Multiple successional pathways of boreal forest stands in central Canada

Predicting forest composition change through time is a key challenge in forest management. While multiple successional pathways are theorized for boreal forests, empirical evidence is lacking, largely because succession has been inferred from chronosequence and dendrochronological methods. We tested the hypotheses that stands of compositionally similar overstory may follow multiple successional pathways depending on time since last stand‐replacing fire (TSF), edaphic conditions, and presence of intermediate disturbances. We used repeated measurements from combining sequential aerial photography and ground surveys for 361 boreal stands in central Canada. Stands were measured in 8–15 yr intervals over a ~ 60 yr period, covering a wide range of initial stand conditions. Multinomial logistic regression was used to analyze stand type transitions. With increasing TSF, stands dominated by shade‐intolerant Pinus banksiana, Populus sp., and Betula papyrifera demonstrated multiple pathways to stands dominated by shade‐tolerant Picea sp., Abies balsamea, and Thuja occidentalis. Their pathways seemed largely explained by neighborhood effects. Succession of stands dominated by shade‐tolerant species, with an exception of stands dominated by Picea sp., was not related to TSF, but rather dependent on edaphic conditions and presence of intermediate disturbances. Varying edaphic conditions caused divergent pathways with resource limited sites being dominated by nutrient‐poor tolerant species, and richer sites permitting invasion of early successional species and promoting species mixtures during succession. Intermediate disturbances promoted deciduous persistence and species diversity in A. balsamea and mixed‐conifer stands, but no evidence was detected to support “disturbance accelerated succession”. Our results demonstrate that in the prolonged absence of stand‐replacing disturbance boreal forest stands undergo multiple succession pathways. These pathways are regulated by neighborhood effects, resource availability, and presence of intermediate disturbance, but the relative importance of these regulators depends on initial stand type. The observed divergence of successional pathways supports the resource‐ratio hypothesis of plant succession.     

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.     

Stability in the patterns of long‐term development and growth of the Canadian spruce–moss forest

Aim The spruce–moss forest is the main forest ecosystem of the North American boreal forest. We used stand structure and fire data to examine the long‐term development and growth of the spruce–moss ecosystem. We evaluate the stability of the forest with time and the conditions needed for the continuing regeneration, growth and re‐establishment of black spruce (Picea mariana) trees. Location The study area occurs in Québec, Canada, and extends from 70°00′ to 72°00′ W and 47°30′ to 56°00′ N. Methods A spatial inventory of spruce–moss forest stands was performed along 34 transects. Nineteen spruce–moss forests were selected. A 500 m² quadrat at each site was used for radiocarbon and tree‐ring dating of time since last fire (TSLF). Size structure and tree regeneration in each stand were described based on diameter distribution of the dominant and co‐dominant tree species [black spruce and balsam fir (Abies balsamea)]. Results The TSLF of the studied forests ranges from 118 to 4870 cal. yr bp . Forests < 325 cal. yr bp are dominated by trees of the first post‐fire cohort and are not yet at equilibrium, whereas older forests show a reverse‐J diameter distribution typical of mature, old‐growth stands. The younger forests display faster height and radial growth‐rate patterns than the older forests, due to factors associated with long‐term forest development. Each of the stands examined established after severe fires that consumed all the soil organic material. Main conclusions Spruce–moss forests are able to self‐regenerate after fires that consume the organic layer, thus allowing seed regeneration at the soil surface. In the absence of fire the forests can remain in an equilibrium state. Once the forests mature, tree productivity eventually levels off and becomes stable. Further proof of the enduring stability of these forests, in between fire periods, lies in the ages of the stands. Stands with a TSLF of 325–4870 cal. yr bp all exhibited the same stand structure, tree growth rates and species characteristics. In the absence of fire, the spruce–moss forests are able to maintain themselves for thousands of years with no apparent degradation or change in forest type.     

