Forest management and species identity shape climate sensitivity and drought resilience in temperate mountain forests.

Consistent growth responses of silver fir (Abies alba Mill.) and European beech (Fagus sylvatica L.) to drought in mixed and monospecific forests: Insights from Central European forests

The increasing frequency and severity of extreme events, such as drought, are expected to disturb forest ecosystems worldwide. Stand structure, including tree size diversity, may play a crucial role in how forests respond to these changes. This study examines the effect of tree size diversity on the drought resilience of silver fir ( Abies alba Mill.) using data from 138 circular plots in even- and uneven-aged mountain stands across Germany, Italy, and Poland. Increment cores from nearly 600 trees were evaluated to calculate complementary resilience indices. Generalized linear mixed-effect models were fitted to assess how tree size diversity affects individual tree growth response to drought stress under varying environmental conditions, mediated by admixture of broadleaf species, stand density, and individual tree size. We found that tree size diversity improves the growth response of silver fir to drought stress, expressed by higher resistance and stress-driven deviation (SDD) indices, in more water-limited sites. However, this benefit diminishes or becomes slightly negative with increasing climate humidity. Similarly, smaller trees demonstrated higher resistance and SDD, although these effects also weakened with more favorable water conditions. The admixture of broadleaf species and stand density did not mediate the impact of tree size diversity on growth resilience. Our results are in line with the stress-gradient hypothesis. Ecophysiological adaptations to prevailing conditions can lead to different competition regimes (symmetric and asymmetric), causing variations in the impact of stand structure on drought resilience. Since tree size diversity is crucial in water-limited environments, it can be considered a strategic forest management tool for adapting silver fir-dominated forests to anticipated global changes. • Analysis of drought response patterns in Abies alba Mill. mountain forests in Europe. • Silver fir drought resilience is modulated by tree size diversity. • Higher benefits of tree size diversity in water-limited sites. • Smaller trees have greater drought resilience in drier sites. • The effect of tree size and size diversity diminished with higher water availability.

ABSTRACTSilver fir (Abies alba Mill.) is one of the most valuable conifer trees in Europe for historical and economic reasons. Growing interest in the species, along with the recognition that silver fir was subjected to inappropriate silvicultural measures during the 20th century due a lack of understanding of its ecology, have spurred recent studies from various disciplines which have yielded new insights into the gene ecology, history, ecology, structure and dynamics of fir populations. In this paper, we provide a multidisciplinary, synoptical review discussing the prospects for silver fir forestry in the 21st century. We describe the main characteristics of the history and ecology of silver fir, with a particular emphasis on gene ecology and the structure and dynamics of fir populations with respect to mixed stands and fir management. Fir is a fundamental species for maintaining high biodiversity in forested ecosystems because of its shade tolerance, plasticity to environmental conditions and ability to coexist with many tree species. Silver fir ecotypes show remarkable variation in features such as frost hardiness, drought resilience and shade tolerance. Fir can create stands of heterogeneous stand structure, from one-storied through to multi-storied structure and finally to selection structure. Natural regeneration of fir will be scarce if fir stands are managed inappropriately through the application of clear cutting, a short regeneration period and abrupt removal of shelter trees. Climate change is expected to reduce the abundance and distribution range of silver fir. Improvements in forest management treatments are recommended to improve silver fir conservation.

Глобальні зміни клімату вже зараз суттєво змінили умови ведення лісового господарства в Українських Карпатах. Зокрема внаслідок масового всихання ялини багато підприємств змушені переорієнтовувати свої лісівничі заходи на вирощування товарної деревини інших порід. Виявлено два основних напрями кліматогенних змін у лісах регіону: зміна едафотопів лісових ділянок у напрямі покращення їхньої родючості з одночасним зменшенням вологості, зміна конкурентоздатності різних деревних видів, зміна породного складу лісів. Найбільші зміни відбуваються в ялинових лісах, де місце ялини, що всихає, природним шляхом займають ялиця і бук, однак під час створення лісових культур вибір головної породи може бути й іншим. Тому актуальним є виявлення закономірностей цих змін для правильного вибору головних порід за типами лісу і висотними рослинними смугами. Тенденції динаміки лісорослинних умов і породного складу лісів регіону були ідентифіковані за відомчими даними метеостанцій та лісовпорядкування. Встановлено, що кліматогенні зміни лісів Українських Карпат відбуваються у таких напрямах: а) дубово-букові ліси трансформуються в буково-дубові зі зміною гігротопу зі «свіжого» на «сухий»; б) буково-ялицеві ліси – в ялицево-букові зі зміною гігротопу з «вологого» на «свіжий»; в) грабово-букові ліси – в грабово-дубово-букові зі зміною гігротопу з «вологого» на «свіжий»; г) смереково-буково-ялицеві ліси – в смереково-ялицево-букові або ялицево-букові зі зміною гігротопу з «вологого» на «свіжий»; д) буково-ялицево-смерекові ліси – в смереково-ялицево-букові або ялицево-букові зі зміною гігротопу з «сирого» і «вологого» на «свіжий»; е) щільнодернинні луки та полонини інтенсивно заростають смерековими, ялівцевими та душекієвими угрупованнями. Констатується нагальна потреба в лісотипологічному картуванні лісогосподарських підприємств Карпатського регіону під час проведення їх повторного лісовпорядкування.

