Carbon and nitrogen cycling in North American boreal forests

A model of boreal forest dynamics was adapted to examine the factors controlling carbon and nitrogen cycling in the boreal forests of interior Alaska. Empirical relationships were used to simulate decomposition and nitrogen availability as a function of either substrate quality, the soil thermal regime, or their interactive effects. Test comparisons included black spruce forests growing on permafrost soils and black spruce, birch, and white spruce forests growing on permafrost-free soils. For each forest, simulated above-ground tree biomass, basal area, density, litterfall, moss biomass, and forest floor mass, turnover, thickness, and nitrogen concentration were compared to observed data. No one decay equation simulated forests entirely consistent with observed data, but over the range of upland forest types in interior Alaska, the equation that combined the effects of litter quality and the soil thermal regime simulated forests that were most consistent with observed data. For black spruce growing on permafrost soils, long-term simulated forest dynamics in the absence of fire resulted in unproductive forests with a thick forest floor and low nitrogen mineralization. Fires were an important means to interrupt this sequence and to restart forest succession.     

Changes in the structure and function of northern Alaskan ecosystems when considering variable leaf‐out times across groupings of species in a dynamic vegetation model

The phenology of arctic ecosystems is driven primarily by abiotic forces, with temperature acting as the main determinant of growing season onset and leaf budburst in the spring. However, while the plant species in arctic ecosystems require differing amounts of accumulated heat for leaf‐out, dynamic vegetation models simulated over regional to global scales typically assume some average leaf‐out for all of the species within an ecosystem. Here, we make use of air temperature records and observations of spring leaf phenology collected across dominant groupings of species (dwarf birch shrubs, willow shrubs, other deciduous shrubs, grasses, sedges, and forbs) in arctic and boreal ecosystems in Alaska. We then parameterize a dynamic vegetation model based on these data for four types of tundra ecosystems (heath tundra, shrub tundra, wet sedge tundra, and tussock tundra), as well as ecotonal boreal white spruce forest, and perform model simulations for the years 1970–2100. Over the course of the model simulations, we found changes in ecosystem composition under this new phenology algorithm compared with simulations with the previous phenology algorithm. These changes were the result of the differential timing of leaf‐out, as well as the ability for the groupings of species to compete for nitrogen and light availability. Regionally, there were differences in the trends of the carbon pools and fluxes between the new phenology algorithm and the previous phenology algorithm, although these differences depended on the future climate scenario. These findings indicate the importance of leaf phenology data collection by species and across the various ecosystem types within the highly heterogeneous Arctic landscape, and that dynamic vegetation models should consider variation in leaf‐out by groupings of species within these ecosystems to make more accurate projections of future plant distributions and carbon cycling in Arctic regions.     

Carbon allocation in boreal black spruce forests across regions varying in soil temperature and precipitation

A common hypothesis for northern ecosystems is that low soil temperatures inhibit plant productivity. To address this hypothesis, we reviewed how separate components of ecosystem carbon (C) cycling varied along a soil temperature gradient for nine well-drained, relatively productive boreal black spruce ( Picea mariana Mill. [B.S.P.]) forests in Alaska, USA, and Saskatchewan and Manitoba, Canada. Annual soil temperature [expressed as soil summed degree days (SDD)] was positively correlated with aboveground net primary productivity (ANPP), while negatively correlated with total belowground carbon flux (TBCF). The partitioning of C to ANPP at the expense of root processes represented a nearly 1 : 1 tradeoff across the soil temperature gradient, which implied that the amount of C cycling through these black spruce ecosystems was relatively insensitive to variation in SDD. Moreover, the rate at which C accumulated in the ecosystem since the last stand replacing fire was unrelated to SDD, but SDD was positively correlated to the ratio of spruce-biomass : forest-floor-mass. Thus, plant partitioning of C and the distribution of ecosystem C were apparently affected by soil temperature, although across regions, precipitation co-varied with soil temperature. These two factors likely correlated with one another because of precipitation's influence on soil heat balance, suggesting that a soil temperature–precipitation interaction could be responsible for the shifts in C allocation. Nonetheless, our results highlight that for this boreal ecosystem, ANPP and TBCF can be negatively correlated. In tropical and temperate forests, TBCF and ANPP have been reported as positively correlated, and our results may reflect the unique interactions between soil temperature, forest floor accumulation, rooting depth, and nutrient availability that characterize the black spruce forest type.     

