Adaptive monitoring in action-what drives arthropod diversity and composition in central European beech forests?

Recent studies suggest that arthropod diversity in German forests is declining. Currently, different national programs are being developed to monitor arthropod trends and to unravel the effects of forest management on biodiversity in forests. To establish effective long-term monitoring programs, a set of drivers of arthropod diversity and composition as well as suitable species groups have to be identified. To aid in answering these questions, we investigated arthropod data collected in four Hessian forest reserves (FR) in the 1990s. To fully utilize this data set, we combined it with results from a retrospective structural sampling design applied at the original trap locations in central European beech (Fagus sylvatica) forests. As expected, the importance of the different forest structural, vegetation, and site attributes differed largely between the investigated arthropod groups: beetles, spiders, Aculeata, and true bugs. Measures related to light availability and temperature such as canopy cover or potential radiation were important to all groups affecting either richness, composition, or both. Spiders and true bugs were affected by the broadest range of explanatory variables, which makes them a good choice for monitoring general trends. For targeted monitoring focused on forestry-related effects on biodiversity, rove and ground beetles seem more suitable. Both groups were driven by a narrower, more management-related set of variables. Most importantly, our study approach shows that it is possible to utilize older biodiversity survey data. Although, in our case, there are strong restrictions due to the long time between species and structural attribute sampling.

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.

Formation of mineral-associated organic matter in temperate soils is primarily controlled by mineral type and modified by land use and management intensity.

Formation of mineral-associated organic matter (MAOM) supports the accumulation and stabilization of carbon (C) in soil, and thus, is a key factor in the global C cycle. Little is known about the interplay of mineral type, land use and management intensity in MAOM formation, especially on subdecadal time scales. We exposed mineral containers with goethite or illite, the most abundant iron oxide and phyllosilicate clay in temperate soils, for 5 years in topsoils of 150 forest and 150 grassland sites in three regions across Germany. Results show that irrespective of land use and management intensity, more C accumulated on goethite than illite (on average 0.23 ± 0.10 and 0.06 ± 0.03 mg m-2 mineral surface respectively). Carbon accumulation across regions was consistently higher in coniferous forests than in deciduous forests and grasslands. Structural equation models further showed that thinning and harvesting reduced MAOM formation in forests. Formation of MAOM in grasslands was not affected by grazing. Fertilization had opposite effects on MAOM formation, with the positive effect being mediated by enhanced plant productivity and the negative effect by reduced plant species richness. This highlights the caveat of applying fertilizers as a strategy to increase soil C stocks in temperate grasslands. Overall, we demonstrate that the rate and amount of MAOM formation in soil is primarily driven by mineral type, and can be modulated by land use and management intensity even on subdecadal time scales. Our results suggest that temperate soils dominated by oxides have a higher capacity to accumulate and store C than those dominated by phyllosilicate clays, even under circumneutral pH conditions. Therefore, adopting land use and management practices that increase C inputs into oxide-rich soils that are under their capacity to store C may offer great potential to enhance near-term soil C sequestration.

Integrated global assessment of the natural forest carbon potential.

Forests are a substantial terrestrial carbon sink, but anthropogenic changes in land use and climate have considerably reduced the scale of this system1. Remote-sensing estimates to quantify carbon losses from global forests2-5 are characterized by considerable uncertainty and we lack a comprehensive ground-sourced evaluation to benchmark these estimates. Here we combine several ground-sourced6 and satellite-derived approaches2,7,8 to evaluate the scale of the global forest carbon potential outside agricultural and urban lands. Despite regional variation, the predictions demonstrated remarkable consistency at a global scale, with only a 12% difference between the ground-sourced and satellite-derived estimates. At present, global forest carbon storage is markedly under the natural potential, with a total deficit of 226 Gt (model range = 151-363 Gt) in areas with low human footprint. Most (61%, 139 Gt C) of this potential is in areas with existing forests, in which ecosystem protection can allow forests to recover to maturity. The remaining 39% (87 Gt C) of potential lies in regions in which forests have been removed or fragmented. Although forests cannot be a substitute for emissions reductions, our results support the idea2,3,9 that the conservation, restoration and sustainable management of diverse forests offer valuable contributions to meeting global climate and biodiversity targets.

