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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]

Boreal forests store substantial amounts of carbon in vegetation and soil pools. The magnitude of these pools is related to fire regime attributes. Climate change is expected to alter boreal fire regimes, leading to changes in the amount of stored carbon. Quantifying these changes is of importance to understanding and managing global carbon budgets. We investigate how fire return interval (FRI) interacts with seasonal variation in fire intensity and severity to affect carbon stocks and fluxes in the boreal black spruce forests of Québec, Canada. A size-class structured population model of stand dynamics was coupled with an established model of boreal carbon dynamics and linked to a simplified representation of fire regime that simulates the occurrence of fires and their direct effects on canopy tree mortality, surface fuels combustion and regeneration. We simulated carbon stocks and fluxes under seven levels of FRI and two fire seasons: spring and summer. We tested for an effect of these fire regime parameters on equilibrium mean C stocks. All dead organic matter and biomass carbon stocks were sensitive to FRI between 60 and 300 years. These C stocks were lower for summer fires that occurred under shorter FRIs. Net primary production was highest at FRIs between 150 and 300 years. Total C stocks were highest for FRIs from 150 to 700 years, varying little over this range. There was a small but significant difference of C pool sizes between stands with even and uneven tree-diameter distributions. This difference was greatest for FRIs of 150 years or less. Reductions in equilibrium C storage are forecasted for nearly 27% of the study region under expected end-of-century climates. [ABSTRACT FROM AUTHOR]

Commercial thinning enables forest managers to meet timber production objectives. Thinning reduces tree density to alleviate competition for resources and favour growth of selected tree species. However, in doing so, thinning can homogenize the composition of mixed-species forests and raise biodiversity issues. There is increasing evidence that species richness can lead to higher productivity through a complementarity effect. Hence, thinning that would maintain species diversity of mixed-species forests could enhance stand productivity and help forest managers to reconcile timber production objectives and biodiversity issues. The objective of this study was to compare post-thinning stand production, experimentally over 10 years, in mixed and monospecific stands of black spruce (Picea mariana [Mill.] B.S.P.) and jack pine (Pinus banksiana Lamb.). The post-thinning stand production curve of the mixed stand converged toward that of the unthinned mixed stand while the production curves of the thinned and unthinned monospecific stands remained parallel. The convergent productivity of the mixed stand could be explained by a positive interaction between effects of thinning and niche complementarity. We propose that thinning that maintains species diversity of mixed stands could help forest managers who are implementing ecosystem management to reconcile timber production objectives with biodiversity issues. [ABSTRACT FROM AUTHOR]

The growth of high-latitude temperature-limited boreal forest ecosystems is projected to become more constrained by soil water availability with continued warming. The purpose of this study was to document ongoing shifts in tree growth sensitivity to the evolving local climate in unmanaged black spruce (Picea mariana (Miller) B.S.P.) forests of eastern boreal North America (49°N-52°N, 58°W-82°W) using a comparative study of field and modeled data. We investigated growth relationships to climate (gridded monthly data) from observed (50 site tree-ring width chronologies) and simulated growth data (stand-level forest growth model) over 1908-2013. No clear strengthening of moisture control over tree growth in recent decades was detected. Despite climate warming, photosynthesis (main driver of the forest growth model) and xylem production (main driver of radial growth) have remained temperature-limited. Analyses revealed, however, a weakening of the influence of growing season temperature on growth during the mid- to late twentieth century in the observed data, particularly in high-latitude (> 51.5°N) mountainous sites. This shift was absent from simulated data, which resulted in clear model-data desynchronization. Thorough investigations revealed that desynchronization was mostly linked to the quality of climate data, with precipitation data being of particular concern. The scarce network of weather stations over eastern boreal North America (> 51.5°N) affects the accuracy of estimated local climate variability and critically limits our ability to detect climate change effects on high-latitude ecosystems, especially at high altitudinal sites. Climate estimates from remote sensing could help address some of these issues in the future. [ABSTRACT FROM AUTHOR]

Under a warming climate, the boreal forest could become one of the largest terrestrial net CO 2 sources, as increasing disturbances and soil organic matter decomposition rates (heterotrophic respiration, Rh) could offset net primary production. Since soil represents the boreal forest's largest C pool, it is critical of correctly predicting future changes in Rh, as well as its sensitivity to temperature (Q 10 of Rh). We simulated a soil warming by transplanting soil cores from boreal balsam fir ( Abies balsamea , BF) and black spruce ( Picea mariana, BS) stands to a more southern Eastern hemlock stand ( Tsuga canadensis , EH). We measured Rh and soil properties over 3 years, from June to October. Over three snow-free seasons, soil temperature (first 10 cm, including the FH organic layers) and Rh increased for BF (+3.2 °C, +60% of Rh) and BS cores (+2.3 °C, +27% of Rh). Microbial C concentration decreased by 54–73% in the FH layers of warmed and control cores relative to initial values, despite unchanged chemically labile C, probably due to excised roots and mycorrhizal hyphae. This suggests a possible underestimation of Rh during the experiment. In BF soils only, the increase in Rh was accompanied by an increase in its sensitivity to temperature. Under a +5 °C soil warming, mean predicted Rh of BF soils would increase by 83% rather than by 56%. Relative to BS soils, such increase in sensitivity could be partly due to a higher fraction of chemically labile C (+52%) in the FH layers and a higher mean warming effect. It suggests that for BF forest soils, predicting decomposition rates for a warmer climate based on current temperature sensitivities could be inadequate. However, longer-term studies are needed to see if this increase in Q 10 of Rh for BF soils would be maintained for longer periods. [ABSTRACT FROM AUTHOR]

