Species- and trait-environment linkages in forest plant communities continue to be a frequent topic in ecological research. We studied the dependence of floristic and functional trait composition on ...environmental factors, namely local soil properties, overstory characteristics, climatic parameters and other abiotic and biotic variables. The study area comprised 50 monitoring plots across Slovenia, belonging to the EU ICP Forests monitoring network. Vegetation was surveyed in accordance with harmonized protocols, and environmental variables were either measured or estimated during vegetation sampling. Significant predictors of species composition were identified by canonical correspondence analysis. Correlations between plant traits, i.e. plant growth habit, life form, flowering features and CSR signature, were examined with fourth-corner analysis and linear regressions. Our results show that variation in floristic composition was mainly explained by climatic parameters (mean annual temperature, mean annual precipitation), soil properties (pH) and tree layer-dependent light conditions. Trait composition was most closely related with tree layer characteristics, such as shade-casting ability (SCA, a proxy for light availability in the understory layer), tree species richness and tree species composition. Amongst soil properties, total nitrogen content and soil texture (proportion of clay) were most frequently correlated with different species traits or trait states. The CSR signature of herb communities was associated with tree layer SCA, soil pH and mean annual temperature. The floristic composition of the studied herb-layer vegetation depended on temperature and precipitation, which are likely to be influenced by ongoing climate change (warming and drying). Trait composition exhibited significant links to tree layer characteristics and soil conditions, which are in turn directly modified by forest management interventions.
•Post-harvest herb layer demography was studied over a 6-year period (2012–2018).•The degree of species compositional changes was largest immediately after forest harvest.•Compositional shifts were ...significantly controlled by pre-harvest biotic (species richness) and abiotic (sinkhole geomorphology) conditions.•The floristic dissimilarity tended to decrease but was modulated by the demographic type of persistent residents.
Timber harvesting constitutes extensive anthropogenic disturbance in temperate forests, producing a broad range of ecological impacts that most often enhance the demographic processes of vegetation. This study monitored post-harvest herb layer demography over a 6-year period in mesic Dinaric fir-beech forests (Slovenia), a vascular plant diversity hotspot among European forests. Three experimental harvesting intensities, i.e. full harvest (FH), partial harvest (PH) and a control treatment (NH), were each applied over a circular area of 4000 m2 and replicated three times at each of three study sites. Vegetation sampling was conducted before harvesting (in 2012), and two (2014) and six (2018) years following it, in a 400 m2 circular plot positioned in the centre of each treatment area. We focused on identifying general demographic patterns and evaluating the effects of various pre-disturbance abiotic and biotic predictors on compositional responses to disturbance. Two years after harvest (2012–2014), compositional shifts were larger than those in the next 4-year period (2014–2018), confirming the general theoretical prediction that species turnover rate decreases along a successional gradient. The degree of compositional shifts in gaps (FH) and thinned stands (PH) was affected by local abiotic factors (geomorphology of karst sinkholes) and community attributes, such as pre-harvest species richness. Our results indicate that compositional stability is positively associated with pre-disturbance species richness. Over the whole study period, increases in plot-level species richness (alpha diversity) and overall enrichment of the species pool (gamma diversity) were accompanied by compositional convergence, i.e. a decline in floristic dissimilarity (beta diversity) between and within study sites. However, the observed tendency towards homogenization was rather weak and would have been even more pronounced if the demographic type of persistent resident species had not shown a high degree of resistance, thus leaving a strong imprint on post-harvest vegetation development by preserving the forest characteristics of the herb layer community.
•Maximum daily temperatures in cutting-induced canopy gaps were up to 5.9°C higher compared to closed stands.•High management intensity resulted in larger temporal variability (daily and seasonal ...fluctuations) of microclimate.•Topography (within-sinkhole position) effects on microclimate were increased by more intensive cutting.•After the cutting, south-facing slopes within karst sinkholes exhibited the highest air temperatures and lowest humidity.•Microsites with extreme microclimatic conditions (high VPD) are likely unfavourable for typical forest plant species.