Fire severity mediates climate‐driven shifts in understorey community composition of black spruce stands of interior Alaska

Question: How do pre‐fire conditions (community composition and environmental characteristics) and climate‐driven disturbance characteristics (fire severity) affect post‐fire community composition in black spruce stands? Location: Northern boreal forest, interior Alaska. Methods: We compared plant community composition and environmental stand characteristics in 14 black spruce stands before and after multiple, naturally occurring wildfires. We used a combination of vegetation table sorting, univariate (ANOVA, paired t‐tests), and multivariate (detrended correspondence analysis) statistics to determine the impact of fire severity and site moisture on community composition, dominant species and growth forms. Results: Severe wildfires caused a 50% reduction in number of plant species in our study sites. The largest species loss, and therefore the greatest change in species composition, occurred in severely burned sites. This was due mostly to loss of non‐vascular species (mosses and lichens) and evergreen shrubs. New species recruited most abundantly to severely burned sites, contributing to high species turnover on these sites. As well as the strong effect of fire severity, pre‐fire and post‐fire mineral soil pH had an effect on post‐fire vegetation patterns, suggesting a legacy effect of site acidity. In contrast, pre‐fire site moisture, which was a strong determinant of pre‐fire community composition, showed no relationship with post‐fire community composition. Site moisture was altered by fire, due to changes in permafrost, and therefore post‐fire site moisture overrode pre‐fire site moisture as a strong correlate. Conclusions: In the rapidly warming climate of interior Alaska, changes in fire severity had more effect on post‐fire community composition than did environmental factors (moisture and pH) that govern landscape patterns of unburned vegetation. This suggests that climate change effects on future community composition of black spruce forests may be mediated more strongly by fire severity than by current landscape patterns. Hence, models that represent the effects of climate change on boreal forests could improve their accuracy by including dynamic responses to fire disturbance.     

Spatial and temporal variation in moss-associated dinitrogen fixation in coniferous- and deciduous-dominated Alaskan boreal forests

Dominant canopy tree species have strong effects on the composition and function of understory species, particularly bryophytes. In boreal forests, bryophytes and their associated microbes are a primary source of ecosystem nitrogen (N) inputs, and an important process regulating ecosystem productivity. We investigated how feather moss-associated N₂-fixation rates and contribution to N budgets vary in time and space among coniferous and broadleaf deciduous forests. We measured N₂-fixation rates using stable isotope (¹⁵N₂) labeling in two moss species (Pleurozium schreberi and Hylocomium splendens) in broadleaf deciduous (Alaska paper birch—Betula neoalaskana) and coniferous (black spruce—Picea mariana) stands near Fairbanks, interior Alaska, from 2013 to 2015. N₂-fixation rates showed substantial inter-annual variation among the 3 years. High N₂-fixation was more strongly associated with high precipitation than air temperature or light availability. Overall, contribution of N₂-fixation to N budgets was greater in spruce than in birch stands. Our results enhance the knowledge of the processes that drive N₂-fixation in boreal forests, which is important for predicting ecosystem consequences of changing forest composition.     

Are post-fire residual forest patches refugia for boreal bryophyte species? Implications for ecosystem based management and conservation

Residual patches of forest remaining after natural or anthropogenic disturbance may facilitate regeneration of fragmented forest. However, residual patch function remains unclear, especially after natural wildfire. We investigate the role of residual boreal forest patches as refugia for bryophytes and ask the question, do they house bryophyte communities similar to those encountered in undisturbed forests? Bryophytes were sampled in three habitat types in black spruce boreal forests illustrating a gradient of disturbance severity: undisturbed forests, residual patches and burned matrices. Temporal, disturbance severity, spatial and structural variables of habitats were also recorded. Bryophyte community composition differed among habitat types with residual patches characterized by higher species richness, the loss of forest specialists and the addition of disturbance-prone species. The bryophyte community found in residual patches is at the interface between the communities of undisturbed forests and burned matrices. As residual patches did not conserve all species, particularly forest specialists, they were not refugia. However, we identify temporal, spatial and structural characteristics that can maintain bryophyte communities most similar to undisturbed forests and enhance residual patch “refugia potential”. Residual patches enhance bryophyte diversity of the landscape housing species that cannot survive in the burned matrix. As conclusion we discuss the use of retention patches in harvested stands, together with the preservation of undisturbed stands that house singular bryophyte communities and especially sensitive forest specialists.     

