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

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

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

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

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

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

Effects of fire severity on plant nutrient uptake reinforce alternate pathways of succession in boreal forests

Fire activity in the North American boreal region is projected to increase under a warming climate and trigger changes in vegetation composition. In black spruce forests of interior Alaska, fire severity impacts residual organic layer depth which is strongly linked to the relative dominance of deciduous versus coniferous trees in early succession. These alternate successional pathways may be reinforced by biogeochemical processes that affect the relative ability of deciduous versus coniferous trees to acquire limiting nutrients. To test this hypothesis, we examined changes in soil inorganic nitrogen (N) supply and in situ ¹⁵N root uptake by aspen (Populus tremuloides) and black spruce (Picea mariana) saplings regenerating in lightly and severely burned sites, 16 years following fire. Fire severity did not impact the composition or magnitude of N supply, and nitrate represented nearly 40 % of total N supply. Both aspen and spruce took up more N in severely burned than in lightly burned sites. Spruce exhibited only a moderately lower rate of nitrate uptake, and a higher ammonium uptake rate than aspen in severely burned sites. At the stand level, differences in species nutrient uptake were magnified, with aspen taking up nearly an order-of-magnitude more N per m² in severely burned than in lightly burned sites. We suggest that differences in nutrient sinks (biomass) established early in succession and effects of post-fire organic layer depth on nutrient uptake, are key mechanisms reinforcing the opposing stand dominance patterns that have developed in response to variations in organic layer depth.     

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

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

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.     

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.     

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

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

黑龙江省大兴安岭林区火烧迹地森林更新及其影响因子

林火干扰是大兴安岭森林更新的影响因子之一,研究火烧迹地森林更新的影响因子(立地条件、火前植被、火干扰特征)对理解生态系统的结构、功能和火后演替轨迹具有重要意义。选取呼中及新林林业局55个代表性火烧样地,利用增强回归树分析法分析了火烧迹地森林更新的影响因素。结果表明:(1)立地条件是影响针、阔叶树更新苗密度的主要因素;海拔对针叶树更新苗密度的影响最大;坡度对阔叶树更新苗密度影响最大;(2)距上次火烧时间对针叶树更新苗比重影响最大,其次是林型;(3)中度林火干扰后森林更新状况好于轻度和重度火烧迹地。根据火烧迹地森林更新调查分析可知:林型影响火后演替模式,火前为针叶树或阔叶树纯林,火后易发生自我更新(火后树种更新组成与火前林型相同),而针阔混交林在火干扰影响下易于发生序列演替(火后初期以早期演替树种更新为主)。     

Carbon and nitrogen cycling in North American boreal forests

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

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.     

Ecosystem controls on nitrogen fixation in boreal feather moss communities

N fixation in feather moss carpets is maximized in late secondary successional boreal forests; however, there is limited understanding of the ecosystem factors that drive cyanobacterial N fixation in feather mosses with successional stage. We conducted a reciprocal transplant experiment to assess factors in both early and late succession that control N fixation in feather moss carpets dominated by Pleurozium schreberi. In 2003, intact microplots of moss carpets (30 cm × 30 cm × 10–20 cm deep) were excavated from three early secondary successional (41–101 years since last fire) forest sites and either replanted within the same stand or transplanted into one of three late successional (241–356 years since last fire) forest sites and the transverse was done for late successional layers of moss. Moss plots were monitored for changes in N-fixation rates by acetylene reduction (June 2003–September 2005) and changes in the presence of cyanobacteria on moss shoots by microscopy (2004). Forest nutrient status was measured using ionic resin capsules buried in the humus layer. Late successional forests exhibit high rates of N fixation and consistently high numbers of cyanobacteria on moss shoots, but low levels of available N. Conversely, early successional forests have higher N availability and have low rates of N fixation and limited presence of cyanobacteria on moss shoots. Transplantation of moss carpets resulted in a significant shift in presence and activity of cyanobacteria 1 year after initiation of the experiment responding to N fertility differences in early versus late successional forests.     

