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.     

Fire disturbance and forest structure in an old‐growth Pinus ponderosa forest,southern Cascades,USA

Questions: Did fire regimes in old‐growth Pinus ponderosa forest change with Euro‐American settlement compared to the pre‐settlement period? Do tree age structures exhibit a pattern of continuous regeneration or is regeneration episodic and related to fire disturbance or fire‐free periods? Are the forests compositionally stable? Do trees have a clumped spatial pattern and are clumps even‐ or mixed‐age? How might information from this old‐growth forest inform current restoration and management practices? Location: A 235‐ha old‐growth forest in the Ishi Wilderness, southern Cascade Mountains, California. Methods: Age, size, and spatial pattern of trees were quantified in seven stands. Fire history was reconstructed using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing fire history with age, size, and spatial structure of trees and identifying and measuring trees killed by two recent fires. Results: Species composition in plots was similar but density and basal area of tree populations varied. Age structure for P. ponderosa and Quercus kelloggii showed periods of episodic recruitment that varied among plots. Fire disturbance was frequent before 1905, with a median period between fires of 12 years. Fire frequency declined after 1905 but two recent fires (1990, 1994) killed 36% and 41% of mostly smaller diameter P. ponderosa and Q. kelloggii. Clusters of similar age trees occurred at scales of 28‐1018 m² but patches were not even‐aged. Interactions between tree regeneration and fire promoted development of uneven age groups of trees. Conclusions: Fire disturbance strongly influenced density, basal area, and spatial structure of tree populations. Fire exclusion over the last 100 years has caused compositional and structural changes. Two recent fires, however, thinned stands and created gaps favorable for Q. kelloggii and P. ponderosa regeneration. The effects of infrequent 20th century fire indicate that a low fire frequency can restore and sustain structural characteristics resembling those of the pre‐fire suppression period forest.     

Effects of Disturbance on Fine Root Dynamics in the Boreal Forests of Northern Ontario, Canada

Wildfire and clearcutting are two main disturbances in North American forests, but whether root systems may respond differently to such disturbances is unknown. Here, we studied how the dynamics of fine roots (<2 mm in diameter) varied with stand origins in a boreal forest in northern Ontario, Canada. Fine root biomass increased with stand age, but did not differ between stands originating from fire and clearcutting. By contrast, fine root production, mortality and turnover rates were lower in 3- and 11-year-old clearcut-origin than fire-origin stands, but did not differ in 29-year-old stands of different stand origins. The lower rates of production, mortality, and turnover rates in 3- and 11-year-old clearcut-origin than fire-origin stands are attributable to a lower density of shrubs and herbs and larger nutrient pools after clearcutting than fire. The similarities among 29-year-old stands indicate that the effects of stand origin on fine root processes tend to converge at this time scale. Our results illustrate that time scale is critical for assessing ecosystem responses to disturbances.     

Influence of Post‐Clearing Treatment on the Recovery of Herbaceous Plant Communities in Amazonian Secondary Forests

Secondary forests are an increasingly common feature in tropical landscapes worldwide and understanding their regeneration is necessary to design effective restoration strategies. It has previously been shown that the woody species community in secondary forests can follow different successional pathways according to the nature of past human activities in the area, yet little is known about patterns of herbaceous species diversity in secondary forests with different histories of land use. We compared the diversity and abundance of herbaceous plant communities in two types of Central Amazonian secondary forests—those regenerating on pastures created by felling and burning trees and those where trees were felled only. We also tested if plant density and species richness in secondary forests are related to proximity to primary forest. In comparison with primary forest sites, forests regenerating on non‐burned habitats had lower herbaceous plant density and species richness than those on burned ones. However, species composition and abundance in non‐burned stands were more similar to those of primary forest, whereas several secondary forest specialist species were found in burned stands. In both non‐burned and burned forests, distance from the forest edge was not related to herbaceous density and species richness. Overall, our results suggest that the natural regeneration of herbaceous species in secondary tropical forests is dependent on a site's post‐clearing treatment. We recommend evaluating the land history of a site prior to developing and implementing a restoration strategy, as this will influence the biological template on which restoration efforts are overlaid.     

