Quantifying the impacts of land cover change on gross primary productivity globally.

Historically, humans have cleared many forests for agriculture. While this substantially reduced ecosystem carbon storage, the impacts of these land cover changes on terrestrial gross primary productivity (GPP) have not been adequately resolved yet. Here, we combine high-resolution datasets of satellite-derived GPP and environmental predictor variables to estimate the potential GPP of forests, grasslands, and croplands around the globe. With a mean GPP of 2.0 kg C m-2 yr-1 forests represent the most productive land cover on two thirds of the total area suitable for any of these land cover types, while grasslands and croplands on average reach 1.5 and 1.8 kg C m-2 yr-1, respectively. Combining our potential GPP maps with a historical land-use reconstruction indicates a 4.4% reduction in global GPP from agricultural expansion. This land-use-induced GPP reduction is amplified in some future scenarios as a result of ongoing deforestation (e.g., the large-scale bioenergy scenario SSP4-3.4) but partly reversed in other scenarios (e.g., the sustainability scenario SSP1-1.9) due to agricultural abandonment. Comparing our results to simulations from state-of-the-art Earth System Models, we find that all investigated models deviate substantially from our estimates and from each other. Our maps could be used as a benchmark to reduce this inconsistency, thereby improving projections of land-based climate mitigation potentials.

Climate-change-driven growth decline of European beech forests.

The growth of past, present, and future forests was, is and will be affected by climate variability. This multifaceted relationship has been assessed in several regional studies, but spatially resolved, large-scale analyses are largely missing so far. Here we estimate recent changes in growth of 5800 beech trees (Fagus sylvatica L.) from 324 sites, representing the full geographic and climatic range of species. Future growth trends were predicted considering state-of-the-art climate scenarios. The validated models indicate growth declines across large region of the distribution in recent decades, and project severe future growth declines ranging from -20% to more than -50% by 2090, depending on the region and climate change scenario (i.e. CMIP6 SSP1-2.6 and SSP5-8.5). Forecasted forest productivity losses are most striking towards the southern distribution limit of Fagus sylvatica, in regions where persisting atmospheric high-pressure systems are expected to increase drought severity. The projected 21st century growth changes across Europe indicate serious ecological and economic consequences that require immediate forest adaptation.

The European Forest Condition Monitor: Using Remotely Sensed Forest Greenness to Identify Hot Spots of Forest Decline.

Forest decline, in course of climate change, has become a frequently observed phenomenon. Much of the observed decline has been associated with an increasing frequency of climate change induced hotter droughts while decline induced by flooding, late-frost, and storms also play an important role. As a consequence, tree mortality rates have increased across the globe. Despite numerous studies that have assessed forest decline and predisposing factors for tree mortality, we still lack an in-depth understanding of (I) underlying eco-physiological mechanisms, (II) the influence of varying environmental conditions related to soil, competition, and micro-climate, and (III) species-specific strategies to cope with prolonged environmental stress. To deepen our knowledge within this context, studying tree performance within larger networks seems a promising research avenue. Ideally such networks are already established during the actual period of environmental stress. One approach for identifying stressed forests suitable for such monitoring networks is to assess measures related to tree vitality in near real-time across large regions by means of satellite-borne remote sensing. Within this context, we introduce the European Forest Condition monitor (EFCM)-a remote-sensing based, freely available, interactive web information tool. The EFCM depicts forest greenness (as approximated using NDVI from MODIS at a spatial resolution of roughly 5.3 hectares) for the pixel-specific growing season across Europe and consequently allows for guiding research within the context of concurrent forest performance. To allow for inter-temporal comparability and account for pixel-specific features, all observations are set in relation to normalized difference vegetation index (NDVI) records over the monitoring period beginning in 2001. The EFCM provides both a quantile-based and a proportion-based product, thereby allowing for both relative and absolute comparison of forest greenness over the observational record. Based on six specific examples related to spring phenology, drought, late-frost, tree die-back on water-logged soils, an ice storm, and windthrow we exemplify how the EFCM may help identifying hotspots of extraordinary forest greenness. We discuss advantages and limitations when monitoring forest condition at large scales on the basis of moderate resolution remote sensing products to guide users toward an appropriate interpretation.

