At regional scale, it is dominantly considered in the European literature that litter decomposition is higher in soils from calcareous than siliceous bedrocks because of higher pH in the former ...forest ecosystems. We tested the alternative hypothesis that this should rather be due to differences in physical characteristics of the bedrock, with likely higher decomposition on calcareous than siliceous bedrocks in wet climates because of more favourable texture in the former and nutrient leaching in the latter, and the opposite in dry climates because of higher drought stress in calcareous soils. We assessed, four consecutive years with contrasting climates, the litter decomposition of a unified litter of Abies alba needles with the litterbag method in 70 forest sites located on both bedrocks and in wet oceanic and dry continental climates of the European Alps and the Pyrenees. Average and annual climate data were analysed with principal component analyses and the effects of bedrock type, average and annual climate drought stresses on litter decomposition were analysed, separately in the Alps and the Pyrenees, with analyses of variance. We found, in both mountain ranges, a highly significant bedrock type by average climate drought stress interaction on litter decomposition, due to a strong decrease in litter decomposition from wet oceanic to dry continental sites on calcareous bedrocks only. Although litter decomposition did not change over all years in siliceous sites with increasing climate drought stress, it increased during wet years in the dry continental sites only, in particular in the Pyrenees where interannual climate variability was higher. Together our results strongly support the physical hypothesis and not the chemical hypothesis. We argue that the chemical hypothesis was proposed based on studies only conducted in low elevation wet temperate sites comparing mull humus types from deciduous forests on calcareous soils to mor and moder humus types from evergreen forests on highly sandy siliceous bedrocks. Our study conducted on a wider range of climate and bedrock conditions bring strong evidence that litter decomposition is rather dependent on the physical characteristics of the bedrocks.
•Litterbag experiment conducted in 70 sites from the Alps and the Pyrenees.•Litter decomposition is higher on siliceous than calcareous soils in dry climates.•Soil pH does not explain differences in litter decomposition.
•Multiple types of evidence yield an overview of the disturbance regime in Dinaric Mountain forests.•Variation in regime components from multiple agents results in a complex disturbance ...regime.•Intermediate severity disturbance events are characteristic of the regime.
Quantitative descriptions of natural disturbance regimes are lacking for temperate forest regions in Europe, primarily because a long history of intensive land-use has been the overriding driver of forest structure and composition across the region. The following contribution is the first attempt to comprehensively describe the natural disturbance regime of the dominant forest communities in the Dinaric Mountain range, with an emphasis on the range of natural variability of regime components for the main disturbance agents. Compared to other forest regions in Europe, the mountain range has a history of less intensive forest exploitation and provides a suitable record of natural disturbance processes. Our synthesis is based on multiple types of evidence, including meteorological information, historical documentation, evidence from old-growth remnants, and salvage logging data from National forest inventories. Taken together, the results show that no single disturbance agent dominates the regime in the dominant forest types (i.e. beech and mixed beech-fir forests), and any given agent exhibits remarkable variation in terms of severity and spatial extent both within and among individual disturbance events. Thunderstorm winds cause the most severe damage (i.e. near stand replacement), but blowdown patches are typically limited to stand-scales (e.g. 10s of ha). Ice storms and heavy snow typically cause intermediate severity damage and affect much larger areas (e.g. 100s of km2). A notable exception was the 2014 ice storm, which was nearly an order of magnitude larger and more severe than any other event recorded in the synthesis. Severe and prolonged periods of drought have occurred several times over the past century, and along with secondary insect damage (e.g. bark beetles), have caused episodes of forest decline. Overall, our synthesis indicates that on top of the background of relatively continuous gap dynamics, stand-scale intermediate severity events are an important part of the regime; these events likely have rotation periods that are less than the lifespan of a tree cohort (e.g. several centuries) and create canopy openings large enough to alter successional trajectories.
The Kunming-Montreal Global Biodiversity Framework recognised the urgency of taking action to conserve intraspecific genetic diversity (IGD) as an insurance against habitat degradation and ...environmental change. Recent work suggests that 90–99 % of IGD should be conserved to safeguard viability of future generations.
