Major tree species of Central European forests differ in their proportion of positive, negative, and nonstationary growth trends.

Temperate forests are undergoing significant transformations due to the influence of climate change, including varying responses of different tree species to increasing temperature and drought severity. To comprehensively understand the full range of growth responses, representative datasets spanning extensive site and climatic gradients are essential. This study utilizes tree-ring data from 550 sites from the temperate forests of Czechia to assess growth trends of six dominant Central European tree species (European beech, Norway spruce, Scots pine, silver fir, sessile and pedunculate oak) over 1990-2014. By modeling mean growth series for each species and site, and employing principal component analysis, we identified the predominant growth trends. Over the study period, linear growth trends were evident across most sites (56% increasing, 32% decreasing, and 10% neutral). The proportion of sites with stationary positive trends increased from low toward high elevations, whereas the opposite was true for the stationary negative trends. Notably, within the middle range of their distribution (between 500 and 700 m a.s.l.), Norway spruce and European beech exhibited a mix of positive and negative growth trends. While Scots pine growth trends showed no clear elevation-based pattern, silver fir and oaks displayed consistent positive growth trends regardless of site elevation, indicating resilience to the ongoing warming. We demonstrate divergent growth trajectories across space and among species. These findings are particularly important as recent warming has triggered a gradual shift in the elevation range of optimal growth conditions for most tree species and has also led to a decoupling of growth trends between lowlands and mountain areas. As a result, further future shifts in the elevation range and changes in species diversity of European temperate forests can be expected.

Lipid biomarkers and stable isotopes uncover paleovegetation changes in extremely species-rich forest-steppe ecosystems, Central Europe.

The historical development of the vegetation of semi-dry grasslands in Central Europe is not satisfactorily understood. Long-term continuity of open vegetation or, conversely, deep-past forest phases are considered possible sources of the current extreme species diversity of these ecosystems. We aimed to reveal the trajectory of paleovegetation development in these ecosystems through detailed analysis of terrestrial in-situ soil geoarchives. We measured the bulk soil carbon and nitrogen contents, lipid molecular distribution, and compound-specific stable carbon and hydrogen isotopic signatures of mid- and long-chain n-alkanes extracted from soil and modern plant material tissues (i.e., deciduous and Pinus leaves and grass/herbaceous species). The C23-C33 n-alkane homologues were identified in soils with different abundances. Normally, C27 and C29 n-alkanes were the most abundant homologues in tree-leaf samples, while grass-derived n-alkanes were mostly C31 and C33 homologues. Soils were largely dominated by C29 and C31 n-alkanes. Odd-numbered C27-C33 soil n-alkane δ13C values ranged from -36.2‰ to -23.2‰, whereas their δ2H values showed a wider range of variability that fluctuated from -224‰ to -172‰. Molecular distribution in combination with radiocarbon analysis of soil organic matter (SOM) and δ13C and δ2H values of n-alkanes revealed a large contribution of C3 trees (both deciduous and coniferous trees/pine trees) as the main source of n-alkanes between the late Pleistocene and early Holocene (ca 15,000-8200 calibrated year before present/cal year BP). A clear shift toward more grassy/herbaceous vegetation was observed from the early Holocene (ca 11,700-8200 cal year BP) onwards. Distribution patterns of lipids and soil geochemical parameters showed that plants are the main source of SOM and that biodegradation and kinetic isotope fractionation are not the main reasons for 13C enrichment in soil profiles. Past C3 vegetation shifts as well as paleoclimate changes (i.e., aridity) can have played a role in the observed 13C depth profiles.

Joint effects of climate, tree size, and year on annual tree growth derived from tree-ring records of ten globally distributed forests.

Tree rings provide an invaluable long-term record for understanding how climate and other drivers shape tree growth and forest productivity. However, conventional tree-ring analysis methods were not designed to simultaneously test effects of climate, tree size, and other drivers on individual growth. This has limited the potential to test ecologically relevant hypotheses on tree growth sensitivity to environmental drivers and their interactions with tree size. Here, we develop and apply a new method to simultaneously model nonlinear effects of primary climate drivers, reconstructed tree diameter at breast height (DBH), and calendar year in generalized least squares models that account for the temporal autocorrelation inherent to each individual tree's growth. We analyze data from 3811 trees representing 40 species at 10 globally distributed sites, showing that precipitation, temperature, DBH, and calendar year have additively, and often interactively, influenced annual growth over the past 120 years. Growth responses were predominantly positive to precipitation (usually over ≥3-month seasonal windows) and negative to temperature (usually maximum temperature, over ≤3-month seasonal windows), with concave-down responses in 63% of relationships. Climate sensitivity commonly varied with DBH (45% of cases tested), with larger trees usually more sensitive. Trends in ring width at small DBH were linked to the light environment under which trees established, but basal area or biomass increments consistently reached maxima at intermediate DBH. Accounting for climate and DBH, growth rate declined over time for 92% of species in secondary or disturbed stands, whereas growth trends were mixed in older forests. These trends were largely attributable to stand dynamics as cohorts and stands age, which remain challenging to disentangle from global change drivers. By providing a parsimonious approach for characterizing multiple interacting drivers of tree growth, our method reveals a more complete picture of the factors influencing growth than has previously been possible.

