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Leading up to the Convention on Biological Diversity Conference of the Parties 15, there is momentum around setting bold conservation targets. Yet, it remains unclear how much of Earth's land area remains without significant human influence and where this land is located. We compare four recent global maps of human influences across Earth's land, Anthromes, Global Human Modification, Human Footprint and Low Impact Areas, to answer these questions. Despite using various methodologies and data, these different spatial assessments independently estimate similar percentages of the Earth's terrestrial surface as having very low (20%–34%) and low (48%–56%) human influence. Three out of four spatial assessments agree on 46% of the non‐permanent ice‐ or snow‐covered land as having low human influence. However, much of the very low and low influence portions of the planet are comprised of cold (e.g., boreal forests, montane grasslands and tundra) or arid (e.g., deserts) landscapes. Only four biomes (boreal forests, deserts, temperate coniferous forests and tundra) have a majority of datasets agreeing that at least half of their area has very low human influence. More concerning, <1% of temperate grasslands, tropical coniferous forests and tropical dry forests have very low human influence across most datasets, and tropical grasslands, mangroves and montane grasslands also have <1% of land identified as very low influence across all datasets. These findings suggest that about half of Earth's terrestrial surface has relatively low human influence and offers opportunities for proactive conservation actions to retain the last intact ecosystems on the planet. However, though the relative abundance of ecosystem areas with low human influence varies widely by biome, conserving these last intact areas should be a high priority before they are completely lost. [ABSTRACT FROM AUTHOR]

Northern boreal peatlands are important ecosystems in modulating global biogeochemical cycles, yet their biological communities and related carbon dynamics are highly sensitive to changes in climate. Despite this, the strength and recent direction of these feedbacks are still unclear. The response of boreal peatlands to climate warming has received relatively little attention compared with other northern peatland types, despite forming a large northern hemisphere‐wide ecosystem. Here, we studied the response of two ombrotrophic boreal peatlands to climate variability over the last c. 200 years for which local meteorological data are available. We used remains from plants and testate amoebae to study historical changes in peatland biological communities. These data were supplemented by peat property (bulk density, carbon and nitrogen content), 14C, 210Pb and 137Cs analyses and were used to infer changes in peatland hydrology and carbon dynamics. In total, six peat cores, three per study site, were studied that represent different microhabitats: low hummock (LH), high lawn and low lawn. The data show a consistent drying trend over recent centuries, represented mainly as a change from wet habitat Sphagnum spp. to dry habitat S. fuscum. Summer temperature and precipitation appeared to be important drivers shaping peatland community and surface moisture conditions. Data from the driest microhabitat studied, LH, revealed a clear and strong negative linear correlation (R2 =.5031; p <.001) between carbon accumulation rate and peat surface moisture conditions: under dry conditions, less carbon was accumulated. This suggests that at the dry end of the moisture gradient, availability of water regulates carbon accumulation. It can be further linked to the decreased abundance of mixotrophic testate amoebae under drier conditions (R2 =.4207; p <.001). Our study implies that if effective precipitation decreases in the future, the carbon uptake capacity of boreal bogs may be threatened. [ABSTRACT FROM AUTHOR]

The boreal biome exchanges large amounts of carbon (C) and greenhouse gases (GHGs) with the atmosphere and thus significantly affects the global climate. A managed boreal landscape consists of various sinks and sources of carbon dioxide (CO2), methane (CH4), and dissolved organic and inorganic carbon (DOC and DIC) across forests, mires, lakes, and streams. Due to the spatial heterogeneity, large uncertainties exist regarding the net landscape carbon balance (NLCB). In this study, we compiled terrestrial and aquatic fluxes of CO2, CH4, DOC, DIC, and harvested C obtained from tall‐tower eddy covariance measurements, stream monitoring, and remote sensing of biomass stocks for an entire boreal catchment (~68 km2) in Sweden to estimate the NLCB across the land–water–atmosphere continuum. Our results showed that this managed boreal forest landscape was a net C sink (NLCB = 39 g C m−2 year−1) with the landscape–atmosphere CO2 exchange being the dominant component, followed by the C export via harvest and streams. Accounting for the global warming potential of CH4, the landscape was a GHG sink of 237 g CO2‐eq m−2 year−1, thus providing a climate‐cooling effect. The CH4 flux contribution to the annual GHG budget increased from 0.6% during spring to 3.2% during winter. The aquatic C loss was most significant during spring contributing 8% to the annual NLCB. We further found that abiotic controls (e.g., air temperature and incoming radiation) regulated the temporal variability of the NLCB whereas land cover types (e.g., mire vs. forest) and management practices (e.g., clear‐cutting) determined their spatial variability. Our study advocates the need for integrating terrestrial and aquatic fluxes at the landscape scale based on tall‐tower eddy covariance measurements combined with biomass stock and stream monitoring to develop a holistic understanding of the NLCB of managed boreal forest landscapes and to better evaluate their potential for mitigating climate change. [ABSTRACT FROM AUTHOR]

Interspecific interactions are crucial in determining species occurrence and community assembly. Understanding these interactions is thus essential for correctly predicting species' responses to climate change. We focussed on an avian forest guild of four hole‐nesting species with differing sensitivities to climate that show a range of well‐understood reciprocal interactions, including facilitation, competition and predation. We modelled the potential distributions of black woodpecker and boreal, tawny and Ural owl, and tested whether the spatial patterns of the more widespread species (excluding Ural owl) were shaped by interspecific interactions. We then modelled the potential future distributions of all four species, evaluating how the predicted changes will alter the overlap between the species' ranges, and hence the spatial outcomes of interactions. Forest cover/type and climate were important determinants of habitat suitability for all species. Field data analysed with N‐mixture models revealed effects of interspecific interactions on current species abundance, especially in boreal owl (positive effects of black woodpecker, negative effects of tawny owl). Climate change will impact the assemblage both at species and guild levels, as the potential area of range overlap, relevant for species interactions, will change in both proportion and extent in the future. Boreal owl, the most climate‐sensitive species in the guild, will retreat, and the range overlap with its main predator, tawny owl, will increase in the remaining suitable area: climate change will thus impact on boreal owl both directly and indirectly. Climate change will cause the geographical alteration or disruption of species interaction networks, with different consequences for the species belonging to the guild and a likely spatial increase of competition and/or intraguild predation. Our work shows significant interactions and important potential changes in the overlap of areas suitable for the interacting species, which reinforce the importance of including relevant biotic interactions in predictive climate change models for increasing forecast accuracy. [ABSTRACT FROM AUTHOR]

Long‐term trends in ecosystem resource use efficiencies (RUEs) and their controlling factors are key pieces of information for understanding how an ecosystem responds to climate change. We used continuous eddy covariance and microclimate data over the period 1999–2017 from a 120‐year‐old black spruce stand in central Saskatchewan, Canada, to assess interannual variability, long‐term trends, and key controlling factors of gross ecosystem production (GEP) and the RUEs of carbon (CUE = net primary production [NPP]/GEP), light (LUE = GEP/absorbed photosynthetic radiation [APAR]), and water (WUE = GEP/evapotranspiration [E]). At this site, annual GEP has shown an increasing trend over the 19 years (p < 0.01), which may be attributed to rising atmospheric CO2 concentration. Interannual variability in GEP, aside from its increasing trend, was most strongly related to spring temperatures. Associated with the significant increase in annual GEP were relatively small changes in NPP, APAR, and E, so that annual CUE showed a decreasing trend and annual LUE and WUE showed increasing trends over the 19 years. The long‐term trends in the RUEs were related to the increasing CO2 concentration. Further analysis of detrended RUEs showed that their interannual variation was impacted most strongly by air temperature. Two‐factor linear models combining CO2 concentration and air temperature performed well (R2~0.60) in simulating annual RUEs. LUE and WUE were positively correlated both annually and seasonally, while LUE and CUE were mostly negatively correlated. Our results showed divergent long‐term trends among CUE, LUE, and WUE and highlighted the need to account for the combined effects of climatic controls and the 'CO2 fertilization effect' on long‐term variations in RUEs. Since most RUE‐based models rely primarily on one resource limitation, the observed patterns of relative change among the three RUEs may have important implications for RUE‐based modeling of C fluxes. [ABSTRACT FROM AUTHOR]

Wildfire is the dominant disturbance in boreal forests and fire activity is increasing in these regions. Soil fungal communities are important for plant growth and nutrient cycling postfire but there is little understanding of how fires impact fungal communities across landscapes, fire severity gradients, and stand types in boreal forests. Understanding relationships between fungal community composition, particularly mycorrhizas, and understory plant composition is therefore important in predicting how future fire regimes may affect vegetation. We used an extreme wildfire event in boreal forests of Canada's Northwest Territories to test drivers of fungal communities and assess relationships with plant communities. We sampled soils from 39 plots 1 year after fire and 8 unburned plots. High‐throughput sequencing (MiSeq, ITS) revealed 2,034 fungal operational taxonomic units. We found soil pH and fire severity (proportion soil organic layer combusted), and interactions between these drivers were important for fungal community structure (composition, richness, diversity, functional groups). Where fire severity was low, samples with low pH had higher total fungal, mycorrhizal, and saprotroph richness compared to where severity was high. Increased fire severity caused declines in richness of total fungi, mycorrhizas, and saprotrophs, and declines in diversity of total fungi and mycorrhizas. The importance of stand age (a surrogate for fire return interval) for fungal composition suggests we could detect long‐term successional patterns even after fire. Mycorrhizal and plant community composition, richness, and diversity were weakly but significantly correlated. These weak relationships and the distribution of fungi across plots suggest that the underlying driver of fungal community structure is pH, which is modified by fire severity. This study shows the importance of edaphic factors in determining fungal community structure at large scales, but suggests these patterns are mediated by interactions between fire and forest stand composition. [ABSTRACT FROM AUTHOR]

