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.     

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.     

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.     

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.     

Characterization of tree macroremains production in a recently burned conifer forest in northern Québec,Canada

The production of plant macroremains was studied in a conifer forest twomonths after it burned in 1996 in northern Québec. The proportions ofvarious types of charred and uncharred pieces (needles, cones andwood) produced by black spruce (Piceamariana) and jack pine (Pinusbanksiana) were determined by sampling around individual trees.Both species produced equivalent masses of charred material, but pieces of woodcharcoal from jack pine are generally larger that those of black spruce. Theproportion of charred versus uncharred needles is the bestindicator of the species dominance in the forest. Although the fall of uncharredneedles is delayed from the time of a fire, they contribute to more than half ofall remains produced. Jack pine cones remain on the tree for a long time after afire, while charred cones of black spruce are dehiscent (cones come off thebranches easily). Trees are poor wood charcoal producers compared toundergrowth shrubs. As a result, the macroremains assemblage associated with afire event is made up of large amount of uncharred material from trees and alarge proportion of charred pieces produced by undergrowth vegetation. Modernassemblages of plant macroremains indicate that in order to reconstruct pastvegetation associated with fire disturbance, it is important to distinguishbetween the various types of remains, because wood charcoal is mainly producedby material that is already dead.     

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.     

Responses of Pinus resinosa in Newfoundland to wildfire

Abstract. Natural Pinus resinosa (red pine) stands in Newfoundland are restricted to 22 small, dry, nutrient-poor sites. A short wildfire cycle (15 - 30 yr) of both surface and crown fire regulates stand perimeters and is the main factor in regulating stand development. At the nucleus of current stands < 100 yr old, a few trees > 200 yr occur, usually showing multiple fire scars. Stem char heights confirmed an increased flammability of the stand and tree mortality for fires moving in the upslope position, as well as for mixed Pinus resinosa-Picea mariana stands. All P. resinosa stands are severely nutrient-deficient. Leaf concentrations of N, P and K were below or near the reported critical values. Nutrient concentrations were highest three months after a surface fire, but dropped considerably one year later. A gradual increase to near post-fire levels is achieved four years after fire. Foliar nutrient concentrations were positively correlated with average width of the annual rings. Aspects of the ericaceous understory dynamics and its relation to P. resinosa regeneration are also discussed.     

A two thousand year history of a northern Swedish boreal forest stand

Abstract. Successional processes within northern Swedish boreal forest are investigated for the last 2000 years by analysis of pollen, charcoal fragments and insect remains preserved in a deep mor humus layer on a small island in a large lake. Frequent disturbances by fire, blow-down, insect attack and flood have been the chief influence on vegetation dynamics. The gradual climatically induced replacement of Betula pubescens by Picea abies is superimposed on the disturbance dynamics. Picea first became established during the Little Ice Age, which was the longest period without fire in the record. A thin layer of volcanic ash from Iceland confirms the chronology at the onset of the Little Ice Age. The present importance of Pinus sylvestris and Ericaceae on the island is a product of a recent increase in disturbance frequency. The disturbance regime ensures that the communities stay in equilibrium with prevailing climate.     

Revegetation following experimental disturbance in a boreal old-growth Picea abies forest

Abstract. We studied revegetation patterns after experimental fine-scale disturbance (e.g. uprooting) in an old-growth Picea abies forest in southeastern Norway. An experimental severity gradient was established by manipulation of the depth of soil disturbance; two types of disturbed areas were used. Species recovery was recorded in the disturbed patches in three successive years after disturbance. The cover of vascular plants and, even more so the cover of bryophytes and lichens, recovered slowly after disturbance. The least severe treatments (removal of vegetation and removal of vegetation and the litter layer) was followed by the fastest recovery. The mean number of vascular plant species was usually higher three years after disturbance than before disturbance, while the opposite was true for bryophytes. Several vascular plant species that were abundant in intact forest floor vegetation (Vaccinium myrtillus, V. vitis-idaea and Deschampsia flexuosa) recovered during a three-year period primarily by resprouting from intact rhizomes and clonal in-growth. Other important recovery mechanisms included germination from a soil-buried propagule bank (e.g. Luzula pilosa, Plagiothecium laetum agg., Pohlia nutans and Polytrichum spp.) and dispersal of propagules into the disturbed patches (e.g. Betula pubescens and Picea abies). Microsite limitation seemed to occur in several species that were abundant in the soil propagule bank (e.g. the ferns Athyrium filix-femina, Gymnocarpium dryopteris and Phegopteris connectilis) but which did not appear in disturbed patches. Disturbance severity influenced revegetation patterns, recorded both as trajectories of vegetation composition in a DCA ordination space and as change in floristic dissimilarity. The length of the successional path (compositional change measured in β-diversity units) increased with increasing disturbance severity, and was also influenced by the area of the disturbed patch and the distance to intact vegetation. The rate of succession depended on the method by which it was measured; decreasing year by year in floristic space, while first decreasing and then increasing in ordination space. The reason for this difference is explained.     