Old-growth (>100 years old) boreal forests are recognized as having accumulated large inventories of terrestrial carbon, but the magnitude of annual net exchange is not well established. Eddy covariance measurements of carbon dioxide exchange were made at a flux tower at the Northern Old Black Spruce forest in Manitoba, Canada, from 1994 to 2008. We evaluated the uncertainty in estimates of annual net ecosystem production by comparing four methods of filling missing measurement periods. Three of the methods were previously developed by groups at Harvard University, the Fluxnet Canada Research Network, and the Max Planck Institute of Biogeochemistry; and the fourth method used the mean of measurements from the dataset selected for the same time and day. Two methods showed carbon losses near the beginning of the period but all four methods estimated carbon gains in the later years. Individual years and methods ranged ±90 g C m −2 y −1 , demonstrating a wide range of estimates if data had been available from only a single year and a single method. When averaged over multiple years, the mean and standard error among methods gave a net forest carbon sink of 29 ± 10 g C m −2 y −1 , ranging from 4 to 48 g C m −2 y −1 . Only the upper (Mean Data and Max Planck Institute of Biogeochemistry methods) and lower (Harvard method) estimates were significantly different, with the Fluxnet Canada Research Network method being the same as all other methods. There was no difference among methods for the January to August period, or for daytime periods. All methods were statistically different from a carbon-neutral forest suggesting that the Northern Old Black Spruce forest was likely a small carbon sink. Methods that estimated high ecosystem respiration also estimated high gross ecosystem production, and the small net differences could not be easily attributed to differences in flux partitioning. For forests that are close to carbon neutral, the use of several gap-filling methods: increases confidence in conclusions regarding the ecosystem carbon balance. [ABSTRACT FROM AUTHOR]

An open black spruce forest, the most common ecosystem in interior Alaska, is characterized by patchy canopy gaps where the forest understory is exposed. This study measured CO 2 , sensible heat, and latent heat fluxes with eddy covariance (EC) in one of those large canopy gaps, and estimated understory fluxes in a black spruce forest in 2011–2014. Then understory fluxes and ecosystem fluxes were compared. The understory fluxes during the snow-free seasons were determined by two approaches. The first approach determined understory fluxes as the fluxes from the canopy gap, assuming that fluxes under the canopy crown also had the same magnitude as the canopy gap fluxes. The second approach determined the understory fluxes by scaling canopy gap fluxes with a canopy gap fraction, assuming that only canopy gaps, which mostly constitutes the forest floor, contribute to fluxes. The true understory fluxes would be in between these two estimates. Overall, the understory accounted for 53% (39–66%), 61% (45–77%), 63% (45–80%), 73% (56–90%), and 79% (59–98%) of the total net ecosystem productivity (NEP), gross primary productivity (GPP), ecosystem respiration (RE), sensible heat flux ( H ), and latent heat flux (LE), respectively. The ratio of understory NEP (NEP U ) to the ecosystem NEP (NEP E ) and similarly calculated LE U /LE E during the daytime increased with vapor pressure deficit (VPD) at low VPD conditions (∼2000 Pa) at half-hourly temporal scale. At high VPD conditions, however, NEP U /NEP E decreased with VPD, whereas LE U /LE E was maintained at the high level even at high VPD conditions. Despite large ranges of the estimates for the understory contributions, we conclude that the understory plays an important role in the carbon and energy balances of the black spruce ecosystem, and their contribution highly depends on the level of VPD. [ABSTRACT FROM AUTHOR]

In the boreal forest of Alaska, increased fire severity associated with climate change is expanding deciduous forest cover in areas previously dominated by black spruce ( Picea mariana). Needle-leaf conifer and broad-leaf deciduous species are commonly associated with differences in tree growth, carbon (C) and nutrient cycling, and C accumulation in soils. Although this suggests that changes in tree species composition in Alaska could impact C and nutrient pools and fluxes, few studies have measured these linkages. We quantified C, nitrogen, phosphorus, and base cation pools and fluxes in three stands of black spruce and Alaska paper birch ( Betula neoalaskana) that established following a single fire event in 1958. Paper birch consistently displayed characteristics of more rapid C and nutrient cycling, including greater aboveground net primary productivity, higher live foliage and litter nutrient concentrations, and larger ammonium and nitrate pools in the soil organic layer (SOL). Ecosystem C stocks (aboveground + SOL + 0-10 cm mineral soil) were similar for the two species; however, in black spruce, 78% of measured C was found in soil pools, primarily in the SOL, whereas aboveground biomass dominated ecosystem C pools in birch forest. Radiocarbon analysis indicated that approximately one-quarter of the black spruce SOL C accumulated prior to the 1958 fire, whereas no pre-fire C was observed in birch soils. Our findings suggest that tree species exert a strong influence over C and nutrient cycling in boreal forest and forest compositional shifts may have long-term implications for ecosystem C and nutrient dynamics. [ABSTRACT FROM AUTHOR]

Disturbance plays an important role in the distributional range of species by affecting their colonization potential and persistence. Short disturbance intervals have been linked to reduced seedbank sizes of some species, but the effects of long intervals are largely unknown. To explore the potential existence of seedbank sizes that may also be limited by long disturbance intervals, we studied an area in boreal eastern North America where time since fire (TSF) coincides with an increase in environmental stress (accumulating organic matter measured as depth of the soil organic layer (SOL)). Along a chronosequence dating back about 710 years, we counted the number of seeds cone of black spruce ( Picea mariana) and then estimated the number of seeds tree and site by upscaling. Younger sites [TSF 60-150 years] with mature first regeneration trees had average-sized seedbanks for black spruce [12.0-17.9 (10) seeds ha], whereas subsequent pulse trees that established in SOL depths greater than 35 cm showed highly reduced seed numbers. Sites with second- to fourth-regeneration pulse individuals [TSF c. 350-710 years] had exceptionally small seedbanks of 0.90 (10) and 0.46 (10) seeds ha, respectively. Radial tree growth rate showed a similarly negative response to SOL depth and could potentially be used as an indicator of seed output in plant species. Because the decline in seedbank size was possibly caused by more general environmental stress factors such as reduced nutrient availability, we suggest exploring whether other examples of ecosystems exist where long time since disturbance may lead to reduced seedbank sizes. [ABSTRACT FROM AUTHOR]

Disturbance plays a key role in driving ecological responses by creating opportunities for new ecological communities to assemble and by directly influencing the outcomes of assembly. Legacy effects (such as seed banks) and environmental filters can both influence community assembly, but their effects are impossible to separate with observational data. Here, we used seeding experiments in sites covering a broad range of postdisturbance conditions to tease apart the effects of seed availability, environmental factors, and disturbance characteristics on early community assembly after fire. We added seed of four common boreal trees to experimental plots in 55 replicate sites in recently burned areas of black spruce forest in northwestern North America. Seed addition treatments increased the probability of occurrence for all species, indicating a widespread potential for seed limitation to affect patterns of recruitment after fire. Small-seeded species (aspen and birch) were most sensitive to environmental factors such as soil moisture and organic layer depth, suggesting a role for niche-based environmental filtering in community assembly. Fire characteristics related to severity and frequency were also important drivers of seedling regeneration, indicating the potential for disturbance to mediate environmental filters and legacy effects on seed availability. Because effects of seed availability are typically impossible to disentangle from environmental constraints on recruitment in observational studies, legacy effects contingent on vegetation history may be misinterpreted as being driven by strong environmental filters. Results from the seeding experiments suggest that vegetation legacies affecting seed availability play a pivotal role in shaping patterns of community assembly after fire in these low-diversity boreal forests. [ABSTRACT FROM AUTHOR]

