Effects of plant diversity on productivity strengthen over time due to trait-dependent shifts in species overyielding.

Plant diversity effects on community productivity often increase over time. Whether the strengthening of diversity effects is caused by temporal shifts in species-level overyielding (i.e., higher species-level productivity in diverse communities compared with monocultures) remains unclear. Here, using data from 65 grassland and forest biodiversity experiments, we show that the temporal strength of diversity effects at the community scale is underpinned by temporal changes in the species that yield. These temporal trends of species-level overyielding are shaped by plant ecological strategies, which can be quantitatively delimited by functional traits. In grasslands, the temporal strengthening of biodiversity effects on community productivity was associated with increasing biomass overyielding of resource-conservative species increasing over time, and with overyielding of species characterized by fast resource acquisition either decreasing or increasing. In forests, temporal trends in species overyielding differ when considering above- versus belowground resource acquisition strategies. Overyielding in stem growth decreased for species with high light capture capacity but increased for those with high soil resource acquisition capacity. Our results imply that a diversity of species with different, and potentially complementary, ecological strategies is beneficial for maintaining community productivity over time in both grassland and forest ecosystems.

Mother trees, altruistic fungi, and the perils of plant personification.

There are growing doubts about the true role of the common mycorrhizal networks (CMN or wood wide web) connecting the roots of trees in forests. We question the claims of a substantial carbon transfer from 'mother trees' to their offspring and nearby seedlings through the CMN. Recent reviews show that evidence for the 'mother tree concept' is inconclusive or absent. The origin of this concept seems to stem from a desire to humanize plant life but can lead to misunderstandings and false interpretations and may eventually harm rather than help the commendable cause of preserving forests. Two recent books serve as examples: The Hidden Life of Trees and Finding the Mother Tree.

Pioneer tree species accelerate restoration of tree-related microhabitats in 50-year-old reserves of Bialowieza Forest, Poland.

Retention of structural elements such as deadwood and habitat trees at the level of forest stands has been promoted to integrate biodiversity conservation into multiple-use forest management. The conservation value of habitat trees is largely determined by the presence, richness, and abundance of tree-related microhabitats (TreMs). Since TreMs are often lacking in intensively managed forests, an important question of forest conservation is how the abundance and richness of TreMs may be effectively restored. Here, we investigated whether the strict protection of forest through cessation of timber harvesting influenced TreM occurrence at tree and stand levels. For that purpose, we compared four managed and four set-aside stands (0.25 ha each) in the Bialowieza Forest, with identical origin following clear-cuts approximately 100 years ago. We found that the abundance and richness of TreMs on living trees were not significantly different between stands that were either conventionally managed or where active forest management ceased 52 years ago. Yet, our analysis of TreMs on tree species with contrasting life-history traits revealed that short-lived, fast-growing species (pioneers) developed TreMs quicker than longer-lived, slower-growing species. Hence, tree species such as Populus or Betula, which supply abundant and diverse TreMs, can play an important role in accelerating habitat restoration.

Exotic tree species have consistently lower herbivore load in a cross-Atlantic tree biodiversity experiment.

It is commonly expected that exotic plants experience reduced herbivory, but experimental evidence for such enemy release is still controversial. One reason for conflicting results might be that community context has rarely been accounted for, although the surrounding plant diversity may moderate enemy release. Here, we tested the effects of focal tree origin and surrounding tree diversity on herbivore abundance and leaf damage in a cross-Atlantic tree-diversity experiment in Canada and Germany. We evaluated six European tree species paired with six North American congeners in both their native and exotic range, expecting lower herbivory for the exotic tree species in each pair at each site. Such reciprocal experiments have long been called for, but have not been realized thus far. In addition to a thorough evaluation of overall enemy release effects, we tested whether enemy release effects changed with the surrounding tree diversity. Herbivore abundance was indeed consistently lower on exotics across all six tree genera (12 comparisons). This effect of exotic status was independent of the continent, phylogenetic relatedness, and surrounding tree diversity. In contrast, leaf damage associated with generalist leaf chewers was consistently higher on North American tree species. Interestingly, several species of European weevils were the most abundant leaf chewers on both continents and the dominant herbivores at the Canadian site. Thus, most observed leaf damage is likely to reflect the effect of generalist herbivores that feed heavily on plant species with which they have not evolved. At the German site, sap suckers were the dominant herbivores and showed a pattern consistent with enemy release. Taken together, the consistently lower herbivory on exotics on both continents is not purely a pattern of enemy release in the strictest sense, but to some degree additionally reflects the susceptibility of native plants to invasive herbivores. In conclusion, our cross-Atlantic study is consistent with the idea that nonnative trees have generally reduced herbivory, regardless of tree community diversity and species identity, but for different reasons depending on the dominant herbivore guild.