Vegetation changes caused by recent fires in the northern boreal forest of eastern Canada

Abstract. From 1980–1989, fires burned 32 440 km² of boreal forest, 200 km south of the forest-tundra border in northern Québec, Canada. An assessment of the impact of fire on tree population densities was carried out by comparing the number of Pinus banksiana and Picea mariana in 83 sites before and after the sites burned in 1981, 1983, 1988 or 1989. Age structure analysis of post-fire populations burned in 1972, 1976 and 1983, along with the rapid exhaustion of the seed bank from burned trees, suggest that the majority of seedlings were established within 3 to 10 yr after fire. Consequently, given the absence of nearby living seed bearers, little (if any) further recruitment can be expected in the even-aged, regenerating populations. According to the tree density comparison (pre-fire vs post-fire), a shift from Picea- to Pinus-dominated communities occurred in most of the sites burned in 1981 or 1983, and in some of the sites burned in 1988 or 1989. The 1988 fire reduced the tree population density by 95% in 10 of the 15 sites; total tree density decreased by at least 75% in 28 out of 40 sites burned in 1989. This suggests that the areas burned in 1988 and 1989 will mainly regenerate as very open forests or lichen-heath communities that are more commonly found in the forest-tundra zone, north of the study area. Fire intensity, short fire interval, and unfavorable climate during and after fires are three plausible mechanisms associated with these post-fire vegetation changes.     

Effects of preconditioning on electrolyte leakasge and lipid composition in black spruce (Picea mariand) stressed with polyethylene giycol

Black spruce ( Picea mariana Mill. B. S. P.) rooted cuttings were grown in solution culture and preconditioned by osmotically stressing plants with polyethylene glycol. After relief from preconditioning stress, water relations, membrane leakiness, and the composition of lipids and fatty acids were compared in preconditioned and control, unconditioned plants. Both groups of plants were subsequently subjected to a severe osmotic stress with polyethylene glycol and examined again. Osmotic stress decreased shoot water potentials and increased the leakage of electrolytes from shoots of stressed, compared with unstressed, plants. However, both unstressed and stressed preconditioned plants leaked less electrolytes compared with unconditioned plants. Changes in sterol, phospholipid and glycolipid composition were observed in preconditioned unstressed and stressed plants. Sterol and phospholipid levels de clined as a result of stress, and preconditioning resulted in a decline in sterol: phospholipid ratios in plants.     

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.     

Effect of increased Populus cover on Abies regeneration in the Picea–feathermoss boreal forest

Question: Does the increase in Populus tremuloides cover within the Picea mariana–feathermoss domain enhance establishment and growth conditions for Abies balsamea regeneration? Location: Boreal forest of northwest Quebec, Canada. Method: To document the effect of Populus tremuloides on A. balsamea regeneration, mixed stands with a heterogeneous presence of P. tremuloides adjacent to Picea mariana‐dominated stands were selected. Abies balsamea regeneration, understorey environment and canopy composition were characterized from 531 sampling units distributed along transects covering the mixed–coniferous gradient. Abundance of understorey A. balsamea regeneration was described using three height groups: seedling (<30 cm), small sapling (30 to <100 cm) and tall sapling (100 to 300 cm). Growth characteristics were measured from 251 selected individuals of A. balsamea (<3 m). Results: Results showed that A. balsamea regeneration was generally more abundant when P. tremuloides was present in the canopy. Differences between seedling and sapling abundance along the mixed–coniferous gradient suggest that while establishment probably occurs over a wide range of substrates, the better growth conditions found under mixed stands ensure a higher survival rate for A. balsamea seedlings. Conclusions: The abundant A. balsamea regeneration observed within mixed stands of the Picea mariana–feathermoss domain suggests that the increase in P. tremuloides cover, favoured by intensive management practices and climatic change, could contribute to acceleration of the northward expansion of the A. balsamea–Betula papyrifera domain into the northern boreal forest dominated by Picea mariana.     