Background and Aims: The carbon (C) balance of boreal terrestrial ecosystems is sensitive to increasing temperature, but the direction and thresholds of responses are uncertain. Annual C uptake in Picea and other evergreen boreal conifers is dependent on seasonal- and cohort-specific photosynthetic and respiratory temperature response functions, so this study examined the physiological significance of maintaining multiple foliar cohorts for Picea mariana trees within an ombrotrophic bog ecosystem in Minnesota, USA. Methods: Measurements were taken on multiple cohorts of needles for photosynthetic capacity, foliar respiration (Rd) and leaf biochemistry and morphology of mature trees from April to October over 4 years. The results were applied to a simple model of canopy photosynthesis in order to simulate annual C uptake by cohort age under ambient and elevated temperature scenarios. Key Results: Temperature responses of key photosynthetic parameters [i.e. light-saturated rate of CO2 assimilation (Asat), rate of Rubisco carboxylation (Vcmax) and electron transport rate (Jmax)] were dependent on season and generally less responsive in the developing current-year (Y0) needles compared with 1-year-old (Y1) or 2-year-old (Y2) foliage. Temperature optimums ranged from 18·7 to 23·7, 31·3 to 38·3 and 28·7 to 36·7°C for Asat, Vcmax and Jmax, respectively. Foliar cohorts differed in their morphology and photosynthetic capacity, which resulted in 64% of modelled annual stand C uptake from Y1&2 cohorts (LAI 0·67m²m-2) and just 36% from Y0 cohorts (LAI 0·52m²m-2). Under warmer climate change scenarios, the contribution of Y0 cohorts was even less; e.g. 31% of annual C uptake for a modelled 9°C rise in mean summer temperatures. Results suggest that net annual C uptake by P. mariana could increase under elevated temperature, and become more dependent on older foliar cohorts. Conclusions: Collectively, this study illustrates the physiological and ecological significance of different foliar cohorts, and indicates the need for seasonal- and cohort-specific model parameterization when estimating C uptake capacity of boreal forest ecosystems under ambient or future temperature scenarios. [ABSTRACT FROM AUTHOR]

Many northern forests are limited by nitrogen (N) availability, slight changes in which can have profound effects on ecosystem function and the activity of ectomycorrhizal (EcM) fungi. Increasing N and phosphorus (P) availability, an analog to accelerated soil organic matter decomposition in a warming climate, could decrease plant dependency on EcM fungi and increase plant productivity as a result of greater carbon use efficiency. However, the impact of altered N and P availability on the growth and activity of EcM fungi in boreal forests remains poorly understood despite recognition of their importance to host plant nutrition and soil carbon sequestration. To address such uncertainty we examined above and belowground ecosystem properties in a boreal black spruce forest following five years of factorial N and P additions. By combining detailed soil, fungal, and plant δ15N measurements with in situ metrics of fungal biomass, growth, and activity, we found both expected and unexpected patterns. Soil nitrate isotope values became 15N enriched in response to both N and P additions; fungal biomass was repressed by N yet both biomass and growth were stimulated by P; and, black spruce dependency on EcM derived N increased slightly when N and P were added alone yet significantly declined when added in combination. These findings contradict predictions that N fertilization would increase plant P demands and P fertilization would further exacerbate plant N demands. As a result, the prediction that EcM fungi predictably respond to plant N limitation was not supported. These findings highlight P as an under appreciated mediator of the activity of denitrifying bacteria, EcM fungi, and the dynamics of N cycles in boreal forests. Further, use of δ15N values from bulk soils, plants, and fungi to understand how EcM systems respond to changing nutrient availabilities will often require additional ecological information. [ABSTRACT FROM AUTHOR]

The article focuses on effects climatic changes, forest fires, and insect migrations are having on boreal forests in the north. It states more intense forest fires are burning away organic layers of moss and duff and allowing aspens and other broadleaf trees to establish themselves in areas once dominated by black spruce. It comments on outbreaks of insect pests such as the mountain pine beetle and the hemlock looper. It talks about declines in photosynthesis in North America's boreal forests.

Climate change is altering insect disturbance regimes via temperature-mediated phenological changes and trophic interactions among host trees, herbivorous insects, and their natural enemies in boreal forests. Range expansion and increase in outbreak severity of forest insects are occurring in Europe and North America. The degree to which northern forest ecosystems are resilient to novel disturbance regimes will have direct consequences for the provisioning of goods and services from these forests and for long-term forest management planning. Among major ecological disturbance agents in the boreal forests of North America is a tortricid moth, the eastern spruce budworm, which defoliates fir (Abies spp.) and spruce (Picea spp.). Northern expansion of this defoliator in eastern North America and climate-induced narrowing of the phenological mismatch between the insect and its secondary host, black spruce (Picea mariana), may permit greater defoliation and mortality in extensive northern black spruce forests. Although spruce budworm outbreak centers have appeared in the boreal black spruce zone historically, defoliation and mortality were minor. Potential increases in outbreak severity and tree mortality raise concerns about the future state of this northern ecosystem. Severe spruce budworm outbreaks could decrease stand productivity compared with their occurrence in more diverse, southern balsam fir forest landscapes that have coevolved with outbreaks. Furthermore, depending on the proportion of balsam fir and deciduous species present and fire recurrence, changes in regeneration patterns and in nutrient cycling could alter ecosystem dynamics and replace black spruce by more productive mixed-wood forest, or by less productive ericaceous shrublands. Long-term monitoring, manipulative experiments, and process modeling of climate-induced phenological changes on herbivorous insect pests, their host tree species, and natural enemies in northern forests are therefore crucial to predicting species range shifts and assessing ecological and economic impacts. [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]