Abstract: The accumulation of soil carbon (C) is regulated by a complex interplay between abiotic and biotic factors. Our study aimed to identify the main drivers of soil C accumulation in the boreal forest of eastern North America. Ecosystem C pools were measured in 72 sites of fire origin that burned 2–314 years ago over a vast region with a range of ∆ mean annual temperature of 3°C and one of ∆ 500 mm total precipitation. We used a set of multivariate a priori causal hypotheses to test the influence of time since fire (TSF), climate, soil physico‐chemistry and bryophyte dominance on forest soil organic C accumulation. Integrating the direct and indirect effects among abiotic and biotic variables explained as much as 50% of the full model variability. The main direct drivers of soil C stocks were: TSF >bryophyte dominance of the FH layer and metal oxide content >pH of the mineral soil. Only climate parameters related to water availability contributed significantly to explaining soil C stock variation. Importantly, climate was found to affect FH layer and mineral soil C stocks indirectly through its effects on bryophyte dominance and organo‐metal complexation, respectively. Soil texture had no influence on soil C stocks. Soil C stocks increased both in the FH layer and mineral soil with TSF and this effect was linked to a decrease in pH with TSF in mineral soil. TSF thus appears to be an important factor of soil development and of C sequestration in mineral soil through its influence on soil chemistry. Overall, this work highlights that integrating the complex interplay between the main drivers of soil C stocks into mechanistic models of C dynamics could improve our ability to assess C stocks and better anticipate the response of the boreal forest to global change. [ABSTRACT FROM AUTHOR]

Abstract: Climate change impact assessments are plagued with uncertainties from many sources, such as climate projections or the inadequacies in structure and parameters of the impact model. Previous studies tried to account for the uncertainty from one or two of these. Here, we developed a triple‐ensemble probabilistic assessment using seven crop models, multiple sets of model parameters and eight contrasting climate projections together to comprehensively account for uncertainties from these three important sources. We demonstrated the approach in assessing climate change impact on barley growth and yield at Jokioinen, Finland in the Boreal climatic zone and Lleida, Spain in the Mediterranean climatic zone, for the 2050s. We further quantified and compared the contribution of crop model structure, crop model parameters and climate projections to the total variance of ensemble output using Analysis of Variance (ANOVA). Based on the triple‐ensemble probabilistic assessment, the median of simulated yield change was −4% and +16%, and the probability of decreasing yield was 63% and 31% in the 2050s, at Jokioinen and Lleida, respectively, relative to 1981–2010. The contribution of crop model structure to the total variance of ensemble output was larger than that from downscaled climate projections and model parameters. The relative contribution of crop model parameters and downscaled climate projections to the total variance of ensemble output varied greatly among the seven crop models and between the two sites. The contribution of downscaled climate projections was on average larger than that of crop model parameters. This information on the uncertainty from different sources can be quite useful for model users to decide where to put the most effort when preparing or choosing models or parameters for impact analyses. We concluded that the triple‐ensemble probabilistic approach that accounts for the uncertainties from multiple important sources provide more comprehensive information for quantifying uncertainties in climate change impact assessments as compared to the conventional approaches that are deterministic or only account for the uncertainties from one or two of the uncertainty sources. [ABSTRACT FROM AUTHOR]

Abstract: Rapid warming and changes in water availability at high latitudes alter resource abundance, tree competition, and disturbance regimes. While these changes are expected to disrupt the functioning of boreal forests, their ultimate implications for forest composition are uncertain. In particular, recent site‐level studies of the Alaskan boreal forest have reported both increases and decreases in productivity over the past few decades. Here, we test the idea that variations in Alaskan forest growth and mortality rates are contingent on species composition. Using forest inventory measurements and climate data from plots located throughout interior and south‐central Alaska, we show significant growth and mortality responses associated with competition, midsummer vapor pressure deficit, and increased growing season length. The governing climate and competition processes differed substantially across species. Surprisingly, the most dramatic climate response occurred in the drought tolerant angiosperm species, trembling aspen, and linked high midsummer vapor pressure deficits to decreased growth and increased insect‐related mortality. Given that species composition in the Alaskan and western Canadian boreal forests is projected to shift toward early‐successional angiosperm species due to fire regime, these results underscore the potential for a reduction in boreal productivity stemming from increases in midsummer evaporative demand. [ABSTRACT FROM AUTHOR]

Abstract: Insects, diseases, fire and drought and other disturbances associated with global climate change contribute to forest decline and mortality in many parts of the world. Forest decline and mortality related to drought or insect outbreaks have been observed in North American aspen forests. However, little research has been done to partition and estimate their relative contributions to growth declines. In this study, we combined tree‐ring width and basal area increment series from 40 trembling aspen (Populus tremuloides Michx.) sites along a latitudinal gradient (from 52° to 58°N) in western Canada and attempted to investigate the effect of drought and insect outbreaks on growth decline, and simultaneously partition and quantify their relative contributions. Results indicated that the influence of drought on forest decline was stronger than insect outbreaks, although both had significant effects. Furthermore, the influence of drought and insect outbreaks showed spatiotemporal variability. In addition, our data suggest that insect outbreaks could be triggered by warmer early spring temperature instead of drought, implicating that potentially increased insect outbreaks are expected with continued warming springs, which may further exacerbate growth decline and death in North America aspen mixed forests. [ABSTRACT FROM AUTHOR]

Across Canada's boreal forest, linear disturbances, including cutlines such as seismic lines and roads, crisscross the landscape to facilitate resource exploration and extraction; many of these linear disturbances cross peatland ecosystems. Changes in tree canopy cover and the compression of the peat by heavy equipment alter local thermal, hydrological, and ecological conditions, likely changing carbon exchange on the disturbance, and possibly in the adjacent peatland. We measured bulk density, water table, soil temperature, plant cover, and CO2 and CH4 flux along triplicate transects crossing a winter road through a wooded fen near Peace River, Alberta, Canada. Sample plots were located 1, 5, and 10 m from the road on both sides with an additional three plots on the road. Productivity of the overstory trees, when present, was also determined. The winter road had higher bulk density, shallower water table, higher graminoid cover, and thawed earlier than the adjacent peatland. Tree productivity and CO2 flux varied between the plots, and there was no clear pattern in relation to distance from the road. The plots on the winter road acted as a greater CO2 sink and greater CH4 source compared to the adjacent peatland with plots on the winter road emitting on average (standard error) 479 (138) compared to 41 (10) mg CH4 m−2 day−1 in the adjacent peatland. Considering both gases, global warming potential increased from 70 to 250 g CO2e m−2 year−1 in the undisturbed area to 2100 g CO2e m−2 year−1 on the winter road. Although carbon fluxes on any given cutline through peatland will vary depending on level of compaction, line width and vegetation community shifts, the large number of linear disturbances in Canada's boreal forest and slow recovery on peatland ecosites suggest they could represent an important anthropogenic greenhouse gas source. [ABSTRACT FROM AUTHOR]

In the sporadic permafrost zone of northwestern Canada, boreal forest carbon dioxide ( CO2) fluxes will be altered directly by climate change through changing meteorological forcing and indirectly through changes in landscape functioning associated with thaw-induced collapse-scar bog ('wetland') expansion. However, their combined effect on landscape-scale net ecosystem CO2 exchange ( NEELAND), resulting from changing gross primary productivity ( GPP) and ecosystem respiration ( ER), remains unknown. Here, we quantify indirect land cover change impacts on NEELAND and direct climate change impacts on modeled temperature- and light-limited NEELAND of a boreal forest-wetland landscape. Using nested eddy covariance flux towers, we find both GPP and ER to be larger at the landscape compared to the wetland level. However, annual NEELAND (−20 g C m−2) and wetland NEE (−24 g C m−2) were similar, suggesting negligible wetland expansion effects on NEELAND. In contrast, we find non-negligible direct climate change impacts when modeling NEELAND using projected air temperature and incoming shortwave radiation. At the end of the 21st century, modeled GPP mainly increases in spring and fall due to reduced temperature limitation, but becomes more frequently light-limited in fall. In a warmer climate, ER increases year-round in the absence of moisture stress resulting in net CO2 uptake increases in the shoulder seasons and decreases during the summer. Annually, landscape net CO2 uptake is projected to decline by 25 ± 14 g C m−2 for a moderate and 103 ± 38 g C m−2 for a high warming scenario, potentially reversing recently observed positive net CO2 uptake trends across the boreal biome. Thus, even without moisture stress, net CO2 uptake of boreal forest-wetland landscapes may decline, and ultimately, these landscapes may turn into net CO2 sources under continued anthropogenic CO2 emissions. We conclude that NEELAND changes are more likely to be driven by direct climate change rather than by indirect land cover change impacts. [ABSTRACT FROM AUTHOR]

At high latitudes, winter climate change alters snow cover and, consequently, may cause a sustained change in soil frost dynamics. Altered winter soil conditions could influence the ecosystem exchange of carbon dioxide ( CO2) and, in turn, provide feedbacks to ongoing climate change. To investigate the mechanisms that modify the peatland CO2 exchange in response to altered winter soil frost, we conducted a snow exclusion experiment to enhance winter soil frost and to evaluate its short-term (1-3 years) and long-term (11 years) effects on CO2 fluxes during subsequent growing seasons in a boreal peatland. In the first 3 years after initiating the treatment, no significant effects were observed on either gross primary production ( GPP) or ecosystem respiration ( ER). However, after 11 years, the temperature sensitivity of ER was reduced in the treatment plots relative to the control, resulting in an overall lower ER in the former. Furthermore, early growing season GPP was also lower in the treatment plots than in the controls during periods with photosynthetic photon flux density ( PPFD) ≥800 μmol m−2 s−1, corresponding to lower sedge leaf biomass in the treatment plots during the same period. During the peak growing season, a higher GPP was observed in the treatment plots under the low light condition (i.e. PPFD 400 μmol m−2 s−1) compared to the control. As Sphagnum moss maximizes photosynthesis at low light levels, this GPP difference between the plots may have been due to greater moss photosynthesis, as indicated by greater moss biomass production, in the treatment plots relative to the controls. Our study highlights the different responses to enhanced winter soil frost among plant functional types which regulate CO2 fluxes, suggesting that winter climate change could considerably alter the growing season CO2 exchange in boreal peatlands through its effect on vegetation development. [ABSTRACT FROM AUTHOR]

