In this review and synthesis paper, we review the resilience of secondary forests to climate change through the lenses of ecosystem legacies and landscape diversity. Ecosystem legacy of secondary ...forests was categorized as continuous forest, non-continuous forest, reassembled after conversion to other land uses, and novel reassembled forests of non-native species. Landscape diversity, including landforms that create varied local climatic and soil conditions, can buffer changing climate to some extent by allowing species from warmer climates to exist on warm microsites, while also providing refugial locations for species that grow in cool climates. We present five frames that allow forest managers to visualize a trajectory of change in the context of projected regional climate change, which are: Frame 1 (persistence), keep the same dominant tree species with little change; Frame 2 (moderate change), keep the same tree species with large changes in relative abundance; Frame 3 (forest biome change), major turnover in dominant tree species to a different forest biome; Frame 4 (forest loss), change from a forest to a non-forest biome; and Frame 5 (planted novel ecosystem), establish a novel ecosystem to maintain forest. These frames interact with ecosystem legacies and landscape diversity to determine levels of ecosystem resilience in a changing climate. Although forest readiness to adapt to Frame 1 and 2 scenarios, which would occur with reduced greenhouse gas emissions, is high, a business as usual climate change scenario would likely overwhelm the capacity of ecosystem legacies to buffer forest response, so that many forests would change to warmer forest biomes or non-forested biomes. Furthermore, the interactions among frames, legacies, and landscape diversity influence the transient dynamics of forest change; only Frame 1 leads to stable endpoints, while the other frames would have transient dynamics of change for the remainder of the 21st century.
The anticipated increase in extreme disturbance events due to climate change is likely to expose Norway spruce (
Picea abies
(L.) Karst.) dominated forests in northern Europe to new conditions. ...Empirical data on the resilience of such natural (unmanaged) forests to disturbance and the long-term patterns of regeneration in its aftermath are currently scarce. We performed a quantitative assessment of natural forest stands in north–western Latvia to identify and characterise the patterns of stand structure 44 years after a stand-replacing disturbance and investigated the effects of legacies on regeneration. The spatial distribution of tree species and their dimensions were assessed in 71 circular sample plots (500 m
2
each) in natural forest areas, where Norway spruce dominated prior to the windthrow and salvage logging was not carried out. Spatial indices (species mingling index, size differentiation index, and aggregation index) were used to characterise stand structure and diversity. The different initial states (age and coverage of surviving trees) of stands affected eventual tree species dominance, size differentiation, degree of mingling and aggregation. Our results demonstrate a close relationship between disturbance legacies and spatial indices. The pre-storm understory and canopy survivors decreased species mingling, whereas survivors increased size differentiation. The size differentiation increased also with a higher degree of species mingling. Leaving differential post-storm legacies untouched promotes a higher structural and species diversity and therefore supports the management approach of preserving canopy survivors.
Trees that survive disturbances are important biological legacies that facilitate forests’ recovery and enhance their structural and species diversity, substantially contributing to the resilience of ...these ecosystems. The dynamic pattern of legacy syndromes sets the understudied aspects of survivors of wind disturbance into focus. Several factors at tree, stand, and landscape scales alter the susceptibility of the remnant trees, and affect their potential to recover and survive subsequent disturbances. The characteristics of the survivors interact with direct stress and mortality drivers such as changed environmental conditions and pressure by pests and pathogens. Climate change further enhances the post-storm vulnerability of the remaining stand. This literature review analyzes the impact of disturbance parameters (e.g., severity, seasonal timing) and characteristics of the affected forest (e.g., tree species composition, successional stage of a forest stand) on the conditions of survivors through post-windthrow stand development. We attempted to reveal the main agents and processes driving the fate of remnant trees and linked delayed mortality patterns to the main stand-scale wind disturbance regimes in Eurasian and North American boreal and temperate forests: (1) stand-replacing, (2) partially stand-replacing, and (3) fine-scale gap disturbance. We found that after stand-replacing wind disturbance, the spatial location of the remaining trees largely determines their onward fate, whereas these survivors are generally more susceptible to subsequent mortality compared to trees that survived less severe events. After partially stand-replacing wind disturbance, the structure of the remnant stand as well as characteristics of the individual remnant trees (e.g., species, age, size) largely determine their survival probability. Following a fine-scale gap disturbance, the trees at the gap edge are more likely to die, compared to the trees situated in the stand interior, but the mortality-causing processes usually operate on a longer time scale. Our findings contribute to the current knowledge on post-windthrow stand development and offer insights into temporal stability of these increasingly important biological legacies.