Enhancing structural complexity: An experiment conducted in the Black Forest National Park, Germany.

We report on a structural complexity enhancement (SCE) experiment that was designed to test ecological restoration measures in the Black Forest National Park, Germany. The main goal was to understand as to whether the creation of standing and downed deadwood within previously managed, single-layered Norway spruce (Picea abies L.) forests accelerates the development of forest structure, richness, and diversity of a range of taxonomic groups. Here we introduce the experimental design and describe the development of stand structure including abundance and richness of tree-related microhabitats (TreMs) within 5 years after initiation of the experiment in October 2016. To enhance structural complexity in treatment plots, 10 trees per plot were toppled using a skidder winch, and another 10 trees were ring barked at a height of around 60 cm above ground level with a chainsaw. To monitor stand structure, we collected data on common forest attributes such as diameter at breast height (DBH), tree height, and TreMs of all trees in the six experimental and six control plots measuring 0.25 ha in size before the treatments were carried out in 2016 and again in 2020/21. We analyzed the abundance and richness of TreMs using generalized linear mixed models with DBH and treatment vs. control as predictors. The SCE treatment resulted in a significant increase in deadwood volumes (4.2 vs. 439.5 m3) as well as in TreM abundance and richness (increase of 0.74 TreMs per tree). This indicates that the SCE treatment was effective to increase biodiversity-relevant structures such as deadwood and TreMs, in previously managed Norway spruce-dominated stands. The ongoing monitoring of a range of taxonomic groups (birds, bats, small mammals, coleoptera, fungi, mosses, and vascular plants) in this experiment will demonstrate to what extent the enhancement in structural complexity will lead to an enrichment in species richness and diversity.

Mutually inclusive mechanisms of drought-induced tree mortality.

Unprecedented tree dieback across Central Europe caused by recent global change-type drought events highlights the need for a better mechanistic understanding of drought-induced tree mortality. Although numerous physiological risk factors have been identified, the importance of two principal mechanisms, hydraulic failure and carbon starvation, is still debated. It further remains largely unresolved how the local neighborhood composition affects individual mortality risk. We studied 9435 young trees of 12 temperate species planted in a diversity experiment in 2013 to assess how hydraulic traits, carbon dynamics, pest infestation, tree height and neighborhood competition influence individual mortality risk. Following the most extreme global change-type drought since record in 2018, one third of these trees died. Across species, hydraulic safety margins (HSMs) were negatively and a shift towards a higher sugar fraction in the non-structural carbohydrate (NSC) pool positively associated with mortality risk. Moreover, trees infested by bark beetles had a higher mortality risk, and taller trees a lower mortality risk. Most neighborhood interactions were beneficial, although neighborhood effects were highly species-specific. Species that suffered more from drought, especially Larix spp. and Betula spp., tended to increase the survival probability of their neighbors and vice versa. While severe tissue dehydration marks the final stage of drought-induced tree mortality, we show that hydraulic failure is interrelated with a series of other, mutually inclusive processes. These include shifts in NSC pools driven by osmotic adjustment and/or starch depletion as well as pest infestation and are modulated by the size and species identity of a tree and its neighbors. A more holistic view that accounts for multiple causes of drought-induced tree mortality is required to improve predictions of trends in global forest dynamics and to identify mutually beneficial species combinations.

Species richness stabilizes productivity via asynchrony and drought-tolerance diversity in a large-scale tree biodiversity experiment.