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.     

Canopy seed bank structure in relation to: fire,tree size and density

To assess the canopy seed bank structure of Pinus halepensis, we measured the level of serotiny and the seed bank size and density of trees in unburned stands and post-fire regenerated stands in Israel. We analysed the effects of tree size, tree density and fire history on the level of serotiny. The level of serotiny decreased with an increase in tree height. The high level of serotiny in short trees could be explained by selection to increase regeneration chances after burning at pre-mature age. Also, limitation of long-distance seed dispersal opportunities in short trees may favour high serotiny levels. The level of serotiny was higher in post-fire stands than in unburned stands, suggesting a fast selection for serotiny by fire. Unburned stands had a higher total stand seed density than post-fire regenerated stands, but the proportion of seeds in serotinous cones of the total stand seed density was higher in post-fire regenerated stands. The fact that P. halepensis bears simultaneously serotinous and non-serotinous cones reflects its dual strategy as both a post-fire obligate seeder, mainly from serotinous cones and an early coloniser during fire-free periods, mainly from non-serotinous cones. The relative investment in these strategies is dependent on fire history and varies with tree height. Furthermore, mature brown cones can contribute to post-fire regeneration in case of spring fires, and serotinous cones are known to open partially also in dry spell events. Thus, post-fire regeneration and invasion are strategies, which seem to complement each other.     

How do slow‐growing,fire‐sensitive conifers survive in flammable eucalypt woodlands?

Question: To what extent do low flammability fuel traits enhance the survival and persistence of fire‐sensitive (slowing‐growing, non‐serotinous, non‐resprouting) dominant trees in highly flammable landscapes, under varying fire‐weather conditions? Location: Mixed forests co‐dominated by flammable Eucalyptus species and fire‐sensitive Callitris glaucophylla in Pilliga State Forest, southeast Australia. Methods: The influence of vegetation composition (relative abundance of Callitris and flammable Eucalyptus) on fire intensity and survival of fire‐sensitive Callitris was assessed across gradients of Callitris abundance in mixed Eucalyptus–Callitris forests that burned under low‐moderate and extreme fire‐weather conditions. Results: In areas that burned under low‐moderate fire‐weather conditions, as Callitris abundance increased, fire intensity declined and Callitris survival increased (46%). By comparison, in extreme fire‐weather conditions, lower fire intensity at higher levels of Callitris abundance, was not sufficient to increase Callitris survival (4%). Callitris survival was also positively related to trunk diameter. Ground fuel type, but not biomass, varied with vegetation composition. Conclusions: These results demonstrate that flammable feedbacks, mediated by low flammability fuel traits of dominant trees, can provide an important mechanism for enhancing the survival and persistence of slow‐growing, non‐serotinous, non‐resprouting, fire‐killed trees in highly flammable landscapes. By modifying vegetation and fuel structure, patches of fire‐sensitive Callitris reduce fire intensity, and thereby reduce Callitris mortality, enhancing population persistence. However, this feedback loop is insufficient to ensure Callitris survival under extreme fire‐weather conditions, when fire intensity is greater. After burning, stands remain vulnerable to future fires, until trees grow large enough to modify fuel levels and reduce stand flammability.     

Disturbance history and stand dynamics in tall open forest and riparian rainforest in the Central Highlands of Victoria