Changes in fire regime break the legacy lock on successional trajectories in Alaskan boreal forest

Predicting plant community responses to changing environmental conditions is a key element of forecasting and mitigating the effects of global change. Disturbance can play an important role in these dynamics, by initiating cycles of secondary succession and generating opportunities for communities of long‐lived organisms to reorganize in alternative configurations. This study used landscape‐scale variations in environmental conditions, stand structure, and disturbance from an extreme fire year in Alaska to examine how these factors affected successional trajectories in boreal forests dominated by black spruce. Because fire intervals in interior Alaska are typically too short to allow relay succession, the initial cohorts of seedlings that recruit after fire largely determine future canopy composition. Consequently, in a dynamically stable landscape, postfire tree seedling composition should resemble that of the prefire forest stands, with little net change in tree composition after fire. Seedling recruitment data from 90 burned stands indicated that postfire establishment of black spruce was strongly linked to environmental conditions and was highest at sites that were moist and had high densities of prefire spruce. Although deciduous broadleaf trees were absent from most prefire stands, deciduous trees recruited from seed at many sites and were most abundant at sites where the fires burned severely, consuming much of the surface organic layer. Comparison of pre‐ and postfire tree composition in the burned stands indicated that the expected trajectory of black spruce self‐replacement was typical only at moist sites that burned with low fire severity. At severely burned sites, deciduous trees dominated the postfire tree seedling community, suggesting these sites will follow alternative, deciduous‐dominated trajectories of succession. Increases in the severity of boreal fires with climate warming may catalyze shifts to an increasingly deciduous‐dominated landscape, substantially altering landscape dynamics and ecosystem services in this part of the boreal forest.     

Susceptibility of burned black spruce (Picea mariana) forests to non-native plant invasions in interior Alaska

As climate rapidly warms at high-latitudes, the boreal forest faces the simultaneous threats of increasing invasive plant abundances and increasing area burned by wildfire. Highly flammable and widespread black spruce (Picea mariana) forest represents a boreal habitat that may be increasingly susceptible to non-native plant invasion. This study assess the role of burn severity, site moisture and time elapsed since burning in determining the invisibility of black spruce forests. We conducted field surveys for presence of non-native plants at 99 burned black spruce forest sites burned in 2004 in three regions of interior Alaska that spanned a gradient of burn severities and site moisture levels, and a chronosequence of sites in a single region that had burned in 1987, 1994, and 1999. We also conducted a greenhouse experiment where we grew invasive plants in vegetation and soil cores taken from a subset of these sites. In both our field survey and the greenhouse experiment, regional differences in soils and vegetation between burn complexes outweighed local burn severity or site moisture in determining the invasibility of burned black spruce sites. In the greenhouse experiments using cores from the 2004 burns, we found that the invasive focal species grew better in cores with soil and vegetation properties characteristic of low severity burns. Invasive plant growth in the greenhouse was greater in cores from the chronosequence burns with higher soil water holding capacity or lower native vascular biomass. We concluded that there are differences in susceptibility to non-native plant invasions between different regions of boreal Alaska based on native species regeneration. Re-establishment of native ground cover vegetation, including rapidly colonizing bryophytes, appear to offer burned areas a level of resistance to invasive plant establishment.     