Succession in the southern part of the Canadian boreal forest

Forest succession following fire in a forest mosaic of northwestern Quebec has been studied in order to: (1) describe the successional pathways using communities of different ages and (2) evaluate convergence of successional pathways and possible effect of fire suppression on the establishment of steady-state communities. As a first step, ordination and classification techniques were used in order to remove changes in forest composition which are related to abiotic conditions. Then, ordinations based on tree diameter distributions were used to study shifts in species composition in relation to time since the last fire.Even under similar abiotic conditions, successional pathways are numerous. However, regardless of forest composition after fire, most stands show convergence toward dominance of Thuja occidentalis and Picea mariana on xeric sites and dominance of Abies balsamea and Thuja occidentalis on more mesic sites. Stable communities of >300 yr occur on xeric sites while on mesic sites directional succession still occurs after 224 yr. Nearly all species involved in succession are present in the first 50 yr following fire. Only Abies balsamea and Thuja occidentalis increase significantly in frequency during succession. Following initial establishment, successional processes can generally be explained by species longevity and shade tolerance. Early successional species may be abundant in the canopy for more than 200 yr while the rapid decrease of Picea glauca, a late successional species could be related to spruce budworm outbreaks. Considering the short fire rotation observed (about 150 yr), a steady-state forest is unlikely to occur under natural conditions, though it may be possible if fire is controlled.     

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.     

High and dry: post‐fire tree seedling establishment in subalpine forests decreases with post‐fire drought and large stand‐replacing burn patches

Aim Climate warming and increased wildfire activity are hypothesized to catalyse biogeographical shifts, reducing the resilience of fire‐prone forests world‐wide. Two key mechanisms underpinning hypotheses are: (1) reduced seed availability in large stand‐replacing burn patches, and (2) reduced seedling establishment/survival after post‐fire drought. We tested for regional evidence consistent with these mechanisms in an extensive fire‐prone forest biome by assessing post‐fire tree seedling establishment, a key indicator of forest resilience. Location Subalpine forests, US Rocky Mountains. Methods We analysed post‐fire tree seedling establishment from 184 field plots where stand‐replacing forest fires were followed by varying post‐fire climate conditions. Generalized linear mixed models tested how establishment rates varied with post‐fire drought severity and distance to seed source (among other relevant factors) for tree species with contrasting post‐fire regeneration adaptations. Results Total post‐fire tree seedling establishment (all species combined) declined sharply with greater post‐fire drought severity and with greater distance to seed sources (i.e. the interior of burn patches). Effects varied among key species groups. For conifers that dominate present‐day subalpine forests (Picea engelmannii, Abies lasiocarpa), post‐fire seedling establishment declined sharply with both factors. One exception was serotinous Pinus contorta, which did not vary with either factor. For montane species expected to move upslope under future climate change (Larix occidentalis, Pseudotsuga menziesii, Populus tremuloides) and upper treeline species (Pinus albicaulis), establishment was unrelated to either factor. Greater post‐fire tree seedling establishment on cooler/wetter aspects suggested local topographic refugia during post‐fire droughts. Main conclusions If future drought and wildfire patterns manifest as expected, post‐fire tree seedling establishment of species that currently characterize subalpine forests could be substantially reduced. Compensatory increases from lower montane and upper treeline species may partially offset these reductions, but our data suggest important near‐ to mid‐term shifts in the composition and structure of high‐elevation forests under continued climate warming and increased wildfire activity.     

Growth trends and dynamics in sub‐alpine forest stands in the Varaita Valley (Piedmont,Italy) and their relationships with human activities and global change

Abstract. A study of the forest lines, tree lines and the structures of the sub‐alpine forest was performed in Vallone Vallanta and in Alevé forest in the Varaita Valley (Cottian Alps, Piedmont, Italy). Forest‐ and tree lines were analysed over 1728 ha while forest structures were studied on six 3000‐m² plots located at the tree line (2), at the forest line (2) and inside the sub‐alpine forest (2). Dendro‐ecological analysis of living plants and stumps showed that Larix decidua was more abundant in the past than today and that Pinus cembra has expanded, both upwards and within sub‐alpine forests. Age structure analysis revealed that the current sub‐alpine forest stands were established 200–220 yr ago, probably following a clearcut. At the forest lines the tree density decreases, and some trees are more than 500 yr old, whereas at the tree lines most of the trees (almost exclusively Pinus cembra) are younger than 100 yr. Growth dynamics were investigated both by observing Basal Area Increment (BAI) in the old and dominant trees, and by comparing the BAIs of classes of trees with a given cambial age range in different time periods. The results showed that the growth rates of mature Pinus cembra and Larix decidua had increased. These increments are more substantial for Pinus than for Larix. The growth rate of young trees (< 100 yr) of both species has decreased over recent decades. This could be due to competition caused by increased tree densities that have resulted from a decrease in grazing.     