Forest microclimate is strongly affected by local topography and management activities, as these directly alter overstory structure. In the present work we analysed the dependence of observed patterns of spatio-temporal microclimatic variations on topographic, canopy- and management-related factors. A forestry experiment was conducted in managed fir-beech forests in the Dinaric Mountains (Slovenia), which are characterized by rugged karstic terrain with numerous sinkholes. In 2012, cutting treatments representing a range in the intensity of overstory removal were performed: uncut controls (CON), 50% cut of stand growing stock (intermediate management intensity – IMI) and 100% cut (high management intensity – HMI) creating 0.4 ha canopy gaps. Fine-scale variation in aspect and slope and its effects on microclimate was assessed by comparing central, south-facing and north-facing within-sinkhole positions. We measured microclimatic variables (air temperature – T, relative humidity – RH) 0.5 m above the ground over three consecutive post-treatment growing seasons. Microclimatic variables showed an increase (T and vapour pressure deficit – VPD) or decrease (RH) with management intensity. Daily Tmax and VPDmax in HMI treatment were up to 5.9°C (on average 3.5°C) and up to 1.4 kPa (on average 0.6 kPa) higher than those in CON treatment, respectively, whereas daily RHmin was up to 22.7 (on average 13.0) percentage points lower. Regarding intra-seasonal patterns, microclimatic differences between treatments were largest during the summer. South-facing plots in the HMI treatment overall exhibited the most extreme conditions, i.e. the highest Tmax and lowest RHmin. Differences in microclimate between treatments were strongly modulated by canopy cover. The results also suggest that overstory removal increases topography-mediated variation in microclimate, as evidenced by significant differences in T, RH and VPD along the fine-scale topographic gradient within the created canopy gaps.
•We studied the relationship between plant functional traits and niche parameters.•Niche optimum and niche width were explained by similar set of traits.•Relative importance of traits differed ...markedly between the two niche parameters.•Traits on average explained 20% (optima) and 8% (widths) of the variation.•Trait–niche relationship needs further quantification using broader spatial scale.
The ecological niche is one of the central concepts in plant ecology. Understanding which biological traits influence plant niches remains limited, preventing large-scale generalizations. Using a representative pool of 94 herb-layer species frequently occurring in the Slovenian forest vegetation types and an extensive suite of 28 plant functional traits, we tested whether traits serve as predictors for the optimum and width of plant species ecological niche. Niche optimum (mean) and niche width (standard deviation) of each species were derived from community-level ecological indicator values for six environmental gradients, i.e., light, temperature, continentality, moisture, soil reaction and nutrients. We investigated relationships between niche parameters and functional traits through a random forest analysis to account for relatively high trait correlations. Our results suggest that niche optimum and width of forest plant species are related to their functional traits. The two niche parameters were best explained by similar set of traits; however, the relative importance of traits differed substantially. Traits associated with disturbances (frequency and severity), plant dispersal (seed mass, dispersal syndrome), leaf economics spectrum (specific leaf area) and life strategy (CSR scores) showed the highest overall significance in predicting niche optimum and width. Functional traits were, on average, better predictors for niche optimum (average variance explained across all six environmental factors: 20.2%) than for niche width (average variance explained: 7.7%). Intraspecific trait variability, not considered in this study, likely plays an important role in case of niche width. The analyses suggest that, while not all traits impact niche parameters to the same degree, it is crucial to consider traits representing different ecological dimensions and revealing leading patterns of trait coordination. We recommend that the relative importance of traits for species niche parameters should be tested on a larger spatial scale using broader pool of forest understory plants across Europe.
Introduction
Herbaceous plant species constitute an essential element of the flora of European beech (
Fagus sylvatica
) forests. There is increasing evidence that rapidly changing climate is likely ...to modify the spatial distribution of plant species. However, we lack understanding of the impact that climate change might have on beech forest herbs across the European continent. We investigated the possible effects of predicted increasing rates of global warming and altered precipitation regimes on 71 forest herbs closely associated with beech forests, but with varying biogeographic and climatic niche attributes.
Methods
By using a total of 394,502 occurrence records and an ensemble of species distribution models (SDMs), we quantified the potential current distribution and future (2061-2080) range shifts in climatic suitability (expressed as occurrence probability, OP) according to two climate change scenarios (moderate SSP2-4.5 and severe SSP5-8.5).
Results
Overall, precipitation of the warmest quarter and temperature seasonality were the most influential predictors in shaping current distribution patterns. For SSP5-8.5 scenario, all studied species experienced significant reductions (52.9% on average) in the total size of highly suitable areas (OP >0.75). However, the magnitude and directions of changes in the climatic suitability were highly species-specific; few species might even increase OP in the future, particularly in case of SSP2-4.5 scenario. The SDMs revealed the most substantial decline of climatic suitability at the trailing edges in southern Europe. We found that climatic suitability is predicted to show unidirectional northward shift and to move toward higher elevations. The gain/loss ratio was generally higher for narrow-ranged species compared to widespread taxa.