The global biogeography of tree leaf form and habit.

Understanding what controls global leaf type variation in trees is crucial for comprehending their role in terrestrial ecosystems, including carbon, water and nutrient dynamics. Yet our understanding of the factors influencing forest leaf types remains incomplete, leaving us uncertain about the global proportions of needle-leaved, broadleaved, evergreen and deciduous trees. To address these gaps, we conducted a global, ground-sourced assessment of forest leaf-type variation by integrating forest inventory data with comprehensive leaf form (broadleaf vs needle-leaf) and habit (evergreen vs deciduous) records. We found that global variation in leaf habit is primarily driven by isothermality and soil characteristics, while leaf form is predominantly driven by temperature. Given these relationships, we estimate that 38% of global tree individuals are needle-leaved evergreen, 29% are broadleaved evergreen, 27% are broadleaved deciduous and 5% are needle-leaved deciduous. The aboveground biomass distribution among these tree types is approximately 21% (126.4 Gt), 54% (335.7 Gt), 22% (136.2 Gt) and 3% (18.7 Gt), respectively. We further project that, depending on future emissions pathways, 17-34% of forested areas will experience climate conditions by the end of the century that currently support a different forest type, highlighting the intensification of climatic stress on existing forests. By quantifying the distribution of tree leaf types and their corresponding biomass, and identifying regions where climate change will exert greatest pressure on current leaf types, our results can help improve predictions of future terrestrial ecosystem functioning and carbon cycling.

Native diversity buffers against severity of non-native tree invasions.

Determining the drivers of non-native plant invasions is critical for managing native ecosystems and limiting the spread of invasive species1,2. Tree invasions in particular have been relatively overlooked, even though they have the potential to transform ecosystems and economies3,4. Here, leveraging global tree databases5-7, we explore how the phylogenetic and functional diversity of native tree communities, human pressure and the environment influence the establishment of non-native tree species and the subsequent invasion severity. We find that anthropogenic factors are key to predicting whether a location is invaded, but that invasion severity is underpinned by native diversity, with higher diversity predicting lower invasion severity. Temperature and precipitation emerge as strong predictors of invasion strategy, with non-native species invading successfully when they are similar to the native community in cold or dry extremes. Yet, despite the influence of these ecological forces in determining invasion strategy, we find evidence that these patterns can be obscured by human activity, with lower ecological signal in areas with higher proximity to shipping ports. Our global perspective of non-native tree invasion highlights that human drivers influence non-native tree presence, and that native phylogenetic and functional diversity have a critical role in the establishment and spread of subsequent invasions.

Abrupt height growth setbacks show overbrowsing of tree saplings, which can be reduced by raising deer harvest.

Intensive ungulate browsing significantly impacts forests worldwide. However, it is usually not single browsing events that lead to sapling mortality, but the little-researched interactions of browsed saplings with their biotic and abiotic environment. (I) Our objective was to assess the impact of ungulate browsing on the growth of young saplings relative to other environmental factors by utilizing their height increment as a sensitive measure of vitality to indicate their status. (II) Furthermore, we aimed to identify factors affecting ungulate browsing at our study sites, assessed as browsing probabilities, and identify effective mitigation measures for browsing impact. We analyzed an extensive sapling dataset of 248 wildlife exclosures, which were erected in 2016 in beech dominated forests across Germany and assessed annually until 2020. (I) Browsing probability and light availability were the most influential parameters for selectively browsed, admixed tree species (e.g., sycamore maple). Height increment showed abrupt setbacks, which caused a permanent collapse of growth when browsing exceeded a certain level. However, light availability enhanced height increment. (II) An increase in deer harvest reduced the browsing probability of selectively browsed species considerably. We conclude that the growth-inhibiting effect of ungulate browsing is a multifactorial phenomenon, which can be mitigated by silvicultural management and efficient hunting strategies.