Laboratory studies show that introduction of fresh and easily decomposable organic carbon (OC) into soil-water systems can stimulate the decomposition of soil OC (SOC) via priming effects in temperate forests, shrublands, grasslands, and agro-ecosystems. However, priming effects are still not well understood in the field setting for temperate ecosystems and virtually nothing is known about priming effects (e.g., existence, frequency, and magnitude) in boreal ecosystems. In this study, a coupled dissolved OC (DOC) transport and microbial biomass dynamics model was developed to simultaneously simulate co-occurring hydrological, physical, and biological processes and their interactions in soil pore-water systems. The developed model was then used to examine the importance of priming effects in two black spruce forest soils, with and without underlying permafrost. Our simulations showed that priming effects were strongly controlled by the frequency and intensity of DOC input, with greater priming effects associated with greater DOC inputs. Sensitivity analyses indicated that priming effects were most sensitive to variations in the quality of SOC, followed by variations in microbial biomass dynamics (i.e., microbial death and maintenance respiration), highlighting the urgent need to better discern these key parameters in future experiments and to consider these dynamics in existing ecosystem models. Water movement carries DOC to deep soil layers that have high SOC stocks in boreal soils. Thus, greater priming effects were predicted for the site with favorable water movement than for the site with limited water flow, suggesting that priming effects might be accelerated for sites where permafrost degradation leads to the formation of dry thermokarst. [ABSTRACT FROM AUTHOR]

Abstract: The intensity of boreal forest wildfires is highly variable, such that natural wood charcoal can be produced at different peak temperatures. This, in turn, may have consequences on the physico-chemical and functional properties of the charcoal that is returned to soil. We report on a microcosm study where black spruce wood charcoal produced at five peak temperatures (450, 550, 650, 750 and 850 °C) was added to forest soils, and subsequently incubated at room temperature, with and without additions of condensed tannins and/or protein. A fourth experimental factor was added to this full factorial design, which comprised the mixing of soil so as to simulate mechanical scarification in the field. Increasing the charcoal production temperature resulted in higher %C and lower %O and %H, suggesting an increase in aromatic structures. Specific surface area was negligible at 450 °C, was about 50 m2 g−1 at 550 °C, and was 260–300 m2 g−1 at the three highest temperatures. At these higher charcoal production temperatures, 85–90 % of total surface area was attributed to pores <20 nm. The incubation demonstrated a decline in soil basal respiration (BR) with charcoal addition, with increasing charcoal production temperature and with tannin additions, whereas BR increased with protein addition and with soil mixing. Charcoal addition reduced soil microbial biomass (MB), however charcoal production temperature had no effect on MB. Consequently, we observed a decrease in microbial metabolic quotient (qCO2) with increasing charcoal production temperature, which reflected the decline in BR. Increasing charcoal production temperature increased net production in the non-mixed soils, but decreased net production in the mixed soils. Protein additions increased net production more so in the mixed than in the non-mixed soils. Our results thus show that charcoal production temperature has a marked and significant effect on soil C and N cycling, and that these effects are substantially modified by soil mixing. This further suggests that changes in boreal forest fire intensity, due to anthropogenic changes in tree species composition or climate, could affect the dynamics of forest regeneration through variations in the chemical and functional properties of the charcoal, and that these effects may be substantially modified by post-fire silvicultural treatments such as mechanical scarification. [Copyright &y& Elsevier]

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]

The role of common mycorrhizal networks (CMNs) in postfire boreal forest successional trajectories is unknown. We investigated this issue by sampling a 50-m by 40-m area of naturally regenerating black spruce (Picea mariana), trembling aspen (Populus tremuloides) , and paper birch (Betula papyrifera) seedlings at various distances from alder (Alnus viridis subsp. crispa), a nitrogen-fixing shrub, 5 years after wildfire in an Alaskan interior boreal forest. Shoot biornasses and stern diameters of 4-year-old seedlings were recorded, and the fungal community associated with ectomycorrhizal (ECM) root tips from each seedling was profiled using molecular techniques. We found distinct assemblages of fungi associated with alder compared with those associated with the other tree species, making the formation of CMNs between them unlikely. However, among the spruce, aspen, and birch seedlings, there were many shared fungi (including members of the Pezoloma ericae [Hymenoscyphus ericael species aggregate, Thelephora terrestris, and Russula spp.), raising the possibility that these regenerating seedlings may form interspecies CMNs. Distance between samples did not influence how similar ECM root tip-associated fungal communities were, and of the fungal groups identified, only one of them was more likely to be shared between seedlings that were closer together, suggesting that the majority of fungi surveyed did not have a clumped distribution across the small scale of this study. The presence of some fungal ribotypes was associated with larger or smaller seedlings, suggesting that these fungi may play a role in the promotion or inhibition of seedling growth. The fungal ribotypes associated with larger seedlings were different between spruce, aspen, and birch, suggesting differential impacts of some host-fungus combinations. One may speculate that wildfire-induced shifts in a given soil fungal community could result in variation in the growth response of different plant species after fire and a shift in regenerating vegetation. [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]

Several attributes might influence the decomposition process of fire-killed trees. Here, we tested various tree- and plot-level variables on the decomposition rate of fire-killed black spruce (Picea mariana (Mill.) BSP) in the northern boreal forest. Data were collected from 474 individuals burned 17 years prior to sampling. Mean decomposition rate was relatively slow (k = 0.013) and was lowest for severely burned snags (k = 0.001) and highest for lightly burned logs (k = 0.027-0.036). Vertical position and fire severity were the most important variables influencing the decomposition rates, while plot-level variables were marginally significant. Both predictors strongly influenced the moisture content of fire-killed trees. Logs with greater contact with the ground and lightly burned trees had higher moisture content and faster decomposition rates. Very severely burned trees had lower moisture content because of faster bark shedding. This hampered the decomposition process by slowing the snag falling rate. Higher decomposition rates in lightly burned trees may result from greater colonization by early xylophagous species. By having a considerable impact on the decomposition of woody debris, fire severity may strongly influence many post-fire biological processes related to the woody necromass as well as carbon emission from burned stands. Plusieurs variables peuvent influencer le processus de décomposition des arbres tués par le feu. Dans cette étude, nous avons testé l'effet de plusieurs variables, à l'échelle de l'arbre et du peuplement, sur la décomposition des débris ligneux d'épinette noire (Picea mariana (Mill.) BSP) après feu. Les données proviennent de 474 individus qui ont brûlé 17 ans avant l'échantillonnage. Le taux de décomposition moyen était relativement bas (k = 0,013) et était minimal chez les chicots très sévèrement brûlés (k = 0,001) et maximal chez les débris au sol légèrement brûlés (k = 0,027-0,036). La position de l'arbre et la sévérité du feu représentaient les plus importantes variables influençant les taux de décomposition alors que les variables mesurées à l'échelle du peuplement n'étaient que marginalement significatives. Ces deux variables ont fortement influencé le taux d'humidité des débris ligneux post-feu, ceux présentant un grand contact avec le sol ainsi que ceux étant légèrement brûlés possédant un taux d'humidité plus élevé et un taux de décomposition plus rapide. Les faibles taux d'humidité relevés sur les arbres très sévèrement brûlés seraient causés par une chute plus rapide de l'écorce. Une forte sévérité du feu pourrait aussi retarder le processus de décomposition en ralentissant la chute des chicots. Les taux de décomposition plus rapides chez les arbres légèrement brûlés pourraient résulter d'une plus forte colonisation initiale par les insectes xylophages. En raison de son impact considérable sur la décomposition des débris ligneux, la sévérité du feu pourrait fortement influencer plusieurs processus biologiques post-feu associés à la nécromasse ligneuse ainsi que les émanations de carbone provenant des peuplements brûlés. [ABSTRACT FROM AUTHOR]

