Summer drought exposure, stand structure, and soil properties jointly control the growth of European beech along a steep precipitation gradient in northern Germany.

Increasing exposure to climate warming-related drought and heat threatens forest vitality in many regions on earth, with the trees' vulnerability likely depending on local climatic aridity, recent climate trends, edaphic conditions, and the drought acclimatization and adaptation of populations. Studies exploring tree species' vulnerability to climate change often have a local focus or model the species' entire distribution range, which hampers the separation of climatic and edaphic drivers of drought and heat vulnerability. We compared recent radial growth trends and the sensitivity of growth to drought and heat in central populations of a widespread and naturally dominant tree species in Europe, European beech (Fagus sylvatica), at 30 forest sites across a steep precipitation gradient (500-850 mm year-1 ) of short length to assess the species' adaptive potential. Size-standardized basal area increment remained more constant during the period of accelerated warming since the early 1980s in populations with >360 mm growing season precipitation (April-September), while growth trends were negative at sites with <360 mm. Climatic drought in June appeared as the most influential climatic factor affecting radial growth, with a stronger effect at drier sites. A decadal decrease in the climatic water balance of the summer was identified as the most important factor leading to growth decline, which is amplified by higher stem densities. Inter-annual growth variability has increased since the early 1980s, and variability is generally higher at drier and sandier sites. Similarly, within-population growth synchrony is higher at sandier sites and has increased with a decrease in the June climatic water balance. Our results caution against predicting the drought vulnerability of trees solely from climate projections, as soil properties emerged as an important modulating factor. We conclude that beech is facing recent growth decline at drier sites in the centre of its distribution range, driven by climate change-related climate aridification.

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.

Drought stress mitigation by nitrogen in boreal forests inferred from stable isotopes.

Forest growth in most parts of the boreal zone is originally limited by low temperatures and low nitrogen availability. Due to the rapid climate warming at high latitudes, an increasing forest area is switching to drought limitation, especially in continental and southern parts of the boreal forest. Studies addressing this issue were mostly dendrochronological and remote-sensing analyses focusing on climatic effects, but not answering the question whether drought is effective alone or in combination with nitrogen shortage at limiting the forests' productivity and vitality. Here we show in a case study from larch forests of Mongolia with a combination of stable isotope analyses, tree-ring analysis and bioindication of the local variability of livestock densities using epiphytic lichens that, in the studied highly drought-prone forests at the southern fringe of the boreal forest in Inner Asia, the trees' vulnerability to drought is modified by nitrogen fertilization from livestock kept in the vicinity and the edge of the forests. The most likely mechanism behind this drought-nitrogen interaction is the reduction of stomatal conductance, which is known to be induced by low nitrogen levels in plants. Nitrogen fertilization by the livestock could, thus, shorten the times of stomatal closure and thereby increase tree growth, which we measured as radial stem increment. Even though the underlying mechanisms, which were so far examined in angiosperms, should be experimentally tested for conifers, our results indicate that focusing on water alone is not enough to understand the climate change response of drought-limited boreal forests.

Modelling the productivity of Siberian larch forests from Landsat NDVI time series in fragmented forest stands of the Mongolian forest-steppe.

The monitoring of the spatial and temporal dynamics of vegetation productivity is important in the context of carbon sequestration by terrestrial ecosystems from the atmosphere. The accessibility of the full archive of medium-resolution earth observation data for multiple decades dramatically improved the potential of remote sensing to support global climate change and terrestrial carbon cycle studies. We investigated a dense time series of multi-sensor Landsat Normalized Difference Vegetation Index (NDVI) data at the southern fringe of the boreal forests in the Mongolian forest-steppe with regard to the ability to capture the annual variability in radial stemwood increment and thus forest productivity. Forest productivity was assessed from dendrochronological series of Siberian larch (Larix sibirica) from 15 plots in forest patches of different ages and stand sizes. The results revealed a strong correlation between the maximum growing season NDVI of forest sites and tree ring width over an observation period of 20 years. This relationship was independent of the forest stand size and of the landscape's forest-to-grassland ratio. We conclude from the consistent findings of our case study that the maximum growing season NDVI can be used for retrospective modelling of forest productivity over larger areas. The usefulness of grassland NDVI as a proxy for forest NDVI to monitor forest productivity in semi-arid areas could only partially be confirmed. Spatial and temporal inconsistencies between forest and grassland NDVI are a consequence of different physiological and ecological vegetation properties. Due to coarse spatial resolution of available satellite data, previous studies were not able to account for small-scaled land-cover patches like fragmented forest in the forest-steppe. Landsat satellite-time series were able to separate those effects and thus may contribute to a better understanding of the impact of global climate change on natural ecosystems.