Extreme climatic events threaten forests and their climate mitigation potential globally. Understanding the drivers promoting ecosystem stability is therefore considered crucial for mitigating adverse climate change effects on forests. Here, we use structural equation models to explain how tree species richness, asynchronous species dynamics, species-level population stability, and drought-tolerance traits relate to the stability of forest productivity along an experimentally manipulated species richness gradient ranging from 1 to 24 tree species. Tree species richness improved community stability by increasing asynchrony. That is, at higher species richness, interannual variation in productivity among tree species buffered the community against stress-related productivity declines. This effect was positively related to variation in stomatal control and resistance-acquisition strategies among species, but not to the community-weighted means of these trait syndromes. The identified mechanisms by which tree species richness stabilizes forest productivity emphasize the importance of diverse, mixed-species forests to adapt to climate change.

Growth resistance and resilience of mixed silver fir and Norway spruce forests in central Europe: Contrasting responses to mild and severe droughts.

Extreme droughts are expected to increase in frequency and severity in many regions of the world, threatening multiple ecosystem services provided by forests. Effective strategies to adapt forests to such droughts require comprehensive information on the effects and importance of the factors influencing forest resistance and resilience. We used a unique combination of inventory and dendrochronological data from a long-term (>30 years) silvicultural experiment in mixed silver fir and Norway spruce mountain forests along a temperature and precipitation gradient in southwestern Germany. We aimed at examining the mechanisms and forest stand characteristics underpinning the resistance and resilience to past mild and severe droughts. We found that (i) fir benefited from mild droughts and showed higher resistance (i.e., lower growth loss during drought) and resilience (i.e., faster return to pre-drought growth levels) than spruce to all droughts; (ii) species identity determined mild drought responses while species interactions and management-related factors strongly influenced the responses to severe droughts; (iii) intraspecific and interspecific interactions had contrasting effects on the two species, with spruce being less resistant to severe droughts when exposed to interaction with fir and beech; (iv) higher values of residual stand basal area following thinning were associated with lower resistance and resilience to severe droughts; and (v) larger trees were resilient to mild drought events but highly vulnerable to severe droughts. Our study provides an analytical approach for examining the effects of different factors on individual tree- and stand-level drought response. The forests investigated here were to a certain extent resilient to mild droughts, and even benefited from such conditions, but were strongly affected by severe droughts. Lastly, negative effects of severe droughts can be reduced through modifying species composition, tree size distribution and stand density in mixed silver fir-Norway spruce forests.

Changes in plant-herbivore network structure and robustness along land-use intensity gradients in grasslands and forests.

Land-use intensification poses major threats to biodiversity, such as to insect herbivore communities. The stability of these communities depends on interactions linking herbivores and host plants. How interaction network structure begets robustness, and thus stability, in different ecosystems and how network structure and robustness are altered along land-use intensity gradients are unclear. We analyzed plant-herbivore networks based on literature-derived interactions and long-term sampling from 289 grasslands and forests in three regions of Germany. Network size and nestedness were the most important determinants of network robustness in both ecosystems. Along land-use intensity gradients, networks in moderately grazed grasslands were more robust than in those managed by frequent mowing or fertilization. In forests, changes of network robustness along land-use intensity gradients relied on changes in plant species richness. Our results expand our knowledge of the stability of plant-herbivore networks and indicate options for management aimed at stabilizing herbivore communities.

Groundwater extraction reduces tree vitality, growth and xylem hydraulic capacity in Quercus robur during and after drought events.

Climate change is expected to pose major direct and indirect threats to groundwater-dependent forest ecosystems. Forests that concurrently experience increased rates of water extraction may face unprecedented exposure to droughts. Here, we examined differences in stem growth and xylem hydraulic architecture of 216 oak trees from sites with contrasting groundwater availability, including sites where groundwater extraction has led to reduced water availability for trees over several decades. We expected reduced growth and xylem hydraulic capacity for trees at groundwater extraction sites both under normal and unfavourable growing conditions. Compared to sites without extraction, trees at sites with groundwater extraction showed reduced growth and hydraulic conductivity both during periods of moderate and extremely low soil water availability. Trees of low vigour, which were more frequent at sites with groundwater extraction, were not able to recover growth and hydraulic capacity following drought, pointing to prolonged drought effects. Long-term water deficit resulting in reduced CO2 assimilation and hydraulic capacity after drought are very likely responsible for observed reductions in tree vitality at extraction sites. Our results demonstrate that groundwater access maintains tree function and resilience to drought and is therefore important for tree health in the context of climate change.

Climate affects neighbour-induced changes in leaf chemical defences and tree diversity-herbivory relationships.