The historical disturbance regime of mountain Norway spruce forests in the Western Carpathians and its influence on current forest structure and composition.

In order to gauge ongoing and future changes to disturbance regimes, it is necessary to establish a solid baseline of historic disturbance patterns against which to evaluate these changes. Further, understanding how forest structure and composition respond to variation in past disturbances may provide insight into future resilience to climate-driven alterations of disturbance regimes. We established 184 plots (mostly 1000 m2) in 14 primary mountain Norway spruce forests in the Western Carpathians. On each plot we surveyed live and dead trees and regeneration, and cored around 25 canopy trees. Disturbance history was reconstructed by examining individual tree growth trends. The study plots were further aggregated into five groups based on disturbance history (severity and timing) to evaluate and explain its influence on forest structure. These ecosystems are characterized by a mixed severity disturbance regime with high spatiotemporal variability in severity and frequency. However, periods of synchrony in disturbance activity were also found. Specifically, a peak of canopy disturbance was found for the mid-19th century across the region (about 60% of trees established), with the most important periods of disturbance in the 1820s and from the 1840s to the 1870s. Current stand size and age structure were strongly influenced by past disturbance activity. In contrast, past disturbances did not have a significant effect on current tree density, the amount of coarse woody debris, and regeneration. High mean densities of regeneration with height >50 cm (about 1400 individuals per ha) were observed. Extensive high severity disturbances have recently affected Central European forests, spurring a discussion about the causes and consequences. We found some evidence that forests in the Western Carpathians were predisposed to recent severe disturbance events as a result of synchronized past disturbance activity, which partly homogenized size and age structure and made recent stands more vulnerable to bark beetle outbreak. Our data suggest that these events are still part of the range of natural variability. The finding that regeneration density and volume of coarse woody debris were not influenced by past disturbance illustrates that vastly different past disturbance histories are not likely to change the future trajectories of these forests. These ecosystems currently have high ecological resilience to disturbance. In conclusion, we suggest that management should recognize disturbances as a natural part of ecosystem dynamics in the mountain forests of Central Europe, account for their stochastic occurrence in management planning, and mimic their patterns to foster biodiversity in forest landscapes.

High-resolution soil sampling reveals the pattern of biological weathering and soil formation under trees.

Trees contribute to bedrock weathering in a variety of ways. However, evaluating their full impact is complicated by a lack of direct observation of unexposed root systems of individual trees, especially when the scale of the analysis goes down to the level of microbiomes. In the present study, we investigated the contribution of tree root systems to bioweathering and soil production at the macro- and microscale. Soil profiles developed under trees on granite bedrock were investigated in two parts of the Sudety Mountains, SW Poland: the Rudawy Janowickie Mountains, and the Stolowe Mountains. Soil profiles were gradually excavated and soil samples collected from pre-defined positions of the root zone: 1) bulk soil, 2) rhizosphere, 3) cracks, 4) topsoil, and 5) control positions. In total, we analyzed 103 samples for soil chemistry and microbiological activity. In addition, we analyzed 19 samples using XRF (X-ray Fluorescence). Four parent rock samples, in the form of thin-sections, were the subject of mineralogical evaluation. Soil analyses included: total organic carbon (C) and nitrogen (N) content, soil pHH2O, soluble iron (Fed), and aluminum (Ald), non-crystalline (amorphous) iron (Feox), and aluminum (Alox). For microbiological analyses, we used a Biolog (EcoPlate) system to determine the functional diversity of soil microorganisms. We evaluated the results on soil chemistry and microbiological activity statistically by principal component analysis (PCA) and redundancy analysis (RDA). Differences between soil sampling positions were assessed using a non-parametric Kruskal-Wallis (K-W) rank sum test and a post-hoc pairwise Dunn test. Trees developed different root architectures, likely shaped by the depth to bedrock and its pre-existing net of fractures and fissures. Tree roots were able to enter bedrock cracks at one study site (at Pstrazna, Stolowe Mountains). The soil profile was too deep for root system penetration at the second study site (Mt Janska, Rudawy Janowickie Mountains, RJM). The rhizospheric soil along the roots had significantly different chemical properties compared to non-rhizospheric soil types. At Mt. Janska, soil differed from the crack soil in terms of Alox (pHolm-adj. < 0.0006) and Feox (pHolm-adj. < 0.004), and from the bulk soil (pHolm-adj. < 0.02) and topsoil (pHolm-adj. < 0.007). In addition, at Pstrazna, the soil differed from the control soil in terms of C (pHolm-adj. < 0.009) and soil pHH2O (pHolm-adj. < 0.0008) and from the topsoil in terms of soil pHH2O. The highest metabolic activity was in cracks at Mt. Janska and in control samples from Pstrazna. In general, the spatial distribution of soil microbial activity, and the weathering that results from that portion of the soil biome, is spatially heterogeneous and appears to be partially determined by the interaction of root growth and bedrock fracture patterns.