Biodiversity conservation in old-growth boreal forest: black spruce and balsam fir snags harbour distinct assemblages of saproxylic beetles

Biodiversity conservation of forest ecosystems strongly relies on effective dead wood management. However, the responses of saproxylic communities to variations in dead wood characteristics remains poorly documented, a lack of knowledge that may impede the development of efficient management strategies. We established the relationship between saproxylic beetles—at the species and community levels—and attributes of black spruce and balsam fir in old-growth boreal forests. The relationship was first evaluated for individual snag bole segments, and then for forest stands. A total of 168 bole sections were collected in summer 2006 along a compositional gradient ranging from black spruce-dominated stands to balsam fir-dominated ones, in a boreal forest dominated by >90-year-old stands. A total of 16,804 beetles belonging to 47 species emerged from bole segments, with 21% of the species being found exclusively in black spruce snags and 36% exclusively in balsam fir snags. Black spruce and balsam fir snags thus contributed differently to forest biodiversity by being inhabited by different saproxylic communities. Wood density was an important attribute in the host-use patterns for several species of saproxylic beetles, but no relationship was found between snag availability within stands and abundance of beetles strongly linked to either black spruce or balsam fir. Our study outlines the relative contribution of tree compositional diversity to saproxylic species, while highlighting the contribution of black spruce and balsam fir to animal diversity in old-growth boreal forests.     

Changes in Soil Fungal Communities, Extracellular Enzyme Activities, and Litter Decomposition Across a Fire Chronosequence in Alaskan Boreal Forests

Wildfires are a pervasive disturbance in boreal forests, and the frequency and intensity of boreal wildfires is expected to increase with climate warming. Boreal forests store a large fraction of global soil organic carbon (C), but relatively few studies have documented how wildfires affect soil microbial communities and soil C dynamics. We used a fire chronosequence in upland boreal forests of interior Alaska with sites that were 1, 7, 12, 24, 55, ~90, and ~100 years post-fire to examine the short- and long-term responses of fungal community composition, fungal abundance, extracellular enzyme activity, and litter decomposition to wildfires. We hypothesized that post-fire changes in fungal abundance and community composition would constrain decomposition following fires. We found that wildfires altered the composition of soil fungal communities. The relative abundance of ascomycetes significantly increased following fire whereas basidiomycetes decreased. Post-fire decreases in basidiomycete fungi were likely attributable to declines in ectomycorrhizal fungi. Fungal hyphal lengths in the organic horizon significantly declined in response to wildfire, and they required at least 24 years to return to pre-fire levels. Post-fire reductions in fungal hyphal length were associated with decreased activities of hydrolytic extracellular enzymes. In support of our hypothesis, the decomposition rate of aspen and black spruce litter significantly increased as forests recovered from fire. Our results indicate that post-fire reductions in soil fungal abundance and activity likely inhibit litter decomposition following boreal wildfires. Slower rates of litter decay may lead to decreased heterotrophic respiration from soil following fires and contribute to a negative feedback to climate warming.     

Predicting Ecosystem Resilience to Fire from Tree Ring Analysis in Black Spruce Forests

Climate change has increased the occurrence, severity, and impact of disturbances on forested ecosystems worldwide, resulting in a need to identify factors that contribute to an ecosystem’s resilience or capacity to recover from disturbance. Forest resilience to disturbance may decline with climate change if mature trees are able to persist under stressful environmental conditions that do not permit successful recruitment and survival after a disturbance. In this study, we used the change in proportional representation of black spruce pre- to post-fire as a surrogate for resilience. We explored links between patterns of resilience and tree ring signals of drought stress across topographic moisture gradients within the boreal forest. We sampled 72 recently (2004) burned stands of black spruce in interior Alaska (USA); the relative dominance of black spruce after fire ranged from almost no change (high resilience) to a 90% decrease (low resilience). Variance partitioning analysis indicated that resilience was related to site environmental characteristics and climate–growth responses, with no unique contribution of pre-fire stand composition. The largest shifts in post-fire species composition occurred in sites that experienced the compounding effects of pre-fire drought stress and shallow post-fire organic layer thickness. These sites were generally located at warmer and drier landscape positions, suggesting they are less resilient to disturbance than sites in cool and moist locations. Climate–growth responses can provide an estimate of stand environmental stress to climate change and as such are a valuable tool for predicting landscape variations in forest ecosystem resilience.     

