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.

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.

Biomass Allocation and Leaf Morphology of Saplings Grown under Various Conditions of Light Availability and Competition Types.

Plant growth is almost always limited by light availability and competition. However, plants are generally plastic and can change their morphology and biomass allocation to optimize growth under suboptimal conditions. We set up a controlled pot experiment with three light availability levels (10%, 20%, and 50%) to study the effect of light and competition on the biomass allocation and leaf morphology in monospecific and mixed pots of recently planted European beech (Fagus sylvatica L.), Norway spruce (Picea abies (L.) Karst.), and Douglas fir (Pseudotsuga menziesii (Mirb.) Franco) saplings using a quantile regression model. Specific leaf area (SLA) showed the strongest reaction and increased with decreasing light availability. Woody aboveground mass fraction (AMF) increased with decreasing light availability, but the effect of light on biomass allocation was less pronounced than on SLA. The SLA, woody AMF, and root mass fraction (RMF) of the two conifer species and European beech varied greatly, with European beech having a higher SLA and RMF than the two conifer species. The associated effect of plant size on biomass allocation was small, and the strength of the association was not meaningful on a practical level. The competitor's effect on biomass allocation was minor overall and only present for some species, suggesting that species' functional dissimilarity does not greatly affect allocational patterns in early tree development stages.

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.

Restoring oak forests through direct seeding or planting: Protocol for a continental-scale experiment.

The choice of revegetating via direct seeding or planting nursery-grown seedlings influences the potential stresses suffered by seedlings such as herbivory and drought. The outcome of the balance between both revegetation methods may ultimately depend on how species identity and traits such as seed and seedling size interact with environmental conditions. To test this, we will conduct a continental-scale experiment consisting of one mini-experiment replicated by multiple participants across Europe. Each participant will establish a site with seeded and planted individuals of one or more native, locally growing oak (Quercus) species; the selection of this genus aims to favour continental-scale participation and to allow testing the response of a widely distributed genus of broad ecological and economic relevance. At each site, participants will follow the present protocol for seed collection, seeding in the field, nursery cultivation, outplanting, protection against herbivores, site maintenance, and measurement of seedling performance and environmental variables. Each measurement on each species at each site will produce one effect size; the data will be analysed through mixed-effects meta-analysis. With this approach we will assess the main effect of revegetation method, species, plant functional traits, and the potential effect of site-specific effect moderators. Overall, we will provide a continental-scale estimate on the seeding vs. planting dilemma and analyse to what extent the differences in environmental conditions across sites, seed size, functional traits, and the phylogenetic relatedness of species can account for the differences in the effect of revegetation method on seedling performance across study sites and species.

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.

Initial Survival and Development of Planted European Beech (Fagus sylvatica L.) and Small-Leaved Lime (Tilia cordata Mill.) Seedlings Competing with Black Cherry (Prunus serotina Ehrh.).

Black cherry (Prunus serotina Ehrh.) is considered one of the most invasive tree species in central Europe and causes problems for both nature conservation and silviculture. Besides mechanical control treatments, a suggested control method to prevent its ongoing spread is to underplant shade-tolerant native tree species. Therefore, we combined two mechanical treatments, with underplanting of European beech (Fagus sylvatica L.) or small-leaved lime (Tilia cordata Mill.) on fenced and unfenced plots. After the first growing season, survival rates were evaluated, and selected seedlings were destructively harvested to analyze their growth performance and leaf morphology in association with the different light regimes resulting from mechanical treatments Survival rates for both seedlings were very high (>95%). Survival rates were higher on fenced plots than on unfenced plots, most likely as result of browsing. The mortality of F. sylvatica decreased with increasing light availability on fenced plots. The mortality of T. cordata did not change along the light gradient. After one vegetation period no differences with respect to biomass allocation could be detected along the light gradient. However, the specific leaf areas of both species responded similarly, decreasing with increasing light availability. In summary, both species were able to establish and survive in the dense P. serotina understory and might have the potential to outcompete the invasive alien species in the long run.

How a measure of tree structural complexity relates to architectural benefit-to-cost ratio, light availability, and growth of trees.

Aboveground tree architecture is neither fully deterministic nor random. It is likely the result of mechanisms that balance static requirements and light-capturing efficiency. Here, we used terrestrial laser scanning data to investigate the relationship between tree architecture, here addressed using the box-dimension (D b), and the architectural benefit-to-cost ratio, the light availability, and the growth of trees. We detected a clear relationship between D b and the benefit-to-cost ratio for the tested three temperate forest tree species (Fagus sylvatica L., Fraxinus excelsior L., and Acer pseudoplatanus L.). In addition, we could also show that D b is positively related to the growth performance of several tropical tree species. Finally, we observed a negative relationship between the strength of competition enforced on red oak (Quercus rubra L.) trees and their D b. We therefore argue that D b is a meaningful and integrative measure that describes the structural complexity of the aboveground compartments of a plant as well as its relation to structural efficiency (benefit-to-cost ratio), productivity, and growing conditions (competition or availability of light).

Advanced Aboveground Spatial Analysis as Proxy for the Competitive Environment Affecting Sapling Development.

