on microbial community structure with a potential inhibition of top predators. In addition, warming caused a decrease in Sphagnum cover and an increase in vascular plant cover. Using structural equation modelling, we show that changes in the microbial food web affected the relationships between plants, soil water chemistry, and microbial communities. These results suggest that warming will destabilize C and nutrient recycling of peatlands via changes in above- and belowground linkages, and therefore, the microbial food web associated with mosses will feedback positively to global warming by destabilizing the carbon cycle. This study confirms that microbial food webs thus constitute a key element in the functioning of peatland ecosystems. Their study can help understand how mosses, as ecosystem engineers, tightly regulate biogeochemical cycling and climate feedback in peatlands.
 
The role of tree mortality in the global carbon balance is complicated by strong spatial and temporal heterogeneity that arises from the stochastic nature of carbon loss through disturbance. Characterizing spatio-temporal variation in mortality (including disturbance) and its effects on forest and carbon dynamics is thus essential to understanding the current global forest carbon sink, and to predicting how it will change in future. We analyzed forest inventory data from the eastern United States to estimate plot-level variation in mortality (relative to a long-term background rate for individual trees) for nine distinct forest regions. Disturbances that produced at least a fourfold increase in tree mortality over an approximately 5 year interval were observed in 1-5% of plots in each forest region. The frequency of disturbance was lowest in the northeast, and increased southwards along the Atlantic and Gulf coasts as fire and hurricane disturbances became progressively more common. Across the central and northern parts of the region, natural disturbances appeared to reflect a diffuse combination of wind, insects, disease, and ice storms. By linking estimated covariation in tree growth and mortality over time with a data-constrained forest dynamics model, we simulated the implications of stochastic variation in mortality for long-term aboveground biomass changes across the eastern United States. A geographic gradient in disturbance frequency induced notable differences in biomass dynamics between the least- and most-disturbed regions, with variation in mortality causing the latter to undergo considerably stronger fluctuations in aboveground stand biomass over time. Moreover, regional simulations showed that a given long-term increase in mean mortality rates would support greater aboveground biomass when expressed through disturbance effects compared with background mortality, particularly for early-successional species. The effects of increased tree mortality on carbon stocks and forest composition may thus depend partly on whether future mortality increases are chronic or episodic in nature.
 
Obligate seeder trees requiring high-severity fires to regenerate may be vulnerable to population collapse if fire frequency increases abruptly. We tested this proposition using a long-lived obligate seeding forest tree, alpine ash (Eucalyptus delegatensis), in the Australian Alps. Since 2002, 85% of the Alps bioregion has been burnt by several very large fires, tracking the regional trend of more frequent extreme fire weather. High-severity fires removed 25% of aboveground tree biomass, and switched fuel arrays from low loads of herbaceous and litter fuels to high loads of flammable shrubs and juvenile trees, priming regenerating stands for subsequent fires. Single high-severity fires caused adult mortality and triggered mass regeneration, but a second fire in quick succession killed 97% of the regenerating alpine ash. Our results indicate that without interventions to reduce fire severity, interactions between flammability of regenerating stands and increased extreme fire weather will eliminate much of the remaining mature alpine ash forest. This article is protected by copyright. All rights reserved.
 
Migration of plant populations is a potential survival response to climate change that depends critically on seed dispersal. Biological and physical factors determine dispersal and migration of wind-dispersed species. Recent field and wind tunnel studies demonstrate biological adaptations that bias seed release towards conditions of higher wind velocity, promoting longer dispersal distances and faster migration. However, another suite of international studies also recently highlighted a global decrease in near-surface wind speeds, or 'global stilling'. This study assessed the implications of both factors on potential plant population migration rates, using a mechanistic modeling framework. Non-random abscission was investigated using models of three seed release mechanisms: (i) a simple drag model; (ii) a seed deflection model; and (iii) a 'wear and tear' model. The models generated a single functional relationship between the frequency of seed release and statistics of the near-surface wind environment, independent of the abscission mechanism. An Inertial-Particle, Coupled Eulerian-Lagrangian Closure model (IP-CELC) was used to investigate abscission effects on seed dispersal kernels and plant population migration rates under contemporary and potential future wind conditions (based on reported global stilling trends). The results confirm that non-random seed abscission increased dispersal distances, particularly for light seeds. The increases were mitigated by two physical feedbacks: (i) although non-random abscission increased the initial acceleration of seeds from rest, the sensitivity of the seed dispersal to this initial condition declined as the wind speed increased; and (ii) while non-random abscission increased the mean dispersal length, it reduced the kurtosis of seasonal dispersal kernels, and thus the chance of long-distance dispersal. Wind stilling greatly reduced the modeled migration rates under biased seed release conditions. Thus, species that require high wind velocities for seed abscission could experience threshold-like reductions in dispersal and migration potential if near-surface wind speeds continue to decline. © 2013 Blackwell Publishing Ltd.
 