Fire Interval Effects on Successional Trajectory in Boreal Forests of Northwest Canada

Although succession may follow multiple pathways in a given environment, the causes of such variation are often elusive. This paper describes how changes in fire interval mediate successional trajectory in conifer-dominated boreal forests of northwestern Canada. Tree densities were measured 5 and 19 years after fire in permanent plots and related to pre-fire vegetation, site and fire characteristics. In stands that were greater than 75 years of age when they burned, recruitment density of conifers was significantly correlated with pre-fire species basal area, supporting the expectation of stand self-replacement as the most common successional pathway in these forests. In contrast, stands that were under 25 years of age at the time of burning had significantly reduced conifer recruitment, but showed no change in recruitment of trembling aspen (Populus tremuloides). As a result, young-burned stands had a much higher probability of regenerating to deciduous dominance than mature-burned stands, despite the dominance of both groups by spruce (Picea mariana and Picea glauca) and pine (Pinus contorta) before the fire. Once initiated, deciduous-dominated stands may be maintained across subsequent fire cycles through mechanisms such as low on-site availability of conifer seed, competition with the aspen canopy, and rapid asexual regeneration of aspen after fire. We suggest that climate-related increases in fire frequency could trigger more frequent shifts from conifer to deciduous-dominated successional trajectories in the future, with consequent effects on multiple ecosystem processes.     

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.     

A test of ecological succession hypotheses using 55‐year time‐series data for 361 boreal forest stands

Aim There has been much work on succession over many decades, but succession fundamentals are still debated because of the reliance on chronosequences and dendrochronological reconstruction, both of which are problematic approaches. Here we use time‐series data to test four hypotheses that lie at the heart of successional theory: (1) the neighbourhood effect hypothesis – tree species abundance is time dependent; (2) the density‐dependence hypothesis – a rare species is more favoured over time; (3) the resource ratio hypothesis – species that can grow at the lowest resource level tend to dominate resource limited sites through succession; and (4) the intermediate disturbance hypothesis – intermediate disturbances increase the abundance of rare species. Location Central boreal forest in Canada (47°50′–50°10′ N; 80°10′–85°50′ W). Methods We used repeated measurements from sequential aerial photography and ground surveys for 361 fire‐origin stands that were measured over a c. 55‐year period. Results Shade‐intolerant Pinus banksiana decreased, tolerant Thuja occidentalis increased, intolerant Populus spp. and Betula papyrifera displayed a U‐shaped trend, and intermediate‐tolerant Picea spp. and tolerant Abies balsamea did not change with time since fire, showing evidence of negative, positive, or neutral neighbourhood effects. Species either persisted for longer or increased more in non‐conspecific stands, and had higher increases in abundance when associated with species of contrasting shade tolerance, supporting the density‐dependence hypothesis and indicating shade‐tolerance complementarity as a mechanism for coexistence. Resource‐poor soils favoured those species capable of tolerating limited resources, whereas rich soils permitted invasion and promoted mixtures, supporting the resource ratio hypothesis. Intermediate disturbances increased the invasibility of rare species. Main conclusions Contrary to previous studies where time since a stand‐replacing disturbance is the sole predictor for succession, our study shows that time alone is either an insufficient predictor or is irrelevant to species dynamics in the boreal forest. Rather, density dependence, site resource and intermediate disturbances are key mechanisms in species dynamics and coexistence over time.     

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.     

Posthurricane Seedling Structure in a Multi‐aged Tropical Dry Forest: Implications for Community Succession

Hurricane‐caused tree mortality in tropical dry forests occurs predominantly in early successional species. Consequently, hurricanes may accelerate succession in these forests. Forest regeneration, however, must be measured over an extended posthurricane time period to demonstrate this pattern. In this study, we recorded tree seedlings in 19 Florida Keys forests during May–August 1995, 3 years after Hurricane Andrew. For these forests—spanning a chronosequence from 14 to over 100 years since the most recent clearing—we used weighted averaging regression on relative abundances of pre‐hurricane trees to calculate a successional age optimum for each species; and used weighted averaging calibration to calculate inferred successional ages for stands based on pre‐hurricane trees and on posthurricane seedlings. To test the hypothesis that successional stage of seedlings exceeded successional stage of pre‐hurricane trees, we compared inferred stand ages based on posthurricane seedlings with those based on pre‐hurricane trees. Across the study area, inferred stand ages based on posthurricane seedlings were greater than those based on pre‐hurricane trees (P < 0.005); however, more seedlings in the youngest stands were early successional than in older stands. Of 29 species present both as pre‐hurricane trees and posthurricane seedlings, 23 had animal‐dispersed seeds. These results provide evidence that: (1) hurricanes do not ‘reset’ succession, and may accelerate succession; and (2) a strong legacy of stand successional age influences seedling assemblages in these forests.     