Forest ecosystems are shaped by disturbances and functional features of vegetation recovery after disturbances. There is considerable variation in basic disturbance characteristics, magnitude, ...severity, and intensity. Disturbance legacies provide possible explanations for ecosystem resilience. The impact (length and strength) of the pool of ecosystem legacies and how they vary at different spatial and temporal scales is a most promising line of further research. Analyses of successional trajectories, ecosystem memory, and novel ecosystems are required to improve modelling in support of forests. There is growing evidence that managing ecosystem legacies can act as a driver in adaptive management to achieve goals in forestry. Managers can adapt to climate change and new conditions through anticipatory or transformational strategies of ecosystem management. The papers presented in this Special Issue covers a wide range of topics, including the impact of herbivores, wind, and anthropogenic factors, on ecosystem resilience.
Natural regeneration of windthrow areas is an important issue when planning forestry measures after forest disturbances. Seedling recruitment was investigated in storm-damaged hemiboreal mixed ...forests in eastern Estonia. The establishment and growth of seedlings from natural regeneration was registered for tree species in soil pits and in mounds of uprooted trees in stands that were either heavily or moderately damaged. Seedling growth is expected to be better in large but shallow soil pits created by uprooted Norway spruce Picea abies (L.) Karst. and poorer in small but deep pits created by the hardwoods in the area, silver birch (Betula pendula Roth.) and European aspen (Populus tremula L.). The most abundant regenerating species was birch. Pits hosted larger seedling numbers than mounds, due to soil instability in mounds. Rowan (Sorbus aucuparia L.) showed significantly faster growth than the other seedling species. Norway spruce pits were preferred to pits of other species by both birch and spruce seedlings. Black alder Alnus glutinosa (L.) J. Gaertn. did not show a preference for pits of a certain species of uprooted tree. Both spruce and rowan preferred hardwood mounds over spruce mounds. Storm severity also affected species composition: birch predominantly occurred on pits and mounds in heavily disturbed areas, while spruce was more abundant in the moderately damaged areas. The effects of advance regeneration and surrounding stands on seedling microsite preferences should be considered in future research and subsequent management recommendations.
We reviewed studies dealing with regeneration under variable conditions in boreal and hemiboreal forests as affected by different microsite types by tree species functional groups. Generally, the ...importance of storm-induced microsites for regeneration dynamics in boreal forests depends on several factors: (1) distribution and type of microsites (generated by storm characteristics and stand conditions); (2) viable seed supply (stand history, species dispersal traits and status of surviving trees) and their species' life history strategy; (3) climatic and site conditions (pre-storm conditions and storm-induced changes); and (4) delayed storm effects, such as retarded falling of trees, favoured vegetation growth, etc. Studies acknowledging the significance of microsites were mostly related to intermediate or severe events, causing sufficient changes in resource levels and growth conditions, and influencing extrinsic factors such as frost heaving, erosion and browsing. Also, the dispersal traits of available tree species, including sprouting and response of surviving trees, such as canopy expansion, should be considered in evaluating microsite importance in individual cases. In intermediate to severe windstorm events, pioneer species are generally profiting most from the additional offer in microsites, requiring bare mineral soil and elevated locations for their establishment and growth. Under gap dynamics, shade-tolerant species benefit from dead wood and elevated locations as these offer safe sites in stands with abundant understorey vegetation.
