•First study in primary forest linking ≥250 yr. disturbance history to bird assemblage.•Bird assemblage composition was driven by disturbance related structure.•Disturbance induced forest change did ...not decrease bird diversity or abundance.
Understanding the processes shaping the composition of assemblages in response to disturbance events is crucial for preventing ongoing biodiversity loss in forest ecosystems. However, studies of forest biodiversity responses to disturbance typically analyze immediate or short-term impacts only, while studies relating long-term disturbance history to biodiversity assemblage dynamics are rare. To address this important knowledge gap, we used a dendroecological approach to link natural disturbance history of 250 years (1750–2000) to structural habitat elements and, in turn, to breeding bird assemblages. We used data collected in 2017 and 2018 from 58 permanent study plots within 10 primary spruce forest stands distributed across the Western Carpathian Mountains of Europe. This dataset contained breeding bird counts and environmental variables describing forest density, tree diameter distribution, tree height, tree microhabitats, deadwood quantity and quality, and regeneration. Bird assemblages were significantly influenced by forest structure which was in turn shaped by disturbance dynamics (disturbance frequency, time since the last disturbance and its severity). Early successional species associated with more open habitats were positively influenced by disturbance-related structure (i.e. deadwood-related variables, canopy cover), while some species responded negatively. At the same time, overall abundance, species richness and Shannon diversity of the bird assemblage remained unchanged under variable disturbance histories. Our results support a view of primary spruce forests as a highly dynamic ecosystem, harbouring populations of bird species at all stages of succession despite significant structural changes and shifting patch mosaics over time due to natural disturbances.
Research Highlights: Past disturbances occurred naturally in primary forests in the Southern Carpathians. High- and moderate-severity disturbances shaped the present structure of these ecosystems, ...which regenerated successfully without forestry interventions. Background and Objectives: Windstorms and bark beetle outbreaks have recently affected large forest areas across the globe, causing concerns that these disturbances lie outside the range of natural variability of forest ecosystems. This often led to salvage logging inside protected areas, one of the main reasons for primary forest loss in Eastern Europe. Although more than two-thirds of temperate primary forests in Europe are located in the Carpathian region of Eastern Europe, knowledge about how natural disturbances shape the forest dynamics in this region is highly essential for future management decisions. Material and Methods: We established our study in a primary forest valley situated in the centre of the largest temperate primary forest landscape in Europe (Făgăraș Mountains). A dendrochronological investigation was carried out to reconstruct the natural disturbance history and relate it to the present forest structure. Results: The dendrochronological analysis revealed high temporal variability in the disturbance patterns both at the patch and stand level. Moderate severity disturbance events were most common (20–40% of canopy disturbed in 60% of the plots) but high severity events did also occur (33% of the plots). Regeneration was spruce-dominated and 71% of the seedlings were found on deadwood microsites. Conclusions: We conclude that the current structure of the studied area is a consequence of the past moderate-severity disturbances and sporadic high-severity events. The peak in disturbances (1880–1910) followed by reduced disturbance rates may contribute to a recent and future increase in disturbances in the Făgăraș Mts. Our findings show that these disturbance types are within the range of natural variability of mountain spruce forests in the Southern Carpathians and should not be a reason for salvage logging in primary forests from this area.
Aims
We examined differences in life span among the dominant tree species (spruce, Picea abies; fir, Abies alba; beech, Fagus sylvatica; and maple, Acer pseudoplatanus) across primary mountain ...forests of Europe. We asked how disturbance history, lifetime growth patterns, and environmental factors influence life span.
Locations
Balkan Mountains, Carpathian Mountains, Dinaric Mountains.
Methods
Annual ring widths from 20,600 cores from primary forests were used to estimate tree life spans, growth trends, and disturbance history metrics. Mixed models were used to examine species‐specific differences in life span (i.e., defined as species‐specific 90th percentiles of age distributions), and how metrics of radial growth, disturbance parameters, and selected environmental factors influence life span.
Results
While only a few beech trees surpassed 500 years, individuals of all four species were older than 400 years. There were significant differences in life span among the four species (beech > fir > spruce > maple), indicating life history differentiation in life span. Trees were less likely to reach old age in areas affected by more severe disturbance events, whereas individuals that experienced periods of slow growth and multiple episodes of suppression and release were more likely to reach old age. Aside from a weak but significant negative effect of vegetation season temperature on fir and maple life span, no other environmental factors included in the analysis influenced life span.
Conclusions
Our results indicate species‐specific biological differences in life span, which may play a role in facilitating tree species coexistence in mixed temperate forests. Finally, natural disturbance regimes were a key driver of life span, which could have implications for forest dynamics if regimes shift under global change.
