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Assessment of forest functionality and the effectiveness of forest management and certification
(2021)
Forest ecosystems are complex systems that develop inherent structures and processes relevant for their functioning and the provisioning of ecosystem services that contribute to human wellbeing. With increasing climate change impacts, especially regulating ecosystem services such as microclimate regulation are ever more relevant to maintain forest functions and services. A key question is how forest management supports or undermines the ecosystems’ capacity to maintain those functions and services. The main objective of this thesis is the development of a concept to assess the functionality of forests and to evaluate the effectiveness of forest ecosystem management including certification. An ecosystem-based and participatory methodology, named ECOSEFFECT, was developed. The method comprises a theoretical and an empirical plausibility analysis. It was applied to the Russian National FSC Standard in the Arkhangelsk Region of the Russian Federation - where boreal forests are exploited to meet Europe's demand for timber. In addition, the influence of forestry interventions on temperature regulation in Scots pine and European beech forests in Germany was assessed during two extreme hot and dry years in 2018 and 2019. Microclimate regulation is a suitable proxy for forest functionality and can be applied easily to evaluate the effectiveness of forest management in safeguarding regulating forest functions relevant under climate change. Thus, the assessment of forest microclimate regulation serves as convenient tool to illustrate forest functionality. In the boreal and temperate forests studied in the frame of this thesis, timber harvesting reduced the capacity to self-regulate forests’ microclimate and thus impair a crucial part of ecosystem functionality. Changes in structural forest characteristics influenced by forest management and silviculture significantly affect microclimatic conditions and therefore forest ecosystems' vulnerability to climate change. Canopy coverage and the number of cut trees were most relevant for cooling maximum summer temperature in pine and beech forests in northern Germany. The Russian FSC standard has the potential to improve forest management and ecological outcomes, but there are shortcomings in the precision of targeting actual problems and ecological commitment. It is theoretically plausible that FSC prevents logging in high conservation value forests and intact forest landscapes, reduces the size and number of clearcuts, and prevents hydrological changes in the landscape. However, the standard was not sufficiently explicit and compulsory to generate a strong and positive influence on the identified problems and their drivers. Moreover, spatial data revealed, that the typical regular clearcut patterns of conventional timber harvesting continue to progress into the FSC-certified boreal forests, also if declared as "Intact Forest Landscape". This results in the need to verify the assumptions and postulates on the ground as it remains unclear and questionable if functions and services of boreal forests are maintained when FSC-certified clearcutting continues.The analysis of satellite-based data on tree cover loss showed that clearcutting causes secondary dieback in the surrounding of the cleared area. FSC-certification does not prevent the various negative impacts of clearcutting and thus fails to safeguard ecosystem functions. The postulated success in reducing identified environmental threats and stresses, e. g. through a smaller size of clearcuts, could not be verified on site. The empirical assessment does not support the hypothesis of effective improvements in the ecosystem. In practice, FSC-certification did not contribute to change clearcutting practices sufficiently to effectively improve the ecological performance. Sustainability standards that are unable to translate principles into effective outcomes fail in meeting the intended objectives of safeguarding ecosystem functioning. Clearcuts that carry sustainability labels are ecologically problematic and ineffective for the intended purpose of ecological sustainability.The overexploitation of provisioning services, i.e. timber extraction, diminishes the ecosystems' capacity to maintain other services of global significance. It also impairs ecosystem functions relevant to cope with and adapt to other stresses and disturbances that are rapidly increasing under climate change.
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, the authors 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 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. 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. The authors 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, the findings indicated a high drought sensitivity of Cantabrian beech populations, but also interaction effects of N and D on growth responses of beech seedlings.