Rising global temperatures are suggested to be drivers of shifts in tree species ranges. The resulting changes in community composition may negatively impact forest ecosystem function. However, long-term shifts in tree species ranges remain poorly documented. We test for shifts in the northern range limits of 16 temperate tree species in Quebec, Canada, using forest inventory data spanning three decades, 15° of longitude and 7° of latitude. Range shifts were correlated with climate warming and dispersal traits to understand potential mechanisms underlying changes. Shifts were calculated as the change in the 95th percentile of latitudinal occurrence between two inventory periods (1970-1978, 2000-2012) and for two life stages: saplings and adults. We also examined sapling and adult range offsets within each inventory, and changes in the offset through time. Tree species ranges shifted predominantly northward, although species responses varied. As expected shifts were greater for tree saplings, 0.34 km yr−1, than for adults, 0.13 km yr−1. Range limits were generally further north for adults compared to saplings, but the difference diminished through time, consistent with patterns observed for range shifts within each life stage. This suggests caution should be exercised when interpreting geographic range offsets between life stages as evidence of range shifts in the absence of temporal data. Species latitudinal velocities were on average <50% of the velocity required to equal the spatial velocity of climate change and were mostly unrelated to dispersal traits. Finally, our results add to the body of evidence suggesting tree species are mostly limited in their capacity to track climate warming, supporting concerns that warming will negatively impact the functioning of forest ecosystems. [ABSTRACT FROM AUTHOR]

Turnover concepts in state-of-the-art global vegetation models ( GVMs) account for various processes, but are often highly simplified and may not include an adequate representation of the dominant processes that shape vegetation carbon turnover rates in real forest ecosystems at a large spatial scale. Here, we evaluate vegetation carbon turnover processes in GVMs participating in the Inter-Sectoral Impact Model Intercomparison Project ( ISI- MIP, including HYBRID4, JeDi, JULES, LPJml, ORCHIDEE, SDGVM, and VISIT) using estimates of vegetation carbon turnover rate ( k) derived from a combination of remote sensing based products of biomass and net primary production ( NPP). We find that current model limitations lead to considerable biases in the simulated biomass and in k (severe underestimations by all models except JeDi and VISIT compared to observation-based average k), likely contributing to underestimation of positive feedbacks of the northern forest carbon balance to climate change caused by changes in forest mortality. A need for improved turnover concepts related to frost damage, drought, and insect outbreaks to better reproduce observation-based spatial patterns in k is identified. As direct frost damage effects on mortality are usually not accounted for in these GVMs, simulated relationships between k and winter length in boreal forests are not consistent between different regions and strongly biased compared to the observation-based relationships. Some models show a response of k to drought in temperate forests as a result of impacts of water availability on NPP, growth efficiency or carbon balance dependent mortality as well as soil or litter moisture effects on leaf turnover or fire. However, further direct drought effects such as carbon starvation (only in HYBRID4) or hydraulic failure are usually not taken into account by the investigated GVMs. While they are considered dominant large-scale mortality agents, mortality mechanisms related to insects and pathogens are not explicitly treated in these models. [ABSTRACT FROM AUTHOR]

Adequate and advance knowledge of the response of forest ecosystems to temperature-induced drought is critical for a comprehensive understanding of the impacts of global climate change on forest ecosystem structure and function. Recent massive decline in aspen-dominated forests and an increased aspen mortality in boreal forests have been associated with global warming, but it is still uncertain whether the decline and mortality are driven by drought. We used a series of ring-width chronologies from 40 trembling aspen ( Populus tremuloides Michx.) sites along a latitudinal gradient (from 52° to 58°N) in western Canada, in an attempt to clarify the impacts of drought on aspen growth by using Standardized Precipitation Index ( SPI) and Standardized Precipitation Evapotranspiration Index ( SPEI). Results indicated that prolonged and large-scale droughts had a strong negative impact on trembling aspen growth. Furthermore, the spatiotemporal variability of drought indices is useful for explaining the spatial heterogeneity in the radial growth of trembling aspen. Due to ongoing global warming and rising temperatures, it is likely that severer droughts with a higher frequency will occur in western Canada. As trembling aspen is sensitive to drought, we suggest that drought indices could be applied to monitor the potential effects of increased drought stress on aspen trees growth, achieve classification of eco-regions and develop effective mitigation strategies to maintain western Canadian boreal forests. [ABSTRACT FROM AUTHOR]

Rich fens are common boreal ecosystems with distinct hydrology, biogeochemistry and ecology that influence their carbon (C) balance. We present growing season soil chamber methane emission ( FCH4), ecosystem respiration ( ER), net ecosystem exchange ( NEE) and gross primary production ( GPP) fluxes from a 9-years water table manipulation experiment in an Alaskan rich fen. The study included major flood and drought years, where wetting and drying treatments further modified the severity of droughts. Results support previous findings from peatlands that drought causes reduced magnitude of growing season FCH4, GPP and NEE, thus reducing or reversing their C sink function. Experimentally exacerbated droughts further reduced the capacity for the fen to act as a C sink by causing shifts in vegetation and thus reducing magnitude of maximum growing season GPP in subsequent flood years by ~15% compared to control plots. Conversely, water table position had only a weak influence on ER, but dominant contribution to ER switched from autotrophic respiration in wet years to heterotrophic in dry years. Droughts did not cause inter-annual lag effects on ER in this rich fen, as has been observed in several nutrient-poor peatlands. While ER was dependent on soil temperatures at 2 cm depth, FCH4 was linked to soil temperatures at 25 cm. Inter-annual variability of deep soil temperatures was in turn dependent on wetness rather than air temperature, and higher FCH4 in flooded years was thus equally due to increased methane production at depth and decreased methane oxidation near the surface. Short-term fluctuations in wetness caused significant lag effects on FCH4, but droughts caused no inter-annual lag effects on FCH4. Our results show that frequency and severity of droughts and floods can have characteristic effects on the exchange of greenhouse gases, and emphasize the need to project future hydrological regimes in rich fens. [ABSTRACT FROM AUTHOR]

Species compositional shifts have important consequences to biodiversity and ecosystem function and services to humanity. In boreal forests, compositional shifts from late-successional conifers to early-successional conifers and deciduous broadleaves have been postulated based on increased fire frequency associated with climate change truncating stand age-dependent succession. However, little is known about how climate change has affected forest composition in the background between successive catastrophic fires in boreal forests. Using 1797 permanent sample plots from western boreal forests of Canada measured from 1958 to 2013, we show that after accounting for stand age-dependent succession, the relative abundances of early-successional deciduous broadleaves and early-successional conifers have increased at the expense of late-successional conifers with climate change. These background compositional shifts are persistent temporally, consistent across all forest stand ages and pervasive spatially across the region. Rising atmospheric CO2 promoted early-successional conifers and deciduous broadleaves, and warming increased early-successional conifers at the expense of late-successional conifers, but compositional shifts were not associated with climate moisture index. Our results emphasize the importance of climate change on background compositional shifts in the boreal forest and suggest further compositional shifts as rising CO2 and warming will continue in the 21st century. [ABSTRACT FROM AUTHOR]

Biomass change of the world's forests is critical to the global carbon cycle. Despite storing nearly half of global forest carbon, the boreal biome of diverse forest types and ages is a poorly understood component of the carbon cycle. Using data from 871 permanent plots in the western boreal forest of Canada, we examined net annual aboveground biomass change (Δ AGB) of four major forest types between 1958 and 2011. We found that Δ AGB was higher for deciduous broadleaf ( DEC) (1.44 Mg ha−1 year−1, 95% Bayesian confidence interval ( CI), 1.22-1.68) and early-successional coniferous forests ( ESC) (1.42, CI, 1.30-1.56) than mixed forests ( MIX) (0.80, CI, 0.50-1.11) and late-successional coniferous ( LSC) forests (0.62, CI, 0.39-0.88). Δ AGB declined with forest age as well as calendar year. After accounting for the effects of forest age, Δ AGB declined by 0.035, 0.021, 0.032 and 0.069 Mg ha−1 year−1 per calendar year in DEC, ESC, MIX and LSC forests, respectively. The Δ AGB declines resulted from increased tree mortality and reduced growth in all forest types except DEC, in which a large biomass loss from mortality was accompanied with a small increase in growth. With every degree of annual temperature increase, Δ AGB decreased by 1.00, 0.20, 0.55 and 1.07 Mg ha−1 year−1 in DEC, ESC, MIX and LSC forests, respectively. With every cm decrease of annual climatic moisture availability, Δ AGB decreased 0.030, 0.045 and 0.17 Mg ha−1 year−1 in ESC, MIX and LSC forests, but changed little in DEC forests. Our results suggest that persistent warming and decreasing water availability have profound negative effects on forest biomass in the boreal forests of western Canada. Furthermore, our results indicate that forest responses to climate change are strongly dependent on forest composition with late-successional coniferous forests being most vulnerable to climate changes in terms of aboveground biomass. [ABSTRACT FROM AUTHOR]

Population expansions of boreal species are among the most substantial ecological consequences of climate change, potentially transforming both structure and processes of northern ecosystems. Despite their importance, little is known about expansion dynamics of boreal species. Red foxes (Vulpes vulpes) are forecasted to become a keystone species in northern Europe, a process stemming from population expansions that began in the 19th century. To identify the relative roles of geographic and demographic factors and the sources of northern European red fox population expansion, we genotyped 21 microsatellite loci in modern and historical (1835-1941) Fennoscandian red foxes. Using Bayesian clustering and Bayesian inference of migration rates, we identified high connectivity and asymmetric migration rates across the region, consistent with source-sink dynamics, whereby more recently colonized sampling regions received immigrants from multiple sources. There were no clear clines in allele frequency or genetic diversity as would be expected from a unidirectional range expansion from south to north. Instead, migration inferences, demographic models and comparison to historical red fox genotypes suggested that the population expansion of the red fox is a consequence of dispersal from multiple sources, as well as in situ demographic growth. Together, these findings provide a rare glimpse into the anatomy of a boreal range expansion and enable informed predictions about future changes in boreal communities. [ABSTRACT FROM AUTHOR]