We examined differences in life spans and their drivers among the dominant tree species across primary mountain forests of Europe. Significant differences were found in life spans among the four studied species (Fagus sylvatica > Abies alba > Picea abies > Acer pseudoplatanus). Disturbance regimes and not environmental conditions were identified as the main driver of life span variability in primary forests.
•We compared Tree-related microhabitats (TreMs) in the Dinarides and Carpathians.•Density and diversity of TreMs were very high in beech dominated primary forests.•Geographical region was not a ...primary driver of TreM profile.•Tree species richness, DBH, and snags were the main drivers of TreM profile.•These findings may provide a benchmark for forest management practices.
Tree-related microhabitats (TreMs) are important features for the conservation of biodiversity in forest ecosystems. Although other structural indicators of forest biodiversity have been extensively studied in recent decades, TreMs have often been overlooked, either due to the absence of a consensual definition or a lack of knowledge. Despite the increased number of TreM studies in the last decade, the role of drivers of TreM profile in primary forests and across different geographical regions is still unknown. To evaluate the main drivers of TreM density and diversity, we conducted the first large-scale study of TreMs across European primary forests. We established 146 plots in eight primary forests dominated by European beech (Fagus sylvatica L.) in the Carpathian and Dinaric mountain ranges. Generalized linear mixed effect models were used to test the effect of local plot characteristics and spatial variability on the density and diversity (alpha, beta, and gamma) of TreMs. Total TreM density and diversity were significantly positively related with tree species richness and the proportion of snags. Root mean square tree diameters were significantly related to alpha and gamma diversity of TreMs. Both regions reached similarly high values of total TreM densities and total TreM densities and diversity were not significantly different between the two regions; however, we observed between the two regions significant differences in the densities of two TreM groups, conks of fungi and epiphytes. The density and diversity of TreMs were very high in beech-dominated mountain primary forests, but their occurrence and diversity was highly variable within the landscapes over relatively short spatial gradients (plot and stand levels). Understanding these profile provides a benchmark for further comparisons, such as with young forest reserves, or for improving forest management practices that promote biodiversity.
Assessing the impacts of natural disturbance on the functioning of complex forest systems are imperative in the context of global change. The unprecedented rate of contemporary species extirpations, ...coupled with widely held expectations that future disturbance intensity will increase with warming, highlights a need to better understand how natural processes structure habitat availability in forest ecosystems. Standardised typologies of tree-related microhabitats (TreMs) have been developed to facilitate assessments of resource availability for multiple taxa. However, natural disturbance effects on TreM diversity have never been assessed. We amassed a comprehensive dataset of TreM occurrences and a concomitant 300-year disturbance history reconstruction that spanned large environmental gradients in temperate primary forests. We used nonlinear analyses to quantify relations between past disturbance parameters and contemporary patterns of TreM occurrence. Our results reveal that natural forest dynamics, characterised by fluctuating disturbance intervals and variable severity levels, maintained structurally complex landscapes rich in TreMs. Different microhabitat types developed over time in response to divergent disturbance histories. The relative abundance of alternate TreMs was maximised by unique interactions between past disturbance severity and elapsed time. Despite an unequal distribution of individual TreMs, total microhabitat diversity was maintained at constant levels, suggesting that spatially heterogeneous disturbances maintained a shifting mosaic of habitat types over the region as a whole. Our findings underscore the fundamental role of natural processes in promoting conditions that maximise biodiversity potential. Strict conservation and management systems that preserve natural disturbance outcomes, including associated biological legacies, may therefore safeguard biodiversity at large scales.
•Historical disturbances shaped the contemporary local and regional richness of fungi.•Natural disturbances had contrasting effects on fungi at different spatial scales.•Local red-listed species ...diversity increased due to higher disturbance frequency.•Regional diversity of all species was decreased by higher disturbance severity.