Discussion
Our findings are contextualized with regards to potential confounding factors (dispersal limitation, microclimatic buffering) that may mitigate or accelerate climate change impacts. Given the low long-distance migration ability, many beech forest herbs are unlikely to track the velocity with which macroclimatic isotherms are moving toward higher latitudes, making this species group particularly vulnerable to climate change.
Forest herb-layer vegetation responds sensitively to environmental conditions. This paper compares drivers of both taxonomic, i.e., species richness, cover and evenness, and functional herb-layer ...diversity, i.e., the diversity of clonal, bud bank and leaf-height-seed plant traits. We investigated the dependence of herb-layer diversity on ecological determinants related to soil properties, climatic parameters, forest stand characteristics, and topographic and abiotic and biotic factors associated with forest floor structure. The study was conducted in different forest types in Slovenia, using vegetation and environmental data from 50 monitoring plots (400 m2 each) belonging to the ICP Forests Level I and II network. The main objective was to first identify significant ecological predictors and then quantify their relative importance. Species richness was strongly determined by forest stand characteristics, such as richness of the shrub layer, tree layer shade-casting ability as a proxy for light availability and tree species composition. It showed a clear positive relation to soil pH. Variation in herb-layer cover was also best explained by forest stand characteristics and, to a lesser extent, by structural factors such as moss cover. Species evenness was associated with tree species composition, shrub layer cover and soil pH. Various ecological determinants were decisive for the diversity of below-ground traits, i.e., clonal and bud bank traits. For these two trait groups we observed a substantial climatic signal that was completely absent for taxonomy-based measures of diversity. In contrast, above-ground leaf-height-seed (LHS) traits were driven exclusively by soil reaction and nitrogen availability. In synthesis, local stand characteristics and soil properties acted as the main controlling factors for both species and trait diversity in herb-layer communities across Slovenia, confirming many previous studies. Our findings suggest that the taxonomic and functional facets of herb-layer vegetation are mainly influenced by a similar set of ecological determinants. However, their relative importance varies among individual taxonomy- and functional trait-based diversity measures. Integrating multi-faceted approaches can provide complementary information on patterns of herb-layer diversity in European forest plant communities.
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•Thick clay-silty Luvisols in the bottom of dolines have high SOC storage capacity.•Past agricultural land-use presents long-term impact on SOC stock in dolines.•Succession plots in ...dolines show the highest SOC storage capacity (130 t/ha).•Soil organic carbon stock in dolines was double the amount on the levelled plateau.
The soil organic carbon (SOC) was determined in soils of enclosed karst depressions (dolines) (NW Dinaric Mts.) to define their potential for organic carbon sequestration. SOC was measured in the forest, succession (scrubland), and grassland plots at the bottom of dolines at four depths (0–40 cm) and for 40 cm soil layer SOC stock was calculated. We demonstrated that the prevailing fine soil fractions, the C/N ratio and soil thickness play a positive role in the storage capacity of SOC in dolines regardless land use type. Grasslands have the lowest SOC storage capacity (106 t/ha/40 cm), while the highest SOC storage capacity is in succession plots (130 t/ha/40 cm). The last are covered by shrub communities dominated by Prunus spinosa, forming dense communities, and are typical of abandoned croplands or meadows that have been impacted by high levels of nutrients during cultivation phase. At this stage, there is no additional nutrient input in studied plots, which lowers the nutrient content and increases the C/N ratio. C/N ratio is the highest in the forest, where SOC stock capacity is 116 t/ha/40 cm. Given the trend towards the abandonment of agricultural land at Kras Plateau (SW Slovenia), we can expect more overgrowth of dolines, and thus an increase in carbon stocks and stabilization of organic carbon in forest soils. In contrary, we noticed the alarming decrease in grasslands and increase in urban land. The SOC storage in 2020 was for 12,538 t/ha/40 cm lower than in 2002. Although grasslands showed the lowest SOC storage, their contribution to total SOC storage in dolines is very important. Since there is a lack of studies on carbon stocks in doline soils, our research is of great importance and a novelty and gives an important background for further research on SOC stock in karst landscapes worldwide.