With climate change, the frequency and duration of droughts are increasing, strongly impacting forest ecosystems. Therefore, a thorough understanding of the factors influencing tree response to drought is needed. Particularly, it remains unclear how competition and species diversity influence the drought sensitivity of tree species. Thinning (i.e., lowering competition) and mixing species (i.e., increasing diversity) are two common forest management practices that are thought to help forests to cope with droughts. However, their actual effects are still controversial. We sampled and measured tree-ring widths of 401 conifer trees with a wide range of competition and species diversity in their neighborhood across a bioclimatic gradient in Switzerland. Based on mixed-effects models and correlations, we examined the climate sensitivity of silver fir, European larch and Douglas fir and analyzed how competition and species diversity affect their radial growth and drought sensitivity. Silver fir was the least sensitive species to temperature, precipitation, and climatic water balance. When we analyzed the combined effects of the climatic water balance, competition and species diversity on tree radial growth over the past 20 years, we found that competition usually had a negative effect on radial growth of the three species, while species diversity had no effect. However, when focusing on drought resilience, competition had a negative effect for silver fir only, and a more mixed neighborhood enhanced the drought resilience of Douglas fir. Larch showed a higher drought sensitivity than silver fir and Douglas fir. At most sites, radial growth of all species recovered within two years after the severe droughts of 2003 and 2018. Overall, our results suggest that competition and species diversity have minor effects on the drought resilience of silver fir, larch and Douglas fir in Switzerland.

Key messageWe found a significant increase in the latewood density of European beech, and a decrease in the latewood and mean wood density of silver fir and Norway spruce in European mountain forests over the period 1901–2016. In the past century, drought did not directly influence the wood density trend of the three studied species. However, for both fir and spruce, drought indirectly affected the mean wood density via changes in the latewood to earlywood ratio, i.e., in the case of extreme drought, trees with high values of latewood to earlywood ratio experienced a slight attenuation in the declining trend of their mean wood density.ContextCentury-long wood density measurements can provide novel information on tree response to climate change and the carbon sequestration potential of forest ecosystems. Still, the knowledge about long-term changes in wood density of European beech (Fagus sylvatica L.), silver fir (Abies alba Mill.), and Norway spruce (Picea abies (L.) H.Karst.) in European mountain forests needs to be further explored.AimsWe assessed long-term changes in tree-ring mean wood density, earlywood density, and latewood density in trees of the three species between 1901 and 2016. We investigated the influence of endogenous factors (i.e., tree-ring width, current tree diameter, and latewood to earlywood ratio) and drought events on wood density.MethodsIn total, 150 tree cores were sampled from mountain forests in Bulgaria, Bosnia and Herzegovina, Slovenia, Switzerland, and Germany. The mean, early, and latewood density of these samples were measured with the LIGNOSTATION™ system. To address our research aims, we applied a linear mixed-effect modelling approach using the data from 101 correctly cross-dated cores that spanned the entire period of analysis.ResultsIn the absence of drought, the latewood density of European beech increased by 7.1%, the late and mean wood density of silver fir decreased by 16.8% and 11.0%, respectively, and the late and mean wood density of Norway spruce decreased by 16.1% and 7.2%, respectively, between 1901–2016. In the past century, drought influenced the trends of wood density through an effect mediated by changes in the latewood to earlywood ratio. Specifically, in cases of extreme drought, silver fir and Norway spruce trees with a latewood to earlywood ratio value 50% higher than the median experience a slight attenuation in the declining trend of their mean wood density, making the negative impact of drought marginally less severe.ConclusionsOur findings have significant implications for the accuracy of carbon stock assessments, national greenhouse gas inventories, and the utilization of wood from the three species. Given the fact that changes in wood density follow species-specific patterns and the expectation of more frequent drought events in Europe, in the future, it is essential to build further tree-ring density time series for other species and sites to improve our understanding of how climate change alters wood density and carbon sequestration of forest ecosystems.