A Canopy Shift in Interior Alaskan Boreal Forests: Consequences for Above- and Belowground Carbon and Nitrogen Pools during Post-fire Succession

Global change models predict that high-latitude boreal forests will become increasingly susceptible to fire activity as climate warms, possibly causing a positive feedback to warming through fire-driven emissions of CO₂ into the atmosphere. However, fire-climate feedbacks depend on forest regrowth and carbon (C) accumulation over the post-fire successional interval, which is influenced by nitrogen (N) availability. To improve our understanding of post-fire C and N accumulation patterns in boreal forests, we evaluated above- and belowground C and N pools within 70 stands throughout interior Alaska, a region predicted to undergo a shift in canopy dominance as fire severity increases. Stands represented gradients in age and successional trajectory, from black spruce (Picea mariana) self-replacement to species replacement by deciduous species of trembling aspen (Populus tremuloides) and Alaska paper birch (Betula neoalaskana). Stands undergoing deciduous trajectories stored proportionally more of their C and N in aboveground stemwood and had 5–7 times faster rates of aboveground net primary productivity of trees compared to stands undergoing a black spruce trajectory, which stored more of their C and N in the soil organic layer (SOL), a thick layer of mostly undecomposed mosses. Thus, as successional trajectories shift, total C and N pool sizes will remain relatively unchanged, but there will be a trade-off in pool location and a potential increase in C and N longevity due to decreased flammability and decomposition rates of deciduous stemwood. Despite often warmer, drier conditions in deciduous compared to black spruce stands, deciduous stemwood has a C:N around 10 times higher than the black spruce SOL and often remains standing for many years with reduced exposure to fungal decomposers. Thus, a fire-driven shift in successional trajectories could cause a negative feedback to climate warming because of increased pool longevity in deciduous trajectories.     

Are Mosses Required to Accurately Predict Upland Black Spruce Forest Soil Carbon in National-Scale Forest C Accounting Models?

The boreal forest plays a key role in the global carbon (C) cycle, and black spruce (Picea mariana (Mill.) BSP) forests are the dominant coniferous forest type in the Canadian boreal forest. National-scale forest C models currently do not account for the contribution of moss-derived organic matter that we hypothesize to be significant in the C budget of black spruce ecosystems. One such model, the Carbon Budget Model of the Canadian Forest Sector (CBM-CFS3), is designed to meet Canada’s forest-related greenhouse gas reporting requirements. In this study our goal was to determine if black spruce forest soil C stocks are significantly underestimated by the CBM-CFS3, and if so, to determine if estimates could be improved by adding moss-derived C. We conclude that in black spruce sites, organic layer C is significantly underestimated by CBM-CFS3 compared to sites with all other leading tree species analyzed. We compiled and used published moss net primary productivity rates for upland forest systems, with decomposition rates, in mass-balance calculations to estimate mean moss-derived C in black spruce forests for feather mosses at 64 Mg C ha^?1, and for sphagnum mosses at 103 Mg C ha^?1. These C pools are similar to the CBM-CFS3 mean underestimation of black spruce soil organic layers (63 Mg C ha^?1). We conclude that the contribution of mosses is sufficiently large that a moss C pool should be added to national-scale models including the CBM-CFS3, to reduce uncertainties in boreal forest C budget estimation. Feather and sphagnum mosses should be parameterized separately.     

Stable carbon isotope analysis reveals widespread drought stress in boreal black spruce forests

Unprecedented rates of climate warming over the past century have resulted in increased forest stress and mortality worldwide. Decreased tree growth in association with increasing temperatures is generally accepted as a signal of temperature‐induced drought stress. However, variations in tree growth alone do not reveal the physiological mechanisms behind recent changes in tree growth. Examining stable carbon isotope composition of tree rings in addition to tree growth can provide a secondary line of evidence for physiological drought stress. In this study, we examined patterns of black spruce growth and carbon isotopic composition in tree rings in response to climate warming and drying in the boreal forest of interior Alaska. We examined trees at three nested scales: landscape, toposequence, and a subsample of trees within the toposequence. At each scale, we studied the potential effects of differences in microclimate and moisture availability by sampling on northern and southern aspects. We found that black spruce radial growth responded negatively to monthly metrics of temperature at all examined scales, and we examined ?¹³C responses on a subsample of trees as representative of the wider region. The negative ?¹³C responses to temperature reveal that black spruce trees are experiencing moisture stress on both northern and southern aspects. Contrary to our expectations, ?¹³C from trees on the northern aspect exhibited the strongest drought signal. Our results highlight the prominence of drought stress in the boreal forest of interior Alaska. We conclude that if temperatures continue to warm, we can expect drought‐induced productivity declines across large regions of the boreal forest, even for trees located in cool and moist landscape positions.     