Relationships between change in fire frequency and mortality due to spruce budworm outbreak in the southeastern Canadian boreal forest

Abstract. We present a simple empirical model that allows an estimation of mortality due to spruce budworm (Choristoneura fumiferana) outbreak in relation to fire frequency and site characteristics. The occurrence of a recent spruce budworm outbreak around Lake Duparquet (48° 30’N, 79° 20’W, ca. 300 m a.s.l.) in northwestern Québec permitted a reconstruction of the stand composition before the outbreak, and also of the mortality of Abies balsamea due to the outbreak. The basal area of A. balsamea increases with time since fire in all site types but with increasing values for (1) rock and shallow till, via (2) till and mesic clay up to (3) hydric clay. Mortality (measured as percentage loss of basal area due to the outbreak) increases with time since fire but did not vary with site type. The increasing abundance of A. balsamea with time since fire is mainly responsible for this increase in mortality. Mortality for a specific basal area is, however, lower for the more recently burned stands suggesting a significant residual effect of time since fire. A landscape model integrating mortality due to the outbreak for stands of different age is developed. Both absolute and relative losses of basal area increased with the length of the fire cycles. According to this model, changes in fire cycle could explain a large portion of the spatio-temporal variations observed in outbreak mortality in the southeastern boreal forest of Canada.     

Contrasting patterns of tree mortality in late‐successional Picea abies stands in two areas in northern Fennoscandia

Question: What are tree mortality rates and how and why do they vary in late‐successional Picea abies‐dominated forests? Do observed tree mortality patterns allow comparative assessment of models of long‐term stand development? Location: Northern boreal Fennoscandia. Methods: We measured stand structure in 10 stands in two different areas. We determined age distributions and constructed a chronology of tree deaths by cross‐dating the years of death of randomly sampled dead trees. Results: The stands in the two areas had contrasting tree age distributions, despite similar live tree structure. In one area, stands were relatively even‐aged and originated following a stand‐replacing fire 317 years earlier. The stands in the second area had an uneven age structure and virtually no signs of past fires, suggesting a very long period since the last major disturbance. The younger stands were characterized by a high mortality rate and inter‐annual variation, which we attributed to senescence of the relatively even‐aged stands approaching the maximum age of P. abies. In contrast, the tree mortality rates in the older stands were low and relatively stable. Conclusions: Patterns of tree mortality were, to a large extent, dependent on the time since the last stand‐replacing disturbance, suggesting that northern boreal P. abies stands eventually reach a shifting mosaic state maintained through small‐scale dynamics, but the time needed to reach this state appears to be lengthy; even 300 years after a forest fire stands showed changes in patterns of tree mortality that were related to the developmental stage of the stands.     

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

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

Tree species dynamics and disturbance in three Swedish boreal forest stands during the last two thousand years

Abstract. Data from three forest stands for the past 2000 yr show how the shade-intolerant species Pinus sylvestris and Betula pubescens maintain significant populations in the Swedish boreal landscape. Age structure data from a northern stand close to the range limits of Picea abies and Pinus complement a local pollen diagram, and reveal cyclic population fluctuations which can be related to periods of climatic stress and fire. Pollen data from two southern stands show that high fire frequencies in the past prevented the expansion of Picea populations. Reduction of the fire frequency during the last 200 yr has favoured Picea. A long time perspective reveals the population dynamics of long-lived species and indicates the controlling factors. Such knowledge permits assessment of the current status and likely future of forest stands.