Associational resistance theory predicts that insect herbivory decreases with increasing tree diversity in forest ecosystems. However, the generality of this effect and its underlying mechanisms are still debated, particularly since evidence has accumulated that climate may influence the direction and strength of the relationship between diversity and herbivory.We quantified insect leaf herbivory and leaf chemical defences (phenolic compounds) of silver birch Betula pendula in pure and mixed plots with different tree species composition across 12 tree diversity experiments in different climates. We investigated whether the effects of neighbouring tree species diversity on insect herbivory in birch, that is, associational effects, were dependent on the climatic context, and whether neighbour-induced changes in birch chemical defences were involved in associational resistance to insect herbivory.We showed that herbivory on birch decreased with tree species richness (i.e. associational resistance) in colder environments but that this relationship faded as mean annual temperature increased.Birch leaf chemical defences increased with tree species richness but decreased with the phylogenetic distinctiveness of birch from its neighbours, particularly in warmer and more humid environments.Herbivory was negatively correlated with leaf chemical defences, particularly when birch was associated with closely related species. The interactive effect of tree diversity and climate on herbivory was partially mediated by changes in leaf chemical defences.Our findings confirm that tree species diversity can modify the leaf chemistry of a focal species, hence its quality for herbivores. They further stress that such neighbour-induced changes are dependent on climate and that tree diversity effects on insect herbivory are partially mediated by these neighbour-induced changes in chemical defences.

Biodiversity response to forest management intensity, carbon stocks and net primary production in temperate montane forests.

Managed forests are a key component of strategies aimed at tackling the climate and biodiversity crises. Tapping this potential requires a better understanding of the complex, simultaneous effects of forest management on biodiversity, carbon stocks and productivity. Here, we used data of 135 one-hectare plots from southwestern Germany to disentangle the relative influence of gradients of management intensity, carbon stocks and forest productivity on different components of forest biodiversity (birds, bats, insects, plants) and tree-related microhabitats. We tested whether the composition of taxonomic groups varies gradually or abruptly along these gradients. The richness of taxonomic groups was rather insensitive to management intensity, carbon stocks and forest productivity. Despite the low explanatory power of the main predictor variables, forest management had the greatest relative influence on richness of insects and tree-related microhabitats, while carbon stocks influenced richness of bats, birds, vascular plants and pooled taxa. Species composition changed relatively abruptly along the management intensity gradient, while changes along carbon and productivity gradients were more gradual. We conclude that moderate increases in forest management intensity and carbon stocks, within the range of variation observed in our study system, might be compatible with biodiversity and climate mitigation objectives in managed forests.

Site-specific risk assessment enables trade-off analysis of non-native tree species in European forests.

Non-native tree species (NNT) are used in European forestry for many purposes including their growth performance, valuable timber, and resistance to drought and pest or pathogen damage. Yet, cultivating NNT may pose risks to biodiversity, ecosystem functioning, and the provisioning of ecosystem services, and several NNT have been classified as invasive in Europe. Typically, such classifications are based on risk assessments, which do not adequately consider site-specific variations in impacts of the NNT or the extent of affected areas. Here, we present a new methodological framework that facilitates both mitigating risks associated with NNT and taking advantage of their ecosystem services. The framework is based on a stratified assessment of risks posed by NNT which distinguishes between different sites and considers effectiveness of available management strategies to control negative effects. The method can be applied to NNT that already occur in a given area or those NNT that may establish in future. The framework consists of eight steps and is partly based on existing knowledge. If adequate site-specific knowledge on NNT does not yet exist, new evidence on the risks should be obtained, for example, by collecting and analyzing monitoring data or modeling the potential distribution of NNT. However, limitations remain in the application of this method, and we propose several policy and management recommendations which are required to improve the responsible use of NNT.

Low root functional dispersion enhances functionality of plant growth by influencing bacterial activities in European forest soils.

