Research highlights: Interaction effects of coniferous on deciduous species have been investigated before the background of climate change. Background and objectives: The cultivation of European beech (Fagus sylvatica L.) in mixed stands has currently received attention, since the future performance of beech in mid-European forest monocultures in a changing climate is under debate. We investigated water relations and nitrogen (N) nutrition of beech in monocultures and mixed with silver-fir (Abies alba Mill.) in the Black Forest at different environmental conditions, and in the Croatian Velebit at the southern distribution limit of beech, over a seasonal course at sufficient water availability. Material and methods: Water relations were analyzed via δ13C signatures, as integrative measures of water supply assuming that photosynthesis processes were not impaired. N nutrition was characterized by N partitioning between soluble N fractions and structural N. Results: In the relatively wet year 2016, water relations of beech leaves, fir needles and roots differed by season, but generally not between beech monocultures and mixed cultivation. At all sites, previous and current year fir needles revealed significantly lower total N contents over the entire season than beech leaves. Fir fine roots exhibited higher or similar amounts of total N compared to needles. Correlation analysis revealed a strong relationship of leaf and root δ13C signatures with soil parameters at the mixed beech stands, but not at pure beech stands. While glutamine (Gln) uptake capacity of beech roots was strongly related to soil N in the monoculture beech stands, arginine (Arg) uptake capacities of beech roots were strongly related to soil N in mixed stands. Conclusions: Leaf N contents indicated a facilitative effect of silver-fir on beech on sites where soil total N concentrations where low, but an indication of competition effect where it was high. This improvement could be partially attributed to protein contents, but not to differences in uptake capacity of an individual N source. From these results it is concluded that despite similar performance of beech trees at the three field sites investigated, the association with silver-fir mediated interactive effects between species association, climate and soil parameters even at sufficient water supply.

Research highlights: The admixture of fir to pure European beech hardly affected soil-atmosphere CH4 and N2O fluxes but increased soil organic carbon (SOC) stocks at a site in the Black Forest, Southern Germany. Background and objectives: Admixing deep-rooting silver fir has been proposed as a measure to increase the resilience of beech forests towards intensified drying-wetting cycles. Hence, the goal of this study was to quantify the effect of fir admixture to beech forests on the soil-atmosphere-exchange of greenhouse gases (GHGs: CO2, CH4 and N2O) and the SOC stocks by comparing pure beech (BB) and mixed beech-fir (BF) stands in the Black Forest, Germany. Materials and methods: To account for the impact of drying-wetting events, we simulated prolonged summer drought periods by rainout shelters, followed by irrigation. Results: The admixture of fir to pure beech stands reduced soil respiration, especially during autumn and winter. This resulted in increased SOC stocks down to a 0.9 m depth by 9 t C ha−1 at BF. The mixed stand showed an insignificantly decreased sink strength for CH4 (−4.0 under BB and −3.6 kg C ha−1 year−1 under BF). With maximal emissions of 25 µg N m−2 h−1, N2O fluxes were very low and remained unchanged by the fir admixture. The total soil GHG balance of forest conversion from BB to BF was strongly dominated by changes in SOC stocks. Extended summer droughts significantly decreased the soil respiration in both BB and BF stands and increased the net CH4 uptake. Conclusions: Overall, this study highlights the positive effects of fir admixture to beech stands on SOC stocks and the total soil GHG balance. In view of the positive impact of increased SOC stocks on key soil functions such as water and nutrient retention, admixing fir to beech stands appears to be a suitable measure to mitigate climate change stresses on European beech stands.