Abstract: A study was carried out to investigate the utility of L-band SAR data for estimating aboveground biomass in sites with low levels of vegetation regrowth. Data to estimate biomass were collected from 59 sites located in fire-disturbed black spruce forests in interior Alaska. PALSAR L-band data (HH and HV polarizations) collected on two dates in the summer/fall of 2007 and one date in the summer of 2009 were used. Significant linear correlations were found between the log of aboveground biomass (range of 0.02 to 22.2tha-1) and σ° (L-HH) and σ° (L-HV) for the data collected on each of the three dates, with the highest correlation found using the L-HV data collected when soil moisture was highest. Soil moisture, however, did change the correlations between L-band σ° and aboveground biomass, and the analyses suggest that the influence of soil moisture is biomass dependent. The results indicate that to use L-band SAR data for mapping aboveground biomass and monitoring forest regrowth will require development of approaches to account for the influence that variations in soil moisture have on L-band microwave backscatter, which can be particularly strong when low levels of aboveground biomass occur. [ABSTRACT FROM AUTHOR]

Abstract: Biomass burning in the Alaskan interior is already a major disturbance and source of carbon emissions, and is likely to increase in response to the warming and drying predicted for the future climate. In addition to quantifying changes to the spatial and temporal patterns of burned areas, observing variations in severity is the key to studying the impact of changes to the fire regime on carbon cycling, energy budgets, and post-fire succession. Remote sensing indices of fire severity have not consistently been well-correlated with in situ observations of important severity characteristics in Alaskan black spruce stands, including depth of burning of the surface organic layer. The incorporation of ancillary data such as in situ observations and GIS layers with spectral data from Landsat TM/ETM+ greatly improved efforts to map the reduction of the organic layer in burned black spruce stands. Using a regression tree approach, the R 2 of the organic layer depth reduction models was 0.60 and 0.55 (p <0.01) for relative and absolute depth reduction, respectively. All of the independent variables used by the regression tree to estimate burn depth can be obtained independently of field observations. Implementation of a gradient boosting algorithm improved the R 2 to 0.80 and 0.79 (p <0.01) for absolute and relative organic layer depth reduction, respectively. Independent variables used in the regression tree model of burn depth included topographic position, remote sensing indices related to soil and vegetation characteristics, timing of the fire event, and meteorological data. Post-fire organic layer depth characteristics are determined for a large (>200,000ha) fire to identify areas that are potentially vulnerable to a shift in post-fire succession. This application showed that 12% of this fire event experienced fire severe enough to support a change in post-fire succession. We conclude that non-parametric models and ancillary data are useful in the modeling of the surface organic layer fire depth. Because quantitative differences in post-fire surface characteristics do not directly influence spectral properties, these modeling techniques provide better information than the use of remote sensing data alone. [Copyright &y& Elsevier]

A synthesis was carried out to examine Alaska's boreal forest fire regime. During the 2000s, an average of 767 000 ha·year-1 burned, 50% higher than in any previous decade since the 1940s. Over the past 60 years, there was a decrease in the number of lightning-ignited fires, an increase in extreme lightning-ignited fire events, an increase in human-ignited fires, and a decrease in the number of extreme human-ignited fire events. The fraction of area burned from human-ignited fires fell from 26% for the 1950s and 1960s to 5% for the 1990s and 2000s, a result from the change in fire policy that gave the highest suppression priorities to fire events that occurred near human settlements. The amount of area burned during late-season fires increased over the past two decades. Deeper burning of surface organic layers in black spruce (Picea mariana (Mill.) BSP) forests occurred during late-growing-season fires and on more well-drained sites. These trends all point to black spruce forests becoming increasingly vulnerable to the combined changes of key characteristics of Alaska's fire regime, except on poorly drained sites, which are resistant to deep burning. The implications of these fire regime changes to the vulnerability and resilience of Alaska's boreal forests and land and fire management are discussed. [ABSTRACT FROM AUTHOR]

Black spruce (Picea mariana) is the most abundant tree species in the boreal biome, but little is known about how climate warming may change recruitment in peatlands, especially those affected by permafrost thaw. We used results from a seven-year study in northern Manitoba, Canada, to address the following questions: (1) What is the relative importance of early life history transitions on P. mariana recruitment? (2) How are these transitions mediated by biological and environmental factors, including competition, facilitation, disease, herbivory, water table depth, and soil nutrients? (3) Do interactions among these factors create additional recruitment limitations beyond those imposed by environmental factors changing with climate warming, such as hydrology? Seed rain was measured over six years on forested permafrost plateaus and in neighboring collapse scar bogs. Seed germination and seedling survival and growth were measured over 4-5 years in collapse scars and assessed across a three-level water table treatment. Survival and growth experiments examined additional combinations of above- and belowground vascular plant competition and fertilizer addition. Results showed that failure of germination and survival on growing moss surfaces and reduced survival of seedlings in wetter microsites were primary constraints. Seed influx was significantly lower in collapse scars but likely did not limit recruitment. Biological and environmental factors mediating these life history transitions also differed in relative importance, and interactions among them tended to amplify recruitment limitation. Seedling survival was most strongly controlled by fast-growing mosses in wet microsites but also was influenced by apparent drowning in wet plots, herbivory, and loss of foliage caused by a fungal pathogen. Seedling growth was strongly controlled by water table depth, nutrient and competition levels, and fungal pathogens. Multiple, interacting factors will affect P. mariana establishment in boreal peatlands during climate warming. Generalizations about recruitment relying on few environmental gradients sensitive to climate change, such as water table, may therefore not fully capture the complexities of establishment. [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]