We investigated whether groundwater abstraction for urban water supply diminishes the storage of carbon (C), nitrogen (N), and organic matter in the soil of rural wetlands. Wetland soil organic matter (SOM) benefits air and water quality by sequestering large masses of C and N. Yet, the accumulation of wetland SOM depends on soil inundation, so we hypothesized that groundwater abstraction would diminish stocks of SOM, C, and N in wetland soils. Predictions of this hypothesis were tested in two types of subtropical, depressional-basin wetland: forested swamps and herbaceous-vegetation marshes. In west-central Florida, >650 ML groundwater d(-1) are abstracted for use primarily in the Tampa Bay metropolis. At higher abstraction volumes, water tables were lower and wetlands had shorter hydroperiods (less time inundated). In turn, wetlands with shorter hydroperiods had 50-60% less SOM, C, and N per kg soil. In swamps, SOM loss caused soil bulk density to double, so areal soil C and N storage per m(2) through 30.5 cm depth was diminished by 25-30% in short-hydroperiod swamps. In herbaceous-vegetation marshes, short hydroperiods caused a sharper decline in N than in C. Soil organic matter, C, and N pools were not correlated with soil texture or with wetland draining-reflooding frequency. Many years of shortened hydroperiod were probably required to diminish soil organic matter, C, and N pools by the magnitudes we observed. This diminution might have occurred decades ago, but could be maintained contemporarily by the failure each year of chronically-drained soils to retain new organic matter inputs. In sum, our study attributes the contraction of hydroperiod and loss of soil organic matter, C, and N from rural wetlands to groundwater abstraction performed largely for urban water supply, revealing teleconnections between rural ecosystem change and urban resource demand. This article is protected by copyright. All rights reserved.
 
Climate changes are assumed to shift the ranges of tree species and forest biomes. Such range shifts result from changes in abundances of tree species or functional types. Owing to global warming, the abundance of a tree species or functional type is expected to increase near the colder edge of its range and decrease near the warmer edge. The present study examined directional changes in abundance and demographic parameters of forest trees along a temperature gradient, as well as a successional gradient, in Japan. Changes in the relative abundance of each of four functional types (evergreen broad-leaved, deciduous broad-leaved, evergreen temperate conifer, and evergreen boreal conifer) and the demography of each species (recruitment rate, mortality, and population growth rate) were analysed in 39 permanent forest plots across the Japanese archipelago. Directional changes in the relative abundance of functional types were detected along the temperature gradient. Relative abundance of evergreen broad-leaved trees increased near their colder range boundaries, especially in secondary forests, coinciding with the decrease in deciduous broad-leaved trees. Similarly, relative abundance of deciduous broad-leaved trees increased near their colder range boundaries, coinciding with the decrease in boreal conifers. These functional type-level changes were mainly due to higher recruitment rates and partly to the lower mortality of individual species at colder sites. This is the first report to show that tree species abundances in temperate forests are changing directionally along a temperature gradient, which might be due to current or past climate changes as well as recovery from past disturbances. This article is protected by copyright. All rights reserved. This article is protected by copyright. All rights reserved.
 
Top-cited authors
Riccardo Valentini
  • Tuscia University
Josep Penuelas
  • Spanish National Research Council (CSIC)-Centre for Ecological Research and Forestry Applications (CREAF)
Markus Reichstein
  • Max Planck Institute for Biogeochemistry Jena
Ivan A. Janssens
  • University of Antwerp
Dennis Baldocchi
  • University of California, Berkeley