Abstract Spatial heterogeneity in the intensity of past disturbances has directly influenced the structure and composition of present‐day forests around the world. In south‐eastern Australia infrequent, high‐intensity wildfires are a major part of the historical disturbance regime. While these fires are often assumed to produce even‐aged stands, spatial heterogeneity in fire intensity due to highly variable topography may lead to more complex forest age structures. Our study describes the influence of disturbance on the age structure and dynamics of a mosaic of tall, open eucalypt forest, cool temperate rainforest and mixed species forest surrounding Bellel Creek in the Central Highlands of Victoria using dendrochronological techniques. We were particularly interested in the impacts of the 1939 Black Friday fire and its effects on forest age structure and subsequent stand development patterns. Within our study site tall open forest displayed two distinct age cohorts: (i) trees that established immediately after the 1939 fire and accounted for the majority of individuals in the forest, and (ii) scattered groups of older trees estimated to be approximately 200–250 years old. Cool temperate rainforest and mixed forest were also dominated by the post‐1939 fire age cohort. However, a greater proportion of trees in these forest types survived the 1939 fire relative to the tall open forest. The impact of the 1939 fire on the growth of surviving trees was highly variable but generally short‐lived. In most cases growth decreased after the 1939 fire, but generally returned to prefire levels within 1–3 years. Non‐fire disturbances were limited to small‐scale branch‐ and tree‐fall events, although the extreme snowstorm of 1977 appears to have caused extensive damage to rainforest communities. Our study demonstrates the opportunities for dendroecological studies to reconstruct historical dynamics and disturbance patterns in Australian forests and provides important insights into variation in landscape‐scale fire impacts and their effect on subsequent forest development patterns.     

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.     

Influence of fire severity on stand development of Araucaria araucana–Nothofagus pumilio stands in the Andean cordillera of south‐central Chile

Fire is the prevalent disturbance in the Araucaria–Nothofagus forested landscape in south‐central Chile. Although both surface and stand‐replacing fires are known to characterize these ecosystems, the variability of fire severity in shaping forest structure has not previously been investigated in Araucaria–Nothofagus forests. Age structures of 16 stands, in which the ages of approximately 650 trees were determined, indicate that variability in fire severity and frequency is key to explaining the mosaic of forest patches across the Araucaria–Nothofagus landscape. High levels of tree mortality in moderate‐ to high‐severity fires followed by new establishment of Nothofagus pumilio typically result in stands characterized by one or two cohorts of this species. Large Araucaria trees are highly resistant to fire, and this species typically survives moderate‐ to high‐severity fires either as dispersed individuals or as small groups of multi‐aged trees. Small post‐fire cohorts of Araucaria may establish, depending on seed availability and the effects of subsequent fires. Araucaria's great longevity (often >700 years) and resistance to fire allow some individuals to survive fires that kill and then trigger new Nothofagus cohorts. Even in relatively mesic habitats, where fires are less frequent, the oldest Araucaria–Nothofagus pumilio stands originated after high‐severity fires. Overall, stand development patterns of subalpine Araucaria–N. pumilio forests are largely controlled by moderate‐ to high‐severity fires, and therefore tree regeneration dynamics is strongly dominated by a catastrophic regeneration mode.     

Influence of rocky landscape features and fire regime on vegetation dynamics in Appalachian Quercus forests

Question : How do interactions between rocky landscape features and fire regime influence vegetation dynamics? Location : Continental Eastern USA. Methods : We measured vegetation, disturbance and site characteristics in 40 pairs of rocky and non‐rocky plots: 20 in recently burned stands, and 20 in stands with no evidence of recent fire (‘unburned’ stands). Two‐way analysis of variance (ANOVA) was used to assess the main and interaction effects of fire and rock cover on plant community composition. Results : In burned stands, rock cover had a strong influence on vegetation. Non‐rocky ‘matrix’ forests were dominated by Quercus, and had abundant ground cover and advance regeneration of early and mid‐successional tree species. Burned rocky patches supported greater density of fire‐sensitive species such as Acer rubrum, Sassafras albidum and Nyssa sylvatica and had little advance regeneration or ground cover. Quercus had fewer fire scars and catfaces (open, basal wounds) on rocky patches, suggesting that rocky features mitigate fire severity. In unburned stands, differences between rocky and non‐rocky patches were less distinct, with both patch types having sparse ground cover, little tree regeneration, and high understorey densities of relatively shade tolerant A. rubrum, N. sylvatica and Betula lenta. Conclusion : Under a sustained fire regime, heterogeneity in rock cover created a mosaic where fire‐adapted species such as Quercus dominate the landscape, but where fire‐sensitive species persisted in isolated pockets of lower fire severity. Without fire, species and landscape richness may decline as early‐mid successional species are replaced by more shade tolerant competitors.     

