Effects of stand composition on fire hazard in mixed‐wood Canadian boreal forest

Abstract. Surface fuels were examined in 48 stands of the Canadian mixed‐wood boreal forest. Tree canopy was characterized with the point‐centred quadrant method and stands were characterized as deciduous, mixed‐deciduous, mixed‐coniferous or coniferous according to the percentage of conifer basal area. Woody debris loadings were measured with the line intersect method and the litter, duff, shrub loads and depths or heights were sampled with various quadrats. No significant difference was found among stand types for total woody debris load, large basal diameter shrub loads and load or depth of litter and duff. However, conifer stands had significantly heavier loads of small diameter elements (twigs and shrubs) and conifer pieces were more numerous within these stands than in deciduous stands. The BEHAVE prediction system was used to evaluate the impact of these differences on the potential of fire ignition in situations where topography and weather were constant. The qualitative and quantitative changes in fuels, resulting from species replacement and fast decay rates, influence fire hazard. Simulations of fire behaviour showed that in the mixed‐wood boreal forest fires were less intense and spread more slowly in deciduous stands than in mixed or coniferous stands. Moreover, spring fires were more intense than summer fires, and differences between seasons increased with the increase of deciduous basal area.     

Massive mortality of aspen following severe drought along the southern edge of the Canadian boreal forest

Drought‐induced, regional‐scale dieback of forests has emerged as a global concern that is expected to escalate under model projections of climate change. Since 2000, drought of unusual severity, extent, and duration has affected large areas of western North America, leading to regional‐scale dieback of forests in the southwestern US. We report on drought impacts on forests in a region farther north, encompassing the transition between boreal forest and prairie in western Canada. A central question is the significance of drought as an agent of large‐scale tree mortality and its potential future impact on carbon cycling in this cold region. We used a combination of plot‐based, meteorological, and remote sensing measures to map and quantify aboveground, dead biomass of trembling aspen (Populus tremuloides Michx.) across an 11.5 Mha survey area where drought was exceptionally severe during 2001–2002. Within this area, a satellite‐based land cover map showed that aspen‐dominated broadleaf forests occupied 2.3 Mha. Aerial surveys revealed extensive patches of severe mortality (>55%) resembling the impacts of fire. Dead aboveground biomass was estimated at 45 Mt, representing 20% of the total aboveground biomass, based on a spatial interpolation of plot‐based measurements. Spatial variation in percentage dead biomass showed a moderately strong correlation with drought severity. In the prairie‐like, southern half of the study area where the drought was most severe, 35% of aspen biomass was dead, compared with an estimated 7% dead biomass in the absence of drought. Drought led to an estimated 29 Mt increase in dead biomass across the survey area, corresponding to 14 Mt of potential future carbon emissions following decomposition. Many recent, comparable episodes of drought‐induced forest dieback have been reported from around the world, which points to an emerging need for multiscale monitoring approaches to quantify drought effects on woody biomass and carbon cycling across large areas.     

Effects of overstory and understory vegetation on the understory light environment in mixed boreal forests

Abstract. The percentage of above-canopy Photosynthetic Photon Flux Density (%PPFD) was measured at 0, 50 and 100 cm above the forest floor and above the main understory vegetation in stands of (1) pure Betula papyrifera (White birch), (2) pure Populus tremuloides (Trembling aspen), (3) mixed broad-leaf-conifer, (4) shade-tolerant conifer and (5) pure Pinus banksiana (Jack pine) occurring on both clay and till soil types. %PPFD was measured instantaneously under overcast sky conditions (nine locations within each of 29 stands) and continuously for a full day under clear sky conditions (five locations within each of eight stands). The percentage cover of the understory layer was estimated at the same locations as light measurements. Mean %PPFD varied from 2% at the forest floor under Populus forests to 15% above the understory vegetation cover under Betula forests. Percent PPFD above the understory vegetation cover was significantly higher under shade intolerant tree species such as Populus, Betula and Pinus than under shade tolerant conifers. No significant differences were found in %PPFD above the understory vegetation cover under similar tree species between clay and till soil types. The coefficient of variation in %PPFD measured in the nine locations within each stand was significantly lower under deciduous dominated forests (mean of 19%) than under coniferous dominated forests (mean of 40%). %PPFD measured at the forest floor was positively correlated with %PPFD measured above the understory vegetation and negatively correlated with cumulative total percent cover of the understory vegetation (R² = 0.852). The proportion of sunflecks above 250 and 500 mmol m^–2 s^–1 was much lower and %PPFD in shade much higher under Populus and Betula forests than under the other forests. Differences in the mean, variability and nature of the light environment found among forest and soil types are discussed in relation to their possible influences on tree succession.     