Symbiotic fungi’s role in providing nitrogen to host plants is well-studied in tundra at Toolik Lake, Alaska, but little-studied in the adjoining boreal forest ecosystem. Along a 570 km north-south transect from the Yukon River to the North Slope of Alaska, the 15N content was strongly reduced in ectomycorrhizal and ericoid mycorrhizal plants including Betula, Salix, Picea mariana (P. Mill.) B.S.P., Picea glauca Moench (Voss), and ericaceous plants. Compared with the 15N content of soil, the foliage of nonmycorrhizal plants (Carex and Eriophorum) was unchanged, whereas content of the ectomycorrhizal fungi was very much higher (e.g., Boletaceae, Leccinum and Cortinarius). It is hypothesized that similar processes operate in tundra and boreal forest, both nitrogen-limited ecosystems: (i) mycorrhizal fungi break down soil polymers and take up amino acids or other nitrogen compounds; (ii) mycorrhizal fungi fractionate against 15N during production of transfer compounds; (iii) host plants are accordingly depleted in 15N; and (iv) mycorrhizal fungi are enriched in 15N. Increased N availability for plant roots or decreased light availability to understory plants may have decreased N allocation to mycorrhizal partners and increased δ15N by 3‰-4‰ for southern populations of Vaccinium vitis-idaea L. and Salix. Fungal biomass, measured as ergosterol, correlated strongly with soil organic matter and attained amounts similar to those in temperate forest soils. [ABSTRACT FROM AUTHOR]

Abstract: This paper will discuss TRIPLEX-Flux, a process-based model that estimates net ecosystem production (NEP) as well as analyzing the level of sensitivity of the model''s response by simulating CO2 flux in an existing old black spruce BOREAS site in central Canada. The research objectives were: (1) to test the TRIPLEX-Flux model simulations against flux tower measurements; and (2) to examine parameter and input variable effects on model response via sensitivity analysis. Validation of NEP data at 30min intervals derived from tower and chamber measurements showed that the NEP data from the model corresponded well with the measured NEP from the BOREAS site (R 2 >0.65). Sensitivity analysis demonstrated different levels of sensitivity between morning and noon periods and from the current to doubled atmospheric CO2 concentrations. Additionally, the comparison of different algorithms to calculate stomatal conductance showed that the NEP predicted by the model using the iteration algorithm was consistent with the results using a constant C i/C a of 0.74 for current and 0.81 for doubled CO2 concentrations. Varying parameter and input variable values by ±10% resulted in a similar model response between morning and noon periods (less than or equal to 27.6% and 27.4%, respectively). Most parameters were more sensitive at noon than they were in the morning except for those that were correlated with air temperature, suggesting that air temperature has considerable influence over model sensitivity for these parameters/variables. Air temperature effect was greater under doubled than it was under current atmospheric CO2 concentrations. In contrast, model sensitivity to CO2 decreased under doubled CO2 concentrations. [Copyright &y& Elsevier]

Abstract: The boreal forest, earth''s second largest terrestrial biome, is currently thought to be an important net carbon sink for the atmosphere. In this study, a newly developed carbon exchange model of TRIPLEX-Flux (with half-hourly time step) is used to simulate the ecosystem carbon exchange of a 75-year-old boreal mixedwood forest stand in northeast Ontario and a 110-year-old pure black spruce stand in southern Saskatchewan, Canada. Results of net ecosystem exchange (NEE) simulated by TRIPLEX-Flux for 2004 are compared with those measured by eddy-flux towers and suggest overall agreement between model simulation and observations. The mean coefficient of determination (R 2) is approximately 0.77 for boreal mixedwood and 0.62 for old black spruce. Differences between model simulation and observation may be attributed to uncertainties not only in model input parameters and calibration, but also in eddy-flux measurements caused by systematic and random errors. The model is able to capture the diurnal variations of NEE for the growing season (from May to August) of 2004 for both sites. Both boreal mixedwood and old black spruce were acting as carbon sinks for the atmosphere during the growing season of 2004. However, the boreal mixedwood stand shows higher ecosystem productivity, carbon sequestration, and carbon use efficiency than the old black spruce stand. [Copyright &y& Elsevier]

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]

Aim Our two main goals are first to evaluate the resilience of the boreal forest according to latitude across the closed-crown forest zone using the post-disturbance distribution and cover of lichen woodlands and closed-crown forests as a metric, and second to identify the disturbance factors responsible for the regeneration and degradation of the closed-crown forest according to latitude since the 1950s. Location The study area extends between 70°00′ and 72°00′ W and throughout the closed-crown forest zone, from its southern limit near 47°30′ N to its northern limit at the contact with the lichen woodland zone at around 52°40′ N. Methods Recent (1972–2002) and old (1954–1956) aerial photos were used to map the distribution of lichen woodlands across the closed-crown forest zone. Forest disturbances such as fire, spruce budworm ( Choristoneura fumiferana (Clemens)) outbreak, and logging were recorded on each set of aerial photos. Each lichen woodland and stand disturbance was validated by air-borne surveys and digitized using GIS software. Results Over the last 50 years, the area occupied by lichen woodlands has increased according to latitude; that is, 9% of the area that was occupied by closed-crown forests has shifted to lichen woodlands. Although logging activities have been concentrated in the same areas during the last 50 years, the area covered by logging has increased significantly. Outbreaks by the spruce budworm occurred predominantly in the southern (47°30′ N to 48°30′ N) and central (48°53′ N to 50°42′ N) parts of the study area, where balsam fir stands are extensive. In the northern part of the study area (51°–52°40′ N), extensive fires affected the distribution and cover of closed-crown forests and lichen woodlands. Main conclusions Over the last 50 years, the area occupied by closed-crown forests has decreased dramatically, and the ecological conditions that allow closed-crown forests to establish and develop are currently less prevalent. Fire is by far the main disturbance, reducing the ability of natural closed-crown forests to self-regenerate whatever the latitude. Given the current biogeographical shift from dense to open forests, the northern part of the closed-crown forest zone is in a process of dramatic change towards the dominance of northern woodlands. [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]