Patch to landscape patterns in post fire recruitment of a serotinous conifer

Obligate seeding species are highly specialized to fire disturbance and many conifers such as cypress, which are adapted to high intensity stand-replacing fires, have canopy seed banks stored in serotinous cones. Resilience of these trees to fire disturbance is a function of disturbance frequency and one focus of this study was to determine the effect of patch age on postfire recruitment. A second focus was to determine the extent to which fire induced a landscape level change in the location of the forest boundary. Prior to a fire in 1994, a large Cupressus sargentii forest was a mosaic landscape of different aged patches of nearly pure cypress bordered by chaparral. Patches less than 60 years of age were relatively dense with roughly one tree every 1–2 m² but older patches had thinned to one tree every 3–15 m². Older trees had substantially greater canopy cone crops but the stand level seed bank size was not significantly correlated with stand age. Fire-dependent obligate seeding species are sensitive to fire return interval because of potential changes in the size of seed banks – facing both a potential `immaturity risk' and a `senescence risk'. At our site, C. sargentii regeneration was substantial in stands as young as 20 years, suggesting that fire return interval would need to be shorter than this to pose any significant risk. Reduced seedling recruitment in stands nearly 100 years of age may indicate risk from senescence is greater, however, even the lowest density seedling recruitment was many times greater than the density of mature forests – thus this cypress would appear to be resilient to a wide range of fire return intervals. Changes in landscape patterning of forest and chaparral are unlikely except after fire. Factors that inhibit tree establishment within the shrubland, as well as factors that affect shrub establishment within the forest border likely affect the `permeability' of this ecotone. After the 1994 fire this boundary appeared to be stable in that cypress recruited best within the shadow of burned canopies and cypress were weak invaders of adjacent shrublands.     

Recovery of late-seral vascular plants in a chronosequence of post-clearcut forest stands in coastal Nova Scotia,Canada

We investigated the impacts of clearcutting on the ground vegetation of remnant late-successional coastal Acadian forests in southwestern Nova Scotia. Vegetation was sampled in 750 1-m² quadrats established in 16 stands belonging to different recovery periods since clearcutting (3–54 years) and 9 late-successional forests (100–165 years) with no signs of significant human disturbance. Our objectives were to: i) describe the changes in species richness, diversity, and abundance of ground vegetation after clearcutting; ii) examine the responses of residual species (i.e., late-successional flora) to clearcutting; and iii) determine whether any forest species were restricted to or dependent upon the late-successional stages of stand development for maximal frequency and/or abundance. Although clearcutting had no immediate impact on overall alpha richness or diversity, the richness and diversity of residual plants declined after canopy removal and showed no evidence of recovery over 54 years of secondary succession. Consequently, compositional differences between secondary and late-seral stands persisted for many decades after clearcutting. Several understory herbs (e.g., Coptis trifolia (L.) , Oxalis montana (L.), Monotropa uniflora (L.)) were restricted to or attained their highest frequency and abundance in late-seral forests. These results suggest that the preservation of remnant old stands may be necessary for the maintenance of some residual plants in highly disturbed and fragmented forest landscapes in eastern Canada.     

Non‐fire induced seed release ina weakly serotinous pine: climatic factors,maintenance costs or both?

The advantages of canopy seed retention (serotiny) for plants inhabiting fire‐prone ecosystems are well documented. However, very few species are completely serotinous and non‐fire induced opening of serotinous fruits is commonly observed (weak serotiny). Two non‐mutually exclusive causes are envisaged to contribute to this process: mechanical changes in serotinous fruits mediated by climatic conditions (e.g. drought) or the costs of maintenance for the plant of these long‐lasting structures. However, their relative contribution to the spontaneous opening of serotinous fruits remains elusive as well as the consequences for the build‐up of the canopy seed bank and inter‐individual differences in serotiny. In this study we monitored the dynamics of cone production and cone opening in the weakly serotinous Pinus halepensis for five years (2004–2008), including two severe drought episodes (2005, 2006). Drought decreased the production of conelets, increased the abortion of immature cones, reduced the seed quality in the cohorts of cones produced during these years, and increased the opening of serotinous cones. During the first drought episode, a higher proportion of serotinous cones opened in those pines bearing a larger crop of younger cones. This suggests that not only passive changes induced by drought but also competition among cones for resources (e.g. water) might be involved in this process. The opening of serotinous cones in pines bearing more cones made inter‐individual differences in the size of the canopy cone bank to narrow or even to reverse from 2004 to 2008. These results may help to understand the decrease in serotiny when pines grow and accumulate more cones and the large inter‐individual variability in the degree of serotiny observed in P. halepensis forests. In addition, the negative effects of drought episodes for the size of the canopy cone bank and the seeds contained can be an unexplored cause of post‐fire regeneration constraint.     