The dissertation deals with the impact of nitrogen deposition on the functioning of heathland ecosystems. Special interests were the displacement of heather (Calluna vulgaris) by the purple moor-grass (Molinia caerulea) as well as the fate of nitrogen loads in dry heathland ecosystems. The results of the studies undertaken in the field and in the greenhouse are presented as five individual journal articles. The nature of nutrient limitation was studied by means of fertilisation experiments with nitrogen (N) and phosphorus for heather and purple moor-grass (Articles I and II). The impact of nitrogen deposition on the outcome of competition between these two species was analysed during a competition experiment in the greenhouse (Article III). The aim of a 15N tracer experiment was to determine the fate of nitrogen deposition as well as allocation patterns (Article IV). In addition, the response of purple moor-grass to the combined effects of nitrogen deposition and summer droughts was investigated in a second greenhouse experiment (Article V). The fertilisation experiments showed that the growth of heather as well as of purple moor-grass is predominantly limited by N (Articles I and II). However, the results of the competition experiment demonstrated that only purple moor-grass has the ability to benefit from additional N loads, which in turn gives the grass the opportunity to displace heather (Article III). Drought treatment resulted in strikingly reduced biomass production of purple moor-grass in N-fertilised pots, mainly as a result of dying aboveground biomass during dry periods (Article V). This striking susceptibility of purple moor-grass to the combination of nitrogen deposition and drought must be taken into account, when predicting future developments of dry heathlands. The results of the 15N tracer experiment showed that the investigated heath is still in an early stage of N saturation, as indicated by a high immobilisation capacity and negligible leaching losses of 15N (Article IV). The findings of the dissertation contribute to a better understanding of the processes underlying the encroachment of purple moor-grass in dry heathlands and can enhance heathland management. The results can also be used to to evaluate the current and future status of this ecosystem particularly with regard to the various stages of N saturation as well as in the determination of “Critical Loads”.
Restoration and management of abandoned, dry Continental heathland and sandy grassland communities
(2018)
Whereas heathlands in the Atlantic biogeographical region have been in the centre of interest, dry heathland communities in the Continental biogeographical region have been widely neglected. Thereby, the conservation status of both habitat types is listed as unfavourable-bad across the Continental biogeographical region. Surprisingly, no detailed studies are available on cost-efficient and sustainable restoration and management schemes to successfully restore and maintain highly degraded, long-abandoned Continental heaths and sandy grasslands, and thus, to counteract the poor conservation status of the habitat types. This shows the great need for research for the Continental biogeographical region. Thus, the present thesis provides substantial knowledge about the population dynamics of the key plant species of dry heaths Calluna vulgaris (L.) HULL by investigating key processes in the biology of the species as well as about the restoration and management of long-abandoned, dry Continental heathland and sandy grassland communities. In order to better understand the process of successful Continental heathland restoration, I analysed the reproductive potential (seed production, soil seed bank, and germination ability of seeds) of degenerate Calluna stands as well as the effects of single and combined management options on the generative rejuvenation (i.e., recruitment and survival) of Calluna. The results are based on a comprehensive three-year field experiment including the management options year-round, low-intensity cattle and horse grazing, one-time mowing and one-time shallow soil disturbances combined with greenhouse investigations on the soil seed bank content and germination ability of Calluna seeds. The results showed that even after long-term abandonment, seed production of degenerate Calluna stands and the germination ability of seeds proved to be high, being similar to Atlantic heathlands, whereas the soil seed bank is considerably reduced probably due to the dry conditions in the Continental region. In addition, low-intensity grazing with free-ranging robust breeds and the combination with one-time mowing at the beginning of the restoration process is an effective means of supporting the generative rejuvenation of this key plant species in degraded Continental heaths. The second study of this thesis focussed on the first-year establishment of Calluna in managed and unmanaged dry heaths and heaths in mosaics with dry sandy grasslands. The germination ability of seeds of different life-history phases of Calluna was analysed to determine if the predominance of the late life-history phase restricts the rejuvenation process of this key plant species. In addition, beside effects of management measures (year-round, low-intensity grazing, one-time mowing, one-time shallow soil disturbances) the author analysed the most important safe site conditions that possibly influenced the germination and the first-year survival of Calluna. The results of the study combine field experiments with growth chamber investigations. The author found that life-history phase of Calluna did not significantly affect seed germination and thus, the predominance of the degenerate life-history phase does not restrict the rejuvenation process. In addition, the results of the study revealed that grazing and thus trampling intensity must be temporarily and locally enhanced at the beginning of the restoration process of highly degraded heaths to increase safe site availability for successful Calluna establishment. Thereby, shadowing is the most important safe site condition. In the third study, the author investigated the impacts of year-round low-intensity cattle and horse grazing on the development of the highly competitive grass Calamagrostis epigejos (henceforth referred to as Calamagrostis), as well as the vegetation structure and plant species richness of long-abandoned but nutrient-poor dry heathland and sandy grassland communities, their mosaics and Calamagrostis stands. Finally, I assessed the local conservation status of the habitat types after seven years of grazing in comparison to long-abandoned sites. The results are based on a comprehensive field study on two spatial scales (plot-level: 25 square meters, macroplot-level: 1 ha). The author found that grazing successfully reduced the coverage and prevented the further spread of Calamagrostis, while simultaneously maintained or improved characteristic species richness and vegetation structure across the different nutrient-poor vegetation types over time, and thus enhanced the local conservation status of habitat types of community interest. In conclusion, the results of the studies considerably improved the understanding of dry, Continental heathland and sandy grassland restoration and management. They provide evidence that even after long-term management abandonment, year-round low-intensity cattle and horse grazing is a suitable management tool for restoring, maintaining and even improving nutrient-poor heathland and sandy grassland communities. However, at the beginning of the restoration process, additional management measures are necessary to faster restore abandoned habitats, especially highly degraded heaths.