Hydraulic architecture and vulnerability to drought-induced embolism in southern boreal tree species of Inner Asia.

The branch xylem of six important Inner Asian southern boreal forest trees was studied for wood-anatomical and hydraulic traits in order to infer the species' drought tolerance from embolism resistance, potential hydraulic conductivity, mean conduit diameters and conduit density. The only studied angiosperm tree, Betula pendula Roth, was much more sensitive to cavitation than all five conifers (evergreen or summer-green), even when using 88% loss of conductivity (P88) in birch, but 50% (P50) in the conifers as critical thresholds. This suggests that pioneer birch forests, which have widely replaced the conifer climax forests after anthropogenic disturbance (e.g., logging, man-made fire), are more vulnerable to climate warming-induced drought than the original conifer forests. In contrast to expectation, the generally more drought-exposed light taiga species (Larix sibirica Ledeb., Pinus sylvestris L.) did not have consistently lower P50 and P88 values than the dark taiga conifers, suggesting that other drought survival traits are equally important. Among the dark-taiga species, only Pinus sibirica Du Tour, but not Abies sibirica Ledeb. and Picea obovata Ledeb., had relatively high P50 values indicating higher vulnerability. In the light-taiga forest, P. sylvestris revealed lower embolism resistance than L. sibirica. In the face of rapid climate warming and drying in Inner Asia, the drought survival strategies of southern boreal tree species deserve further intensive study, which should include other drought survival traits.

Higher climate warming sensitivity of Siberian larch in small than large forest islands in the fragmented Mongolian forest steppe.

Forest fragmentation has been found to affect biodiversity and ecosystem functioning in multiple ways. We asked whether forest size and isolation in fragmented woodlands influences the climate warming sensitivity of tree growth in the southern boreal forest of the Mongolian Larix sibirica forest steppe, a naturally fragmented woodland embedded in grassland, which is highly affected by warming, drought, and increasing anthropogenic forest destruction in recent time. We examined the influence of stand size and stand isolation on the growth performance of larch in forests of four different size classes located in a woodland-dominated forest-steppe area and small forest patches in a grassland-dominated area. We found increasing climate sensitivity and decreasing first-order autocorrelation of annual stemwood increment with decreasing stand size. Stemwood increment increased with previous year's June and August precipitation in the three smallest forest size classes, but not in the largest forests. In the grassland-dominated area, the tree growth dependence on summer rainfall was highest. Missing ring frequency has strongly increased since the 1970s in small, but not in large forests. In the grassland-dominated area, the increase was much greater than in the forest-dominated landscape. Forest regeneration decreased with decreasing stand size and was scarce or absent in the smallest forests. Our results suggest that the larch trees in small and isolated forest patches are far more susceptible to climate warming than in large continuous forests pointing to a grim future for the forests in this strongly warming region of the boreal forest that is also under high land use pressure.

Carbon pool densities and a first estimate of the total carbon pool in the Mongolian forest-steppe.

The boreal forest biome represents one of the most important terrestrial carbon stores, which gave reason to intensive research on carbon stock densities. However, such an analysis does not yet exist for the southernmost Eurosiberian boreal forests in Inner Asia. Most of these forests are located in the Mongolian forest-steppe, which is largely dominated by Larix sibirica. We quantified the carbon stock density and total carbon pool of Mongolia's boreal forests and adjacent grasslands and draw conclusions on possible future change. Mean aboveground carbon stock density in the interior of L. sibirica forests was 66 Mg C ha(-1) , which is in the upper range of values reported from boreal forests and probably due to the comparably long growing season. The density of soil organic carbon (SOC, 108 Mg C ha(-1) ) and total belowground carbon density (149 Mg C ha(-1) ) are at the lower end of the range known from boreal forests, which might be the result of higher soil temperatures and a thinner permafrost layer than in the central and northern boreal forest belt. Land use effects are especially relevant at forest edges, where mean carbon stock density was 188 Mg C ha(-1) , compared with 215 Mg C ha(-1) in the forest interior. Carbon stock density in grasslands was 144 Mg C ha(-1) . Analysis of satellite imagery of the highly fragmented forest area in the forest-steppe zone showed that Mongolia's total boreal forest area is currently 73 818 km(2) , and 22% of this area refers to forest edges (defined as the first 30 m from the edge). The total forest carbon pool of Mongolia was estimated at ~ 1.5-1.7 Pg C, a value which is likely to decrease in future with increasing deforestation and fire frequency, and global warming.