Current studies show that multispecies forests are beneficial regarding biodiversity and ecosystem functionality. However, there are only little efforts to understand the ecological mechanisms behind these advantages of multispecies forests. Bacteria are among the key plant growth-promoting microorganisms that support tree growth and fitness. Thus, we investigated links between bacterial communities, their functionality and root trait dispersion within four major European forest types comprising multispecies and monospecific plots. Bacterial diversity revealed no major changes across the root functional dispersion gradient. In contrast, predicted gene profiles linked to plant growth activities suggest an increasing bacterial functionality from monospecific to multispecies forest. In multispecies forest plots, the bacterial functionality linked to plant growth activities declined with the increasing functional dispersion of the roots. Our findings indicate that enriched abundant bacterial operational taxonomic units are decoupled from bacterial functionality. We also found direct effects of tree species identity on bacterial community composition but no significant relations with root functional dispersion. Additionally, bacterial network analyses indicated that multispecies forests have a higher complexity in their bacterial communities, which points towards more stable forest systems with greater functionality. We identified a potential of root dispersion to facilitate bacterial interactions and consequently, plant growth activities.

Fungal guilds and soil functionality respond to tree community traits rather than to tree diversity in European forests.

At the global scale, most forest research on biodiversity focuses on aboveground organisms. However, understanding the structural associations between aboveground and belowground communities provides relevant information about important functions linked to biogeochemical cycles. Microorganisms such as soil fungi are known to be closely coupled to the dominant tree vegetation, and we hypothesize that tree traits affect fungal guilds and soil functionality in multiple ways. By analysing fungal diversity of 64 plots from four European forest types using Illumina DNA sequencing, we show that soil fungal communities respond to tree community traits rather than to tree species diversity. To explain changes in fungal community structure and measured soil enzymatic activities, we used a trait-based ecological approach and community-weighted means of tree traits to define 'fast' (acquisitive) versus 'slow' (conservative) tree communities. We found specific tree trait effects on different soil fungal guilds and soil enzymatic activities: tree traits associated with litter and absorptive roots correlated with fungal, especially pathogen diversity, and influenced community composition of soil fungi. Relative abundance of the symbiotrophic and saprotrophic guilds mirrored the litter quality, while the root traits of fast tree communities enhanced symbiotrophic abundance. We found that forest types of higher latitudes, which are dominated by fast tree communities, correlated with high carbon-cycling enzymatic activities. In contrast, Mediterranean forests with slow tree communities showed high enzymatic activities related to nitrogen and phosphorous. Our findings highlight that tree trait effects of either 'fast' or 'slow' tree communities drive different fungal guilds and influence biogeochemical cycles.

What do tree-related microhabitats tell us about the abundance of forest-dwelling bats, birds, and insects?

Retaining trees during harvesting to conserve biodiversity is becoming increasingly common in forestry. To assess, select and monitor these habitat trees, ecologists and practitioners often use Tree-related Microhabitats (TreMs), which are assumed to represent the abundance and diversity of environmental resources for a wide range of forest-dwelling taxa. However, the relationship between TreMs and forest organisms is not fully understood. In this context, we attempted to identify and quantify the links between TreMs and three groups of forest organisms: insects, bats, and birds. Specifically, we tested whether species abundance is influenced by TreM abundance, either as direct predictor or as mediator of environmental predictors. We collected data in 86 temperate, 1-ha mixed forest plots and employed a hierarchical generalized mixed model to assess the influence of seven environmental predictors (aspect, number and height of standing dead trees, cover of herb and shrub layer, volume of lying deadwood, and terrain ruggedness index (TRI)) on the abundance of TreMs (15 groups) on potential habitat trees, insects (10 orders), bats (5 acoustic groups) and birds (29 species) as a function of seven environmental predictors: aspect, number and height of standing dead trees, cover of herb and shrub layer, volume of lying deadwood, and terrain ruggedness index (TRI). This allowed us to generate a correlation matrix with potential links between abundances of TreMs and co-occurring forest organisms. These correlations and the environmental predictors were tested in a structural equation model (SEM) to disentangle and quantify the effects of the environment from direct effects of TreMs on forest organisms. Four TreM groups showed correlations > |0.30| with forest organisms, in particular with insects and bats. Rot holes and concavities were directly linked with three insect groups and two bat groups. Their effect was smaller than effects of environmental predictors, except for the pairs "rot holes - Sternorrhyncha" and "rot holes - bats" of the Pipistrellus group. In addition, TreMs had indirect effects on forest organisms through mediating the effects of environmental predictors. We found significant associations between two out of fifteen TreM groups and five out of 44 forest organism groups. These results indicate that TreM abundance on potential habitat trees is not suited as a general indicator of the species abundance across broad taxonomic groups but possibly for specific target groups with proven links.