Changes of vegetation in coniferous monocultures in the context of conversion to mixed forests in 30 years - Implications for biodiversity restoration.

The understorey vegetation of temperate forests harbours a major proportion of terrestrial biodiversity and fulfills an important role in ecosystem functioning. Over the past decades, temperate forest understoreys were found to change in species diversity and composition due to several anthropogenic and natural drivers. Currently, the conversion and restoration of even-aged coniferous monocultures into more diverse and mixed broad-leaved forests are major objectives of sustainable forest management in Central Europe. This forest conversion alters understorey communities and abiotic site conditions but the underlying patterns and processes are not yet fully understood. Therefore, we investigated changes in the Bavarian Spessart mountains in southwest Germany, where we re-sampled 108 semi-permanent plots from four different coniferous stand types (i.e., Norway spruce, Scots pine, Douglas fir, European larch) about 30 years after the initial assessment. On these plots, we recorded understorey vegetation and forest structure, and derived abiotic site conditions based on ecological indicator values of understorey vegetation, followed by multivariate analysis. We found changes in plant communities that point towards a decrease of soil acidity and a "thermophilization" of forest understoreys. Understorey species richness remained constant, while understorey's Shannon and Simpson diversity increased. The observed changes in forest structure explained the temporal shifts in understorey species composition. The understorey species composition did not experience a significant floristic homogenization since the 1990s. However, plant communities exhibited a reduction in species characteristic of coniferous forests and a simultaneous increase in species associated with broad-leaved forests. The increase of specialist species (closed forests and open sites) may have compensated for the detected decrease in generalist species. We conclude that the forest conversion towards mixed broad-leaved forest in the Spessart mountains of the past decades might have masked homogenization trends that are increasingly reported from Central European forest understoreys.

Regeneration in European beech forests after drought: the effects of microclimate, deadwood and browsing.

With progressing climate change, increasing weather extremes will endanger tree regeneration. Canopy openings provide light for tree establishment, but also reduce the microclimatic buffering effect of forests. Thus, disturbances can have both positive and negative impacts on tree regeneration. In 2015, three years before an extreme drought episode hit Central Europe, we established a manipulation experiment with a factorial block design in European beech (Fagus sylvatica L.)-dominated forests. At five sites located in southeastern Germany, we conducted three censuses of tree regeneration after implementing two different canopy disturbances (aggregated and distributed canopy openings), and four deadwood treatments (retaining downed, standing, downed + standing deadwood and removing all deadwood), as well as in one untreated control plot. In addition, we measured understory light levels and recorded local air temperature and humidity over five years. We (i) tested the effects of experimental disturbance and deadwood treatments on regeneration and (ii) identified the drivers of regeneration density as well as seedling species and structural diversity. Regeneration density increased over time. Aggregated canopy openings supported species and structural diversity, but reduced regeneration density. Tree regeneration was positively associated with understory light levels, while maximum vapor pressure deficit influenced tree regeneration negatively. Deadwood and browsing impacts on regeneration varied and were inconclusive. Our study indicates that despite the drought episode regeneration in beech-dominated forests persisted under moderately disturbed canopies. However, the positive effect of increased light availability on tree regeneration might have been offset by harsher microclimate after canopies have been disturbed. Supplementary Information: The online version contains supplementary material available at 10.1007/s10342-022-01520-1.

Enhancing the structural diversity between forest patches-A concept and real-world experiment to study biodiversity, multifunctionality and forest resilience across spatial scales.