Radial growth response of four dominant boreal tree species to climate along a latitudinal gradient in the eastern Canadian boreal forest

To address the central question of how climate change influences tree growth within the context of global warming, we used dendroclimatological analysis to understand the reactions of four major boreal tree species –Populus tremuloides, Betula papyrifera, Picea mariana, and Pinus banksiana– to climatic variations along a broad latitudinal gradient from 46 to 54°N in the eastern Canadian boreal forest. Tree‐ring chronologies from 34 forested stands distributed at a 1° interval were built, transformed into principal components (PCs), and analyzed through bootstrapped correlation analysis over the period 1950–2003 to identify climate factors limiting the radial growth and the detailed radial growth–climate association along the gradient. All species taken together, previous summer temperature (negative influences), and current January and March–April temperatures (positive influences) showed the most consistent relationships with radial growth across the gradient. Combined with the identified species/site‐specific climate factors, our study suggested that moisture conditions during the year before radial growth played a dominant role in positively regulating P. tremuloides growth, whereas January temperature and growing season moisture conditions positively impacted growth of B. papyrifera. Both P. mariana and P. banksiana were positively affected by the current‐year winter and spring or whole growing season temperatures over the entire range of our corridor. Owing to the impacts of different climate factors on growth, these boreal species showed inconsistent responsiveness to recent warming at the transition zone, where B. papyrifera, P. mariana, and P. banksiana would be the most responsive species, whereas P. tremuloides might be the least. Under continued warming, B. papyrifera stands located north of 49°N, P. tremuloides at northern latitudes, and P. mariana and P. banksiana stands located north of 47°N might benefit from warming winter and spring temperatures to enhance their radial growth in the coming decades, whereas other southern stands might be decreasing in radial growth.     

Differences in forest composition in two boreal forest ecoregions of Quebec

Abstract. In order to describe and compare the post‐fire succession patterns of the two ecological regions (mixed‐wood and coniferous ecoregions) of northwestern Quebec, 260 forest stands were sampled with the point‐centred plot method. The mixed‐wood ecological region belongs to the Abies balsamea‐Betula papyrifera bioclimatic domain whereas the coniferous ecological region belongs to the Picea mariana‐moss bioclimatic domain. In each plot, tree composition was described, surficial deposits and drainage were recorded, and fire history was reconstructed using standard dendro‐ecological methods. Ordination techniques (Correspondence Analysis and Canonical Correspondence Analysis) were used to describe the successional patterns of forest vegetation and to correlate them with the explanatory variables. The results showed the importance of surficial deposits, the time since fire and the ecoregion in explaining the variation of stand composition. Abies balsamea tends to increase in importance with an increase in time since fire, and this trend is more pronounced in the mixed‐wood region. Even when controlling both for surficial deposits and time since fire, differences in successional trends were observed between the two ecoregions. As all the species are present in both ecoregions and as they are all observed further north, our results suggest that both the landscape configuration and fire regime parameters such as fire size and fire intensity are important factors involved in these differences.     

Long‐term post‐fire changes in the northeastern boreal forest of Quebec

Abstract. Natural dynamics in the boreal forest is influenced by disturbances. Fire recurrence affects community development and landscape diversity. Forest development was studied in the northeastern boreal forest of Quebec. The objective was to describe succession following fire and to assess the factors related to the changes in forest composition and structure. The study area is located in northeastern Quebec, 50 km north of Baie‐Comeau. We used the forest inventory data gathered by the Ministère des Ressources naturelles du Québec (MRNQ). In circular plots of 400 m², the diameter at breast height (DBH) of all stems of tree species greater than 10 cm was recorded and in 40 m² subplots, stems smaller than 10 cm were measured. A total of 380 plots were sampled in an area of 6000 km². The fire history reconstruction was done based on historical maps, old aerial photographs and field sampling. A time‐since‐fire class, a deposit type, slope, slope aspect and altitude were attributed to each plot. Each plot was also described according to species richness and size structure characteristics. Traces of recent disturbance were also recorded in each plot. Changes in forest composition were described using ordination analyses (NMDS and CCA) and correlated with the explanatory variables. Two successional pathways were observed in the area and characterized by the early dominance of intolerant hardwood species or Picea mariana. With time elapsed since the last fire, composition converged towards either Picea mariana, Abies balsamea or a mixture of both species and the size structure of the coniferous dominated stands got more irregular. The environmental conditions varied between stands and explained part of the variability in composition. Their effect tended to decrease with increasing time elapsed since fire, as canopy composition was getting more similar. Gaps may be important to control forest dynamics in old successional communities.     