Evaluating the effectiveness of retention forestry to enhance biodiversity in production forests of Central Europe using an interdisciplinary, multi-scale approach.

Retention forestry, which retains a portion of the original stand at the time of harvesting to maintain continuity of structural and compositional diversity, has been originally developed to mitigate the impacts of clear-cutting. Retention of habitat trees and deadwood has since become common practice also in continuous-cover forests of Central Europe. While the use of retention in these forests is plausible, the evidence base for its application is lacking, trade-offs have not been quantified, it is not clear what support it receives from forest owners and other stakeholders and how it is best integrated into forest management practices. The Research Training Group ConFoBi (Conservation of Forest Biodiversity in Multiple-use Landscapes of Central Europe) focusses on the effectiveness of retention forestry, combining ecological studies on forest biodiversity with social and economic studies of biodiversity conservation across multiple spatial scales. The aim of ConFoBi is to assess whether and how structural retention measures are appropriate for the conservation of forest biodiversity in uneven-aged and selectively harvested continuous-cover forests of temperate Europe. The study design is based on a pool of 135 plots (1 ha) distributed along gradients of forest connectivity and structure. The main objectives are (a) to investigate the effects of structural elements and landscape context on multiple taxa, including different trophic and functional groups, to evaluate the effectiveness of retention practices for biodiversity conservation; (b) to analyze how forest biodiversity conservation is perceived and practiced, and what costs and benefits it creates; and (c) to identify how biodiversity conservation can be effectively integrated in multi-functional forest management. ConFoBi will quantify retention levels required across the landscape, as well as the socio-economic prerequisites for their implementation by forest owners and managers. ConFoBi's research results will provide an evidence base for integrating biodiversity conservation into forest management in temperate forests.

Retention as an integrated biodiversity conservation approach for continuous-cover forestry in Europe.

Retention forestry implies that biological legacies like dead and living trees are deliberately selected and retained beyond harvesting cycles to benefit biodiversity and ecosystem functioning. This model has been applied for several decades in even-aged, clearcutting (CC) systems but less so in uneven-aged, continuous-cover forestry (CCF). We provide an overview of retention in CCF in temperate regions of Europe, currently largely focused on habitat trees and dead wood. The relevance of current meta-analyses and many other studies on retention in CC is limited since they emphasize larger patches in open surroundings. Therefore, we reflect here on the ecological foundations and socio-economic frameworks of retention approaches in CCF, and highlight several areas with development potential for the future. Conclusions from this perspective paper, based on both research and current practice on several continents, although highlighting Europe, are also relevant to other temperate regions of the world using continuous-cover forest management approaches.

Arthropod decline in grasslands and forests is associated with landscape-level drivers.

Recent reports of local extinctions of arthropod species1, and of massive declines in arthropod biomass2, point to land-use intensification as a major driver of decreasing biodiversity. However, to our knowledge, there are no multisite time series of arthropod occurrences across gradients of land-use intensity with which to confirm causal relationships. Moreover, it remains unclear which land-use types and arthropod groups are affected, and whether the observed declines in biomass and diversity are linked to one another. Here we analyse data from more than 1 million individual arthropods (about 2,700 species), from standardized inventories taken between 2008 and 2017 at 150 grassland and 140 forest sites in 3 regions of Germany. Overall gamma diversity in grasslands and forests decreased over time, indicating loss of species across sites and regions. In annually sampled grasslands, biomass, abundance and number of species declined by 67%, 78% and 34%, respectively. The decline was consistent across trophic levels and mainly affected rare species; its magnitude was independent of local land-use intensity. However, sites embedded in landscapes with a higher cover of agricultural land showed a stronger temporal decline. In 30 forest sites with annual inventories, biomass and species number-but not abundance-decreased by 41% and 36%, respectively. This was supported by analyses of all forest sites sampled in three-year intervals. The decline affected rare and abundant species, and trends differed across trophic levels. Our results show that there are widespread declines in arthropod biomass, abundance and the number of species across trophic levels. Arthropod declines in forests demonstrate that loss is not restricted to open habitats. Our results suggest that major drivers of arthropod decline act at larger spatial scales, and are (at least for grasslands) associated with agriculture at the landscape level. This implies that policies need to address the landscape scale to mitigate the negative effects of land-use practices.