Intensification of land use by humans has led to a homogenization of landscapes and decreasing resilience of ecosystems globally due to a loss of biodiversity, including the majority of forests. Biodiversity-ecosystem functioning (BEF) research has provided compelling evidence for a positive effect of biodiversity on ecosystem functions and services at the local (α-diversity) scale, but we largely lack empirical evidence on how the loss of between-patch ß-diversity affects biodiversity and multifunctionality at the landscape scale (γ-diversity). Here, we present a novel concept and experimental framework for elucidating BEF patterns at α-, ß-, and γ-scales in real landscapes at a forest management-relevant scale. We examine this framework using 22 temperate broadleaf production forests, dominated by Fagus sylvatica. In 11 of these forests, we manipulated the structure between forest patches by increasing variation in canopy cover and deadwood. We hypothesized that an increase in landscape heterogeneity would enhance the ß-diversity of different trophic levels, as well as the ß-functionality of various ecosystem functions. We will develop a new statistical framework for BEF studies extending across scales and incorporating biodiversity measures from taxonomic to functional to phylogenetic diversity using Hill numbers. We will further expand the Hill number concept to multifunctionality allowing the decomposition of γ-multifunctionality into α- and ß-components. Combining this analytic framework with our experimental data will allow us to test how an increase in between patch heterogeneity affects biodiversity and multifunctionality across spatial scales and trophic levels to help inform and improve forest resilience under climate change. Such an integrative concept for biodiversity and functionality, including spatial scales and multiple aspects of diversity and multifunctionality as well as physical and environmental structure in forests, will go far beyond the current widely applied approach in forestry to increase resilience of future forests through the manipulation of tree species composition.

Co-limitation towards lower latitudes shapes global forest diversity gradients.

The latitudinal diversity gradient (LDG) is one of the most recognized global patterns of species richness exhibited across a wide range of taxa. Numerous hypotheses have been proposed in the past two centuries to explain LDG, but rigorous tests of the drivers of LDGs have been limited by a lack of high-quality global species richness data. Here we produce a high-resolution (0.025° × 0.025°) map of local tree species richness using a global forest inventory database with individual tree information and local biophysical characteristics from ~1.3 million sample plots. We then quantify drivers of local tree species richness patterns across latitudes. Generally, annual mean temperature was a dominant predictor of tree species richness, which is most consistent with the metabolic theory of biodiversity (MTB). However, MTB underestimated LDG in the tropics, where high species richness was also moderated by topographic, soil and anthropogenic factors operating at local scales. Given that local landscape variables operate synergistically with bioclimatic factors in shaping the global LDG pattern, we suggest that MTB be extended to account for co-limitation by subordinate drivers.

Negative effects of forest gaps on dung removal in a full-factorial experiment.

Ecosystem functioning may directly or indirectly-via change in biodiversity-respond to land use. Dung removal is an important ecosystem function central for the decomposition of mammal faeces, including secondary seed dispersal and improved soil quality. Removal usually increases with dung beetle diversity and biomass. In forests, dung removal can vary with structural variables that are, however, often interrelated, making experiments necessary to understand the role of single variables on ecosystem functions. How gaps and deadwood, two main outcomes of forest management influence dung removal, is unknown. We tested if dung removal responds to gap creation and deadwood provisioning or if treatment effects are mediated via responses of dung beetles. We expected lower removal rates in gaps due to lower dung beetle biomass and diversity. We sampled dung beetles and measured dung removal in a highly-replicated full-factorial forest experiment established at 29 sites in three regions of Germany (treatments: Gap, Gap + Deadwood, Deadwood, Control). All gaps were experimentally created and had a diameter of around 30 m. Dung beetle diversity, biomass and dung removal were each lower in gaps than in controls. Dung removal decreased from 61.9% in controls to 48.5% in gaps, irrespective of whether or not the gap had deadwood. This treatment effect was primarily driven by dung beetle biomass but not diversity. Furthermore, dung removal was reduced to 56.9% in the deadwood treatment. Our findings are not consistent with complementarity effects of different dung beetle species linked to biodiversity-ecosystem functioning relationships that have been shown in several ecosystems. In contrast, identity effects can be pronounced: gaps reduced the abundance of a large-bodied key forest species (Anoplotrupes stercorosus), without compensatory recruitment of open land species. While gaps and deadwood are important for many forest organisms, dung beetles and dung removal respond negatively. Our results exemplify how experiments can contribute to test hypotheses on the interrelation between land use, biodiversity and ecosystem functioning.

Soil Inoculation With Beneficial Microbes Buffers Negative Drought Effects on Biomass, Nutrients, and Water Relations of Common Myrtle.