Rising temperatures caused by climate change could negatively alter plant ecosystems if temperatures exceed optimal temperatures for carbon gain. Such changes may threaten temperature-sensitive species, causing local extinctions and range migrations. This study examined the optimal temperature of net photosynthesis ( Topt) of two boreal and four temperate deciduous tree species grown in the field in northern Minnesota, United States under two contrasting temperature regimes. We hypothesized that Topt would be higher in temperate than co-occurring boreal species, with temperate species exhibiting greater plasticity in Topt, resulting in better acclimation to elevated temperatures. The chamberless experiment, located at two sites in both open and understory conditions, continuously warmed plants and soils during three growing seasons. Results show a modest, but significant shift in Topt of 1.1 ± 0.21 °C on average for plants subjected to a mean 2.9 ± 0.01 °C warming during midday hours in summer, and shifts with warming were unrelated to species native ranges. The 1.1 °C shift in Topt with 2.9 °C warming might be interpreted as suggesting limited capacity to shift temperature response functions to better match changes in temperature. However, Topt of warmed plants was as well-matched with prior midday temperatures as Topt of plants in the ambient treatment, and Topt in both treatments was at a level where realized photosynthesis was within 90-95% of maximum. These results suggest that seedlings of all species were close to optimizing photosynthetic temperature responses, and equally so in both temperature treatments. Our study suggests that temperate and boreal species have considerable capacity to match their photosynthetic temperature response functions to prevailing growing season temperatures that occur today and to those that will likely occur in the coming decades under climate change. [ABSTRACT FROM AUTHOR]

Conservation strategies are often established without consideration of the impact of climate change. However, this impact is expected to threaten species and ecosystem persistence and to have dramatic effects towards the end of the 21st century. Landscape suitability for species under climate change is determined by several interacting factors including dispersal and human land use. Designing effective conservation strategies at regional scales to improve landscape suitability requires measuring the vulnerabilities of specific regions to climate change and determining their conservation capacities. Although methods for defining vulnerability categories are available, methods for doing this in a systematic, cost-effective way have not been identified. Here, we use an ecosystem model to define the potential resilience of the Finnish forest landscape by relating its current conservation capacity to its vulnerability to climate change. In applying this framework, we take into account the responses to climate change of a broad range of red-listed species with different niche requirements. This framework allowed us to identify four categories in which representation in the landscape varies among three IPCC emission scenarios (B1, low; A1B, intermediate; A2, high emissions): (i) susceptible (B1 = 24.7%, A1B = 26.4%, A2 = 26.2%), the most intact forest landscapes vulnerable to climate change, requiring management for heterogeneity and resilience; (ii) resilient (B1 = 2.2%, A1B = 0.5%, A2 = 0.6%), intact areas with low vulnerability that represent potential climate refugia and require conservation capacity maintenance; (iii) resistant (B1 = 6.7%, A1B = 0.8%, A2 = 1.1%), landscapes with low current conservation capacity and low vulnerability that are suitable for restoration projects; (iv) sensitive (B1 = 66.4%, A1B = 72.3%, A2 = 72.0%), low conservation capacity landscapes that are vulnerable and for which alternative conservation measures are required depending on the intensity of climate change. Our results indicate that the Finnish landscape is likely to be dominated by a very high proportion of sensitive and susceptible forest patches, thereby increasing uncertainty for landscape managers in the choice of conservation strategies. [ABSTRACT FROM AUTHOR]

In cold climates, the expected global warming will lead to earlier cambial resumptions in spring, with a resultant lengthening of the growing season but unknown consequences on forest productivity. The phenological traits of cambium activity and xylem formation were analyzed at a short time scale along a thermal gradient represented by an alti-latitudinal range from the 48th to 53rd parallels and covering the whole closed black-spruce [ Picea mariana (Mill.) BSP] forest in Quebec, Canada. A hypothesis was tested that warmer temperatures influence cambium phenology, allowing longer duration and higher intensity of growth, and resulting in proportionally increased xylem production. From April to October 2012, cell division in cambium and post-cambial differentiation of xylem were observed on anatomical sections obtained from microcores collected weekly from the stem of fifty trees. The southern and warmer site was characterized by the highest radial growth, which corresponded to both the highest rates and longest durations of cell production. The differences in terms of xylem phenology and growth were marginal between the other sites. Xylem growth was positively correlated with rate and duration of cell production, with the latter explaining most variability in growth. Within the range analyzed, the relationship between temperature and most phenological phases of xylogenesis was linear. On the contrary, temperature was related with cell production according to an exponential pattern. Periods of xylogenesis of 14 days longer (+13.1%) corresponded to a massive increase in cell production (33 cells, +109%). This disproportionate change occurred at a May-September average temperature of ca. 14 °C and a snow-free period of 210-235 days. At the lower boundary of the distribution of black spruce, small environmental changes allowing marginal lengthening of the period of cell division could potentially lead to disproportionate increases in xylem cell production, with substantial consequences for the productivity of this boreal species. [ABSTRACT FROM AUTHOR]

Changes in climate are influencing the distribution and abundance of the world's biota, with significant consequences for biological diversity and ecosystem processes. Recent work has raised concern that populations of moths and butterflies (Lepidoptera) may be particularly susceptible to population declines under environmental change. Moreover, effects of climate change may be especially pronounced in high latitude ecosystems. Here, we examine population dynamics in an assemblage of subarctic forest moths in Finnish Lapland to assess current trajectories of population change. Moth counts were made continuously over a period of 32 years using light traps. From 456 species recorded, 80 were sufficiently abundant for detailed analyses of their population dynamics. Climate records indicated rapid increases in temperature and winter precipitation at our study site during the sampling period. However, 90% of moth populations were stable (57%) or increasing (33%) over the same period of study. Nonetheless, current population trends do not appear to reflect positive responses to climate change. Rather, time-series models illustrated that the per capita rates of change of moth species were more frequently associated negatively than positively with climate change variables, even as their populations were increasing. For example, the per capita rates of change of 35% of microlepidoptera were associated negatively with climate change variables. Moth life-history traits were not generally strong predictors of current population change or associations with climate change variables. However, 60% of moth species that fed as larvae on resources other than living vascular plants (e.g. litter, lichen, mosses) were associated negatively with climate change variables in time-series models, suggesting that such species may be particularly vulnerable to climate change. Overall, populations of subarctic forest moths in Finland are performing better than expected, and their populations appear buffered at present from potential deleterious effects of climate change by other ecological forces. [ABSTRACT FROM AUTHOR]

Empirical models alongside remotely sensed and station measured meteorological observations are employed to investigate both the local and global direct climate change impacts of alternative forest management strategies within a boreal ecosystem of eastern Norway. Stand-level analysis is firstly executed to attribute differences in daily, seasonal, and annual mean surface temperatures to differences in surface intrinsic biophysical properties across conifer, deciduous, and clear-cut sites. Relative to a conifer site, a slight local cooling of −0.13 °C at a deciduous site and −0.25 °C at a clear-cut site were observed over a 6-year period, which were mostly attributed to a higher albedo throughout the year. When monthly mean albedo trajectories over the entire managed forest landscape were taken into consideration, we found that strategies promoting natural regeneration of coniferous sites with native deciduous species led to substantial global direct climate cooling benefits relative to those maintaining current silviculture regimes - despite predicted long-term regional warming feedbacks and a reduced albedo in spring and autumn months. The magnitude and duration of the cooling benefit depended largely on whether management strategies jointly promoted an enhanced material supply over business-as-usual levels. Expressed in terms of an equivalent CO2 emission pulse at the start of the simulation, the net climate response at the end of the 21st century spanned −8 to −159 Tg- CO2-eq., depending on whether near-term harvest levels increased or followed current trends, respectively. This magnitude equates to approximately −20 to −300% of Norway's annual domestic (production) emission impact. Our analysis supports the assertion that a carbon-only focus in the design and implementation of forest management policy in boreal and other climatically similar regions can be counterproductive - and at best - suboptimal if boreal forests are to be used as a tool to mitigate global warming. [ABSTRACT FROM AUTHOR]

It is proposed that increases in anthropogenic reactive nitrogen (Nr) deposition may cause temperate and boreal forests to sequester a globally significant quantity of carbon (C); however, long-term data from boreal forests describing how C sequestration responds to realistic levels of chronic Nr deposition are scarce. Using a long-term (14-year) stand-scale (0.1 ha) N addition experiment (three levels: 0, 12.5, and 50 kg N ha−1 yr−1) in the boreal zone of northern Sweden, we evaluated how chronic N additions altered N uptake and biomass of understory communities, and whether changes in understory communities explained N uptake and C sequestration by trees. We hypothesized that understory communities (i.e. mosses and shrubs) serve as important sinks for low-level N additions, with the strength of these sinks weakening as chronic N addition rates increase, due to shifts in species composition. We further hypothesized that trees would exhibit nonlinear increases in N acquisition, and subsequent C sequestration as N addition rates increased, due to a weakening understory N sink. Our data showed that understory biomass was reduced by 50% in response to the high N addition treatment, mainly due to reduced moss biomass. A 15N labeling experiment showed that feather mosses acquired the largest fraction of applied label, with this fraction decreasing as the chronic N addition level increased. Contrary to our hypothesis, the proportion of label taken up by trees was equal (ca. 8%) across all three N addition treatments. The relationship between N addition and C sequestration in all vegetation pools combined was linear, and had a slope of 16 kg C kg−1 N. While canopy retention of Nr deposition may cause C sequestration rates to be slightly different than this estimate, our data suggest that a minor quantity of annual anthropogenic CO2 emissions are sequestered into boreal forests as a result of Nr deposition. [ABSTRACT FROM AUTHOR]