The role of tree diversity for individual tree growth, crown architecture and branch demography
(2012)
In the light of the concurrent loss of biodiversity, biodiversity and ecosystem functioning (BEF) research attracted a great deal of attention and emerged as one of the important fields of research in ecology. Since important ecological interactions such as competition occur between individuals, the understanding of individual tree growth was considered to be fundamental for forest related BEF research. Individual tree growth is determined by the above- and belowground interactions of a tree individual with its local neighbourhood. To obtain a deeper understanding of BEF relationships, I broadened the focus from individual tree growth (usually measured as diameter or biomass increment) to the arrangement and dynamics of the above-ground modules of trees in dependence of their local neighbourhood. More precisely, the main objective of the present thesis was to analyse the impact of tree diversity on individual tree growth, crown architectural and branch demographic variables. Thereby I considered crown architectural variables as important indicators of the competition for light. In addition, crown architectural variables impacted ecosystem services such as erosion control. Furthermore, the results of the present thesis contributed to the current discussion on species coexistence theories, which may be differentiated by two opposing views: one that relies on neutral processes and one that implicates a role for meaningful differences in the ecological strategy (niche) of co-occurring species. The studied forest ecosystems were the subtropical broad-leaved evergreen forests of southeast China, which have been under high human pressure due to a long history of intensive land-use. The area is of particular interest for BEF research due to the high species richness of woody plants, including many, yet poorly studied species, and due to the rough terrain with steep slopes, which cause severe soil erosion. The present thesis combines three observational with two experimental studies, applying the local neighbourhood approach along an age gradient from tree saplings to mature trees. In the Gutianshan National Nature Reserve (GNNR), I conducted two observational studies on permanent plots which were chosen according to a space-for-time substitution design. The aim of the first study was to reveal the effects of diversity (species richness, functional diversity) together with other biotic and abiotic variables on morphological growth parameters (crown area, crown displacement and stem inclination) of target trees of four tree species (Castanea henryi, Castanopsis eyrei, Quercus serrata and Schima superba). In the second study, the same target trees together with their neighbours were used to analyse the relation between stand related functional diversity and the horizontal and vertical structure of the canopy. The third study was conducted in a young secondary broad-leaved evergreen forest. Using two target species (Castanopsis fargesii and Quercus fabri), the role of diversity, intra- vs. inter-specific competition and the mode of competition (symmetric vs. asymmetric) on the target individuals was tested by analysing five-year radial growth increments. The two other studies were carried out in an experimentally established plantation, using saplings of four tree species (C. henryi, Elaeocarpus decipiens, Q. serrata and S. superba), which were planted in monoculture, twoand four-species combinations and in three densities. The fourth study focused on mechanisms of coexistence and the role of species richness, species composition, species identity and density on sapling growth. The fifth study tested the effect of sapling density and identity on the througfall kinetic energy, which represents a measure for the erosive power of rain. It was found that functional diversity does affect crown architectural and canopy related parameters of forests in the GNNR. However, no effects of species richness on radial-growth were detected in the younger forest. Since I also did not find strong effects of species richness on saplings in the experimental plantation, diversity effects may evolve at a later age stage. The importance of the diversity effect may be related reversely to that of species identity in an age gradient of forest stands. The findings suggest that different mechanisms of coexistence operate simultaneously but that their relative importance may shift through the life stages of trees. During the sapling stage, species-specific differences in growth and architectural traits support niche theory. In older forest stands, no species-specific differences in growth parameters could be detected. However, I did find effects of functional diversity on horizontal canopy structure. I conclude that mechanisms of coexistence may not only change with forest stand age, but may also differ for distinct traits. The present thesis, being the first to apply the local neighbourhood approach with regard to crown architecture and branch demography within the BEF field of research, stresses the importance of this individual based approach. Although the observed forest systems are very complex, crown architectural and canopy structural variables were found to be affected by diversity. The finding that the degree of erosive power of rain could be elucidated by crown architectural variables, encourages further studies to reveal possible relations between biodiversity and other ecosystem functions or services, which might be mediated by crown architectural and canopy structural variables.
