Andreas Fichtner

Does Forest Continuity Enhance the Resilience of Trees to Environmental Change

There is ample evidence that continuously existing forests and afforestations on previously agricultural land differ with regard to ecosystem functions and services such as carbon sequestration, nutrient cycling and biodiversity. However, no studies have so far been conducted on possible long-term (>100 years) impacts on tree growth caused by differences in the ecological continuity of forest stands. In the present study we analysed the variation in tree-ring width of sessile oak (Quercus petraea (Matt.) Liebl.) trees (mean age 115–136 years) due to different land-use histories (continuously existing forests, afforestations both on arable land and on heathland). We also analysed the relation of growth patterns to soil nutrient stores and to climatic parameters (temperature, precipitation). Tree rings formed between 1896 and 2005 were widest in trees afforested on arable land. This can be attributed to higher nitrogen and phosphorous availability and indicates that former fertilisation may continue to affect the nutritional status of forest soils for more than one century after those activities have ceased. Moreover, these trees responded more strongly to environmental changes – as shown by a higher mean sensitivity of the tree-ring widths – than trees of continuously existing forests. However, the impact of climatic parameters on the variability in tree-ring width was generally small, but trees on former arable land showed the highest susceptibility to annually changing climatic conditions. We assume that incompletely developed humus horizons as well as differences in the edaphon are responsible for the more sensitive response of oak trees of recent forests (former arable land and former heathland) to variation in environmental conditions. We conclude that forests characterised by a long ecological continuity may be better adapted to global change than recent forest ecosystems.

From competition to facilitation: how tree species respond to neighbourhood diversity

Studies on tree communities have demonstrated that species diversity can enhance forest productivity, but the driving mechanisms at the local neighbourhood level remain poorly understood. Here, we use data from a large-scale biodiversity experiment with 24 subtropical tree species to show that neighbourhood tree species richness generally promotes individual tree productivity. We found that the underlying mechanisms depend on a focal trees functional traits: For species with a conservative resource-use strategy diversity effects were brought about by facilitation, and for species with acquisitive traits by competitive reduction. Moreover, positive diversity effects were strongest under low competition intensity (quantified as the total basal area of neighbours) for acquisitive species, and under high competition intensity for conservative species. Our findings demonstrate that net biodiversity effects in tree communities can vary over small spatial scales, emphasising the need to consider variation in local neighbourhood interactions to better understand effects at the community level.

High Trophic Similarity in the Sympatric North European Trawling Bat Species Myotis daubentonii and Myotis dasycneme

Most European bat species are insectivorous and share foraging areas to some extent. Where similar species rely on similar resources in the same foraging habitat, they are likely to interact. This study addresses the trophic niche of the Northern European trawling bat species Myotis dasycneme (Boie, 1825) and Myotis daubentonii (Kuhl, 1817), occurring in the same habitat, and possible interactions or differences within their dietary behaviour. Dietary data of both species were analysed to draw conclusions on their ecology, possible dietary overlap, hints for coexistence mechanisms and community structure. In this study, M. dasycneme and M. daubentonii fed mainly on Chironomidae (M. dasycneme: 44.4%; M. daubentonii: 32.8%) and Trichoptera (M. dasycneme: 20.4%; M. daubentonii: 22.2%), showing a high trophic niche overlap and similar niche breadth. Nevertheless, there were differences in the diet of the two species concerning the predation of chironomids. Differences also occur regarding the prey types, referring to the terrestrial or aquatic life-cycle of prey groups. This could be evidence for different foraging habitats and a spatial segregation of both species. High resource abundance is also likely to allow the coexistence of both species within the same hunting habitat.

Nitrogen Addition Enhances Drought Sensitivity of Young Deciduous Tree Species

Understanding how trees respond to global change drivers is central to predict changes in forest structure and functions. Although there is evidence on the mode of nitrogen (N) and drought (D) effects on tree growth, our understanding of the interplay of these factors is still limited. Simultaneously, as mixtures are expected to be less sensitive to global change as compared to monocultures, we aimed to investigate the combined effects of N addition and D on the productivity of three tree species (Fagus sylvatica, Quercus petraea, Pseudotsuga menziesii) in relation to functional diverse species mixtures using data from a 4-year field experiment in Northwest Germany. Here we show that species mixing can mitigate the negative effects of combined N fertilization and D events, but the community response is mainly driven by the combination of certain traits rather than the tree species richness of a community. For beech, we found that negative effects of D on growth rates were amplified by N fertilization (i.e., combined treatment effects were non-additive), while for oak and fir, the simultaneous effects of N and D were additive. Beech and oak were identified as most sensitive to combined N+D effects with a strong size-dependency observed for beech, suggesting that the negative impact of N+D becomes stronger with time as beech grows larger. As a consequence, the net biodiversity effect declined at the community level, which can be mainly assigned to a distinct loss of complementarity in beech-oak mixtures. This pattern, however, was not evident in the other species-mixtures, indicating that neighborhood composition (i.e., trait combination), but not tree species richness mediated the relationship between tree diversity and treatment effects on tree growth. Our findings point to the importance of the qualitative role (‘trait portfolio’) that biodiversity play in determining resistance of diverse tree communities to environmental changes. As such, they provide further understanding for adaptive management strategies in the context of global change.