Respective importance of ecological conditions and stand composition on Abies alba Mill. dominant height growth

Importance of tree species size dominance and heterogeneity on the productivity of spruce-fir-beech mountain forest stands in Europe

Silver fir (Abies alba Mill.) is one of the most valuable and productive tree species across European mountains, that accomplish multiple economic, protective and ecologic functions. Alongside spruce (Picea abies (L.) Karst) and beech (Fagus sylvatica L.), silver fir is a characteristic species for the Romanian Carpathians. Although silver fir tree is recommended for the diversification of forests in order to increase the resistance to climate change, it is very sensitive to climatic excesses, especially those that proceed rapidly. Therefore, the aim of this study is to investigate both the environmental conditions and stand characteristics of fir from five mountain ranges of the Romanian Carpathians. The study is based on data recorded over a period of 10 years (1990–2000). As such, a total of 77,251 stands that occupy 211,954 hectares have been investigated in regard to silver fir behaviour. MATLAB scripts were used for analysing consistent data volumes as well as the impact of eight factors on the silver fir productivity (altitude, field aspect, field slope, soil type, participation percentage, road distance, structure and consistency). Our analysis has revealed that higher silver fir productivity is found at altitudes of up to 1200 m, on mid and upper slopes, on NW field aspects, on eutric cambisols and dystric cambisols, with a 10–20% participation in stand composition and in relatively-even aged stands with a full consistency. This study offers valuable insights for forest managers that require comprehensive information in adopting effective strategies to enhance forest resilience under 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.

Beech (Fagus sylvatica) and silver fir (Abies alba) are often cultivated in mixed stands and, hence, compete for water and nutrients. Besides nitrogen (N), also phosphorus (P) is an important nutrient for growth and development. Beech trees in Central Europe grow on both P-poor and P-rich soils, thereby showing similar growth and low variation in foliar P. The central aim of the present study was to test the hypothesis that variations in foliar P contents of beech are driven by seasonal changes rather than by the competition with silver fir. It was further hypothesized that P contents in silver fir needles depend on needle age and forest site. To test these hypotheses, P contents and P fractions, i.e. organic-bound P (Porg) and inorganic phosphate P (Pi), were measured in the foliage of beech trees from pure beech and mixed beech/silver fir plots as well as in needles of silver fir of the mixed plots. The forest sites investigated are located in Central Europe in the Black Forest, Germany, and in Croatia near the south-eastern distribution limit of beech and are all poor in plant-available soil P. The analyses showed that the main driver of P contents and P fractions in beech leaves at all forest sites is the season and that competition with silver fir had no effect. Hence, the present results demonstrate the high plasticity of beech trees to adapt to both poor plant-available soil P and competition with silver fir. Total P contents of silver fir needles were higher at the Croatian site compared to the Black Forest sites and originated from higher foliar Pi contents. One third of the P present in current-year needles in late summer was remobilized and exported until the needles reached the age of 1 year. The difference in P contents between current-year and 1-year-old needles can be seen as the amount of P resorbed from 1-year-old needles in summer during the generation of new needles to support the P demand of current-year needles for growth and development.

Summary There is a rising interest in the role of species diversity in ecosystem functioning and services, including productivity. Yet, how the diversity–productivity relationship depends on species identity and abiotic conditions remains a challenging issue. We analysed mixture effects on species productivity along site productivity gradients, calculated from a set of abiotic factors, in two biogeographic contexts (highlands and lowlands). We compared the productivity of 5 two‐species mixtures (i.e. 10 cases of mixed species) with that of monocultures of the same species. Five main European tree species were considered: sessile oak (Quercus petraea Liebl.), Scots pine (Pinus sylvestris L.), European beech (Fagus sylvatica L.), silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) H. Karst). Our data set was compiled from the 2006 to 2010 French National Forest Inventory data base and covers 2361 plots including pure and mixed stands. Overall productivity of mixtures in highlands, that is European beech–Norway spruce, European beech–silver fir and to a lesser extent, silver fir–Norway spruce, was found to be higher than expected from the productivity of corresponding monospecific stands. Overyielding was mainly due to European beech for the first two mixtures and to silver fir for the third one. No effect of mixture was found for sessile oak–Scots pine and sessile oak–European beech stands in lowlands. Overyielding of sessile oak mixed with Scots pine was not strong enough to significantly increase overall stand productivity. Overyielding of European beech was balanced by an underyielding of sessile oak. The mixture effect changed along site productivity gradients for six cases out of the 10 studied, with a stronger and positive effect on sites with low productivity. The magnitude of this change along site productivity gradients varied up to 89% depending on the tree species. Synthesis. The nature of species interaction in mixtures with regard to productivity changes with species assemblage and abiotic conditions. Overyielding is strongest when species grow in highlands on less productive sites. A negative link between mixture effect and site productivity was found, in line with the stress‐gradient hypothesis.