The objective of this study was to characterize the effects of soil burn severity and initial tree composition on long-term forest floor dynamics and ecosystem biomass partitioning within the Picea mariana [Mill.] BSP-feathermoss bioclimatic domain of northwestern Quebec. Changes in forest floor organic matter and ecosystem biomass partitioning were evaluated along a 2,355-year chronosequence of extant stands. Dendroecological and paleoecological methods were used to determine the time since the last fire, the soil burn severity of the last fire (high vs. low severity), and the post-fire tree composition of each stand ( P. mariana vs. Pinus banksiana Lamb). In this paper, soil burn severity refers to the thickness of the organic matter layer accumulated above the mineral soil that was not burned by the last fire. In stands originating from high severity fires, the post-fire dominance by Pinus banksiana or P. mariana had little effect on the change in forest floor thickness and tree biomass. In contrast, stands established after low severity fires accumulated during the first century after fire 73% thicker forest floors and produced 50% less tree biomass than stands established after high severity fires. Standing tree biomass increased until approximately 100 years after high severity fires, and then decreased at a logarithmic rate in the millennial absence of fire. Forest floor thickness also showed a rapid initial accumulation rate, and continued to increase in the millennial absence of fire at a much slower rate. However, because forest floor density increased through time, the overall rate of increase in forest floor biomass (58 g m−2 y−1) remained constant for numerous centuries after fire (700 years). Although young stands (< 200 years) have more than 60% of ecosystem biomass locked-up in living biomass, older stands (> 200 years) sequester the majority (> 80%) of it in their forest floor. The results from this study illustrate that, under similar edaphic conditions, a single gradient related to time since disturbance is insufficient to account for the full spectrum of ecosystem biomass dynamics occurring in eastern boreal forests and highlights the importance of considering soil burn severity. Although fire severity induces diverging ecosystem biomass dynamics in the short term, the extended absence of fire brings about a convergence in terms of ecosystem biomass accumulation and partitioning. [ABSTRACT FROM AUTHOR]

1 The northward expansion of the boreal forest vegetation zone is generally predicted under a warmer doubled CO2, but the delay associated with vegetation development processes often has been overlooked. In the subarctic forest-tundra of northern Québec, reforestation of tundra uplands appears currently limited by the poor reproductive capacity of shrubby black spruce (Picea mariana), and the development of erect stems through accelerated height growth should be the first registered response to 20th century climate warming. The subarctic forest-tundra is characterized by small- and large-scale heterogeneity in topography, vegetation structure and climate. This spatial heterogeneity, added to the complexity of tree growth–climate relationships, can cause various growth responses of subarctic tree line black spruce to 20th century climate change.2 Twenty spruce populations at subarctic tree lines and seven isolated clones at the species limit were sampled along a> 300-km latitudinal transect from the southern forest-tundra to the shrub tundra. Height growth patterns of black spruce at tree line and above tree line were examined (i) over their life span, using dendrochronological dating of stem cross-sections, and (ii) for the recent decades, using leader shoot elongation measurements. Indexed elongation chronologies were compared with regional climate data.3 Height growth of tree line trees generally decreased with increasing latitude. However, tree line trees in the northern forest-tundra have experienced an acceleration of height growth since the 1970s, with their growth comparable to that of trees in the southern forest-tundra. Height growth response of spruce trees appeared increasingly delayed from the northern forest-tundra to the species limit. Above the subarctic tree line, wind-exposed conditions obscured the decrease in height growth with latitude observed for tree line trees.4 Leader shoot elongation of spruce trees established on tundra hilltops appeared more controlled by summer heat sums than those at tree line all over the forest-tundra, except at the arctic tree line. Winter precipitation also was linked to leader shoot elongation in some forest-tundra sites. The increasing snow cover associated with recent warming appeared to have reduced the shoot elongation of spruce at forest margins showing the steepest slopes, hence subjected to snow overloading.5 In the northern forest-tundra sites, the recent increase in height growth and positive trend in leader shoot elongation, consistent with a 1990s’ increase in heat sums, point to the development of spruce krummholz into erect growth forms. In the southern forest-tundra, reforestation of tundra hilltops and northward expansion of the boreal forest predicted under doubled CO2 conditions could be delayed, as suggested by suppressed height growth of spruce above tree line.Journal of Ecology(2004)92, 835 –845 [ABSTRACT FROM AUTHOR]

Net primary production (NPP) was measured in seven black spruce ( Picea mariana (Mill.) BSP)-dominated sites comprising a boreal forest chronosequence near Thompson, Man., Canada. The sites burned between 1998 and 1850, and each contained separate well- and poorly drained stands. All components of NPP were measured, most for 3 consecutive years. Total NPP was low (50–100 g C m−2 yr−1) immediately after fire, highest 12–20 years after fire (332 and 521 g C m−2 yr−1 in the dry and wet stands, respectively) but 50% lower than this in the oldest stands. Tree NPP was highest 37 years after fire but 16–39% lower in older stands, and was dominated by deciduous seedlings in the young stands and by black spruce trees (>85%) in the older stands. The chronosequence was unreplicated but these results were consistent with 14 secondary sites sampled across the landscape. Bryophytes comprised a large percentage of aboveground NPP in the poorly drained stands, while belowground NPP was 0–40% of total NPP. Interannual NPP variability was greater in the youngest stands, the poorly drained stands, and for understory and detritus production. Net ecosystem production (NEP), calculated using heterotrophic soil and woody debris respiration data from previous studies in this chronosequence, implied that the youngest stands were moderate C sources (roughly, 100 g C m−2 yr−1), the middle-aged stands relatively strong sinks (100–300 g C m−2 yr−1), and the oldest stands about neutral with respect to the atmosphere. The ecosystem approach employed in this study provided realistic estimates of chronosequence NPP and NEP, demonstrated the profound impact of wildfire on forest–atmosphere C exchange, and emphasized the need to account for soil drainage, bryophyte production, and species succession when modeling boreal forest C fluxes. [ABSTRACT FROM AUTHOR]