Predictors of moss and liverwort species diversity of microsites in conifer‐dominated boreal forest

Patterns of moss and liverwort species diversity — species richness and species turnover (β‐diversity) — in three conifer‐dominated boreal forest stands of northern Alberta, Canada are described. We examined the relationship between bryophyte species diversity and micro‐environment at two sample grains, the microsite — substrate types for moss colonization: logs, stumps, tree bases, undisturbed patches of forest floor (dominated by feather moss species), and disturbed patches of forest floor — and the mesosite (25 m × 25 m plots). Microsite type and properties (e.g. decay class, hardwood vs softwood, pH) were the principal predictors of bryophyte species diversity and not micro‐environment variation among mesosites. Microsite type was the strongest predictor of microsite species richness and β‐diversity was higher among microsites and types and within microsites than among mesosites or stands. Microsite properties were significant predictors of species richness for all microsite types. Log and stump decay classes, influenced also by hardwood vs softwood predicted species richness of woody microsite types and soil pH and moisture predicted species richness of forest floor microsites. β‐diversity was highest for tree bases and disturbed patches of forest floor and lowest for logs. Mesosite β‐diversity was lower than that among microsites, and mesosite species richness was not well explained by measured environmental parameters. Results suggest that in conifer‐dominated boreal stands, species richness of microsites is only negligibly influenced by within‐stand variation at the mesosite grain and that substrate characteristics are the most important predictors of bryophyte species diversity in this ecosystem.     

峨眉山冷杉森林群落研究

依据 5 1个 40 0 m2乔木层和 1 0 2个 1 6m2灌木层、草本层与地被层样方调查资料 ,分析了峨嵋山冷杉森林的林型、环境、植物地理成分、群落结构和乔、灌层植被数量特征。结果表明 :当地冷杉森林分布区内生境多样 ;植物区系复杂 ,有一定的热性地理成分入侵和残遗性 ;各层次水平盖度具有精巧的消长关系 ,垂直结构在亚层水平上的发育与海拔和“混交”程度有关 ;冷杉在乔木层占绝对优势 ,但由于其种群衰退 ,冷杉所构成的森林有退化演替的趋势 ;灌木层的优势植物以竹类为主 ,亚优势种的变化多样。生物地史、生境和植物区系的种种特点 ,共同决定了当地冷杉林型的多样性及其某些少见的种群组合式样。     

Predicting the composition of Canadian southern boreal forest in different fire cycles

Abstract. Post-fire succession was reconstructed for a sector located in the southern part of the Québec boreal forest. Forest composition for different periods since fire was evaluated using a stand initiation map together with ecological maps representing both site conditions and stand types. Nine fires covering at least 100 ha and representing a chronosequence of more than 230 yr were used. Although a relatively clear successional pattern from deciduous to coniferous composition relating to time-since-fire was observed, Pinus banksiana stands showed an erratic distribution not related to succession but possibly to the pre-fire stand composition. A comparison with forest cover maps produced after a recent spruce budworm outbreak, showed that succession toward coniferous dominance appeared to be interrupted by spruce budworm (Choristoneura fumiferana) outbreaks which, by killing Abies balsamea, lead to a mixed deciduous forest composition. A simple empirical model based on a negative exponential distribution of age classes was developed to evaluate how changes in the fire cycle would affect the composition of the forest mosaic. The transition between deciduous dominance and coniferous dominance occurs in a fire cycle > 200 yr. Although pure deciduous stands tend to disappear during long fire cycles, the proportion of mixed stands remains relatively constant. Prediction of the forest composition for longer fire cycles is complicated by the interaction between post-fire composition and stand vulnerability to spruce budworm outbreaks.     