Investigations of biophysical changes on earth caused by anthropogenic disturbance provide governments with tools to generate sustainable development policy. Canada currently experiences one of the fastest rates of boreal forest disturbance in the world. Plans to conserve the 330 000 km2 boreal forest in the province of Alberta exist but conservation targets and schedules must be aligned with rates of forest disturbance. We explore how disturbance rate, and the accuracy with which we detect it, may affect conservation success. We performed a change detection analysis from 1992 to 2008 using Landsat and SPOT satellite image data processing. Canada's recovery strategy for boreal caribou ( Rangifer tarandus caribou) states that ≤35% of a caribou range can be either burned or within 500 m of a man-made feature for caribou to recover. Our analyses show that by 2008 78% of the boreal forest was disturbed and that, if the current rate continues, 100% would be disturbed by 2028. Alberta plans to set aside 22% for conservation in a region encompassing oil sands development to balance economic, environmental, and traditional indigenous land-use goals. Contrary to the federal caribou recovery strategy, provincial conservation plans do not consider wildfire a disturbance. Based on analyses used in the provincial plan, we apply a 250 m buffer around anthropogenic footprints. Landsat image analysis indicates that the yearly addition of disturbance is 714 km2 (0.8%). The higher resolution SPOT images show fine-scale disturbance indicating that actual disturbance was 1.28 times greater than detected by Landsat. If the SPOT image based disturbance rates continue, the 22% threshold may be exceeded within the next decade, up to 20 years earlier than indicated by Landsat-based analysis. Our results show that policies for sustainable development will likely fail if governments do not develop time frames that are grounded by accurate calculations of disturbance rates. [ABSTRACT FROM AUTHOR]

The boreal Northeast Atlantic is strongly affected by current climate change, and large shifts in abundance and distribution of many organisms have been observed, including the dominant copepod Calanus finmarchicus, which supports the grazing food web and thus many fish populations. At the same time, large-scale declines have been observed in many piscivorous seabirds, which depend on abundant small pelagic fish. Here, we combine predictions from a niche model of C. finmarchicus with long-term data on seabird breeding success to link trophic levels. The niche model shows that environmental suitability for C. finmarchicus has declined in southern areas with large breeding seabird populations (e.g. the North Sea), and predicts that this decline is likely to spread northwards during the 21st century to affect populations in Iceland and the Faroes. In a North Sea colony, breeding success of three common piscivorous seabird species [black-legged kittiwake ( Rissa tridactyla), common guillemot ( Uria aalge) and Atlantic puffin ( Fratercula arctica)] was strongly positively correlated with local environmental suitability for C. finmarchicus, whereas this was not the case at a more northerly colony in west Norway. Large seabird populations seem only to occur where C. finmarchicus is abundant, and northward distributional shifts of common boreal seabirds are therefore expected over the coming decades. Whether or not population size can be maintained depends on the dispersal ability and inclination of these colonial breeders, and on the carrying capacity of more northerly areas in a warmer climate. [ABSTRACT FROM AUTHOR]

To predict the long-term effects of climate change - global warming and changes in precipitation - on the diameter (radial) growth of jack pine ( Pinus banksiana Lamb.) and black spruce ( Picea mariana [ Mill.] B.S.P.) trees in boreal Ontario, we modified an existing diameter growth model to include climate variables. Diameter chronologies of 927 jack pine and 1173 black spruce trees, growing in the area from 47°N to 50°N and 80°W to 92°W, were used to develop diameter growth models in a nonlinear mixed-effects approach. Our results showed that the variables long-term average of mean growing season temperature, precipitation during wettest quarter, and total precipitation during growing season were significant (alpha = 0.05) in explaining variation in diameter growth of the sample trees. Model results indicated that higher temperatures during the growing season would increase the diameter growth of jack pine trees, but decrease that of black spruce trees. More precipitation during the wettest quarter would favor the diameter growth of both species. On the other hand, a wetter growing season, which may decrease radiation inputs, increase nutrient leaching, and reduce the decomposition rate, would reduce the diameter growth of both species. Moreover, our results indicated that future (2041-2070) diameter growth rate may differ from current (1971-2000) growth rates for both species, with conditions being more favorable for jack pine than black spruce trees. Expected future changes in the growth rate of boreal trees need to be considered in forest management decisions. We recommend that knowledge of climate-growth relationships, as represented by models, be combined with learning from adaptive management to reduce the risks and uncertainties associated with forest management decisions. [ABSTRACT FROM AUTHOR]

Many ecosystems are currently undergoing dramatic changes in biodiversity due to habitat loss and climate change. Responses to global change at the community level are poorly understood, as are the impacts of community disassembly on ecosystem-level processes. Uncertainties remain regarding the patterns of extirpation and persistence under single vs. multiple forms of environmental change. Here, we use a trait-based and food web approach to examine the effects of experimentally changing moisture, temperature and habitat 'openness' on a functionally important group of microarthropods associated with a boreal forest floor bryosphere (detrital moss) system. Overall, the outcome of community disassembly was mediated by the correlation between our environmental factors and species traits, particularly body size. Minor increases in summer temperatures maintained greater species richness, whereas drought stress had a significant negative effect on community-level abundance and richness. These effects were reflected in modifications to the community-wide body-size spectra. Habitat openness alleviated biodiversity loss in the larger-bodied species of the most abundant taxonomic group, but did not fully mitigate the effects of drought. The most striking result of this experiment was an overall contraction of the food web among persistent species under drought stress (i.e. those not extirpated by environmental change). These results suggest that major changes in boreal microarthropod community structure are likely to occur in response to common forms of global change. Moreover, the contraction in trophic structure even amongst tolerant species suggests that ecosystem function within the bryosphere can be altered by environmental change. [ABSTRACT FROM AUTHOR]

Nitrogen (N) deposition has increased globally over the last 150 years and further increases are predicted. Epiphytic lichens decline in abundance and diversity in areas with high N loads, and the abundance of lichens decreases along gradients of increased deposition. Thus, although N is an essential nutrient for lichens, excessive loads may be detrimental for them. However, these gradients include many correlated pollutants and the mechanisms behind the decline are thus poorly known. The aim of this study was to assess effects of N deposition, alone, on the epiphytic lichen community composition in a naturally N-poor boreal forest. For this purpose, whole spruce trees were fertilized daily with N at five levels, equivalent to 0.6, 6, 12.5, 25, and 50 kg N ha−1 yr−1, during four consecutive growing seasons (2006-2009), and changes in the abundance of lichens were monitored each autumn from the preceding year (2005). The studied lichen communities were highly dynamic and responded strongly to the environmental perturbation. N deposition detectably altered the direction of succession and reduced the species richness of the epiphytic lichen communities, even at the lowest fertilization application (6 kg N ha−1 yr−1). The simulated N deposition caused significant changes in the abundance of Alectoria sarmentosa, Bryoria spp., and Hypogymnia physodes, which all increased at low N loads and decreased at high loads, but with species-specific optima. The rapid decline of A. sarmentosa may have been caused by the added nitrogen reducing the stability of the lichen thalli, possibly due to increases in the photobiont: mycobiont ratio or parasitic fungal attacks. We conclude that increases in nitrogen availability, per se, could be responsible for the reductions in lichen abundance and diversity observed along deposition gradients, and those community responses may be due to physiological responses of the individual species rather than changes in competitive interactions. [ABSTRACT FROM AUTHOR]

During the past ∼50 years, the number and area of lakes have declined in several regions in boreal forests. However, there has been substantial finer-scale heterogeneity; some lakes decreased in area, some showed no trend, and others increased. The objective of this study was to identify the primary mechanisms underlying heterogeneous trends in closed-basin lake area. Eight lake characteristics (δ18O, electrical conductivity, surface : volume index, bank slope, floating mat width, peat depth, thaw depth at shoreline, and thaw depth at the forest boundary) were compared for 15 lake pairs in Alaskan boreal forest where one lake had decreased in area since ∼ 1950, and the other had not. Mean differences in characteristics between paired lakes were used to identify the most likely of nine mechanistic scenarios that combined three potential mechanisms for decreasing lake area (talik drainage, surface water evaporation, and terrestrialization) with three potential mechanisms for nondecreasing lake area (subpermafrost groundwater recharge through an open talik, stable permafrost, and thermokarst). A priori expectations of the direction of mean differences between decreasing and nondecreasing paired lakes were generated for each scenario. Decreasing lakes had significantly greater electrical conductivity, greater surface : volume indices, shallower bank slopes, wider floating mats, greater peat depths, and shallower thaw depths at the forest boundary. These results indicated that the most likely scenario was terrestrialization as the mechanism for lake area reduction combined with thermokarst as the mechanism for nondecreasing lake area. Terrestrialization and thermokarst may have been enhanced by recent warming which has both accelerated permafrost thawing and lengthened the growing season, thereby increasing plant growth, floatingmat encroachment, transpiration rates, and the accumulation of organic matter in lake basins. The transition to peatlands associated with terrestrialization may provide a transient increase in carbon storage enhancing the role of northern ecosystems as major stores of global carbon. [ABSTRACT FROM AUTHOR]