Indirect effects of black spruce (Picea mariana) cover on community structure and function in sheep laurel (Kalmia angustifolia) dominated heath of eastern Canada

Recent studies on phenotypic plasticity of plant traits indicate that within-species variation in litter quality may be a significant factor that feeds back on litter decomposition and nutrient cycling rates at the stand level. These findings may be especially significant for understanding biodiversity-stability relationships in species-poor ecosystems that have little functional redundancy among primary producers. We tested the null hypothesis that black spruce and Kalmia were functional equivalents with respect to their structuring roles of subordinate vegetation and their influence on site biogeochemistry. The purpose of the study was to determine the degree to which forest cover exerts top-down control on community structure and function of Kalmia-black spruce communities. This community type dominates much of the forest understory and unforested heathlands in Atlantic Canada. We intensively studied a representative stand of Kalmia heath in Terra Nova National Park in eastern Newfoundland. Thirty-two 0.5 m × 0.5 m sample plots were randomly distributed among five transects bisecting gradients in dominance of black spruce and Kalmia. Light levels, species composition, vascular plant cover and soil respiration rate were determined for each plot. Tissue samples of litter, mature and current year leaves of Kalmia were collected and analyzed for nutrient status. Herbaceous species richness and cover peaked at intermediate light levels. Kalmia foliar N concentration and above-ground biomass increased with increasing shade. Soil respiration rates were strongly related to the light gradient and increased with increasing quality of Kalmia litter inputs. Our data indicate that Kalmias vigour and foliar nitrogen concentrations are greater under black spruce canopy as opposed to heath condition and that the shaded phenotype has relatively benign feedbacks on soil productivity compared to the open-habitat phenotype. In the absence of functional diversity at the species level in these species-poor habitats, phenotypic plasticity in Kalmia appears to be an important dimension of the biodiversity-stability relationship in these communities since our data suggest that this species has the potential either to inhibit or facilitate carbon cycling and the pathway is strongly linked to the presence or absence of overstory cover. The role of forest regeneration as an indirect control of forest soil processes such as carbon and nitrogen cycling in this ecosystem is discussed.     

Soil CO2 efflux in contrasting boreal deciduous and coniferous stands and its contribution to the ecosystem carbon balance

Similar nonsteady‐state automated chamber systems were used to measure and partition soil CO₂ efflux in contrasting deciduous (trembling aspen) and coniferous (black spruce and jack pine) stands located within 100 km of each other near the southern edge of the Boreal forest in Canada. The stands were exposed to similar climate forcing in 2003, including marked seasonal variations in soil water availability, which provided a unique opportunity to investigate the influence of climate and stand characteristics on soil CO₂ efflux and to quantify its contribution to the net ecosystem CO₂ exchange (NEE) as measured with the eddy‐covariance technique. Partitioning of soil CO₂ efflux between soil respiration (including forest‐floor vegetation) and forest‐floor photosynthesis showed that short‐ and long‐term temporal variations of soil CO₂ efflux were related to the influence of (1) soil temperature and water content on soil respiration and (2) below‐canopy light availability, plant water status and forest‐floor plant species composition on forest‐floor photosynthesis. Overall, the three stands were weak to moderate sinks for CO₂ in 2003 (NEE of ?103, ?80 and ?28 g C m^?2 yr^?1 for aspen, black spruce and jack pine, respectively). Forest‐floor respiration accounted for 86%, 73% and 75% of annual ecosystem respiration, in the three respective stands, while forest‐floor photosynthesis contributed to 11% and 14% of annual gross ecosystem photosynthesis in the black spruce and jack pine stands, respectively. The results emphasize the need to perform concomitant measurements of NEE and soil CO₂ efflux at longer time scales in different ecosystems in order to better understand the impacts of future interannual climate variability and vegetation dynamics associated with climate change on each component of the carbon balance.     