Conifer seedling recruitment in a southeastern Canadian boreal forest: the importance of substrate

Abstract. In order to explain conifer species recruitment in Canada's southeastern boreal forest, we characterized conifer regeneration microsites and determined how these microsites vary in abundance during succession. Microsite abundance was evaluated in deciduous, mixed and coniferous stands along a 234-yr postfire chronosequence. Conifers were most often found in relatively well-illuminated microsites, devoid of litter, especially broad-leaf litter, and with a reduced cover of lower vegetation (< 50 cm tall). Although associated with moss-rich forest floor substrates, Abies balsamea was the most ubiquitously distributed species. Picea glauca and especially Thuja occidentalis seedlings were frequently found on rotten logs. Light measurements did not show differences among seedling species nor between stand types. The percentage cover of broad-leaf litter decreased significantly during succession. Also, rotten logs covered with moss occupied a significantly larger area in the mid-successionnal stands than in early successional deciduous or late successional coniferous stands. The results suggest that the presence of specific forest floor substrate types is a factor explaining low conifer recruitment under deciduous stands, conifer codominance in the mid-successional stage, and delayed Thuja recolonization after fire. Results also suggest that some facilitation mechanism is responsible for the observed directional succession.     

Changes in the forest floor of Canadian southern boreal forest after disturbance

Abstract. The concentrations and contents of organic matter and nutrients in organic deposits on the forest floor were estimated along a 231-yr chronosequence following fire at the southern limit of the boreal forest in eastern Canada. The sampling design was stratified to take into account the variability related to the presence of the principal tree species as well as to the presence of large gaps created by a recent spruce budworm (Choristoneura fumiferana) outbreak. The forest floor showed a steady accumulation of organic matter and total nutrients with time-since-fire and a 50 % decrease in the concentrations of available P and K, but not N (as determined by aerobic incubation). The increase in forest-floor weight was accompanied by an increased storage of available N, Ca and Mg. The availability of N and Ca was more strongly affected by tree species and gaps than by time-since-fire. A high N-availability was observed under Betula papyrifera and in gaps, while high a Ca-availability was found near Populus tremuloides and Thuja occidentalis. In old sites, the forest floor of gaps, created by a recent spruce budworm outbreak, had a necromass similar to that of a young forest, but the low concentrations of available P and K of an old forest.     

Changes in the understory of Canadian southern boreal forest after fire

Abstract. We investigated changes in the composition and abundance of understory species after fire in the southern boreal forest around Lake Duparquet, Québec. Ten plots of 100 m² were sampled in each of eight sites varying in post-fire age from 26 to 230 yr, with 20 1-m² quadrats in each of these 80 plots. Variation in the understory was described by DCA ordination and interpreted as a regeneration succession series. Thickness of the organic layers, stand age and canopy composition were all correlated with vegetational change. This change was not constant throughout succession; some old sites showed an increase in the diversity and abundance of certain pioneer species. This was partly related to openings in the canopy resulting from a major outbreak of spruce budworm, which affected sites dominated by Abies balsamea. The ordinations were performed on both the 100-m² plots and the 1-m² quadrats. Heterogeneity within sites was larger at the 1-m² scale and there was a great deal of overlap in the position of the quadrats in ordination space. At the smaller scale of analysis, stand age and thickness of the organic layers were not correlated with the changes observed in the understory.     

Fifty‐seven years of composition change in the eastern boreal forest of Canada

Question: In the boreal forest of eastern Canada, how does forest vegetation change in the sustained absence of fire? Location: Eastern boreal forest in Quebec's North Shore region, Canada (49°30′–50°00′N; 67°30′–68°35′W). Methods: Aerial photos from three different periods (1930, 1965 and 1987) were used to characterize changes in vegetation composition in 23 scenes of 200 ha. Time since fire, presence of secondary disturbances and data on soil and topographic variables were obtained. Ordination and clustering techniques were used to define compositional trajectories of change over the 57‐yr period. These trajectories were further grouped into pathways based on compositional changes, time since fire and preferential deposit‐drainage types. Results: Among the 26 compositional trajectories, three successional pathways were distinguished. Two start post‐fire succession with a dominance of intolerant hardwood. In one of these, this is followed by an increase in Abies balsamea, while in the second the importance of Picea mariana increases with time. In the third pathway P. mariana is an important component from the outset. In this pathway, we observed modest fluctuation in the relative dominance of P. mariana and A. balsamea and variation in stand structure. Conclusion: The boreal forest vegetation of Eastern Canada is diverse and dynamic even in the absence of fire, notably under the influence of partial disturbances. Such disturbances can be associated with changes in composition or stand structure. The development of management strategies aimed at maintaining stand diversity by emulating a broader variety of partial and secondary disturbances should be encouraged.     