Interactive Effects of Fire, Soil Climate, and Moss on CO2 Fluxes in Black Spruce Ecosystems of Interior Alaska

Fire is an important control on the carbon (C) balance of the boreal forest region. Here, we present findings from two complementary studies that examine how fire modifies soil organic matter properties, and how these modifications influence rates of decomposition and C exchange in black spruce (Picea mariana) ecosystems of interior Alaska. First, we used laboratory incubations to explore soil temperature, moisture, and vegetation effects on CO₂ and DOC production rates in burned and unburned soils from three study regions in interior Alaska. Second, at one of the study regions used in the incubation experiments, we conducted intensive field measurements of net ecosystem exchange (NEE) and ecosystem respiration (ER) across an unreplicated factorial design of burning (2 year post-fire versus unburned sites) and drainage class (upland forest versus peatland sites). Our laboratory study showed that burning reduced the sensitivity of decomposition to increased temperature, most likely by inducing moisture or substrate quality limitations on decomposition rates. Burning also reduced the decomposability of Sphagnum-derived organic matter, increased the hydrophobicity of feather moss-derived organic matter, and increased the ratio of dissolved organic carbon (DOC) to total dissolved nitrogen (TDN) in both the upland and peatland sites. At the ecosystem scale, our field measurements indicate that the surface organic soil was generally wetter in burned than in unburned sites, whereas soil temperature was not different between the burned and unburned sites. Analysis of variance results showed that ER varied with soil drainage class but not by burn status, averaging 0.9 ± 0.1 and 1.4 ± 0.1 g C m⁻² d⁻¹ in the upland and peatland sites, respectively. However, a more complex general linear model showed that ER was controlled by an interaction between soil temperature, moisture, and burn status, and in general was less variable over time in the burned than in the unburned sites. Together, findings from these studies across different spatial scales suggest that although fire can create some soil climate conditions more conducive to rapid decomposition, rates of C release from soils may be constrained following fire by changes in moisture and/or substrate quality that impede rates of decomposition. Author contributions: JAO: performed research, analyzed data, contributed new methods, wrote the paper; MRT: designed laboratory study, performed research, analyzed data; JWH: designed field study, performed research; KLM: performed research; LEP: performed research, contributed new method; GS: performed research; JCN: performed research.     

Early successional pathways in the Tatra Mountains (Slovakia) forest ecosystems following natural disturbances

Large scale windstorms disturbed forest ecosystems in the Tatra Mountains in 2004, and were followed by a severe fire in 2005. A long-term study on the vegetation successional dynamics of the area was launched immediately after the 2005 event. Relevé plots were established under five different disturbance and management treatments: windthrow left, windthrow removed, hydrologically managed, burnt and reference. We used weighted Ellenberg’s indicator plant values for ordination analyses of the following environmental gradients: light, temperature, continentality, moisture, acidity, nitrogen. Successional patterns depended on the management treatment. Heavily burnt areas were colonized by plants disseminated by airborn diasporas, mainly by Chamaerion angustifolium, less burnt or unburnt localities were settled by plants germinating from the soil seed bank or by plants surviving by root system. Nitrophilous weed vegetation invaded plots with increased moisture (fallen or standing overstory vegetation or irrigated by man-made system) and nitrogen (burnt or windthrow removal). The felled-area species were recorded in each plot. Abundant natural regeneration was observed in plots with increased moisture. The moss layer disappeared soon after the disturbance events. The results presented here refer to a very early successional stage, new insights into initial successional paterns are gained.     

Amino acid uptake in deciduous and coniferous taiga ecosystems

We measured in situ uptake of amino acids and ammonium across deciduous and coniferous taiga forest ecosystems in interior Alaska to examine the idea that late successional (coniferous) forests rely more heavily on dissolved organic nitrogen (DON), than do early successional (deciduous) ecosystems. We traced ¹⁵N-NH₄⁺ and ¹³C-¹⁵N-amino acids from the soil solution into plant roots and soil pools over a 24 h period in stands of early successional willow and late successional black spruce. Late successional soils have much higher concentrations of amino acid in soil solution and a greater ratio of DON to dissolved inorganic N (DIN) (ammonium plus nitrate) than do early successional soils. Moreover, late successional coniferous forests exhibit higher rates of soil proteolytic activity, but lower rates of inorganic N turnover. Differences in ammonium and amino acid uptake by early successional willow stands were insignificant. By contrast, the in situ uptake of amino acid by late successional black spruce forests were approximately 4-fold greater than ammonium uptake. The relative difference in uptake of ammonium and amino acids in these forests was approximately proportional to the relative difference of these N forms in the soil solution. Thus, we suggest that differences in uptake of different N forms across succession in these boreal forests largely reflect edaphic variation in available soil N (composition), rather than any apparent physiological specialization to absorb particular forms of N. These finding are relevant to our understanding of how taiga ecosystems may respond to increases in temperature, fire frequency, N deposition, and other potential consequences of global change.     