Common myrtle (Myrtus communis L.) occurs in (semi-)arid areas of the Palearctic region where climate change, over-exploitation, and habitat destruction imperil its existence. The evergreen shrub is of great economic and ecological importance due to its pharmaceutical value, ornamental use, and its role in urban greening and habitat restoration initiatives. Under greenhouse conditions, we investigated the effect of soil inoculation with arbuscular mycorrhizal fungi (AMF) and plant growth-promoting rhizobacteria (PGPR) on biomass allocation, water relations, and nutritional status of drought-stressed myrtle seedlings. Single and dual AMF (Funneliformis mosseae and Rhizophagus irregularis) and PGPR (Pseudomonas fluorescens and P. putida) soil inoculations were applied to myrtle seedlings growing under different soil water regimes (100, 60, and 30% of field capacity) for 6 months using a full factorial, completely randomized design. AMF and PGPR treatments, especially dual inoculations, alleviated negative drought effects on biomass and morpho-physiological traits, except for water-use efficiency, which peaked under severe drought conditions. Under the greatest soil water deficit, dual inoculations promoted leaf biomass (104%-108%), root biomass (56%-73%), mesophyll conductance (58%), and relative water content (1.4-fold) compared to non-inoculated controls. Particularly, dual AMF and PGPR inoculations stimulated nutrient dynamics in roots (N: 138%-151%, P: 176%-181%, K: 112%-114%, Ca: 124%-136%, and Mg: 130%-140%) and leaves (N: 101%-107%, P: 143%-149%, K: 83%-84%, Ca: 98%-107%, and Mg: 102%-106%). Our findings highlight soil inoculations with beneficial microbes as a cost-effective way to produce highly drought resistant seedling stock which is vital for restoring natural myrtle habitats and for future-proofing myrtle crop systems.

Tree regeneration characteristics in limestone forests of the Cat Ba National Park, Vietnam.

BACKGROUND: The ability of overstory tree species to regenerate successfully is important for the preservation of tree species diversity and its associated flora and fauna. This study investigated forest regeneration dynamics in the Cat Ba National Park, a biodiversity hotspot in Vietnam. Data was collected from 90 sample plots (500 m2) and 450 sub-sample plots (25 m2) in regional limestone forests. We evaluated the regeneration status of tree species by developing five ratios relating overstory and regeneration richness and diversity. By examining the effect of environmental factors on these ratios, we aimed to identify the main drivers for maintaining tree species diversity or for potential diversity gaps between the regeneration and the overstory layer. Our results can help to increase the understanding of regeneration patterns in tropical forests of Southeast Asia and to develop successful conservation strategies. RESULTS: We found 97 tree species in the regeneration layer compared to 136 species in the overstory layer. The average regeneration density was 3764 ± 1601 per ha. Around 70% of the overstory tree species generated offspring. According to the International Union for Conservation of Nature's Red List, only 36% of threatened tree species were found in the regeneration layer. A principal component analysis provided evidence that the regeneration of tree species was slightly negatively correlated to terrain factors (percentage of rock surface, slope) and soil properties (cation exchange capacity, pH, humus content, soil moisture, soil depth). Contrary to our expectations, traces of human impact and the prevailing light conditions (total site factor, gap fraction, openness, indirect site factor, direct site factor) had no influence on regeneration density and composition, probably due to the small gradient in light availability. CONCLUSION: We conclude that the tree species richness in Cat Ba National Park appears to be declining at present. We suggest similar investigations in other biodiversity hotspots to learn whether the observed trend is a global phenomenon. In any case, a conservation strategy for the threatened tree species in the Cat Ba National Park needs to be developed if tree species diversity is to be maintained.

Global data on earthworm abundance, biomass, diversity and corresponding environmental properties.