Effects of leaf litter on the growth of boreal feather mosses: Implication for forest floor development

Question: What is the effect of leaf litter on growth and mortality of feather mosses. Three experiments were conducted to isolate the shading effect from the effect of leaching of soluble compounds from the leaf litter effect on feather moss mortality and growth. Location: Edmonton, Alberta and the northern boreal coniferous forest, west‐central Alberta, Canada. Methods: In a field experiment Populus tremuloides (aspen) leaf litter was applied to beds of feather mosses dominated by Hylocomium splendens. Treatments were one layer of leaves held in place with netting; one layer of leaves coarsely ground, sprinkled over the moss layer; a shade cloth equivalent to one layer of leaves; and a control. In two growth chamber experiments, the application of aqueous extracts of P. tremuloides and Pinus contorta leaf litter and the effect of shade and soluble carbohydrates were tested on the growth and mortality of Ptilium crista‐castrensis. Results: Mortality of Hylocomium was greatest under the intact leaves, followed by the treatments using shade cloth and ground leaves and finally the control. The application of aqueous extracts of aspen leaf litter resulted in senescence or death of nearly all Ptilium shoots compared to no effects for the control or for similar extracts of pine needle litter. Extracts from aspen litter had greater concentrations of phenolic compounds and soluble sugars than pine extracts. Addition of sugars to Ptilium allowed it to grow and accumulate carbohydrates, even in low light conditions. Conclusions: Results suggest that broad‐leaved deciduous overstory species can limit the growth of feather mosses through their leaf litter and by implication affect the humus form, nutrient cycling, and understory composition of these forests.     

Effect of defoliation on tracheid production and the level of indole-3-acetic acid in Abies balsamea shoots

To simulate feeding by the spruce budworm ( Choristoneura fumiferana Clem.), the apical current-year shoots on 1-year-old branches in the uppermost whorl of 6-year-old balsam fir [ Abies balsamea (L.) Mill.] trees were either removed completely by debudding before the start of the growing season or defoliated 0, 50, 90 or 100% shortly after budbreak. Debudded branches were treated at the apical end with 0, 0.1 or 1.0 mg of indole-3-acetic acid (IAA) (g lanolin)⁻¹. Ninety % of the 1-year-old needles were also removed from some of the experimental branches. After ca 4 weeks of growth, the radial width of new xylem and the level of IAA were determined in the 1-year-old internode. The IAA content was measured by radioimmunoassay.
The removal or defoliation of current-year shoots inhibited tracheid production and decreased the IAA level. Exogenous IAA stimulated tracheid production and increased the IAA level in debudded branches. Current-year shoot defoliation also inhibited current-year shoot elongation. The inhibitory effect of current-year needle removal on all parameters generally increased with increasing intensity of defoliation. The removal of 1-year-old needles did not affect the IAA level or current-year shoot elongation, nor did it influence tracheid production in branches with current-year shoots. However, removal of 1-year-old needles inhibited tracheid production in debudded branches supplied with exogenous IAA. The results indicate that (1) IAA is involved in the control of tracheid production in the 1-year-old internode, (2) IAA is supplied primarily by current-year shoots, and (3) defoliation by the spruce budworm inhibits tracheid production partly by decreasing the supply of IAA.     

沙地土壤种植杨树-刺槐混交林后持水特性变化的研究

通过对比法研究了沙地土壤种杨树刺槐混交林后持水特性的变化。结果表明,沙地土壤种植杨树,刺槐纯林和混交林后,土壤持水特性,孔隙分布状况和水容量得到了改善和提高,提高和改善的幅度均以混交林最大,无林地植被覆盖后土壤持水性能改善的根本原因在于土壤有机质含量的增加,从而改善了土壤的孔隙状况。     

Tree growth and soil acidification in response to 30 years of experimental nitrogen loading on boreal forest

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

Effects of fire on regional evapotranspiration in the central Canadian boreal forest

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 (E_C) 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 E_C, and a younger and more deciduous forest. Well‐ and poorly drained areas had similar rates of evaporation from the canopy and soil, but E_C 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.