Productivity in boreal ecosystems is primarily limited by available soil nitrogen (N), and there is substantial interest in understanding whether deposition of anthropogenically derived reactive nitrogen (Nr) results in greater N availability to woody vegetation, which could result in greater carbon (C) sequestration. One factor that may limit the acquisition of Nr by woody plants is the presence of bryophytes, which are a significant C and N pool, and a location where associative cyanobacterial N-fixation occurs. Using a replicated stand-scale N-addition experiment (five levels: 0, 3, 6, 12, and 50 kgNha-1 yr-1; n56) in the boreal zone of northern Sweden, we tested the hypothesis that sequestration of Nr into bryophyte tissues, and downregulation of N-fixation would attenuate Nr inputs, and thereby limit anthropogenic Nr acquisition by woody plants. Our data showed that N-fixation per unit moss mass and per unit area sharply decreased with increasing N addition. Additionally, the tissue N concentrations of Pleuorzium schreberi increased and its biomass decreased with increasing N addition. This response to increasing N addition caused the P. schreberi N pool to be stable at all but the highest N addition rate, where it significantly decreased. The combined effects of changed N-fixation and P. schreberi biomass N accounted for 56.7% of cumulative Nr additions at the lowest Nr addition rate, but only a minor fraction for all other treatments. This 'bryophyte effect' can in part explain why soil inorganic N availability and acquisition by woody plants (indicated by their δ15N signatures) remained unchanged up to N addition rates of 12 kg ha-1 yr-1 or greater. Finally, we demonstrate that approximately 71.8% of the boreal forest experiences Nr deposition rates at or below 3 kg ha-1 yr-1, suggesting that bryophytes likely limit woody plant acquisition of ambient anthropogenic Nr inputs throughout a majority of the boreal forest. [ABSTRACT FROM AUTHOR]

The Northern Hemisphere's boreal forests, particularly the Siberian boreal forest, may have a strong effect on Earth's climate through changes in dominant vegetation and associated regional surface albedo. We show that warmer climate will likely convert Siberia's deciduous larch ( Larix spp.) to evergreen conifer forests, and thus decrease regional surface albedo. The dynamic vegetation model, FAREAST, simulates Russian boreal forest composition and was used to explore the feedback between climate change and forest composition at continental, regional, and local scales. FAREAST was used to simulate the impact of changes in temperature and precipitation on total and genus-level biomass at sites across Siberia and the Russian Far East (RFE), and for six high- and low-diversity regions. Model runs with and without European Larch ( Larix decidua) included in the available species pool were compared to assess the potential for this species, which is adapted to warmer climate conditions, to mitigate the effects of climate change, especially the shift to evergreen dominance. At the continental scale, when temperature is increased, larch-dominated sites become vulnerable to early replacement by evergreen conifers. At the regional and local scales, the diverse Amur region of the RFE does not show a strong response to climate change, but the low-diversity regions in central and southern Siberia have an abrupt vegetation shift from larch-dominated forest to evergreen-conifer forest in response to increased temperatures. The introduction of L. decidua prevents the collapse of larch in these low-diversity areas and thus mitigates the response to warming. Using contemporary MODIS albedo measurements, we determined that a conversion from larch to evergreen stands in low-diversity regions of southern Siberia would generate a local positive radiative forcing of 5.1±2.6 W m. This radiative heating would reinforce the warming projected to occur in the area under climate change. [ABSTRACT FROM AUTHOR]

High-latitude regions store large amounts of organic carbon (OC) in active-layer soils and permafrost, accounting for nearly half of the global belowground OC pool. In the boreal region, recent warming has promoted changes in the fire regime, which may exacerbate rates of permafrost thaw and alter soil OC dynamics in both organic and mineral soil. We examined how interactions between fire and permafrost govern rates of soil OC accumulation in organic horizons, mineral soil of the active layer, and near-surface permafrost in a black spruce ecosystem of interior Alaska. To estimate OC accumulation rates, we used chronosequence, radiocarbon, and modeling approaches. We also developed a simple model to track long-term changes in soil OC stocks over past fire cycles and to evaluate the response of OC stocks to future changes in the fire regime. Our chronosequence and radiocarbon data indicate that OC turnover varies with soil depth, with fastest turnover occurring in shallow organic horizons (∼60 years) and slowest turnover in near-surface permafrost (>3000 years). Modeling analysis indicates that OC accumulation in organic horizons was strongly governed by carbon losses via combustion and burial of charred remains in deep organic horizons. OC accumulation in mineral soil was influenced by active layer depth, which determined the proportion of mineral OC in a thawed or frozen state and thus, determined loss rates via decomposition. Our model results suggest that future changes in fire regime will result in substantial reductions in OC stocks, largely from the deep organic horizon. Additional OC losses will result from fire-induced thawing of near-surface permafrost. From these findings, we conclude that the vulnerability of deep OC stocks to future warming is closely linked to the sensitivity of permafrost to wildfire disturbance. [ABSTRACT FROM AUTHOR]

The fate of carbon (C) contained within permafrost in boreal forest environments is an important consideration for the current and future carbon cycle as soils warm in northern latitudes. Currently, little is known about the microbiology or chemistry of permafrost soils that may affect its decomposition once soils thaw. We tested the hypothesis that low microbial abundances and activities in permafrost soils limit decomposition rates compared with active layer soils. We examined active layer and permafrost soils near Fairbanks, AK, the Yukon River, and the Arctic Circle. Soils were incubated in the lab under aerobic and anaerobic conditions. Gas fluxes at −5 and 5 °C were measured to calculate temperature response quotients ( Q10). The Q10 was lower in permafrost soils (average 2.7) compared with active layer soils (average 7.5). Soil nutrients, leachable dissolved organic C (DOC) quality and quantity, and nuclear magnetic resonance spectroscopy of the soils revealed that the organic matter within permafrost soils is as labile, or even more so, than surface soils. Microbial abundances (fungi, bacteria, and subgroups: methanogens and Basidiomycetes) and exoenzyme activities involved in decomposition were lower in permafrost soils compared with active layer soils, which, together with the chemical data, supports the reduced Q10 values. CH4 fluxes were correlated with methanogen abundance and the highest CH4 production came from active layer soils. These results suggest that permafrost soils have high inherent decomposability, but low microbial abundances and activities reduce the temperature sensitivity of C fluxes. Despite these inherent limitations, however, respiration per unit soil C was higher in permafrost soils compared with active layer soils, suggesting that decomposition and heterotrophic respiration may contribute to a positive feedback to warming of this eco region. [ABSTRACT FROM AUTHOR]

As atmospheric CO2 levels rise, temperate and boreal forests in the Northern Hemisphere are gaining importance as carbon sinks. Quantification of that role, however, has been difficult due to the confounding effects of climate change. Recent large-scale experiments with quaking aspen ( Populus tremuloides), a dominant species in many northern forest ecosystems, indicate that elevated CO2 levels can enhance net primary production. Field studies also reveal that droughts contribute to extensive aspen mortality. To complement this work, we analyzed how the growth of wild aspen clones in Wisconsin has responded to historical shifts in CO2 and climate, accounting for age, genotype (microsatellite heterozygosity), and other factors. Aspen growth has increased an average of 53% over the past five decades, primarily in response to the 19.2% rise in ambient CO2 levels. CO2-induced growth is particularly enhanced during periods of high moisture availability. The analysis accounts for the highly nonlinear changes in growth rate with age, and is unaffected by sex or location sampled. Growth also increases with individual heterozygosity, but this heterozygote advantage has not changed with rising levels of CO2 or moisture. Thus, increases in future growth predicted from previous large-scale, common-garden work are already evident in this abundant and ecologically important tree species. Owing to aspen's role as a foundation species in many North American forest ecosystems, CO2-stimulated growth is likely to have repercussions for numerous associated species and ecosystem processes. [ABSTRACT FROM AUTHOR]

Natural fires annually decimate up to 1% of the forested area in the boreal region of Québec, and represent a major structuring force in the region, creating a mosaic of watersheds characterized by large variations in vegetation structure and composition. Here, we investigate the possible connections between this fire-induced watershed heterogeneity and lake metabolism and CO2 dynamics. Plankton respiration, and water–air CO2 fluxes were measured in the epilimnia of 50 lakes, selected to lie within distinct watershed types in terms of postfire terrestrial succession in the boreal region of Northern Québec. Plankton respiration varied widely among lakes (from 21 to 211 μg C L−1 day−1), was negatively related to lake area, and positively related to dissolved organic carbon (DOC). All lakes were supersaturated in CO2 and the resulting carbon (C) flux to the atmosphere (150 to over 3000 mg C m2 day−1) was negatively related to lake area and positively to DOC concentration. CO2 fluxes were positively related to integrated water column respiration, suggesting a biological component in this flux. Both respiration and CO2 fluxes were strongly negatively related to years after the last fire in the basin, such that lakes in recently burnt basins had significantly higher C emissions, even after the influence of lake size was removed. No significant differences were found in nutrients, chlorophyll, and DOC between lakes in different basin types, suggesting that the fire-induced watershed features influence other, more subtle aspects, such as the quality of the organic C reaching lakes. The fire-induced enhancement of lake organic C mineralization and C emissions represents a long-term impact that increases the overall C loss from the landscape as the result of fire, but which has never been included in current regional C budgets and future projections. The need to account for this additional fire-induced C loss becomes critical in the face of predictions of increasing incidence of fire in the circumboreal landscape. [ABSTRACT FROM AUTHOR]