Changes in two Minnesota forests during 14 years following catastrophic windthrow

Abstract. We measured tree damage and mortality following a catastrophic windthrow in permanent plots in an oak forest and a pine forest in central Minnesota. We monitored changes in forest structure and composition over the next 14 years. Prior to the storm, the oak forest was dominated by Quercus ellipsoidalis, and the pine forest by Pinus strobus. The immediate impacts of the storm were to differentially damage and kill large, early‐successional hardwoods and pines. Subsequent recovery was characterized by the growth of late‐successional hardwoods. In both forests the disturbance acted to accelerate succession. Ordination of tree species composition confirmed the trend of accelerated succession, and suggested a convergence of composition between the two forests.     

Canopy and age structures of some old sub-boreal Picea stands in British Columbia

Abstract. 14 old, unlogged, Picea-dominated stands in the moist cool Sub-Boreal Spruce biogeoclimatic subzone of central British Columbia, Canada, were sampled to describe canopy heterogeneity, regeneration patterns and tree population age structures. These stands are composed of Picea engelmannii × glauca hybrids, Abies lasiocarpa and lesser amounts of Pinus contorta and Populus tremuloides, and had survived 124–343 yr since the last stand-destroying wildfire. Canopy cover was patchy and highly variable (ranging from 30.5 % to 86.4 %) but was not significantly related to stand age. Vertical canopy structure was less variable, reflecting the shade-tolerance and live crown ratios (length of live canopy expressed relative to tree height) of component species: 18.8 % for Populus, 20.2 % for Pinus, 46.7 % for Picea and 51.4 % for Abies. Individual stands varied considerably in their population structures and in their stand development trajectories, yet some patterns are evident. Survivors of the initial post-disturbance cohort of trees took 51 to 118 yr (mean = 80, s.d. = 20) to establish. Some stands had all tree species present during stand initiation, while other stands indicated early successional roles for Populus and Pinus, or a late successional role for Abies. Abies recruitment, while often slow in the beginning, occurs uniformly throughout the history of most stands, reflecting the high shade-tolerance of this species. Picea is often recruited in high densities early in stand development, and then (after long periods of exclusion) may be displaced by Abies in some stands but maintains itself in others. Minor, single-tree disturbances (due to bark beetles, root rot, and windthrow) were important in accelerating the reinitiation of Picea in the understory. Results thus suggest that stands from this region can be self-perpetuating in the absence of fire. Yet, post-fire tree populations still clearly dominate these spruce-fir forests, for only the oldest stand had greater basal area in the replacement cohort than in the initial cohort.     

Understorey vegetation development in North Finnish Picea forests after disturbance: re‐analysis of Sirén's data

Abstract. Sirén (1955) studied understorey species composition, tree stand properties and humus‐layer thickness in 64 unlogged forest stands on topographically and pedologically comparable sites. The stands were of even age (6 – 300 yr), stocked with the first or second tree generation after wildfire. The view of Sirén and several authors after him, that the vegetation of old‐growth boreal Picea forests is homogeneous on a broad scale, was examined by applying, in parallel, the partial variants of two ordination methods (DCA and PCA) to Sirén's vegetation data. Two main vegetation gradients were found: a major gradient running from recently burnt plots with prominence of pioneer species to plots with stand age > 100 yr, a well stocked tree layer and a thick humus layer, dominance of feather‐mosses and ample occurrence of shade‐tolerant as well as light‐preferring vascular plant species, and a second gradient along which first‐ and second‐generation plots segregate. The more prominent element of Betula trees in first‐ than in second‐generation stands < 100 yr contributed to the latter. A minor third gradient related to humus‐layer thickness was recovered by partial DCA only. The main vegetation gradient reappeared in separate ordinations of data from 47 mature forest stands (> 100 yr), but without being correlated with forest age. Variation among mature‐forest stands in the importance of pioneer species is considered mainly to be brought about by fine‐scale disturbance processes such as tree uprooting. Increasing importance of factors operating on within‐stand scales [development of a varied gap structure and stronger gradients in tree influence (radiation at ground level), soil moisture, soil depth and nutrient availability] with time is also reflected in the second and third mature‐forest ordination axes. Possible implications of the results for conservation of biological diversity and monitoring of changes in boreal forests are discussed.     