Excess forest mortality is consistently linked to drought across Europe.

Pulses of tree mortality caused by drought have been reported recently in forests around the globe, but large-scale quantitative evidence is lacking for Europe. Analyzing high-resolution annual satellite-based canopy mortality maps from 1987 to 2016 we here show that excess forest mortality (i.e., canopy mortality exceeding the long-term mortality trend) is significantly related to drought across continental Europe. The relationship between water availability and mortality showed threshold behavior, with excess mortality increasing steeply when the integrated climatic water balance from March to July fell below -1.6 standard deviations of its long-term average. For -3.0 standard deviations the probability of excess canopy mortality was 91.6% (83.8-97.5%). Overall, drought caused approximately 500,000 ha of excess forest mortality between 1987 and 2016 in Europe. We here provide evidence that drought is an important driver of tree mortality at the continental scale, and suggest that a future increase in drought could trigger widespread tree mortality in Europe.

A Dynamic Model for Strategies and Dynamics of Plant Water-Potential Regulation Under Drought Conditions.

Vegetation responds to drought through a complex interplay of plant hydraulic mechanisms, posing challenges for model development and parameterization. We present a mathematical model that describes the dynamics of leaf water-potential over time while considering different strategies by which plant species regulate their water-potentials. The model has two parameters: the parameter λ describing the adjustment of the leaf water potential to changes in soil water potential, and the parameter Δψww describing the typical 'well-watered' leaf water potentials at non-stressed (near-zero) levels of soil water potential. Our model was tested and calibrated on 110 time-series datasets containing the leaf- and soil water potentials of 66 species under drought and non-drought conditions. Our model successfully reproduces the measured leaf water potentials over time based on three different regulation strategies under drought. We found that three parameter sets derived from the measurement data reproduced the dynamics of 53% of an drought dataset, and 52% of a control dataset [root mean square error (RMSE) < 0.5 MPa)]. We conclude that, instead of quantifying water-potential-regulation of different plant species by complex modeling approaches, a small set of parameters may be sufficient to describe the water potential regulation behavior for large-scale modeling. Thus, our approach paves the way for a parsimonious representation of the full spectrum of plant hydraulic responses to drought in dynamic vegetation models.

Standardized drought indices in ecological research: Why one size does not fit all.

While we generally agree with Slette et al. (Global Change Biol, 2019), that ecologists 'should do better' when defining drought in ecological studies, we argue against the uncritical use of a standardized drought index (such as the Standardized Precipitation and Evapotranspiration Index, SPEI; Vicente-Serrano et al. J Climate, 23: 1696-1718, 2010), as a stand-alone criterium for quantifying and reporting drought conditions. Specifically, we raise the following issues: (a) standardization can lead to a misrepresentation of actual water supply, especially for moist climates; (b) standardized values are not directly comparable between different reference periods; and finally, (c) spatially coarsely resolved data sources are unlikely to represent site-level water supply. This article is a commentary on Slette et al., 25, 3193-3200; See also the response to this Letter to the Editor by Slette et al., 26, e1-e3.

Geographical adaptation prevails over species-specific determinism in trees' vulnerability to climate change at Mediterranean rear-edge forests.