A large area of boreal jack pine ( Pinus banksiana Lamb.) forest in Canada is recovering from clear-cut harvesting, and the carbon (C) balance of these regenerating forests remains uncertain. Net ecosystem CO2 exchange was measured using the eddy-covariance technique at four jack pine sites representing different stages of stand development: three postharvest sites (HJP02, HJP94, and HJP75) and one preharvest site (OJP). The four sites, located in the southern Canadian boreal forest, Saskatchewan, Canada, are typical of low productivity jack pine stands and were 2, 10, 29, and 90 years old in 2004, respectively. Mean annual net ecosystem production (NEP) for 2004 and 2005 was -137±11, 19±16, 73±28, and 22±30 g C m-2 yr-1 at HJP02, HJP94, HJP75 and OJP, respectively, showing the postharvest jack pine stands to be moderate C sources immediately after harvesting, weak sinks at 10 years, moderate C sinks at 30 years, then weak C sinks at 90 years. Mean annual gross ecosystem photosynthesis (GEP) for the 2 years was 96±10, 347±20, 576±34, and 583±35 g C m-2 yr-1 at HJP02, HJP94, HJP75, and OJP, respectively. The ratio of annual ecosystem respiration ( R) to annual GEP was 2.51±0.15, 0.95±0.04, 0.87±0.03, and 0.96±0.03. Seasonally, NEP peaked in May or June at all four sites but GEP and R were highest in July. R at a reference soil temperature of 10 °C, ecosystem quantum yield and photosynthetic capacity were lowest for the 2-year-old stand. R was most sensitive to soil temperature for the 90-year-old stand. The primary source of variability in NEP over the course of succession of the jack pine ecosystem following harvesting was stand age due to the changes in leaf area index. Intersite variability in GEP and R was an order of magnitude greater than interannual variability at OJP. For both young and old stands, GEP had greater interannual variability than R and played a more important role than R in interannual variation in NEP. Based on year-round flux measurements from 2000 to 2005, the 10-year stand had larger interannual variability in GEP and R than the 90-year stand. Interannual variability in NEP was driven primarily by early-growing-season temperature and growing-season length. Photosynthesis played a dominant role in the rapid rise in NEP early in stand development. Late in stand development, however, the subtle decrease in NEP resulted primarily from increasing respiration. [ABSTRACT FROM AUTHOR]

The boreal forest is predicted to experience the greatest warming of any forest biome during the next 50--100 years, but the effects of warming on vegetation phenology are not well known. The objectives of this study were to (1) examine the effects of whole ecosystem warming on bud burst and annual shoot growth of black spruce trees in northern Manitoba, Canada and (2) correlate bud burst to cumulative degree-days (CDD). The experimental design was a complete randomized block design that consisted of four replicated blocks. Each replicate block contained four treatments: soil warming only (heated outside, HO), soil and air warming (heated inside, HI), control outside (no chamber, no heating, CO), and inside a chamber maintained at ambient conditions (no soil or air warming, control inside, CI). Bud burst was measured during the first and second years of the experiment, starting in 2004, and annual shoot growth was measured for the first 3 years (2004--2006) of the study. On average, shoot bud burst occurred 11 and 9 days earlier in 2004 and 2005, respectively, for HI than for other treatments. However, mean CDD required for bud burst for HI was within the standard deviation of CO for both years. In year 1 of the treatments, shoot bud burst occurred earlier for HI than other treatments (CI, CO, HO), but final shoot length of HI trees was less than in CO trees. In the second year of warming, final shoot length was not different for HI than CO. By the third year of warming final shoot length was significantly greater for HI than all other treatments. Empirical results from this study suggest that soil and air warming causes an earlier bud burst for all years of observation and greater shoot lengths by the third season of warming. A longer growing season and greater annual shoot growth should increase carbon uptake by boreal black spruce trees in a warmer climate. [ABSTRACT FROM AUTHOR]

Changes in fire regimes are driving the carbon balance of much of the North American boreal forest, but few studies have examined fire-driven changes in evapotranspiration (ET) at a regional scale. This study used a version of the Biome-BGC process model with dynamic and competing vegetation types, and explicit spatial representation of a large (106 km2) region, to simulate the effects of wildfire on ET and its components from 1948 to 2005 by comparing the fire dynamics of the 1948–1967 period with those of 1968–2005. Simulated ET averaged, over the entire temporal and spatial modeling domain, 323 mm yr−1; simulation results indicated that changes in fire in recent decades decreased regional ET by 1.4% over the entire simulation, and by 3.9% in the last 10 years (1996–2005). Conifers dominated the transpiration ( EC) flux (120 mm yr−1) but decreased by 18% relative to deciduous broadleaf trees in the last part of the 20th century, when increased fire resulted in increased soil evaporation, lower canopy evaporation, lower EC, and a younger and more deciduous forest. Well- and poorly drained areas had similar rates of evaporation from the canopy and soil, but EC was twice as high in the well-drained areas. Mosses comprised a significant part of the evaporative flux to the atmosphere (22 mm yr−1). Modeled annual ET was correlated with net primary production, but not with temperature or precipitation; ET and its components were consistent with previous field and modeling studies. Wildfire is driving significant changes in hydrological processes by affecting mean stand age, forest species, and energy balance. These changes, particularly in poorly drained areas, may control the future carbon balance of the boreal forest. [ABSTRACT FROM AUTHOR]

The influence of climatic and local nonclimatic factors on the fire regime of the eastern Canadian boreal forest over the last 8000 years is investigated by examining charred particles preserved in four lacustrine deposits. Herein, we compare the distribution of fire-free intervals (FFIs) and the synchronicity of fire events among sites, using Ripley's K-function to determine the extent of the role of local-scale vs. large-scale processes with respect to fire control. Between 8000 and 5800 cal.bp (calibrated years before present) the climatic and ecological conditions were less conducive to fire events than after this date. After 5800 cal.bp, the number of fires per 1000 years (fire frequency) progressively increased, reaching a maximum ca. 3400 cal.bp. There was a sharp decrease in fire frequency during the last 800 years. Between 8000 and 4000 cal.bp, comparable FFIs and synchronous fire episodes were determined for the study sites. During this period, the fire frequency was predominantly controlled by climate. After 4000 cal.bp, two sites displayed independent fire histories (different FFI distributions or asynchronous fire events), underlining the important influence of local factors, including short-term fuel wetness, characteristics of the watershed and landscape connectivity, in determining fire occurrence. We conclude that climatic changes occurred during the last 4000 years that induced a rise in the water table; this may explain the high spatial heterogeneity in fire history. Current and projected global climatic changes may cause similar spatial variability in fire frequency. [ABSTRACT FROM AUTHOR]

We used measurements of CO2 exchange by six Canadian boreal forest stands to determine how sites of contrasting age respond to interannual temperature and precipitation variation. The stands ranged in age (time since last burn) from 4 to 155 years. The study included years that were anomalously cool and moist, warm and dry, cool and dry, and warm and moist. Warmer than average springs accelerated the onset of older stand (> 20 years) photosynthesis by as much as 37 days and younger stand (< 20 years) photosynthesis by as much as 25 days. The warm–dry year resulted in a marked reduction of mid-summer CO2 uptake by the younger, but not older, stands. The mid-summer decline in young stand photosynthesis reflected the combination of warmth and drought; similar declines were not observed during the cool–dry or warm–moist years. The annual carbon gain by the oldest stands was greatest during the warm–dry year as a result of the expanded growing season. The annual carbon gain by the youngest stands was consistent from year to year, largely as a result of offsets between increased spring photosynthesis and reduced mid-summer photosynthesis during the warm–dry year. Night-time CO2 efflux increased by 2–29% during the warm–moist year relative to the warm–dry year. This increase was not systematically related to age. We conclude that the major source of interannual CO2 exchange variation at the landscape scale is the ability of older, evergreen canopies to respond to warm springs by advancing the onset of photosynthesis. Drought-related reductions in photosynthesis, moisture-driven respiratory losses, and the effects of spring warmth on young-stand photosynthesis are of secondary importance. The advantage conferred on older, evergreen stands during warmer years carries implications for the possible effects of climate change on boreal forest succession. Warmer temperatures may increase the competitive advantage of late successional species relative to early successional species, and also the incidence of fire, thereby accelerating plant succession and shortening the fire-return interval. [ABSTRACT FROM AUTHOR]

We explored the effect of high-growth temperatures on a dominant North American boreal tree, black spruce [ Picea mariana (Mill.) B.S.P.]. In 2004 and 2005, we grew black spruce at either 22 °C/16 °C day/night temperatures [low temperature (LT)] or 30°/24 °C [high temperature (HT)] and determined how temperature affected growth, leaf morphology, photosynthesis, respiration and thermotolerance. HT spruce were 20% shorter, 58% lighter, and had a 58% lower root : shoot ratio than LT trees. Mortality was negligible in the LT treatment, but up to 14% of HT seedlings died by the end of the growing season. HT seedlings had a higher photosynthetic temperature optimum, but net photosynthesis at growth temperatures was 19–35% lower in HT than LT trees. HT seedlings had both a lower apparent maximum ribulose-1,5-bisphosphate carboxylation capacity ( Vcmax) and a lower apparent maximum electron transport rate ( Jmax) than LT trees, indicating reduced allocation to photosynthetic components. Consistently, HT needles had 26% lower leaf nitrogen content than LT needles. At each measurement temperature, HT seedlings had 20–25% lower respiration rates than LT trees; however, this did not compensate for reduced photosynthetic rates at growth temperature, leading to a greater ratio of dark respiration to net carbon dioxide assimilation rate in HT trees. HT needles had 16% lower concentrations of soluble sugars than LT needles, but similar starch content. Growth at high temperatures increased the thermotolerance of black spruce. While trees that develop at high temperatures have enhanced tolerance for brief, extreme heat events, the reduction in root allocation indicates that seedlings will be more susceptible to episodic soil drying and less competitive for belowground resources in future climates of the boreal region. [ABSTRACT FROM AUTHOR]

Projected temperature increases under global warming could benefit southern tree species by providing them the optimal growing temperature and could be detrimental to northern species by exposing them to the supra optimal growing temperatures. This benefit-detriment trade-off could increase the competitive advantage of southern species in the northern species range and cause the increase or even dominance of southern species in the northern domain. However, the optimum temperature for photosynthesis of C3 plants may increase due to CO2 enrichment. An increase in the optimum temperature could greatly reduce the benefit-detriment effect. In this study, we coupled a forest ecosystem process model (PnET-II) and a forest GAP model (LINKAGES) with a spatially dynamic forest landscape model (LANDIS-II) to study how an optimum temperature increase could affect forest landscape response due to global warming. We simulated 360 years of forest landscape change in the Boundary Water Canoe Area (BWCA) in northern Minnesota, which is transitional between boreal and temperate forest. Our results showed that, under the control scenario of continuing the historic 1984–1993 mean climate (mainly temperature, precipitation and CO2), the BWCA will become a spruce-fir dominated boreal forest. However, under the scenario of predicted climatic change [the 2000–2099 climates are predicted by Canadian Climate Center (CCC), followed by 200 years of continuing the predicted 2090–2099 mean climate], the BWCA will become a pine-dominated mixed forest. If the optimum temperature increases gradually with [CO2] (the increase in optimum temperature is assumed to change gradually from 0 °C in year 2000 to 5 °C in year 2099 when [CO2] reaches 711 ppm and stabilizes at 5 °C after year 2099), the BWCA would remain a fir-dominated boreal forest in areas with relatively high water-holding capacity, but not in areas with relatively low water-holding capacity. Our results suggest that the [CO2] induced increases in optimum temperature could substantially reduce forest landscape change caused by global warming. However, not all tree species would be able to successfully adapt to future warming as predicted by CCC, regardless of optimum temperature acclimations. [ABSTRACT FROM AUTHOR]