Here, we addressed such a conservation issue in three forest tree species in Italy: silver fir (Abies alba Mill.), Heldreich's pine (Pinus heldreichii H. Christ), and pedunculate oak (Quercus robur L.). We used microsatellite markers to measure IGD of 36 (A. alba), 15 (P. heldreichii) and 25 (Q. robur) natural sites, including several putative glacial refugia. We developed a spatial conservation planning (SCP) analysis to quantify the genetic irreplaceability of each site and identify the minimum set coverage ensuring IGD protection. Finally, we compared SCP results with the contributions to allelic diversity within and between sites, total allelic diversity and private allelic richness.
We found that between 44 % and 73 % of sites were required to conserve 90–99 % of the alleles, and that this conservation effort held even when targeting lower percentages of alleles to protect (50–75 %). Glacial refugia were often included in the minimum set coverage, confirming biogeographical expectations. Finally, sites with high genetic irreplaceability were found to have higher private allelic richness on average. These results are discussed in the light of the biogeographic history of the species studied and the current policies for the conservation of forest genetic resources.
•Intraspecific genetic diversity (IGD) must be protected to avoid extirpation.•Spatial conservation planning of IGD in forest trees from a multi-refugial area.•½ to ¾ of sites are to be conserved to achieve adequate protection for IGD.•Setting unambitious conservation targets has limited impact on conservation costs.•Glacial refugia are priority candidates for maximising IGD conservation.
In the context of global decline in old‐growth forest, historical ecology is a valuable tool to derive insights into vegetation legacies and dynamics and develop new conservation and restoration ...strategies. In this cross‐disciplinary study, we integrate palynology (Lago del Pesce record), history, dendrochronology, and historical and contemporary land cover maps to assess drivers of vegetation change over the last millennium in a Mediterranean mountain forest (Pollino National Park, southern Italy) and discuss implications in conservation ecology. The study site hosts a remnant beech–fir (Fagus sylvatica–Abies alba) mixed forest, a priority habitat for biodiversity conservation in Europe. In the 10th century, the pollen record showed an open environment that was quickly colonized by silver fir when sociopolitical instabilities reduced anthropogenic pressures in mountain forests. The highest forest cover and biomass was reached between the 14th and the 17th centuries following land abandonment due to recurring plague pandemics. This rewilding process is also reflected in the recruitment history of Bosnian pine (Pinus heldreichii) in the subalpine elevation belt. Our results show that human impacts have been one of the main drivers of silver fir population contraction in the last centuries in the Mediterranean, and that the removal of direct human pressure led to ecosystem renovation. Since 1910, the Rubbio State Forest has locally protected and restored the mixed beech–fir forest. The institutions in 1972 for the Rubbio Natural Reserve and in 1993 for Pollino National Park have guaranteed the survival of the silver fir population, demonstrating the effectiveness of targeted conservation and restoration policies despite a warming climate. Monitoring silver fir populations can measure the effectiveness of conservation measures. In the last decades, the abandonment of rural environments (rewilding) along the mountains of southern Italy has reduced the pressure on ecosystems, thus boosting forest expansion. However, after four decades of natural regeneration and increasing biomass, pollen influx and forest composition are still far from the natural attributes of the medieval forest ecosystem. We conclude that long‐term forest planning encouraging limited direct human disturbance will lead toward rewilding and renovation of carbon‐rich and highly biodiverse Mediterranean old‐growth forests, which will be more resistant and resilient to future climate change.
•Silver fir occupy sites with raised water table at north-eastern limit of its distribution.•Black alder outcompete silver fir from the most wet parts of forest stand.•In mixed lowland forest fir ...seedling establishment is limited by seeds availability.•Less intensive cuttings aimed at promotion of fir sapling are better than salvage cuttings.