Drought-sensitive European beech forests are increasingly challenged by climate change. Admixing other, preferably more deep-rooting, tree species has been proposed to increase the resilience of beech forests to drought. This diversification of beech forests might also affect soil organic carbon (SOC) and total nitrogen (TN) stocks that are relevant for a wide range of soil functions and ecosystem services, such as water and nutrient retention, filter functions and erosion control. Since information of these effects is scattered, our aim was to synthesize results from studies that compared SOC/TN stocks of beech monocultures with those of beech stands mixed with other tree species as well as monocultures of other tree species. We conducted a meta-analysis including 38 studies with 203, 220, and 160 observations for forest floor (i.e., the organic surface layer), mineral soil (0.5 m depth) and the total soil profile, respectively. Monoculture conifer stands had higher SOC stocks compared to monoculture beech in general, especially in the forest floor (up to 200% in larch forests). In contrast, other broadleaved tree species (oak, ash, lime, maple, hornbeam) showed lower SOC stocks in the forest floor compared to beech, with little impact on total SOC stocks. Comparing mixed beech-conifer stands (average mixing ratio with regard to number of trees 50:50) with beech monocultures revealed significantly higher total SOC stocks of around 9% and a smaller increase in TN stocks of around 4%. This equaled a SOC accrual of 0.1 Mg ha−1 yr−1. In contrast, mixed beech-broadleaved stands did not show significant differences in total SOC stocks. Conifer admixture effects on beech forest SOC were of additive nature. Admixing other tree species to beech monoculture stands was most effective to increase SOC stocks on low carbon soils with a sandy texture and nitrogen limitation (i.e., a high C/N ratio and low nitrogen deposition). We conclude that, with targeted admixture measures of coniferous species, an increase in SOC stocks in beech forests can be achieved as part of the necessary adaptation of beech forests to climate change.

Hydraulic redistribution (HR) of water from wet- to dry-soil zones is suggested as an important process in the resilience of forest ecosystems to drought stress in semiarid and tropical climates. Scenarios of future climate change predict an increase of severe drought conditions in temperate climate regions. This implies the need for adaptations of locally managed forest systems, such as European beech (Fagus sylvatica L.) monocultures, for instance, through the admixing of deep-rooting silver fir (Abies alba Mill.). We designed a stable-isotope-based split-root experiment under controlled conditions to test whether silver fir seedlings could perform HR and therefore reduce drought stress in neighboring beech seedlings. Our results showed that HR by silver fir does occur, but with a delayed onset of three weeks after isotopic labelling with 2H2O (δ2H ≈ +6000‰), and at low rates. On average, 0.2% of added ²H excess could be recovered via HR. Fir roots released water under dry-soil conditions that caused some European beech seedlings to permanently wilt. On the basis of these results, we concluded that HR by silver fir does occur, but the potential for mitigating drought stress in beech is limited. Admixing silver fir into beech stands as a climate change adaptation strategy needs to be assessed in field studies with sufficient monitoring time.

Heatwaves combined with drought affect tree functioning with as yet undetermined legacy effects on carbon (C) and nitrogen (N) allocation.We continuously monitored shoot and root gas exchange, δ 13 CO 2 of respiration and stem growth in well-watered and drought-treated Pinus sylvestris (Scots pine) seedlings exposed to increasing daytime temperatures (max. 42°C) and evaporative demand. Following stress release, we used 13 CO 2 canopy pulse-labeling, supplemented by soil-applied 15 N, to determine allocation to plant compartments, respiration and soil microbial biomass (SMB) over 2.5 wk.Previously heat-treated seedlings rapidly translocated 13 C along the long-distance transport path, to root respiration (R root ; 7.1 h) and SMB (3 d). Furthermore, 13 C accumulated in branch cellulose, suggesting secondary growth enhancement. However, in recovering drought-heat seedlings, the mean residence time of 13 C in needles increased, whereas C translocation to R root was delayed (13.8 h) and 13 C incorporated into starch rather than cellulose. Concurrently, we observed stress-induced low N uptake and aboveground allocation.C and N allocation during early recovery were affected by stress type and impact. Although C uptake increased quickly in both treatments, drought-heat in combination reduced the above-belowground coupling and starch accumulated in leaves at the expense of growth. Accordingly, C allocation during recovery depends on phloem translocation capacity.