Fine root processes play a prominent role in the carbon and nutrient cycling of boreal ecosystems due to the high proportion of biomass allocated belowground and the rapid decomposition of fine roots relative to aboveground tissues. To examine these issues in detail, major components of ecosystem carbon flux were studied in three mature black spruce forests in interior Alaska, where fine root production, respiration, mortality and decomposition, and aboveground production of trees, shrubs, and mosses were measured relative to soil CO[sub 2] fluxes. Fine root production, measured over a two-year period using minirhizotrons, varied from 0.004 ± 0.001 mm·cm[sup -2]·d[sup -1] over winter, to 0.051 ± 0.015 mm·cm[sup -2]·d[sup -1] during July, with peak growing season values comparable to those reported for many temperate forests using similar methods. On average, 84% of this production occurred within 20 cm of the moss surface, although the proportion occurring in deeper profiles increased as soils gradually warmed throughout the summer. Monthly rates of production and mortality were somewhat asynchronous because mortality tended to peak during fall and be minimal during periods of peak production. Production and mortality were, however, positively correlated across all tubes and time periods. Annual fine root production averaged 2.45 ± 0.31, 8.01 ± 1.39, and 2.53 ± 0.27 mm·cm[sup -2]·yr[sup -1] (means ± 1 SE) among the three sites, when averaged across years. Fine root survival and decomposition were measured by tracking and analyzing the fate of individual fine roots using mark-recapture techniques. Fine root survival was greatest during periods of peak root growth, and least over winter (φ[sub time]). Roots first appearing in the middle of the growing season had higher survival rates than those first appearing early or late in the growing season, or over winter (φ[sub cohort]), and risk of mortality decreased with root age (φ[sub age]). Survival estimates translate to mean life spans of 108 ± 4 d during the growing season. While these values are in striking contrast to needle longevity and rates of aboveground litter decomposition, they are similar to many values found for temperate systems, supporting the notion that there are basic morphological and physiological traits of first-order roots that are common to most woody plant root systems. During the growing season, monthly fine root decomposition rates averaged 0.46 ± 0.01 per month, while decomposition rates over winter averaged 0.73 ± 0.01 per winter. These growing season estimates translate to 49 ± 2 d from the time a root was first observed as dead, to the time it disappeared. For roots that decomposed during the growing season, those with longer life spans decomposed more slowly after death. Comparing these results with other minirhizotron studies suggests that life-history traits of black spruce first-order roots are similar to those from temperate (and perhaps most) forest ecosystems. Annual production of fine roots averaged 228 ± 75 g biomass·m[sup -2]·yr[sup -1], constituting ∼56% of total stand production. Aboveground production of trees (50 ± 14 g biomass·m[sup -2]·yr[sup -1], 13%) and shrubs (40 ± 2 g biomass·m[sup -2]·yr[sup -1], 11%) contributed similarly to total production, while mosses (73 ± 14 g biomass·m[sup -2]·yr[sup -1], 20%) accounted for the largest component of aboveground production. Soil temperature had a strong control over both soil respiration (Q[sub 10] = 2.21 ± 0.31) and root respiration (Q[sub 10] = 2.30 ± 0.37). During the growing season (15 May to 15 September), ∼56% of soil CO[sub 2] efflux (580 ± 40 g C/m²) was derived from fine root respiration (329 ± 54 g C/m²). Although apparent rates of heterotrophic respiration (May through September) and total production did not differ, definitive estimates of net ecosystem production are impossible given the potentially large, unmeasured components of NPP (net primary production), such as root exudation and mycorrhizal production. Nevertheless, rates of fine root production, mortality, and decomposition indicate that in these black spruce ecosystems, fine roots are much more dynamic than would be predicted from patterns of aboveground processes, and that carbon, and presumably nutrients, are cycling through fine roots at rates several orders of magnitude faster than through aboveground tissues. [ABSTRACT FROM AUTHOR]

Determines the nature of the negative influence of Kalmia angustifolia L. on black spruce (Picea mariana) seedling growth and foliar nutrient concentrations, and the effect of proximity to Kalmia on spruce seedlings was studied on two Kalmia-dominated sites of contrasting soil characteristics in central Newfoundland. Results of sampling of spruce seedlings and the soil at the base of their stems; Use of path analysis; How path diagrams were generated based on current knowledge of nutrient cycling in boreal ecosystems and mechanisms previously hypothesized to account for the influence of Kalmia on black spruce; Results of analysis on the wetter and richer site suggesting nutrient competition; Results on the drier and poorer site suggesting negative effect of Kalmia on the spruce.

Presents the results of a study to retrospectively evaluate, with the help of stem analysis, the growth increases of natural and planted black spruce (Picea mariana (Mill.) BSP) seedlings in 39 drainage networks located in Quebec public boreal forest and drained operationally after cutting. Ecological conditions of the site; Effect of the drainage on 9-year periodic growth in diameter and height of natural black spruce seedlings; Conclusion that height growth increment of planted black spruce seedlings was weaker than that of the seedlings.

The authors discuss the effects of warming to the drought tolerance of boreal tree. They state that drought and increased in temperature possibly threaten the species of black spruce in boreal ecology, and discovered to be the main cause of escalating boreal trees' mortality rate. They mention that in the development of black spruce seedlings, its carbon balance is lowered by elevated temperatures.

Net ecosystem productivity (NEP) of boreal coniferous forests is believed to rise with climate warming, thereby offsetting some of the rise in atmospheric CO2 concentration (Ca) by which warming is caused. However, the response of conifer NEP to warming may vary seasonally, with rises in spring and declines in summer. To gain more insight into this response, we compared changes in CO2 exchange measured by eddy covariance and simulated by the ecosystem process model ecosys under rising mean annual air temperatures (Ta) during 2004–2006 at black spruce stands in Saskatchewan, Manitoba and Quebec. Hourly net CO2 uptake was found to rise with warming at Ta < 15 °C and to decline with warming at Ta > 20 °C. As mean annual Ta rose from 2004 to 2006, increases in net CO2 uptake with warming at lower Ta were greater than declines with warming at higher Ta so that annual gross primary productivity and hence NEP increased. Increases in net CO2 uptake measured at lower Ta were explained in the model by earlier recovery of photosynthetic capacity in spring, and by increases in carboxylation activity, using parameters for the Arrhenius temperature functions of key carboxylation processes derived from independent experiments. Declines in net CO2 uptake measured at higher Ta were explained in the model by sharp declines in mid-afternoon canopy stomatal conductance (gc) under higher vapor pressure deficits (D). These declines were modeled from a hydraulic constraint to water uptake imposed by low axial conductivity of conifer roots and boles that forced declines in canopy water potential (c), and hence in gc under higher D when equilibrating water uptake with transpiration. In a model sensitivity study, the contrasting responses of net CO2 uptake to specified rises in Ta caused annual NEP of black spruce in the model to rise with increases in Ta of up to 6 °C, but to decline with further increases at mid-continental sites with lower precipitation. However, these contrasting responses to warming also indicate that rises in NEP with climate warming would depend on the seasonality (spring versus summer) as well as the magnitude of rises in Ta. [ABSTRACT FROM AUTHOR]