Climate change may reduce forest growth and increase forest mortality, which is connected to high carbon costs through reductions in gross primary production and net ecosystem exchange. Yet, the spatiotemporal patterns of vulnerability to both short-term extreme events and gradual environmental changes are quite uncertain across the species' limits of tolerance to dryness. Such information is fundamental for defining ecologically relevant upper limits of species tolerance to drought and, hence, to predict the risk of increased forest mortality and shifts in species composition. We investigate here to what extent the impact of short- and long-term environmental changes determines vulnerability to climate change of three evergreen conifers (Scots pine, silver fir, Norway spruce) and two deciduous hardwoods (European beech, sessile oak) tree species at their southernmost limits of distribution in the Mediterranean Basin. Finally, we simulated future forest growth under RCP 2.6 and 8.5 emission scenarios using a multispecies generalized linear mixed model. Our analysis provides four key insights into the patterns of species' vulnerability to climate change. First, site climatic marginality was significantly linked to the growth trends: increasing growth was related to less climatically limited sites. Second, estimated species-specific vulnerability did not match their a priori rank in drought tolerance: Scots pine and beech seem to be the most vulnerable species among those studied despite their contrasting physiologies. Third, adaptation to site conditions prevails over species-specific determinism in forest response to climate change. And fourth, regional differences in forests vulnerability to climate change across the Mediterranean Basin are linked to the influence of summer atmospheric circulation patterns, which are not correctly represented in global climate models. Thus, projections of forest performance should reconsider the traditional classification of tree species in functional types and critically evaluate the fine-scale limitations of the climate data generated by global climate models.

Climatically controlled reproduction drives interannual growth variability in a temperate tree species.

Climatically controlled allocation to reproduction is a key mechanism by which climate influences tree growth and may explain lagged correlations between climate and growth. We used continent-wide datasets of tree-ring chronologies and annual reproductive effort in Fagus sylvatica from 1901 to 2015 to characterise relationships between climate, reproduction and growth. Results highlight that variable allocation to reproduction is a key factor for growth in this species, and that high reproductive effort ('mast years') is associated with stem growth reduction. Additionally, high reproductive effort is associated with previous summer temperature, creating lagged climate effects on growth. Consequently, understanding growth variability in forest ecosystems requires the incorporation of reproduction, which can be highly variable. Our results suggest that future response of growth dynamics to climate change in this species will be strongly influenced by the response of reproduction.

Two centuries of masting data for European beech and Norway spruce across the European continent.

Tree masting is one of the most intensively studied ecological processes. It affects nutrient fluxes of trees, regeneration dynamics in forests, animal population densities, and ultimately influences ecosystem services. Despite a large volume of research focused on masting, its evolutionary ecology, spatial and temporal variability, and environmental drivers are still matter of debate. Understanding the proximate and ultimate causes of masting at broad spatial and temporal scales will enable us to predict tree reproductive strategies and their response to changing environment. Here we provide broad spatial (distribution range-wide) and temporal (century) masting data for the two main masting tree species in Europe, European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) H. Karst.). We collected masting data from a total of 359 sources through an extensive literature review and from unpublished surveys. The data set has a total of 1,747 series and 18,348 yearly observations from 28 countries and covering a time span of years 1677-2016 and 1791-2016 for beech and spruce, respectively. For each record, the following information is available: identification code; species; year of observation; proxy of masting (flower, pollen, fruit, seed, dendrochronological reconstructions); statistical data type (ordinal, continuous); data value; unit of measurement (only in case of continuous data); geographical location (country, Nomenclature of Units for Territorial Statistics NUTS-1 level, municipality, coordinates); first and last record year and related length; type of data source (field survey, peer reviewed scientific literature, gray literature, personal observation); source identification code; date when data were added to the database; comments. To provide a ready-to-use masting index we harmonized ordinal data into five classes. Furthermore, we computed an additional field where continuous series with length >4 yr where converted into a five classes ordinal index. To our knowledge, this is the most comprehensive published database on species-specific masting behavior. It is useful to study spatial and temporal patterns of masting and its proximate and ultimate causes, to refine studies based on tree-ring chronologies, to understand dynamics of animal species and pests vectored by these animals affecting human health, and it may serve as calibration-validation data for dynamic forest models.

Patterns of late spring frost leaf damage and recovery in a European beech (Fagus sylvatica L.) stand in south-eastern Germany based on repeated digital photographs.