Does excess nitrogen supply increase the drought sensitivity of European beech (Fagus sylvatica L.) seedlings

Climate change and atmospheric deposition of nitrogen affect biodiversity patterns and functions of forest ecosystems worldwide. Many studies have quantified tree growth responses to single global change drivers, but less is known about the interaction effects of these drivers at the plant and ecosystem level. In the present study, we conducted a full-factorial greenhouse experiment to analyse single and combined effects of nitrogen fertilization (N treatment) and drought (D treatment) on 16 morphological and chemical response variables (including tissue δ13C signatures) of one-year-old Fagus sylvatica seedlings originating from eight different seed families from the Cantabrian Mountains (NW Spain). Drought exerted the strongest effect on response variables, reflected by decreasing biomass production and increasing tissue δ13C signatures. However, D and N treatments interacted for some of the response variables, indicating that N fertilization has the potential to strengthen the negative effects of drought (with both antagonistic and amplifying interactions). For example, combined effects of N and D treatments caused a sevenfold increase of necrotic leaf biomass. We hypothesize that increasing drought sensitivity was mainly attributable to a significant reduction of the root biomass in combined N and D treatments, limiting the plants’ capability to satisfy their water demands. Significant seed family effects and interactions of seed family with N and D treatments across response variables suggest a high within-population genetic variability. In conclusion, our findings indicated a high drought sensitivity of Cantabrian beech populations, but also interaction effects of N and D on growth responses of beech seedlings.

Local variability in the diet of Daubenton's bat (Myotis daubentonii) in a lake landscape of northern Germany

Abstract. Daubentons bat Myotis daubentonii is one of the most common bat species of Europe, hunting its prey in the surroundings of water bodies, with different microhabitats. To explore the local adaptability to different environmental conditions, we compared the diet of Daubentons bats at four different sites in a lake landscape in northern Germany with a main focus on prey diversity. Bats were caught (n = 85) in mist nets for collecting individual faecal pellets (n = 276). Pellets were dissected and the occurrence of identifiable pieces of each prey group was evaluated and grouped in five different frequency groups. We found 17 different groups of arthropods among the prey, with a clear dominance of Chironomidae and Trichoptera. There were significant differences among the sampling sites in prey diversity but not in prey richness.The changes in prey diversity were associated with sample sites. We conclude that on a local scale there is low variability in diet of Daubentons bat caused by hunting in various habitat structures in the surroundings of water bodies. Our results highlight the ecological flexibility of M. daubentonii, which could be an explanation for the commonness of M. daubentonii across Europe in comparison to the rather rare pond bat (Myotis dasycneme), which has similar habitats and main prey group preferences.

Facilitative-Competitive Interactions in an Old-Growth Forest: The Importance of Large-Diameter Trees as Benefactors and Stimulators for Forest Community Assembly

The role of competition in tree communities is increasingly well understood, while little is known about the patterns and mechanisms of the interplay between above- and belowground competition in tree communities. This knowledge, however, is crucial for a better understanding of community dynamics and developing adaptive near-natural management strategies. We assessed neighbourhood interactions in an unmanaged old-growth European beech (Fagus sylvatica) forest by quantifying variation in the intensity of above- (shading) and belowground competition (crowding) among dominant and co-dominant canopy beech trees during tree maturation. Shading had on average a much larger impact on radial growth than crowding and the sensitivity to changes in competitive conditions was lowest for crowding effects. We found that each mode of competition reduced the effect of the other. Increasing crowding reduced the negative effect of shading, and at high levels of shading, crowding actually had a facilitative effect and increased growth. Our study demonstrates that complementarity in above- and belowground processes enable F. sylvatica to alter resource acquisition strategies, thus optimising tree radial growth. As a result, competition seemed to become less important in stands with a high growing stock and tree communities with a long continuity of anthropogenic undisturbed population dynamics. We suggest that growth rates do not exclusively depend on the density of potential competitors at the intraspecific level, but on the conspecific aggregation of large-diameter trees and their functional role for regulating biotic filtering processes. This finding highlights the potential importance of the rarely examined relationship between the spatial aggregation pattern of large-diameter trees and the outcome of neighbourhood interactions, which may be central to community dynamics and the related forest ecosystem services.