Stand‐replacing fires reduce susceptibility of lodgepole pine to mountain pine beetle outbreaks in Colorado

Aim As climate change is increasing the frequency, severity and extent of wildfire and bark beetle outbreaks, it is important to understand how these disturbances interact to affect ecological patterns and processes, including susceptibility to subsequent disturbances. Stand‐replacing fires and outbreaks of mountain pine beetle (MPB), Dendroctonus ponderosae, are both important disturbances in the lodgepole pine, Pinus contorta, forests of the Rocky Mountains. In the current study we investigated how time since the last stand‐replacing fire affects the susceptibility of the stand to MPB outbreaks in these forests. We hypothesized that at a stand‐scale, young post‐fire stands (< c. 100–150 years old) are less susceptible to past and current MPB outbreaks than are older stands. Location Colorado, USA. Methods We used dendroecological methods to reconstruct stand‐origin dates and the history of outbreaks in 23 lodgepole pine stands. Results The relatively narrow range of establishment dates among the oldest trees in most sampled stands suggested that these stands originated after stand‐replacing or partially stand‐replacing fires over the past three centuries. Stands were affected by MPB outbreaks in the 1940s/1950s, 1980s and 2000s/2010s. Susceptibility to outbreaks generally increased with stand age (i.e. time since the last stand‐replacing fire). However, this reduced susceptibility of younger post‐fire stands was most pronounced for the 1940s/1950s outbreak, less so for the 1980s outbreak, and did not hold true for the 2000s/2010s outbreak. Main conclusions Younger stands may not have been less susceptible to the most recent outbreak because: (1) after stands reach a threshold age of > 100–150 years, stand age does not affect susceptibility to outbreaks, or (2) the high intensity of the most recent outbreak reduces the importance of pre‐disturbance conditions for susceptibility to disturbance. If the warm and dry conditions that contribute to MPB outbreaks concurrently increase the frequency and/or extent of severe fires, they may thereby mitigate the otherwise increased landscape‐scale susceptibility to outbreaks. Potential increases in severe fires driven by warm and dry climatic trends may lead to a negative feedback by making lodgepole pine stands less susceptible to future MPB outbreaks.     

Minireviews: Neighborhood Effects, Disturbance Severity, and Community Stability in Forests

A theoretical framework and conceptual model for temporal stability of forest tree-species composition was developed based on a synthesis of existing studies. The model pertains primarily to time periods of several tree lifetimes (several hundred to a few thousand years) at the neighborhood and stand spatial scales (0.01–10 ha), although a few extensions to the landscape scale are also made. The cusp catastrophe was chosen to illustrate compositional dynamics at the stand level for jack pine, northern hardwood, and white pine forests in the Great Lakes Region of the United States and for tropical rainforests in the northern Amazon basin. The models feature a response surface (degree of dominance by late-successional species) that depends on two variables: type of neighborhood effects of the dominant tree species and severity of disturbances. Neighborhood effects are processes that affect the chance of a species replacing itself at the time of disturbance (they can be positive, neutral, or negative) and are of two types: overstory–undestory effects, such as the presence of advanced reproduction; and disturbance-activated effects, such as serotinous seed rain. Disturbance severity is the proportion of trees killed during a disturbance. Interactions between neighborhood effects and disturbance severity can lead to either punctuated stability (dramatic but infrequent change in composition, in those forests dominated by species with positive neighborhood effects) or succession (continuous change, in those forests dominated by species with neutral-negative neighborhood effects). We propose that neighborhood effects are a major organizing factor in forest dynamics that provide a link across spatial scales between individual trees and disturbance/patch dynamics at the stand and landscape scales. Received 23 June 1998; accepted 16 December 1998.     