Rapid recovery of stem increment in Norway spruce at reduced SO2 levels in the Harz Mountains, Germany.

Tree-ring width of Picea abies was studied along an altitudinal gradient in the Harz Mountains, Germany, in an area heavily affected by SO(2)-related forest decline in the second half of the 20th century. Spruce trees of exposed high-elevation forests had earlier been shown to have reduced radial growth at high atmospheric SO(2) levels. After the recent reduction of the SO(2) load due to clean air acts, we tested the hypothesis that stem growth recovered rapidly from the SO(2) impact. Our results from two formerly damaged high-elevation spruce stands support this hypothesis suggesting that the former SO(2)-related spruce decline was primarily due to foliar damage and not to soil acidification, as the deacidification of the (still acidic) soil would cause a slow growth response. Increasing temperatures and deposited N accumulated in the topsoil are likely additional growth-promoting factors of spruce at high elevations after the shortfall of SO(2) pollution.

Diverging climate trends in Mongolian taiga forests influence growth and regeneration of Larix sibirica.

Central and semiarid north-eastern Asia was subject to twentieth century warming far above the global average. Since forests of this region occur at their drought limit, they are particularly vulnerable to climate change. We studied the regional variations of temperature and precipitation trends and their effects on tree growth and forest regeneration in Mongolia. Tree-ring series from more than 2,300 trees of Siberian larch (Larix sibirica) collected in four regions of Mongolia's forest zone were analyzed and related to available weather data. Climate trends underlie a remarkable regional variation leading to contrasting responses of tree growth in taiga forests even within the same mountain system. Within a distance of a few hundred kilometers (140-490 km), areas with recently reduced growth and regeneration of larch alternated with regions where these parameters remained constant or even increased. Reduced productivity could be correlated with increasing summer temperatures and decreasing precipitation; improved growth conditions were found at increasing precipitation, but constant summer temperatures. An effect of increasing winter temperatures on tree-ring width or forest regeneration was not detectable. Since declines of productivity and regeneration are more widespread in the Mongolian taiga than the opposite trend, a net loss of forests is likely to occur in the future, as strong increases in temperature and regionally differing changes in precipitation are predicted for the twenty-first century.

Water relations and photosynthetic performance in Larix sibirica growing in the forest-steppe ecotone of northern Mongolia.

Shoot water relations were studied in Siberian larch (Larix sibirica Ledeb.) trees growing at the borderline between taiga and steppe in northern Mongolia. Larix sibirica is the main tree species in these forests covering 80% of Mongolia's forested area. Minimum shoot water potentials (Psi(m)) close to the point of zero turgor (Psi(0)) repeatedly recorded throughout the growing season suggest that the water relations in L. sibirica were often critical. The Psi(m) varied in close relation to the atmospheric vapor pressure deficit, whereas Psi(0) was correlated with monthly precipitation. Young larch trees growing at the forest line to the steppe were more susceptible to drought than mature trees at the same sites. Furthermore, isolated trees growing on the steppe exhibited lower Psi(m) and recovered to a lower degree from drought overnight than the trees at the forest line. Indications of drought stress in L. sibirica were obtained in two study areas in Mongolia's forest-steppe ecotone: one in the mountain taiga of the western Khentey in northernmost Mongolia, the other in the forest-steppe at the southern distribution limit of L. sibirica on Mt. Bogd Uul, southern Khentey. Larix sibirica growing in riverine taiga with contact to the groundwater table was better water-supplied than the larch trees growing at the forest line to the steppe. Larch trees from the interior of light taiga forests on north-facing slopes, however, exhibited more critical water relations than the trees at the forest line. Frequent drought stress in mature trees and even more in young larch trees at the forest-steppe borderline suggests that L. sibirica does not have the potential to encroach on the steppe under the present climate, except in a sequence of exceptionally moist and cool years. A regression of the present borderline between forest and steppe is likely to occur, as average temperatures are increasing everywhere and precipitation is decreasing regionally in Mongolia's taiga forest region. Higher stomatal conductance concomitant to lower Psi(m) in trees of northern-slope forests compared to trees from the forest line to the steppe may be the result of a recent increase in drought intensity that affects better drought-adapted trees at the forest edge less than the trees in the forest interior. We conclude that drought is a key factor explaining the forest-steppe borderline in northern Mongolia. The proportion of forests within the present vegetation pattern of forests on north-facing slopes and the grasslands on south-facing slopes in Mongolia's forest-steppe ecotone is not likely to increase under the present climate, but may decrease with increasing aridity due to global warming.