The impact of insect herbivory on biogeochemical cycling in broadleaved forests varies with temperature.

Herbivorous insects alter biogeochemical cycling within forests, but the magnitude of these impacts, their global variation, and drivers of this variation remain poorly understood. To address this knowledge gap and help improve biogeochemical models, we established a global network of 74 plots within 40 mature, undisturbed broadleaved forests. We analyzed freshly senesced and green leaves for carbon, nitrogen, phosphorus and silica concentrations, foliar production and herbivory, and stand-level nutrient fluxes. We show more nutrient release by insect herbivores at non-outbreak levels in tropical forests than temperate and boreal forests, that these fluxes increase strongly with mean annual temperature, and that they exceed atmospheric deposition inputs in some localities. Thus, background levels of insect herbivory are sufficiently large to both alter ecosystem element cycling and influence terrestrial carbon cycling. Further, climate can affect interactions between natural populations of plants and herbivores with important consequences for global biogeochemical cycles across broadleaved forests.

Weathering and soil production under trees growing on sandstones - The role of tree roots in soil formation.

Rock weathering drives both landform formation and soil production/evolution. The less studied biological component of weathering and soil production caused by tree root systems is the main focus of the present study. Weathering by trees, which likely has been important in soil formation since the first trees emerged in the middle and late Devonian, is accomplished through both physical and biological means, like acids excreted by plants and exudates from associated bacterial communities. However, these processes are relatively poorly known. We assessed the impact of tree roots and associated microbiota on the potential level of biological weathering. Three research plots were selected in two sandstone regions in Poland. Two plots were in the Stolowe Mountains (Zlotno, Batorów), a tableland built of Cretaceous sandstones. The third plot (Zegiestów) was in the Sacz Beskidy Mountains, the Carpathians. Soil samples were taken from tree root zones of Norway spruces from predefined sampling positions. Soils from non-tree control positions were also sampled. Soil samples were a subject of laboratory analyses which included the content of Fe and Al (amorphous and labile forms), carbon (C), nitrogen (N), and soil pH. The microbial functional diversity of soil microorganisms was determined using the Biolog (EcoPlate) system. Rock fragments were collected for mineralogical and a subject of optical microscopy and cathodoluminescence analyses in order to examine their mineralogical composition. Significant differences (pHolm-corrected < 0.05) between sample locations were found mostly for the Zegiestów plot: Soils at control positions differed from the crack and bulk soil sample positions in terms of C, N, C/N, and pH. Tree roots were able to develop a great variety of sizes and forms by following the existing net of bedrock discontinuities and hillslope microrelief. They developed along the most accessible surfaces, and caused rockcliff retreat and scree slope formation. These two features can be considered as initial stages of soil production. Trees add to the complexity of the soil system and allow formation of rhizospheric soils, and horizons rich in organic matter which are zones of a high microbial activity. However, as our study shows, rock cracks with roots cannot be considered as zones of microbial weathering. In addition, C content and microbial activity decreases with depth but can stay on a high level along living and dead roots. When entering rock fractures, they change the intensity of biomechanical weathering and soil properties. The highest biological activity of microorganisms was found in the control samples. Overall, tree roots do change the pattern of soil formation and explain the existing pattern of soil chemical properties, microbial activity, and potentially biological weathering intensity, and the intensity of those processes in correlation with root presence varies in space.

Hillslope Processes Affect Vessel Lumen Area and Tree Dimensions.

The height growth of the trees depends on sufficient mechanical support given by the stem and an effective hydraulic system. On unstable slopes, tree growth is affected by soil pressure from above and potential soil erosion from below the position of tree. The necessary stabilization is then provided by the production of mechanically stronger wood of reduced hydraulic conductivity. Unfortunately, the interaction between tree growth (both radial and axial) and stabilization in the soil is still insufficiently understood. Therefore, in this study, we aimed to quantify the impact of hillslope dynamics on the degree of tree growth and hydraulic limitation, and the potential effect on tree height growth and growth plasticity. To evaluate this effect, we took four cores from 80 individuals of Quercus robur and Fraxinus excelsior and measured tree-ring widths (TRWs) and vessel lumen areas (VLAs). The tree heights were evaluated using a terrestrial laser scanner, and local soil depth was measured by a soil auger. Our data showed a significant limitation of the tree hydraulic system related with the formation of eccentric tree-rings. The stem eccentricity decreased with increasing stem diameter, but at the same time, the negative effect of stem eccentricity on conduit size increased with the increasing stem diameter. Even though this anatomical adaptation associated with the effect of stem eccentricity differed between the tree species (mainly in the different degree of limitations in conduit size), the trees showed an increase in the proportion of hydraulically inactive wood elements and a lowered effectiveness of their hydraulic system. In addition, we observed a larger negative effect of stem eccentricity on VLA in Quercus. We conclude that the stabilization of a tree in unstable soil is accompanied by an inability to create sufficiently effective hydraulic system, resulting in severe height-growth limitation. This affects the accumulation of aboveground biomass and carbon sequestration.