Understanding the processes shaping the composition of assemblages at multiple spatial scales in response to disturbance events is crucial for preventing ongoing biodiversity loss and for improving current forest management policies aimed at mitigating climate change and enhancing forest resilience. Deadwood-inhabiting fungi represent an essential component of forest ecosystems through their association with deadwood decomposition and the cycling of nutrients and carbon. Although we have sufficient evidence for the fundamental role of deadwood availability and variability of decay stages for fungal species diversity, the influence of long-term natural disturbance regimes as the main driver of deadwood quantity and quality has not been sufficiently documented. We used a dendroecological approach to analyse the effect of 250-years of historical natural disturbance and structural habitat elements on local (plot-level) and regional (stand-level) species richness of deadwood-inhabiting fungi. We used data collected from 51 study plots within nine best-preserved primary spruce forest stands distributed across the Western Carpathian Mountains. Historical disturbances shaped the contemporary local and regional species richness of fungi, with contrasting impacts of disturbance regime components at different spatial scales. While local diversity of red-listed species has increased due to higher disturbance frequency, regional diversity of all species has decreased due to higher severity historical disturbances. The volume of deadwood positively influenced the species richness of deadwood-inhabiting fungi while canopy openness had a negative impact. The high number of observed rare species highlights the important role of primary forests for biodiversity conservation. From a landscape perspective, we can conclude that the distribution of species from the regional species pool is - at least to some extent - driven by past spatiotemporal patterns of disturbance events. Natural disturbances occurring at higher frequencies that create a mosaic forest structure are necessary for fungal species - especially for rare and endangered taxa. Thus, both the protection of intact forest landscapes and forest management practises that emulate natural disturbance processes are recommended to support habitats of diverse fungal communities and their associated ecosystem functions.
•Current DBH distribution shapes are driven by historical disturbance regimes.•Low-severity disturbances showed a reverse J-shaped diameter distribution.•High-severity disturbance lead to a unimodal ...diameter distribution.•Time and severity of last disturbance are good predictors of DBH distribution shape.
Natural disturbances strongly influence forest structural dynamics, and subsequently stand structural heterogeneity, biomass, and forest functioning. The impact of disturbance legacies on current forest structure can greatly influence how we interpret drivers of forest dynamics. However, without clear insight into forest history, many studies default to coarse assumptions about forest structure, for example, whether forests are even or unevenly aged. The aim of this study was to analyze the effects of past disturbances on the current diameter distributions of Norway spruce (Picea abies (L.) Karst.)-dominated landscapes throughout the Carpathian Mountains. Our dendroecological dataset comprises tree cores from 339 plots (7,845 total tree cores), nested within 28 primary forest stands, known to vary greatly in the severity of historical disturbances. Our analyses revealed that historical disturbances had a strong and significant effect on the current diameter distribution shapes at the plot level. We demonstrated that mixed-severity disturbance regimes were more frequent and create a complex pattern of diameter distributions at the plot and stand scale. Here, we show that high severity disturbance was associated with unimodal diameter distributions, while low and moderate severity was associated with the reverse J-shaped distribution. This is a result of complex disturbance patterns, with structural biological legacies. Our results will have important management implication in the context of tree size heterogeneity, biomass storage, and productivity as influenced by natural disturbances. Lastly, these results demonstrate that structural changes may arise as consequences of changing disturbance regime associated with global change.
In a world of accelerating changes in environmental conditions driving tree growth, tradeoffs between tree growth rate and longevity could curtail the abundance of large old trees (LOTs), with ...potentially dire consequences for biodiversity and carbon storage. However, the influence of tree‐level tradeoffs on forest structure at landscape scales will also depend on disturbances, which shape tree size and age distribution, and on whether LOTs can benefit from improved growing conditions due to climate warming. We analyzed temporal and spatial variation in radial growth patterns from ~5000 Norway spruce (Picea abies L. H. Karst) live and dead trees from the Western Carpathian primary spruce forest stands. We applied mixed‐linear modeling to quantify the importance of LOT growth histories and stand dynamics (i.e., competition and disturbance factors) on lifespan. Finally, we assessed regional synchronization in radial growth variability over the 20th century, and modeled the effects of stand dynamics and climate on LOTs recent growth trends. Tree age varied considerably among forest stands, implying an important role of disturbance as an age constraint. Slow juvenile growth and longer period of suppressed growth prolonged tree lifespan, while increasing disturbance severity and shorter time since last disturbance decreased it. The highest age was not achieved only by trees with continuous slow growth, but those with slow juvenile growth followed by subsequent growth releases. Growth trend analysis demonstrated an increase in absolute growth rates in response to climate warming, with late summer temperatures driving the recent growth trend. Contrary to our expectation that LOTs would eventually exhibit declining growth rates, the oldest LOTs (>400 years) continuously increase growth throughout their lives, indicating a high phenotypic plasticity of LOTs for increasing biomass, and a strong carbon sink role of primary spruce forests under rising temperatures, intensifying droughts, and increasing bark beetle outbreaks.