Transplantation of organic surface horizons of boreal soils into warmer regions alters microbiology but not the temperature sensitivity of decomposition

Changes in soil carbon, the largest terrestrial carbon pool, are critical for the global carbon cycle, atmospheric CO₂ levels and climate. Climate warming is predicted to be most pronounced in the northern regions and therefore the large soil carbon pool residing in boreal forests will be subject to larger global warming impact than soil carbon pools in the temperate or the tropical forest. A major uncertainty in current estimates of the terrestrial carbon balance is related to decomposition of soil organic matter (SOM). We hypothesized that when soils are exposed to warmer climate the structure of the ground vegetation will change much more rapidly than the dominant tree species. This change will alter the quality and amount of litter input to the soil and induce changes in microbial communities, thus possibly altering the temperature sensitivity of SOM decomposition. We transferred organic surface soil sections from the northern borders of the boreal forest zone to corresponding forest sites in the southern borders of the boreal forest zone and studied the effects of warmer climate after an adaptation period of 2 years. The results showed that initially ground vegetation and soil microbial community structure and community functions were different in northern and southern forest sites and that 2 years of exposure to warmer climate was long enough to cause changes in these ecological indicators. The rate of SOM decomposition was approximately equally sensitive to temperature irrespective of changes in vegetation or microbial communities in the studied forest sites. However, as temperature sensitivity of the decomposition increases with decreasing temperature regime, the proportional increase in the decomposition rate in northern latitudes could lead to significant carbon losses from the soils.     

Macrocharcoal-Based Chronosequences Reveal Shifting Dominance of Conifer Boreal Forests Under Changing Fire Regime

Balsam fir (Abies balsamea) and black spruce (Picea mariana) forests are the main conifer forest types in the North American boreal zone. The coexistence of the two species as well as their respective canopy dominance in distinct stands raises questions about the long-term evolution from one forest type to the other in relation to environmental factors including climate and stand disturbance. We tested the hypothesis that repetitive fire events promote the succession of balsam fir forest to black spruce forest and vice versa. Postfire chronosequences of one black spruce (BSP) and one balsam fir (BFI) sites were reconstructed based on the botanical composition and ¹⁴C-dated soil macrocharcoals. The results support the hypothesis of a successional dynamics. The BSP site has been affected by fires for the last 7600 years, whereas the BFI site, after having been impacted by several fires during the first half of the Holocene, evolved in a fire-free environment for the last 4400 years. Periods of fire activity facilitated the dominance of black spruce forests. The cessation of fires around 4400 cal. years BP on BFI site marks the beginning of the transition from black spruce to balsam fir stands. This succession is a long process, due to the ability of black spruce to regenerate by layering in the absence of fire. The resulting balsam fir stands are ancient and precarious ecosystems, since fire generally leads to the return of black spruce. The increase in balsam fir to the detriment of black spruce in boreal forests is a response to a decrease in fire frequency.     

The Origin and Dynamics of Subalpine White Spruce and Balsam Fir Stands in Boreal Eastern North America

Associations among the few tree species in the North American boreal landscape are the result of complex interactions between climate, biota, and historical disturbances during the Holocene. The closed-crown boreal forest of eastern North America is subdivided into two ecological regions having distinct tree species associations; the balsam fir zone and the black spruce zone, south and north of 49°N, respectively. Subalpine old-growth stands dominated by trees species typical of the balsam fir forest flora (either balsam fir or white spruce) are found on high plateaus, some of which are isolated within the black spruce zone. Here we identified the ecological processes responsible for the distinct forest associations in the subalpine belt across the eastern boreal landscape. Extensive radiocarbon dating, species composition, and size structure analyses indicated contrasted origin and dynamics of the subalpine forests between the two ecological regions. In the black spruce zone, the subalpine belt is a mosaic of post-fire white spruce or balsam fir stands coexisting at similar elevation on the high plateaus. With increasing time without wildfire, the subalpine forests become structurally similar to the balsam fir forest of the fir zone. These results concur with the hypothesis that the subalpine forests of this area are protected remnants of an historical northern expansion of the fir zone. Its replacement by the fire-prone black spruce forest flora was caused by recurrent fires. In the subalpine belt of the fir zone, no fire was recorded for several millennia. Harsh climate at high altitude is the primary factor explaining white spruce dominance over balsam fir forming a distinct subalpine white spruce belt above the balsam fir dominated forest.