Earthworms are an important soil taxon as ecosystem engineers, providing a variety of crucial ecosystem functions and services. Little is known about their diversity and distribution at large spatial scales, despite the availability of considerable amounts of local-scale data. Earthworm diversity data, obtained from the primary literature or provided directly by authors, were collated with information on site locations, including coordinates, habitat cover, and soil properties. Datasets were required, at a minimum, to include abundance or biomass of earthworms at a site. Where possible, site-level species lists were included, as well as the abundance and biomass of individual species and ecological groups. This global dataset contains 10,840 sites, with 184 species, from 60 countries and all continents except Antarctica. The data were obtained from 182 published articles, published between 1973 and 2017, and 17 unpublished datasets. Amalgamating data into a single global database will assist researchers in investigating and answering a wide variety of pressing questions, for example, jointly assessing aboveground and belowground biodiversity distributions and drivers of biodiversity change.

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.

Spring understory herbs flower later in intensively managed forests.

Many organisms respond to anthropogenic environmental change through shifts in their phenology. In plants, flowering is largely driven by temperature, and therefore affected by climate change. However, on smaller scales climatic conditions are also influenced by other factors, including habitat structure. A group of plants with a particularly distinct phenology are the understory herbs in temperate European forests. In these forests, management alters tree species composition (often replacing deciduous with coniferous species) and homogenizes stand structure, and as a consequence changes light conditions and microclimate. Forest management should thus also affect the phenology of understory herbs. To test this, we recorded the flowering phenology of 16 early-flowering herbs on 100 forest plots varying in management intensity, from near-natural to intensely managed forests, in central and southern Germany. We found that in forest stands with a high management intensity, such as Norway spruce plantations, the plants flowered on average about 2 weeks later than in unmanaged forests. This was largely because management also affected microclimate (e.g., spring temperatures of 5.9°C in managed coniferous, 6.7 in managed deciduous, and 7.0°C in unmanaged deciduous plots), which in turn affected phenology, with plants flowering later on colder and moister forest stands (+4.5 d per -1°C and 2.7 d per 10% humidity increase). Among forest characteristics, the percentage of conifers had the greatest influence on microclimate, but also the age, overall crown projection area, structural complexity and spatial distribution of the forest stands. Our study indicates that forest management alters plant phenology, with potential far-reaching consequences for the ecology and evolution of understorey communities. More generally, our study demonstrates that besides climate change other drivers of environmental change, too, can influence the phenology of organisms.

Predicting Tree Species From 3D Laser Scanning Point Clouds Using Deep Learning.

Automated species classification from 3D point clouds is still a challenge. It is, however, an important task for laser scanning-based forest inventory, ecosystem models, and to support forest management. Here, we tested the performance of an image classification approach based on convolutional neural networks (CNNs) with the aim to classify 3D point clouds of seven tree species based on 2D representation in a computationally efficient way. We were particularly interested in how the approach would perform with artificially increased training data size based on image augmentation techniques. Our approach yielded a high classification accuracy (86%) and the confusion matrix revealed that despite rather small sample sizes of the training data for some tree species, classification accuracy was high. We could partly relate this to the successful application of the image augmentation technique, improving our result by 6% in total and 13, 14, and 24% for ash, oak and pine, respectively. The introduced approach is hence not only applicable to small-sized datasets, it is also computationally effective since it relies on 2D instead of 3D data to be processed in the CNN. Our approach was faster and more accurate when compared to the point cloud-based "PointNet" approach.

Global patterns and climatic controls of forest structural complexity.

The complexity of forest structures plays a crucial role in regulating forest ecosystem functions and strongly influences biodiversity. Yet, knowledge of the global patterns and determinants of forest structural complexity remains scarce. Using a stand structural complexity index based on terrestrial laser scanning, we quantify the structural complexity of boreal, temperate, subtropical and tropical primary forests. We find that the global variation of forest structural complexity is largely explained by annual precipitation and precipitation seasonality (R² = 0.89). Using the structural complexity of primary forests as benchmark, we model the potential structural complexity across biomes and present a global map of the potential structural complexity of the earth´s forest ecoregions. Our analyses reveal distinct latitudinal patterns of forest structure and show that hotspots of high structural complexity coincide with hotspots of plant diversity. Considering the mechanistic underpinnings of forest structural complexity, our results suggest spatially contrasting changes of forest structure with climate change within and across biomes.