Fire-induced shifts in overstory tree species composition and associated understory plant composition in Glacier National Park,Montana

In Rocky Mountain forests, fire can act as a mechanism of change in plant community composition if postfire conditions favor establishment of species other than those that dominated prefire tree communities. We sampled pre and postfire overstory and postfire understory species following recent (1988–2006) stand-replacing fires in Glacier National Park (GNP), Montana. We identified changes in relative density of tree species and groups of species (xerophytes vs. mesophytes and reseeders vs. resprouters) in early succession. Postfire tree seedling densities were adequate to maintain prefire forest structure, but relative densities among species were variously changed. Changes were directly related to individual species’ response to severe fires. Most notably, relative density of the mesophytic resprouter quaking aspen (Populus tremuloides) and the xerophytic reseeder lodgepole pine (Pinus contorta) increased substantially following fire, with a concomitant decline in proportional abundance of other tree species that, in some cases, dominated stands before fire. Trends identified in our study suggest that forest community shifts toward those dominated by lodgepole pine and quaking aspen are occurring in GNP. Cover of understory species was not affected by tree species composition or density. These forest communities will likely change throughout succession with the addition of shade-intolerant species in early seral stages and shade-tolerant species later in succession. However, with increased fire frequency, the lodgepole pine-dominated postfire communities observed in our study may become more common throughout time.     

Changes in woody vegetation abundance and diversity after natural disturbances causing different levels of mortality

Questions: How does woody vegetation abundance and diversity differ after natural disturbances causing different levels of mortality? Location: Abies balsamea–Betula papyrifera boreal mixed‐wood stands of southeast Quebec, Canada. Methods: Woody vegetation abundance and diversity were quantified and compared among three disturbance‐caused mortality classes, canopy gap, moderate‐severity disturbances, and catastrophic fire, using redundancy analysis, a constrained linear ordination technique, and diversity indices. Results: Substantial changes in canopy tree species abundance and diversity only occurred after catastrophic fire. Shade‐tolerant, late‐successional conifer species remained dominant after canopy gap and moderate‐severity disturbances, whereas shade‐intolerant, early‐successional colonizers dominated canopy tree regeneration after catastrophic fire. Density and diversity of mid‐tolerant and shade‐intolerant understory tree and shrub species increased as the impact of disturbance increased. Highest species richness estimates were observed after catastrophic fire, with several species establishing exclusively under these conditions. Relative abundance of canopy tree regeneration was most similar after canopy gap and moderate‐severity disturbances. For the sub‐canopy tree and shrub community, relative species abundances were most similar after moderate‐severity disturbances and catastrophic fire. Vegetation responses to moderate‐severity disturbances thus had commonalities with both extremes of the disturbance‐caused mortality gradient, but for different regeneration layers. Conclusions: Current spatio‐temporal parameters of natural disturbances causing varying degrees of mortality promote the development of a complex, multi‐cohort forest condition throughout the landscape. The projected increase in time intervals between catastrophic fires may lead to reduced diversity within the system.     

Decline in Net Ecosystem Productivity Following Canopy Transition to Late-Succession Forests

Boreal forests are critical to the global carbon (C) cycle. Despite recent advances in our understanding of boreal C budgets, C dynamics during compositional transition to late-succession forests remain unclear. Using a carefully replicated 203-year chronosequence, we examined long-term patterns of forest C stocks and net ecosystem productivity (NEP) following stand-replacing fire in the boreal forest of central Canada. We measured all C pools, including understorey vegetation, belowground biomass, and soil C, which are often missing from C budgets. We found a slight decrease in total ecosystem C stocks during early stand initiation, between 1 and 8 years after fire, at ?0.90 Mg C ha^?1 y^?1. As stands regenerated, live vegetation biomass increased rapidly, with total ecosystem C stocks reaching a maximum of 287.72 Mg C ha^?1 92 years after fire. Total ecosystem C mass then decreased in the 140- and 203-year-old stands, losing between ?0.50 and ?0.74 Mg C ha^?1 y^?1, contrasting with views that old-growth forests continue to maintain a positive C balance. The C decline corresponded with canopy transition from dominance of Populus tremuloides, Pinus banksiana, and Picea mariana in the 92-year-old stands to Betula papyrifera, Picea glauca, and Abies balsamea in the 203-year-old stands. Results from this study highlight the role of succession in long-term forest C dynamics and its importance when modeling terrestrial C flux.     

Fire Severity and Long-term Ecosystem Biomass Dynamics in Coniferous Boreal Forests of Eastern Canada

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.