Silver fir (Abies alba Mill.) is a typical mountain tree species, but it can also grow in lowlands below 300 m a.s.l., where it must coexist with a variety of competitors. Much less were studied in the northeastern part of its natural range, where annual precipitation varies between 500 and 600 mm, distinctly less than in the mountains. The general objective of this study was to investigate the regeneration process of silver fir in mixed lowland forests in regions with water shortage. The evaluation of spatial relationships between old, dominant and newly established trees was used to clarify the mechanisms of tree species coexistence. The study was conducted in managed and protected stands in Janów Forests in the southeastern part of Poland. We selected mature forest stands (oldest trees older than 100 years) representing three types of mixed stands in nature reserves, as well as in comparable managed forests. Data was collected on 500 m2 sample plots established randomly in each stand category (20 sample plots × 6 stands). Silver fir as the target tree showed statistically significant segregation from other species in 15 cases. Small silver fir trees were associated with large pines in two forest types and segregated form large firs in one. The spatial relationships between fir and black alder were negative. Black alder was the second species with the most statistically significant spatial relationships with other tree species (11 cases). Alders separated from large pines and large firs and from medium and small silver firs. Strong segregation of medium-sized hornbeam from the small class of silver fir was observed. Our study showed that silver fir natural regeneration and colonization of new areas at the northeastern limit of its range are limited by low seed rain density, interactions with other competing species, and local site conditions. Areas with moderately high water table are potentially suitable for fir regeneration, but black alder has a competitive advantage in wetter parts of forest stand. The major advantage of such sites is the absence of beech populations, which are an important competitor to fir in the central European mountains, but a likely equally important limitation would be the presence of hornbeam populations. Our results suggest that less intensive cuttings at carefully selected places are more conducive to fir regeneration than more intensive salvage cuttings.
•Silver fir forests show different degrees of dieback in the western Pyrenees.•Dieback and growth decline were caused by late-summer drought stress.•The Vaganov-Shashkin (VS) model simulated ...silver-fir growth and identified its climatic constraints.•Simulated growth series showed a decoupling with respect the climate baseline.•The VS model allows identifying vulnerable forests and early signals of dieback.
Assessing tree growth patterns and deviations from expected climate baselines across wide environmental gradients is fundamental to determine forest vulnerability to drought. This need is particularly compelling for the southernmost limit of the tree species distribution where hot droughts often trigger forest dieback processes. This is the case of some silver fir (Abies alba) populations located in southwestern Europe (Spanish Pyrenees) which present ongoing dieback processes since the 1980s. We sampled 21 silver fir stands showing different dieback intensity, assessed using defoliation levels, quantified their growth patterns and characterized their responses to climate. Then, we assessed growth deviations from climatic predictions using the process-based Vaganov-Shashkin (VS) growth model. The forests showing most intense dieback, i.e. highest defoliation levels, were mainly located in low-elevation sites of the western Pyrenees. Trees in these stands displayed the lowest growth rates and the highest year-to-year variability in growth and their growth was limited by late-summer evaporative demand. In eastern and central Pyrenees, we detected a mild growth limitation by low soil moisture during the late growing season and positive growth recovery in recent years with respect to a climate baseline. Decreasing growth trajectories were the most common pattern, while rising trends were common in stands with low dieback in eastern and central Pyrenees. Our results portend systematic spatial variability of growth trends across the Pyrenean silver fir populations forming the south-western distribution limit of the species in Europe. Decoupling of growth between eastern and western populations observed in the recent decades suggests contrasting vulnerability to climate change, and more importantly, the decoupling of growth patterns in western clusters could be used as an early-warning signal of impending dieback. Consequently, we foresee future dieback events to have more detrimental effects in the western compared with the eastern Pyrenees.