We present a decadal (1994–2004) record of carbon dioxide flux in a 160-year-old black spruce forest/veneer bog complex in central Manitoba, Canada. The ecosystem shifted from a source (+41 g C m−2, 1995) to a sink (−21 g C m−2, 2004) of CO2 over the decade, with an average net carbon balance near zero. Annual mean temperatures increased 1–2° during the period, consistent with the decadal trend across the North American boreal biome. We found that ecosystem carbon exchange responded strongly to air temperature, moisture status, potential evapotranspiration, and summertime solar radiation. The seasonal cycle of ecosystem respiration significantly lagged that of photosynthesis, limited by the rate of soil thaw and the slow drainage of the soil column. Factors acting over long time scales, especially water table depth, strongly influenced the carbon budget on annual time scales. Net uptake was enhanced and respiration inhibited by multiple years of rainfall in excess of evaporative demand. Contrary to expectations, we observed no correlation between longer growing seasons and net uptake, possibly because of offsetting increases in ecosystem respiration. The results indicate that the interactions between soil thaw and water table depth provide critical controls on carbon exchange in boreal forests underlain by peat, on seasonal to decadal time scales, and these factors must be simulated in terrestrial biosphere models to predict response of these regions to future climate. [ABSTRACT FROM AUTHOR]

Mycorrhizal fungi can contribute to soil carbon sequestration by immobilizing carbon in living fungal tissues and by producing recalcitrant compounds that remain in the soil following fungal senescence. We hypothesized that nitrogen (N) fertilization would decrease these carbon stocks, because plants should reduce investment of carbon in mycorrhizal fungi when N availability is high. We measured the abundance of two major groups of mycorrhizal fungi, arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) fungi, in the top 10 cm of soil in control and N-fertilized plots within three Alaskan boreal ecosystems that represented different recovery stages following severe fire. Pools of mycorrhizal carbon included root-associated AM and ECM structures; soil-associated AM hyphae; and glomalin, a glycoprotein produced by AM fungi. Total mycorrhizal carbon pools decreased by approximately 50 g C m−2 in the youngest site under N fertilization, and this reduction was driven mostly by glomalin. Total mycorrhizal carbon did not change significantly in the other sites. Root-associated AM structures were more abundant under N fertilization across all sites, and root-associated ECM structures increased marginally significantly. We found no significant N effects on AM hyphae. Carbon sequestered within living mycorrhizal structures (0. 051–0. 21 g m−2) was modest compared with that of glomalin (33–203 g m−2). We conclude that our hypothesis was only supported in relation to glomalin stocks within one of the three study sites. As N effects on glomalin were inconsistent among sites, an understanding of the mechanisms underlying this variation would improve our ability to predict ecosystem feedbacks to global change. [ABSTRACT FROM AUTHOR]

Stable isotopes of CO2 contain unique information on the biological and physical processes that exchange CO2 between terrestrial ecosystems and the atmosphere. In this study, we developed an integrated modeling system to simulate dynamics of stable carbon isotope of CO2, as well as moisture, energy, and momentum, between a boreal forest ecosystem and the atmosphere, as well as their transport/mixing processes through the convective boundary layer (CBL), using remotely sensed surface parameters to characterize the surface heterogeneity. It has the following characteristics: (i) it accounts for the influences of the CBL turbulent mixing and entrainment of the air aloft; (ii) it scales individual leaf-level photosynthetic discrimination up to the whole canopy (Δcanopy) through the separation of sunlit and shaded leaf groups; (iii) it has the capacity to examine the detailed interrelationships among plant water-use efficiency, isotope discrimination, and vapor pressure deficit; and (iv) it has the potential to investigate how an ecosystem discriminates against 13C at various time and spatial scales. The monthly mean isotopic signatures of ecosystem respiration (i.e. δ13CR) used for isotope flux calculation are retrieved from the nighttime flask data from the intensive campaigns (1998–2000) at 20 m level on Fraserdale tower, and the data from the growing season in 1999 are used for model validation. Both the simulated CO2 mixing ratio and δ13C of CO2 at the 20 m level agreed with the measurements well in different phases of the growing season. On a diurnal basis, the greatest photosynthetic discrimination at canopy level (i.e. Δcanopy) occurred early morning and late afternoon with a varying range of 10–26‰. The diurnal variability of Δcanopy was also associated with the phases of growing season and meteorological variables. The annual mean Δcanopy in 1999 was computed to be 19.58‰. The monthly averages of Δcanopy varied between 18.55‰ and 20.84‰ with a seasonal peak during the middle growing season. Because of the strong opposing influences of respired and photosynthetic fluxes on forest air (both CO2 and 13CO2) on both the diurnal and seasonal time scales, CO2 was consistently enriched with the heavier 13C isotope (less negative δ13C) from July to October and depleted during the remaining months, whereas on a diurnal basis, CO2 was enriched with the heavier 13C in the late afternoon and depleted in early morning. For the year 1999, the model results reveal that the boreal ecosystem in the vicinity of Fraserdale tower was a small sink with net uptake of 29.07 g 12C m−2 yr−1 and 0.34 g 13C m−2 yr−1. [ABSTRACT FROM AUTHOR]

Relations among nitrogen load, soil acidification and forest growth have been evaluated based on short-term (<15 years) experiments, or on surveys across gradients of N deposition that may also include variations in edaphic conditions and other pollutants, which confound the interpretation of effects of N per se. We report effects on trees and soils in a uniquely long-term (30 years) experiment with annual N loading on an un-polluted boreal forest. Ammonium nitrate was added to replicated ( N=3) 0.09 ha plots at two doses, N1 and N2, 34 and 68 kg N ha−1 yr−1, respectively. A third treatment, N3, 108 kg N ha−1 yr−1, was terminated after 20 years, allowing assessment of recovery during 10 years. Tree growth initially responded positively to all N treatments, but the longer term response was highly rate dependent with no gain in N3, a gain of 50 m3 ha−1 stemwood in N2 and a gain of 100 m3 ha−1 stemwood in excess of the control (N0) in N1. High N treatments caused losses of up to 70% of exchangeable base cations (Ca2+, Mg2+, K+) in the mineral soil, along with decreases in pH and increases in exchangeable Al3+. In contrast, the organic mor-layer (forest floor) in the N-treated plots had similar amounts per hectare of exchangeable base cations as in the N0 treatment. Magnesium was even higher in the mor of N-treated plots, providing evidence of up-lift by the trees from the mineral soil. Tree growth did not correlate with the soil Ca/Al ratio (a suggested predictor of effects of soil acidity on tree growth). A boron deficiency occurred on N-treated plots, but was corrected at an early stage. Extractable NH4+ and NO3−were high in mor and mineral soils of on-going N treatments, while NH4+ was elevated in the mor only in N3 plots. Ten years after termination of N addition in the N3 treatment, the pH had increased significantly in the mineral soil; there were also tendencies of higher soil base status and concentrations of base cations in the foliage. Our data suggest the recovery of soil chemical properties, notably pH, may be quicker after removal of the N-load than predicted. Our long-term experiment demonstrated the fundamental importance of the rate of N application relative to the total amount of N applied, in particular with regard to tree growth and C sequestration. Hence, experiments adding high doses of N over short periods do not mimic the long-term effects of N deposition at lower rates. [ABSTRACT FROM AUTHOR]

Mixedwood forests are an ecologically and economically important forest type in central Canada, but the ecology of these forests is not as well studied as that of single-species dominated stands in the boreal forest. Northern boreal mixedwood forests have only recently been harvested and the effects of harvesting on carbon content in these stands are unknown. We quantified the carbon content and aboveground net primary production (NPP) for four different-aged mixedwood boreal forest stands in northern Manitoba, Canada. The stands included 11-, 18-, and 30-year-old stands that originated from harvesting and a 65-year-old fire-originated stand that typifies the origin of all northern boreal mixed-wood forests that are coming under management. Trees included black spruce ( Picea mariana (Mill.) B.S.P.), jack pine ( Pinus banksiana Lamb.), balsam poplar ( Populus balsamifera L.), and quaking aspen ( Populus tremuloides Michx.). Overstory biomass was estimated using species-specific allometric models that generally explained greater than 95% of the observed variation in biomass. Carbon content of the overstory vegetation was greatest in the 65-year-old stand and was 74% larger than the 11-year-old stand and showed a positive relationship with stand age ( F1, 2=122.62, P=0.0081 R2=0.99). The slope of mineral soil carbon did not differ significantly among stands ( F1, 2=0.39, P=0.5956, R2=0.16). Coarse woody debris carbon content followed a U-shaped pattern among stands. Aboveground NPP differed by 24% between the youngest and oldest stand. Mean annual carbon accumulation and aboveground NPP rates of the mixedwood forests were on average two times greater than nearby relatively pure stands studied during the BOREAS (BOReal Ecosystem Atmospheric Study) project. The trends in the results, along with other field studies, suggest that harvesting does not significantly affect the total soil carbon content. The results of this study suggest that scientists should be cautious about extrapolating results from BOREAS stands to a broader region until more data on other forest types and regions are available. [ABSTRACT FROM AUTHOR]