Biodiversity loss could jeopardize ecosystem functioning. Yet, the evidences that support this demonstration have been mostly obtained in aquatic and grassland ecosystems. Howbiodiversity affects ecosystem functioning still remain largely unanswered in forests, particularly in subtropical broad-leaved evergreen forests (EBLF). Tree productivity, among a wealth of forest ecosystem functioning, is of particular interest because it reflects the carbon sink capacity and wood productivity. Biodiversity-productivity relationships have been usually investigated at community level. However, tree-tree interactions occur at small scale. Thus, local neighborhood approach may allow a better understanding of tree-tree interactions and their contributions to the effects of biodiversity on tree productivity / growth rates. This thesis aims to analyze the effects of biodiversity and the abiotic environmental factors on the tree growth rates using both local neighborhood and community-based approaches. Furthermore, tree growth rates vary among different tree species. Functional traits have been related to the species-specific growth rates to understand the effects of species identity. Therefore, I also evaluated the crown- and leaf traits to predict the interspecific difference in growth rates. For a better understanding of the mechanisms that underline the relationships of biodiversity and tree growth rates, data of high solution and along time series is required to scrutinize the tree-tree interactions. Thereupon, I evaluated the applicability of terrestrial laser scanning (TLS) in assessing the tree dendrometrics. This thesis was conducted in the Biodiversity Ecosystem Functioning (BEF)–China experiment, which is located in a mountainous subtropical region in southeast China. A total of 40 native broad-leaved tree species were planted. In the first study, I used the local neighborhood approach to analyze how local abiotic conditions (i.e. topographic and edaphic conditions) and local neighborhood (i.e. species diversity and competition by neighborhood) affect the annual growth rates of 6723 individual trees. The second study used the community approach to partition the effects of environmental factors (i.e. topographic and edaphic), functional diversity according to Rao’s quadratic entropy (FDQ) and community weight mean (CWM) of 41 functional traits on community tree growth rates. The main question of the third study was how the species-specific growth rates are related to five crown- and 12 leaf traits.
In the fourth study, I investigated 438 tree individuals for the congruence between the conventional direct field measurements and TLS measurements. It was found that tree growth rates were strongly influenced by the local topographic and edaphic conditions but not affected by the diversity of local neighborhood. In contrast, results obtained by using the community-based approach showed that FDQ and CWMs of various leaf traits rather than abiotic environmental factors had significant impact on the community means of growth rates. Tree-tree interactions already occur in early life stages of trees, which were evidenced by the significant effect of competition by local neighborhood. These findings imply that the effects of abiotic environmental factors may be more evident at local scale and biodiversity effects may vary at different spatial scales. The species-specific growth rates were found to be related to specific leaf traits but not to crown traits and were best explained by both types of traits in combination. This finding supports the niche theory and provides the evidence for using functional diversity to examine the BEF relationships. The TLS-retrieved total tree height, stem diameter at 5 cm above ground, and length and height of the longest branch were highly congruent with those obtained from direct measurements. It indicates that TLS is a promising tool for high resolution, non-destructive analyses of tree structures in young tree plantations. Being one of very few studies to incorporate the individual tree scale in examining the biodiversity-productivity relationships within the BEF researches, this thesis stresses the importance of using individual-tree based approach, functional diversity and TLS to find the evidences of explanatory mechanisms of the observed biodiversity and ecosystem functioning (e.g. tree growth rates) relationships. Biodiversity effects may evolve along the successional stages. Therefore, incorporating the interaction between biodiversity and time in analyzing BEF relationship is also encouraged.