Drought-induced forest dieback is causing reductions in productivity, increasing tree mortality and impairing terrestrial carbon uptake worldwide. However, the role played by long-term nutrient ...imbalances during drought-induced dieback is still unknown. To improve our knowledge on the relationships between dieback and nutrient imbalances, we analysed wood anatomical traits (tree-ring width and wood density), soil properties and long-term chemical information in tree-ring wood (1900–2010) by non-destructive Micro X-ray fluorescence (μXRF) and destructive (ICP-OES) techniques. We studied two major European conifers with ongoing drought-induced dieback in mesic (Abies alba, silver fir) and xeric (Pinus sylvestris, Scots pine) sites. In each site we compared coexisting declining (D) and non-declining (ND) trees. We used dendrochronology and generalized additive and linear mixed models to analyse trends in tree-ring nutrients and their relationships with wood traits. The D trees presented lower growth and higher minimum wood density than ND trees, corresponding to a smaller lumen area of earlywood tracheids and thus a lower theoretical hydraulic conductivity. These differences in growth and wood-anatomy were more marked in silver fir than in Scots pine. Moreover, most of the chemical elements showed higher concentrations in D than in ND trees during the last two-five decades (e.g., Mn, K and Mg), while Ca and Na increased in the sapwood of ND trees. The Mn concentrations, and related ratios (Ca:Mn, Mn:Al and P:Mn) showed the highest differences between D and ND trees for both tree species. These findings suggest that a reduced hydraulic conductivity, consistent with hydraulic impairment, is affecting the use of P in D trees, making them more prone to drought-induced damage. The retrospective quantifications of Mn ratios may be used as early-warning signals of impending dieback.
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•110-Year long series of wood-chemistry to understand drought-induced dieback.•Declining trees showed lower growth and higher minimum wood density than non-declining trees.•Alterations in wood nutrient concentrations are coherent with hydraulic failure hypothesis.•Mn, Ca:Mn and P:Mn rings content may serve as early-warning signals of dieback.
Some disturbances can drive ecological systems to abrupt shifts between alternative stages (tipping points) when critical transitions occur. Drought‐induced tree death can be considered as a ...nonlinear shift in tree vigour and growth. However, at what point do trees become predisposed to drought‐related dieback and which factors determine this (tipping) point? We investigated these questions by characterizing the responses of three tree species, silver fir (Abies alba), Scots pine (Pinus sylvestris) and Aleppo pine (Pinus halepensis), to a severe drought event. We compared basal area increment (BAI) trends and responses to climate and drought in declining (very defoliated and dying) vs. non‐declining (slightly or not defoliated) trees by using generalized additive mixed models. Defoliation, BAI and sapwood production were related to functional proxies of tree vigour measured at the onset and end of the drought (non‐structural carbohydrate concentrations, needle N content and C isotopic discrimination, presence of wood‐inhabiting fungi). We evaluated whether early warning signals (increases in synchronicity among trees or in autocorrelation and standard deviation) could be extracted from the BAI series prior to tree death. Declining silver fir and Scots pine trees showed less growth than non‐declining trees one to three decades, respectively, before the drought event, whereas Aleppo pines showed growth decline irrespective of tree defoliation. At the end of the drought period, all species showed increased defoliation and a related reduction in the concentration of sapwood soluble sugars. Defoliation was constrained by the BAI of the previous 5 years and sapwood production. No specific wood‐inhabiting fungi were found in post‐drought declining trees apart from blue‐stain fungi, which extensively affected damaged Scots pines. Declining silver firs showed increases in BAI autocorrelation and variability prior to tree death. Synthesis. Early warning signals of drought‐triggered mortality seem to be species specific and reflect how different tree species cope with drought stress. Highly correlated declining growth patterns during drought can serve as a signal in silver fir, whereas changes in the content of sapwood soluble sugars are suitable vigour proxies for Scots and Aleppo pines. Longer growth and defoliation series, additional vigour parameters and multi‐species comparisons are required to understand and predict drought‐induced tree death.
•Recruitment of beech, spruce and fir in uneven-aged stands studied on 200 m2 plots.•Optimal and critical levels of 18 influential factors determined with the Tobit model.•Stand basal area and the ...proportion of target tree species are key factors.•Stocking control is efficient for regulating recruitment of spruce and beech.•Fir recruitment is weakly limited by stand density and largely stochastic.