Understanding growth dynamics and development of large old trees in montane primary forests amidst intensifying environmental pressures can help predict future forest development and ecosystem functioning. Slow growth early in a trees' life is essential for long lifespan, however, disturbance dynamics carry a defining role for achieving old age in primary forests. In response to the late 20th century rising temperatures, Western Carpathian primary spruce forests have shown a synchronous increase in productivity, indicating a high adaptive capacity to improving growing conditions, which could carry broad landscape consequences for the carbon sink potential of primary montane forests under changing climate.
•Basal area (BA) was correlated with macroclimatic gradients along the Carpathians.•Fine-scale variation of BA is linked to local disturbances.•Recent stationarity in stand BA implied despite ...disturbance related variability.•Instances of high recent mortality may indicate a departure from stationarity.
Disentangling the importance of developmental vs. environmental drivers of variation in forest biomass is key to predicting the future of forest carbon sequestration. At coarse scales, forest biomass is likely to vary along major climatic and physiographic gradients. Natural disturbance occurs along these broad biophysical gradients, and depending on their extent, severity and frequency, could either amplify or dampen spatial heterogeneity in forest biomass. Here we evaluate spatial variation in the basal area of late-successional Picea abies (L./Karst.) forests across the Carpathian Mountain Range of central Europe and compare the roles of coarse-scale biophysical gradients and natural disturbances in driving that variation across a hierarchy of scales (landscapes, stands, and plots). We inventoried forest composition and structure, and reconstructed disturbance histories using tree cores collected from 472 plots nested within 30 late-successional stands, spanning the Carpathian Mountains (approximately 4.5 degrees of latitude). We used linear mixed-effects models to compare the effect of disturbance regimes and site conditions on stand basal area at three hierarchical scales. We found that the basal area of late-successional Picea abies forests varied across a range of spatial scales, with climatic drivers being most important at coarse scales and natural disturbances acting as the primary driver of forest heterogeneity at fine scales. For instance, the stand-level basal area varied among landscapes, with the highest values (48–68 m2 ha−1) in the warmer southern Carpathian Mountains, and lower values (37–52 m2 ha−1 on average) in cooler areas of the eastern and western Carpathians. Finer-scale variation was driven by local disturbances (mainly bark beetle and windstorms) and the legacies of disturbances that occurred more than a century ago. Our findings suggest that warming could increase the basal area of northern sites, but potential increasing disturbances could disrupt these environmental responses.
•Overall annual mortality rate was within published range of background mortality.•Plot-level variability in mortality rate was high both within and between stands.•Norway spruce mortality was evenly ...distributed among size classes.•Size class distribution of mortality varies for different mortality agents.•Bark beetle was the most frequent mortality agent, even in stands with low mortality.
Mortality, driven by both climate and disturbance legacies, is a key process shaping forest dynamics. Understanding the mortality patterns in primary forests in the absence of severe disturbances provides information on background natural dynamics of a given forest type under ongoing climate change. This can then be compared to mortality rates in severely-disturbed stands. Using a large number of sample plots along a gradient from low to high disturbance, we examined the mortality rates and composition of mortality agents in primary mountain Norway spruce (Picea abies (L.) Karst.) forests on different spatial scales. We evaluated the mortality rates and causes of mortality in 28 stands across a large geographical gradient spanning over 1000 km. We resampled (five-year period) 371 plots (16,287 living trees) in primary Norway spruce forests along the Carpathian mountain chain. The estimated overall annual mortality rate was within the previously reported range of background (ambient) mortality, however, stand-level and plot-level mortality rates varied substantially. Over 18% of plots displayed more than 2% annual mortality and 6% of plots even exceeded 10% per year. Stands in the Western Carpathians showed the highest variability in the mortality rate, with 30% of the stands in this region showing annual mortality rates over 5%. At the plot level, mixed-severity disturbances increased variability of mortality rates within most localities. Overall mortality was evenly distributed among size classes up to 50 cm diameter at breast height (DBH). However, the distributions differ for individual mortality agents. Mortality modes were classified into six categories (broken crown, broken stem, uprooted, competition, bark beetle/fungi, climatic extremes). Bark beetle (Ips typographus L.) infestation was the most frequent mortality agent in all stands, whereas the influence of competition as a mortality agent varied substantially. Mortality from abiotically-caused physical damage was similar to that from competition, yet the distribution among modes of physical damage (uprooted, crown, or stem breakage) varied. The lack of clear evidence of mortality agents in some locations implies that many tree deaths are caused by a combination of contributing factors. The results suggest the role of bark beetle as a mortality agent does not equate to severe mortality at large scales. Prevalence of different size classes affected by individual mortality agents underline the high complexity of the mortality process in primary forests.
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