Climate change accompanied by altered natural disturbance regimes threatens the resilience of boreal and hemiboreal forests. Bryophytes, an important part of plant biomass and diversity, fulfill ...numerous important functions in forests, thus markedly contributing to the resilience of these ecosystems. We studied abundance, richness, diversity and community composition of bryophyte species 20–21 years after wind disturbance in moderately damaged, heavily damaged and heavily damaged and subsequently salvage-logged stands; overmature forest stands with similar tree species composition were included in the study as a reference group. The community characteristics were linked with treatment and accompanying environmental variables. Altogether, 108 bryophyte taxa (81 mosses and 27 liverworts) were identified in study plots. We found that bryophytes responded to windthrow severity level and salvage logging in terms of species richness, diversity, and composition; diversity of microhabitats was the most important environmental variable explaining the variation in diversity metrics. Besides exhibiting greater overall species richness, uncleared wind-disturbed plots contained more bryophyte species with high conservational value as well, compared to salvage-logged and control plots. The largest number of bryophyte taxa was associated with deadwood substrates. Ordination analysis further emphasized the importance of deadwood substrate for liverworts. In order to fulfill the objective of biodiversity conservation, wind-disturbed forests should be left to recover naturally, to ensure a favorable status of all naturally occurring species and to improve the status of endangered and rare species. Retaining a proportion of surviving and/or wind-damaged trees while salvaging might also alleviate the unfavorable environmental conditions for bryophyte species while “lifeboating” some species through the following successional stages.
•Uncleared wind-disturbed stands host highly diverse bryophyte communities.•Bryophytes responded strongly to variation in substrate diversity.•Abundance of deadwood substrate is especially important for liverworts.•Salvaged and control plots had similar mean richness, but distinct compositions.•Retention of deadwood and live tree legacies during salvaging is recommended.
Disturbance regimes are changing in forests across the world in response to global climate change. Despite the profound impacts of disturbances on ecosystem services and biodiversity, assessments of ...disturbances at the global scale remain scarce. Here, we analyzed natural disturbances in boreal and temperate forest ecosystems for the period 2001–2014, aiming to 1) quantify their within‐ and between‐biome variation and 2) compare the climate sensitivity of disturbances across biomes. We studied 103 unmanaged forest landscapes with a total land area of 28.2 × 106 ha, distributed across five continents. A consistent and comprehensive quantification of disturbances was derived by combining satellite‐based disturbance maps with local expert knowledge of disturbance agents. We used Gaussian finite mixture models to identify clusters of landscapes with similar disturbance activity as indicated by the percent forest area disturbed as well as the size, edge density and perimeter–area‐ratio of disturbed patches. The climate sensitivity of disturbances was analyzed using Bayesian generalized linear mixed effect models and a globally consistent climate dataset. Within‐biome variation in natural disturbances was high in both boreal and temperate biomes, and disturbance patterns did not vary systematically with latitude or biome. The emergent clusters of disturbance activity in the boreal zone were similar to those in the temperate zone, but boreal landscapes were more likely to experience high disturbance activity than their temperate counterparts. Across both biomes high disturbance activity was particularly associated with wildfire, and was consistently linked to years with warmer and drier than average conditions. Natural disturbances are a key driver of variability in boreal and temperate forest ecosystems, with high similarity in the disturbance patterns between both biomes. The universally high climate sensitivity of disturbances across boreal and temperate ecosystems indicates that future climate change could substantially increase disturbance activity.
Several storms have damaged Estonian forests in recent years. Individual tree properties such as diameter at breast height (dbh) and tree height affect the type of damage (stem breakage or uprooting) ...and influence the formation of postdisturbance forest structure. The aim of this study was to analyse how the risk of uprooting and stem breakage varied with respect to tree diameter, height, species and soil conditions. The data were collected from plots located in storm-damaged forests in eastern Estonia. The probability of uprooting increased with increasing diameter for Norway spruce, European aspen and birch. Black alders with large stem diameter usually suffered stem breakage. A high proportion of stem breakage also occurred among small Norway spruce in the understorey, probably damaged by falling canopy trees. Soil conditions influenced the ratio between uprooting and stem breakage of Norway spruce, the ratio being much higher on gley soils than on podzolic soils. The root plate size of Norway spruce was only slightly correlated with dbh, tree height and volume.
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