Forest understory vegetation is largely influenced by disturbances and given local abiotic conditions. Our research focuses on the early response of understory vegetation to various forest management ...intensities in Dinaric fir-beech forests in Slovenia: (i) control, (ii) 50% cut of stand growing stock, and (iii) 100% cut of stand growing stock. Apart from identifying overstory removal effects, we were interested in fine-scale variation of understory vegetation and environmental determinants of its species composition. Vegetation was sampled within 27 karst sinkholes, which represent a dominant landform in studied forests. Within each sinkhole, five sampling plots, varying in slope aspect (centre, north, east, south, west), were established (135 in total), where pre-treatment (in 2012) and post-treatment (in 2014) floristic surveys were conducted. The sampled understory species were characterized in terms of Ellenberg’s indicator values (EIVs) and plant functional traits (plant height, seed mass, specific leaf area, leaf dry matter content). Diversity metrics (species richness, total cover, Shannon index) increased in plots where the silvicultural measures were applied. Tree species richness also increased in 100% cutting. A redundancy analysis revealed that species composition was related to environmental variables, which are directly influenced by management interventions (overstory canopy cover, microclimate—maximum daily temperature, soil properties—thickness of organic soil layer) as well as by topographic factors (slope inclination and surface rockiness). EIVs for light were significantly affected by treatment intensity, whereas soil-related EIVs (moisture, reaction, nutrients) depended more on the within-sinkhole position. Canopy gaps, compared with uncut control plots, hosted a higher number of colonizing species with a higher plant height and smaller seeds, while leaf traits did not show a clear response. We found a negative correlation between pre-treatment species (functional) richness and post-treatment shifts in floristic (functional) composition. Plots with higher richness exhibited smaller changes compared with species-poor communities. Incorporating different perspectives, the results of this study offer valuable insights into patterns of understory vegetation response to forest management in fir-beech forests.
Forest soils harbor hyper-diverse microbial communities which fundamentally regulate carbon and nutrient cycling across the globe. Directly testing hypotheses on how microbiome diversity is linked to ...forest carbon storage has been difficult, due to a lack of paired data on microbiome diversity and in situ observations of forest carbon accumulation and storage. Here, we investigated the relationship between soil microbiomes and forest carbon across 238 forest inventory plots spanning 15 European countries. We show that the composition and diversity of fungal, but not bacterial, species is tightly coupled to both forest biotic conditions and a seven-fold variation in tree growth rates and biomass carbon stocks when controlling for the effects of dominant tree type, climate, and other environmental factors. This linkage is particularly strong for symbiotic endophytic and ectomycorrhizal fungi known to directly facilitate tree growth. Since tree growth rates in this system are closely and positively correlated with belowground soil carbon stocks, we conclude that fungal composition is a strong predictor of overall forest carbon storage across the European continent.
Background and purpose: The objectives of our study were to identify invasive alien plant species (IAS) in the main Natura 2000 forest habitat types (FHT) along the Mura River in Slovenia, and to ...estimate their abundance and cover. The aim of our study was to find out a) What kinds of IAS appear in the research forests? b) What is their frequency and cover percentage? c) Are individual IAS more attached to some FHT than others? d) What is the correlation between the cover of IAS and the tree layer cover?Materials and methods: We analysed the fidelity of invasive plant species to individual FHT. The studied FHTs along the Mura River were following: 91E0* (Alluvial forests with Alnus glutinosa and Fraxinus excelsior), 91F0 (Riparian mixed forests of Quercus robur, Ulmus laevis and Ulmus minor, Fraxinus excelsior or Fraxinus angustifolia, along the great rivers) and 91L0 (Illyrian oak-hornbeam forests). Two forest areas of about 600 ha were studied in total.Results: 16 IAS were recorded in studied FHTs. Some species, like Robinia pseudacacia, Impatiens glandulifera, I. parviflora, Fallopia japonica (incl. F. x bohemica), Erigeron annuus, Ambrosia artemisiifolia, Amorpha fruticosa, Conyza canadensis and Juncus tenuis occur only in one or two FHTs, while some species can be found in all studied FHTs (like Solidago sp.). We have found that the most threatened forests are those with prevailing Salix alba, Alnus glutinosa, Fraxinus angustifolia and Ulmus laevis, tree species, which grow closest to the river and on the wettest sites.Conclusions: The increasing presence of IAS in the study areas seriously affect natural regeneration, stability, and continuity of floodplain FHTs. Therefore, some measures and guidelines for managing of these forests are suggested in this study.
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