林火干扰对北方针叶林林下植被的影响

林下植被在北方针叶林植被群落中的物种多样性最高, 且具有较高的生物量周转率和地上部分净初级生产力, 对北方针叶林生态系统功能起着重要作用。火干扰是决定北方针叶林林下植被结构与功能的一个重要景观过程。该文综述了火干扰是如何通过与地形、火前林冠组成的交互作用而影响环境资源和林下植被的。最近的研究表明: 林下植被能够影响火后树木更新苗的定植、重建速率及森林演替轨迹; 林下植被还会通过影响元素的生物地球化学过程(凋落物降解和养分循环)影响林下环境资源的数量与异质性。因此, 研究火后初期北方针叶林林下植被的动态变化, 对于物种多样性保护和森林管理具有重要意义。     

The Role of Balsam Poplar Secondary Chemicals in Controlling Soil Nutrient Dynamics through Succession in the Alaskan Taiga

The vegetation mosaic of the Alaskan taiga is produced by patterns of disturbance coupled to well-defined successional patterns. In primary succession on river floodplains, one of the critical transitions in succession is that from thinleaf alder (Alnus tenuifolia) to balsam poplar (Populus balsamifera). This is the shift from a N2-fixing shrub to a deciduous tree. Through this transition there are major changes in N cycling including a decrease in N2-fixation, mineralization, and nitrification. Most models of plant effects on soil processes assume that these changes are caused by shifts in litter quality and C/N ratio. This paper reviews several studies examining the effects of balsam poplar secondary chemicals on soil nutrient cycling. Balsam poplar tannins inhibited both N2-fixation in alder, and decomposition and N-mineralization in alder soils. Other poplar compounds, including low-molecular-weight phenolics, were microbial substrates and increased microbial growth and immobilization, thereby reducing net soil N availability. Thus, substantial changes in soil N cycling through succession appear to have been mediated by balsam poplar secondary chemicals.     

Understory Vegetation Dynamics of North American Boreal Forests

Understory vegetation is the most diverse and least understood component of North American boreal forests. Understory communities are important as they act as drivers of overstory succession and nutrient cycling. The objective of this review was to examine how understory vegetation abundance, composition, and diversity change with stand development after a major stand replacing disturbance. Understory vegetation abundance and diversity increase rapidly after fire, in response to abundant resources and an influx of disturbance adapted species. The highest diversity occurs within the first 40 years following fire, and declines indefinitely thereafter as a result of decreasing productivity and increased dominance of a small number of late successional feather mosses and woody plant species. Vascular plant and bryophyte/lichen communities undergo very different successional changes. Vascular plant communities are dynamic and change more dramatically with time after fire, whereas bryophyte and lichen communities are much slower to establish and change over time. Considerable variations in these processes exist depending on canopy composition, site condition, regional climate, and frequently occurring non-stand-replacing disturbances. Forest management practices represent a unique disturbance process and can result in different understory vegetation communities from those observed for natural processes, with potential implications for overstory succession and long-term productivity. Because of the importance of understory vegetation on nutrient cycling and overstory composition, post-harvest treatments emulating stand-replacing fire are required to maintain understory diversity, composition, and promote stand productivity in boreal forests.     

Secondary succession in coastal dune fynbos: variation due to site and disturbance

Succession after fire and bushcutting in coastal dune fynbos was monitored for two and a half years and comparisons were made with adjacent, mature (13 year-old) fynbos. Sixty-two to 68% of pre-disturbance species, including all the dominants, were found in the successional communities 1.5 yr after disturbance: the patterns thus fitted the initial floristic composition model. On the more mesic south facing slope, post-fire succession differed from the north-facing (burnt) and bushcut sites in that ordinations showed a clear separation between the mature and successional communities. This difference was due to the post-fire abundance on the southfacing slope site of short-and medium-lived species not present in the mature fynbos. One and a half years after disturbance, species richness and equitability had increased relative to mature vegetation. This increase was greatest for the south facing slope where short-and medium-lived species and juveniles of pre-disturbance dominants co-occurred. In general, successional patterns were consistent with those described for other fynbos and fire-prone mediterranean shrublands.