Tree recruitment models are important for predicting the dynamics of uneven-aged forests. Previous studies of recruitment of European beech, Norway spruce and silver fir have shown different ecological amplitudes of these species. However, recruitment in uneven-aged stands and the values of environmental factors at which the greatest and poorest recruitment can be expected remain poorly explained. The main objectives of this study were to 1) explain how 39 stand, site and climatic factors and their interactions influence the number of recruited trees in uneven-aged forests; 2) determine the optimal and critical ranges of influential factors, including stand basal area, number of trees, proportion of tree species, shade casting, soil pH, site productivity, temperature and precipitation; and 3) estimate the maximum expected response of recruitment to changes in stand density while controlling for the effect of other limiting factors. A Tobit censored regression model was used to consider that the observed range of the number of recruited trees is censored at zero. The models were parametrized and validated using 30,963 forest inventory plots (200 m2 each) in uneven-aged forests in Slovenia. The models, which used 9 stand, 6 site and 3 climatic factors, explained 15 %, 10 % and 8 % of the total variation of the number of recruited spruce, fir and beech, respectively. Stand structure was the most important factor, with stand basal area (BA) and the proportion of the studied species having the greatest effect. Site factors including soil pH and rockiness were important for fir recruitment. The number of recruited beech and spruce was positively influenced by decadal precipitation. Higher temperatures decreased recruitment of spruce. Beech was the only species sensitive to shade casting. Recruitment of beech was higher if shade was imposed by tree species other than beech. The optimal and critical ranges of limiting factors differ between species. The model suggests that the optimal stand basal area for recruitment of beech is ≤ 19 m2/ha, which is higher than that for spruce (≤ 16 m2/ha) but lower than that for fir (≤ 28 m2/ha). The maximum predicted response of the studied species to changes in stand basal area shows that stand density control is efficient for regulating recruitment of spruce and beech, but not for fir. The suggested sensitivities and threshold values may be used in individual tree growth models or simulation–optimization studies in support of forest management decisions.
Improving our understanding of the potential of forest adaptation is an urgent task in the light of predicted climate change. Long‐term alternatives for susceptible yet economically important tree ...species such as Norway spruce (Picea abies) are required, if the frequency and intensity of summer droughts will continue to increase. Although Silver fir (Abies alba) and Douglas fir (Pseudotsuga menziesii) have both been described as drought‐tolerant species, our understanding of their growth responses to drought extremes is still limited. Here, we use a dendroecological approach to assess the resistance, resilience, and recovery of these important central Europe to conifer species the exceptional droughts in 1976 and 2003. A total of 270 trees per species were sampled in 18 managed mixed‐species stands along an altitudinal gradient (400–1200 m a.s.l.) at the western slopes of the southern and central Black Forest in southwest Germany. While radial growth in all species responded similarly to the 1976 drought, Norway spruce was least resistant and resilient to the 2003 summer drought. Silver fir showed the overall highest resistance to drought, similarly to Douglas fir, which exhibited the widest growth rings. Silver fir trees from lower elevations were more drought prone than trees at higher elevations. Douglas fir and Norway spruce, however, revealed lower drought resilience at higher altitudes. Although the 1976 and 2003 drought extremes were quite different, Douglas fir maintained consistently the highest radial growth. Although our study did not examine population‐level responses, it clearly indicates that Silver fir and Douglas fir are generally more resistant and resilient to previous drought extremes and are therefore suitable alternatives to Norway spruce; Silver fir more so at higher altitudes. Cultivating these species instead of Norway spruce will contribute to maintaining a high level of productivity across many Central European mountain forests under future climate change.
The two extreme droughts in 1976 and 2003 affected negatively the radial growth response of Norway spruce, Silver and Douglas fir in the Black forest at all elevations. The 1976 drought had a less pronounced effect than the 2003 summer drought; however, firs were noticeably more resistant and resilient to extreme drought than spruce. Spruce was the most affected species when comparing performances of drought indices, and Silver fir the least affected. Douglas fir showed consistently the highest growth rates.
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