The reproductive potential and importance of key management aspects for successful Calluna vulgaris rejuvenation on abandoned Continental heaths

Abstract The abandonment of traditional pastoralism as well as the use of heath areas for military purposes has had a major impact on dry heaths in the Continental biogeographical region of Europe, causing severe degradation of its key species Calluna vulgaris (L.) HULL. The reproductive potential of this species in a Continental climate is assumed to be low, although there is yet no observational or experimental evidence for this. More knowledge is also needed about cost‐effective and sustainable measures to restore abandoned dry heaths in this biogeographical region, because traditional management options are often too expensive (e.g., sod‐cutting) or restricted due to environmental laws and the danger of unexploded ammunition (e.g., burning). Using as an example an 800 ha Continental heathland in Germany that has been abandoned for about two decades, we studied the reproductive potential (seed production, soil seed bank, and germination ability) of degenerate C. vulgaris stands. In addition, we conducted a comprehensive field experiment to test the effects of low‐intensity, year‐round grazing by Heck cattle and Konik horses as well as one‐time mowing and patchy exposure of bare soil on the generative rejuvenation (i.e., recruitment and survival) of degenerate C. vulgaris stands over 3 years. We used generalized linear mixed models for statistical analyses. Seed production of degenerate C. vulgaris stands was high as well as the germination ability of their seeds, being similar to Atlantic heathlands. However, soil seed‐bank densities were lower than those found in managed or abandoned Atlantic heaths. Overall seedling recruitment in the field was considerably lower in comparison with Atlantic heaths. Low‐intensity grazing or one‐time mowing did not induce a substantial increase in C. vulgaris recruitment, whereas an additional one‐time creation of bare soil patches or the one‐time creation of bare soil without subsequent management significantly facilitated seedling recruitment and survival in the first year. However, from the second year on, the positive effect of the creation of bare soil without subsequent management was no longer present. In the third year, survival of juveniles was significantly supported by low‐intensity grazing in combination with shallow soil disturbances as well as in combination with one‐time mowing and shallow soil disturbances, whereas mowing alone resulted in marginally significant lower survival. The extremely low seedling recruitment requires a careful choice of suitable management measures to promote the survival of sufficient numbers of Calluna individuals. Therefore, we recommend low‐intensity grazing with free‐ranging robust breeds and the combination of this with one‐time mowing as an effective means of supporting generative rejuvenation of C. vulgaris in degraded heaths. However, at the beginning of the restoration process, the creation of bare soil patches for seedling recruitment is crucial. For implementation into practice, we present different strategies to enhance the proportion of bare soil after long‐term abandonment of heaths when traditional management options are no longer feasible.

Neighbourhood interactions drive overyielding in mixed-species tree communities

Theory suggests that plant interactions at the neighbourhood scale play a fundamental role in regulating biodiversity–productivity relationships (BPRs) in tree communities. However, empirical evidence of this prediction is rare, as little is known about how neighbourhood interactions scale up to influence community BPRs. Here, using a biodiversity–ecosystem functioning experiment, we provide insights into processes underlying BPRs by demonstrating that diversity-mediated interactions among local neighbours are a strong regulator of productivity in species mixtures. Our results show that local neighbourhood interactions explain over half of the variation in observed community productivity along a diversity gradient. Overall, individual tree growth increased with neighbourhood species richness, leading to a positive BPR at the community scale. The importance of local-scale neighbourhood effects for regulating community productivity, however, distinctly increased with increasing community species richness. Preserving tree species diversity at the local neighbourhood scale, thus seems to be a promising way for promoting forest productivity.Though biodiversity is expected to be important in productivity in tree communities, there is little empirical evidence of this at local scales. Here, Fichtner et al. show that higher neighbourhood species richness increased tree growth, explaining over half of the variation in community productivity.

Legacy effects of land-use modulate tree growth responses to climate extremes

Climate change can impact forest ecosystem processes via individual tree and community responses. While the importance of land-use legacies in modulating these processes have been increasingly recognised, evidence of former land-use mediated climate-growth relationships remain rare. We analysed how differences in former land-use (i.e. forest continuity) affect the growth response of European beech to climate extremes. Here, using dendrochronological and fine root data, we show that ancient forests (forests with a long forest continuity) and recent forests (forests afforested on former farmland) clearly differ with regard to climate–growth relationships. We found that sensitivity to climatic extremes was lower for trees growing in ancient forests, as reflected by significantly lower growth reductions during adverse climatic conditions. Fine root morphology also differed significantly between the former land-use types: on average, trees with high specific root length (SRL) and specific root area (SRA) and low root tissue density (RTD) were associated with recent forests, whereas the opposite traits were characteristic of ancient forests. Moreover, we found that trees of ancient forests hold a larger fine root system than trees of recent forests. Our results demonstrate that land-use legacy-mediated modifications in the size and morphology of the fine root system act as a mechanism in regulating drought resistance of beech, emphasising the need to consider the ‘ecological memory’ of forests when assessing or predicting the sensitivity of forest ecosystems to global environmental change.