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.     

Fire disturbance and forest structure in old‐growth mixed conifer forests in the northern Sierra Nevada,California

Question: This study evaluates how fire regimes influence stand structure and dynamics in old‐growth mixed conifer forests across a range of environmental settings. Location: A 2000‐ha area of mixed conifer forest on the west shore of Lake Tahoe in the northern Sierra Nevada, California. Methods: We quantified the age, size, and spatial structure of trees in 12 mixed conifer stands distributed across major topographic gradients. Fire history was reconstructed in each stand using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing the fire history with the age, size, and spatial structure of trees in a stand. Results: There was significant variation in species composition among stands, but not in the size, age and spatial patterning of trees. Stands had multiple size and age classes with clusters of similar aged trees occurring at scales of 113 ‐ 254 m². The frequency and severity of fires was also similar, and stands burned with low to moderate severity in the dormant season on average every 9–17 years. Most fires were not synchronized among stands except in very dry years. No fires have burned since ca. 1880. Conclusions: Fire and forest structure interact to perpetuate similar stand characteristics across a range of environmental settings. Fire occurrence is controlled primarily by spatial variation in fuel mosaics (e.g. patterns of abundance, fuel moisture, forest structure), but regional drought synchronizes fire in some years. Fire exclusion over the last 120 years has caused compositional and structural shifts in these mixed conifer forests.     

The importance of seed trees in the dioecious conifer Pilgerodendron uviferum for passive restoration of fire disturbed southern bog forests

Biological legacies are important for ecosystem recovery following disturbance as demonstrated by studies in the northern hemisphere. Southern bog forests dominated by the conifer Pilgerodendron uviferum in Northern Patagonia are a typical case of an ecosystem with low resilience to disturbance by fire, which kills most trees and seeds, on which the species depends for regeneration. In this study, we hypothesize that the natural recovery of P. uviferum populations in burned areas is limited by seed availability and this limitation may be exacerbated by the dioecy of the species. Using a multi‐scaled approach, we quantified the seed dissemination potential from P. uviferum seed trees, assessed the suitability of substrates for the germination of seedlings, and finally analysed the spatial distribution of seed trees of the species at the landscape level. Our results indicate that 70 years after a fire on Chiloé Island (43°S), natural regeneration from seed trees can assist the recovery of P. uviferum populations following large‐scale fire disturbance, but their effect is limited at a landscape level owing to a low number of reproductive female trees (0.3 trees ha^?1) and limited seed dispersal (<20 m). In this context, a mixed passive‐active restoration approach that takes into account the spatial pattern and sex of seed trees could be the most effective and efficient option to restore not only P. uviferum forests in North Patagonia, but also other heavily disturbed forests with few remnant seed trees, in particular of dioecious species.     

Removal of Encroaching Conifers to Regenerate Degraded Aspen Stands in the Sierra Nevada

Aspen is considered a keystone species, and aspen communities are critical for maintaining biodiversity in western landscapes. Inventories of aspen stand health across the Eagle Lake Ranger District (ELRD), Lassen National Forest, California, U.S.A., indicate that 77% of stands are in decline and at risk of loss as defined by almost complete loss of mature aspen with little or no regeneration. This decline is due to competition from conifers establishing within aspen stands as a result of modification of natural fire regimes coupled with excessive browsing by livestock. Restoration treatments were implemented in four aspen stands in 1999 using mechanical equipment to remove competing conifers to enhance the growth environment for aspen. Recruitment and establishment of aspen stems were measured in treated stands (removal of competing conifers) and non‐treated stands (control) immediately prior to treatment and 2 and 4 years post‐treatment. There was a significant increase in total aspen stem density and in two of three aspen regeneration size classes for treated stands compared to controls. Pre‐treatment total aspen density was positively associated with total aspen density and density in all size classes of aspen (p < 0.001). The results demonstrate that mechanical removal of conifers is an effective treatment for restoring aspen.