Climate growth limitations of European beech and silver fir along the Carpathian arc – the recent state and future prospects

Key messageDrought severely worsened till 2100 and eventually outplayed growth-enhancing CO2 fertilization turning productivity gains into losses for beech and fir. Most scenarios generated notable losses in profitability but economic tipping points were later than for productivity due to lag effects related to discounting. Time mixture of fir and shortening rotation can counteract economic risks under climate change, but requires early admixture and moderate establishment costs.ContextAdaptation strategies to climate change (CC) such as establishing mixed forests are often based on ecological understanding while economic rationale is often disregarded.AimsThis paper studies CC uncertainty on productivity and profitability of European beech (Fagus sylvatica L.) and Silver fir (Abies alba Mill.). Besides, the economic consequences to actively adapt beech forests by admixing Silver fir are investigated.MethodsWe used the process-based forest growth model GOTILWA + to simulate RCP2.6, RCP4.5 and RCP8.5 climatic projection by the MPI-ESM-LR global circulation model (MPI-ESM-LR) with the CO2 fertilization effect (eCO2) switched on and off. We analysed the sensitivity of the land expectation value (LEV) on CC and economic parameters.ResultsCC initially increased productivity, but declined after a tipping point (2040–2070) and later also profitability (2045–2100). RCP8.5 had positive, RCP2.6 negative and RCP4.5 neutral effects on LEV. Switching off eCO2 turned RCP8.5 from the most profitable to the least profitable scenario and the opposite for RCP2.6. CC generally reduced optimal rotation (Ropt) being scenario dependant, but comparatively more for fir than beech. Admixing fir created an economic benefit when implemented before stand age 50 of beech. This benefit was nullified with protection costs for browsing control (fencing or tree shelters).ConclusionsEconomic parameters (not CC) were the major source of uncertainty stemming from discounting factors and establishment costs. Admixture of fir and shortening rotation can provide a solution to tackle economic and climate uncertainties, but requires early admixture and browsing control.

Ongoing climate change and associated extreme events strongly impact the growth and vitality of forest ecosystems in Europe. Because of its’ high drought sensitivity, European beech, which is considered as climax tree species in large parts of Central Europe, may specifically suffer. Hence, recent studies increasingly focus on the resistance and resilience of beech growth to climate change. Intra-specific variations in growth responses by comparing different beech provenances, however, received less attention, as did the question whether provenance selection can be used to mitigate potential future negative impacts of climate change. Therefore, we here investigated 24 provenances belonging to the International Beech Provenance Trial growing at three sites in Germany along a latitudinal gradient (study sites are referred to as ’North’, ‘Center’, ‘South’). Specifically, we compared tree-ring width (TRW), diameter breast height (DBH), climate-growth relationships, as well as drought resistance and resilience in the extreme years 2003 and 2018. Large differences in growth performance were observed between the three study sites. At site North, beech trees showed the highest DBH and TRW. Tree growth was predominantly driven by previous-year October and current-year winter temperature, whereas growth at sites Center and South was significantly impacted by summer SPEI and constrained by precipitation in late winter and early June, respectively. Overall, drought responses in 2003 were less variable than in 2018. We found increasing resistance and decreasing resilience from the wetter North to the drier South, but with minimal differences between the Center and the South. Whereas differences between study sites were large, provenance differentiation within sites was comparably low, substantiating that beech is a highly plastic tree species. Even though some provenances were found to perform slightly better or worse, differences were not statistically significant and show unclear patterns. Hence, we conclude that climate change will affect beech forests in Europe mainly depending upon site conditions, and that provenance selection may not ensure superior growth performance.