Tree saplings are exposed to a competitive growth environment in which resources are limited and the ability to adapt determines general vitality and specific growth performance. In this study we analyzed the aboveground spatial neighborhood of oak [Quercus petraea (Matt.) Liebl.] and beech (Fagus sylvatica L.) saplings growing in Germany, by using hemispherical photography and terrestrial laser scanning as proxy for the competitive pressure saplings were exposed to. The hemispherical images were used to analyze the light availability and the three-dimensional (3D) point clouds from the laser scanning were used to assess the space and forest structure around the saplings. The aim was to increase the precision with which the biomass allocation, growth, and morphology of the saplings could be predicted by including more detailed information of their environment. The predictive strength of the models was especially increased through direct neighborhood variables (e.g., relative space filling), next to the light availability being the most important predictor variable. The biomass allocation patterns within the more light demanding oak were strongly driven by the space availability around the saplings. Diameter and height growth variables of both species reacted significantly to changes in light availability, and partly also to the neighborhood variables. The leaf morphology [as leaf-area ratio (LAR)] was also driven by light availability and decreased with increasing light availability. However, the branch morphology (as mean branch weight) could not be explained for oak and the model outcome for beech was hard to interpret. The results could show that individuals of the same species perform differently under constant light conditions but differing neighborhoods. Assessing the neighborhood of trees with highly precise measurement devices, like terrestrial laser scanners, proved to be useful. However, the primary response to a dense neighborhood seemed to be coping with a reduction of the lateral light availability aboveground, rather than responding to an increase of competition belowground. The results suggest continuing efforts to increase the precision with which plant environments can be described through innovative and efficient methods, like terrestrial laser scanning.

Assessing Understory Complexity in Beech-dominated Forests (Fagus sylvatica L.) in Central Europe-From Managed to Primary Forests.

Understory vegetation influences several ecosystem services and functions of European beech (Fagus sylvatica L.) forests. Despite this knowledge on the importance of understory vegetation, it is still difficult to measure its three-dimensional characteristics in a quantitative manner. With the recent advancements in terrestrial laser scanning (TLS), we now have the means to analyze detailed spatial patterns of forests. Here, we present a new measure to quantify understory complexity. We tested the approach for different management types, ranging from traditionally and alternatively managed forests and national parks in Germany to primary forests of Eastern Europe and the Ukraine, as well as on an inventory site with more detailed understory reference data. The understory complexity index (UCI) was derived from point clouds from single scans and tested for its relationship with forest management and conventional inventory data. Our results show that advanced tree regeneration is a strong driver of the UCI. Furthermore, the newly developed index successfully measured understory complexity of differently managed beech stands and was able to distinguish scanning positions located on and away from skid-trails in managed stands. The approach enables a deeper understanding of the complexity of understory structures of forests and their drivers and dependents.

Site-adapted admixed tree species reduce drought susceptibility of mature European beech.

Some forest-related studies on possible effects of climate change conclude that growth potential of European beech (Fagus sylvatica L.) might be impaired by the predicted increase in future serious drought events during the growing season. Other recent research suggests that not only multiyear increment rates but also growth resistance and recovery of beech during, respectively, after dry years may differ between pure and mixed stands. Thus, we combined dendrochronological investigations and wood stable isotope measurements to further investigate the impact of neighborhood diversity on long-term performance, short-term drought response and soil water availability of European beech in three major geographic regions of Germany. During the last four decades, target trees whose competitive neighborhood consisted of co-occurring species exhibited a superior growth performance compared to beeches in pure stands of the same investigation area. This general pattern was also found in exceptional dry years. Although the summer droughts of 1976 and 2003 predominantly caused stronger relative growth declines if target trees were exposed to interspecific competition, with few exceptions they still formed wider annual rings than beeches growing in close-by monocultures. Within the same study region, recovery of standardized beech target tree radial growth was consistently slower in monospecific stands than in the neighborhood of other competitor species. These findings suggest an improved water availability of beech in mixtures what is in line with the results of the stable isotope analysis. Apparently, the magnitude of competitive complementarity determines the growth response of target beech trees in mixtures. Our investigation strongly suggest that the sensitivity of European beech to environmental constrains depends on neighborhood identity. Therefore, the systematic formation of mixed stands tends to be an appropriate silvicultural measure to mitigate the effects of global warming and droughts on growth patterns of Fagus sylvatica.

Regeneration patterns of European oak species (Quercus petraea (Matt.) Liebl., Quercus robur L.) in dependence of environment and neighborhood.

Quercus robur L. (pedunculate oak) and Quercus petraea (Matt.) Liebl. (sessile oak) are two European oak species of great economic and ecological importance. Even though both oaks have wide ecological amplitudes of suitable growing conditions, forests dominated by oaks often fail to regenerate naturally. The regeneration performance of both oak species is assumed to be subject to a variety of variables that interact with one another in complex ways. The novel approach of this research was to study the effect of many ecological variables on the regeneration performance of both oak species together and identify key variables and interactions for different development stages of the oak regeneration on a large scale in the field. For this purpose, overstory and regeneration inventories were conducted in oak dominated forests throughout southern Germany and paired with data on browsing, soil, and light availability. The study was able to verify the assumption that the occurrence of oak regeneration depends on a set of variables and their interactions. Specifically, combinations of site and stand specific variables such as light availability, soil pH and iron content on the one hand, and basal area and species composition of the overstory on the other hand. Also browsing pressure was related to oak abundance. The results also show that the importance of variables and their combinations differs among the development stages of the regeneration. Light availability becomes more important during later development stages, whereas the number of oaks in the overstory is important during early development stages. We conclude that successful natural oak regeneration is more likely to be achieved on sites with lower fertility and requires constantly controlling overstory density. Initially sufficient mature oaks in the overstory should be ensured. In later stages, overstory density should be reduced continuously to meet the increasing light demand of oak seedlings and saplings.