Damage by late spring frost is a risk deciduous trees have to cope with in order to optimize the length of their growing season. The timing of spring phenological development plays a crucial role, not only at the species level, but also at the population and individual level, since fresh new leaves are especially vulnerable. For the pronounced late spring frost in May 2011 in Germany, we studied the individual leaf development of 35 deciduous trees (mainly European beech Fagus sylvatica L.) at a mountainous forest site in the Bayerischer Wald National Park using repeated digital photographs. Analyses of the time series of greenness by a novel Bayesian multiple change point approach mostly revealed five change points which almost perfectly matched the expected break points in leaf development: (i) start of the first greening between day of the year (DOY) 108-119 (mean 113), (ii) end of greening, and (iii) visible frost damage after the frost on the night of May 3rd/4th (DOY 123/124), (iv) re-sprouting 19-38 days after the frost, and (v) full maturity around DOY 178 (166-184) when all beech crowns had fully recovered. Since frost damage was nearly 100%, individual susceptibility did not depend on the timing of first spring leaf unfolding. However, we could identify significant patterns in fitness linked to an earlier start of leaf unfolding. Those individuals that had an earlier start of greening during the first flushing period had a shorter period of recovery and started the second greening earlier. Thus, phenological timing triggered the speed of recovery from such an extreme event. The maximum greenness achieved, however, did not vary with leaf unfolding dates. Two mountain ashes (Sorbus aucuparia L.) were not affected by the low temperatures of -5°C. Time series analysis of webcam pictures can thus improve process-based knowledge and provide valuable insights into the link between phenological variation, late spring frost damage, and recovery within one stand.

Uniform climate sensitivity in tree-ring stable isotopes across species and sites in a mid-latitude temperate forest.

Tree-ring stable isotopes, providing insight into drought-induced eco-physiological mechanisms, are frequently used to reconstruct past changes in growing season temperature and precipitation. Their climatic response is, however, still not fully understood, particularly for data originating from non-extreme, mid-latitude environments with differing ecological conditions. Here, we assess the response of δ(13)C, δ(18)O and tree-ring width (TRW) from a temperate mountain forest in the Austrian pre-Alps to climate and specific drought events. Variations in stem growth and isotopic composition of Norway spruce, common beech and European larch from dry, medium and moist sites are compared with records of sunshine, temperature, moisture, precipitation and cloud cover. Results indicate uniform year-to-year variations in δ(13)C and δ(18)O across sites and species, but distinct differences in TRW according to habitat and species. While the climate sensitivity of TRW is overall weak, the δ(13)C and δ(18)O chronologies contain significant signals with a maximum sensitivity to cloud cover changes (r = -0.72 for δ(18)O). The coherent inter-annual isotopic variations are accompanied by substantial differences in the isotopic signatures with offsets up to ∼3‰ for δ(13)C, indicating species-specific physiological strategies and varying water-use efficiencies. During severe summer drought, beech and larch benefit from access to deeper and moist soils, allowing them to keep their stomata open. This strategy is accompanied by an increased water loss through transpiration, but simultaneously enables enhanced photosynthesis. Our findings indicate the potential of tree-ring stable isotopes from temperate forests to reconstruct changes in cloud cover, and to improve knowledge on basic physiological mechanisms of tree species growing in different habitats to cope with soil moisture deficits.

Patterns of drought tolerance in major European temperate forest trees: climatic drivers and levels of variability.

The future performance of native tree species under climate change conditions is frequently discussed, since increasingly severe and more frequent drought events are expected to become a major risk for forest ecosystems. To improve our understanding of the drought tolerance of the three common European temperate forest tree species Norway spruce, silver fir and common beech, we tested the influence of climate and tree-specific traits on the inter and intrasite variability in drought responses of these species. Basal area increment data from a large tree-ring network in Southern Germany and Alpine Austria along a climatic cline from warm-dry to cool-wet conditions were used to calculate indices of tolerance to drought events and their variability at the level of individual trees and populations. General patterns of tolerance indicated a high vulnerability of Norway spruce in comparison to fir and beech and a strong influence of bioclimatic conditions on drought response for all species. On the level of individual trees, low-growth rates prior to drought events, high competitive status and low age favored resilience in growth response to drought. Consequently, drought events led to heterogeneous and variable response patterns in forests stands. These findings may support the idea of deliberately using spontaneous selection and adaption effects as a passive strategy of forest management under climate change conditions, especially a strong directional selection for more tolerant individuals when frequency and intensity of summer droughts will increase in the course of global climate change.