The timber shortage led to large scale afforestations on previously agriculturally used land in Central Europe during the 19th and 20th century. Widespread afforestation programs created recent forest ecosystems (i.e. young forest systems in terms of their development history). Despite the positive effect of increasing the forest area of Central Europe, the ecological effects of these land-use changes on forest ecosystems remain poorly understood. In order to increase the understanding of ecosystem processes in forests, an assessment of conceivable shifts in ecosystem functions caused by former land-use changes and forest management is required. By analysing aboveground growth rates of European beech (Fagus sylvatica L.) in response to environmental change drivers, such as climate extremes and nitrogen (N) deposition, the presented thesis aims to assess the role of land-use and management legacies in modulating present responses to drivers of environmental change. To this end, annual radial growth rates of individual trees were measured in mature beech stands. The investigated stands differed either in their land-use history (i.e. ancient forest sites versus recent forests) or their forest management history (i.e. managed forest sites versus short-term and long-term unmanaged forest sites). Measurements of radial growth rates were complemented by analyses of the fine root systems, soil chemical properties and crown projection areas to gain insights into the mechanisms underlying alterations in tree growth. Within the projects of the presented thesis, shifts in the climate-growth relationships driven by land-use and management legacies were analysed. In addition, land-use legacy mediated differences in the climate-nitrogen-growth relationships were assessed. The key findings are: (I) Soil legacy driven alterations in the fine root systems cause a higher sensitivity of radial increment rates to water deficits in summer for trees growing on recent forest sites than for trees growing on ancient forest sites. (II) Management legacies (in terms of tree release) enhance the sensitivity of beech’s radial growth to water deficits in spring through changes in crown sizes. (III) Interacting effects of spring water deficits and co-occurring high deposition of reactive N compounds lead to stronger radial growth declines in trees growing in ancient forests. This is likely caused by resource allocation processes towards seed production, which is, in turn, mirrored by decreasing radial growth rates. In this context, high N deposition likely boosts mass fructification in beech trees. Overall, it has been demonstrated that the ecological continuity plays a crucial role in modulating both climate sensitivity and the growth response to interacting effects of water deficits and nitrogen deposition in beech trees. The presented thesis identified a trade-off between the climate sensitivity and maximised growth rates within beech trees, depending on forest history. The results show that the growth of beech in ancient, unmanaged beech forests is less sensitive to water deficits than in recent and managed beech forests. Additionally, interacting effects of spring water deficits and N deposition likely increase the reproductive effort of beech trees, particularly in ancient forests. Thus, the results of this thesis once again underpin the uniqueness of ancient, unmanaged beech forests, whose importance for the conservation of biodiversity has been widely acknowledged. In summary, the presented thesis highlights the need to consider the "ecological memory" of forest ecosystems when predicting responses to current and future environmental changes.
Excessive fertilizer use leads to nutrient imbalances and losses of these to the environment through leaching, runoff and gaseous emissions. Nutrient use efficiency (NUE) in agriculture is often low and improving it could increase the sustainability of agricultural systems. The main aims of this thesis were to gain a better understanding of plant-soil-microbe interactions in order to improve agricultural NUEs. The studies included experimentally tested how crops respond to addition of high carbon amendments, fertilizer application rates and timing, and crop rotations. Furthermore, methods for measurement of roots were compared and a protocol for measurement of roots was developed. The first experiment simulated an agricultural field using mesocosms. In this setting, the researchers tested the effect of 4 previous crops (precrops), which either had or did not have a symbiosis with arbuscular mycorrhizal fungi (AMF)/rhizobia, on the focal crop (winter barley). They also tested the addition of high carbon amendments (wheat straw/sawdust) for immobilization of residual soil nitrogen (N) at harvest of the previous crop. Overall, the findings were that non-AMF precrops had a positive effect on winter barley yield compared to AMF precrops. Wheat straw reduced N leaching, whereas sawdust addition had a negative effect on the yield of winter barley. The second experiment tested the effect of different fertilizer (N/phosphorus (P)) application timings on plant traits grown in rhizoboxes. Overall, delaying N application had a more detrimental effect on plant biomass than delaying P application. The root system increased its root length initially due to N-deficiency, but was quickly thus N-limited that root length was relatively lower than the control group. Because of the many root related measurements in the second experiment, a step-by-step method for measuring root traits under controlled and field conditions was developed and included in this thesis. This method paper describes precisely how root traits of interest can be measured, and helps with deciding which approach should be taken depending on the experimental design. Additionally, the authors compared the bias and accuracy of several popular root measurement methods. Overall, these results highlight the importance of crop choice in crop rotations and the plasticity of root systems in relation to nutrient application. The results show high carbon amendments could reduce nitrate leaching after the harvest of crops, especially those with high risk of nitrate leaching, although they had only small impacts on yield.
European species-rich grasslands are threatened both by land use intensification as well as land abandonment. The studies shown in this thesis tested the possible use of ecological knowledge to ensure hay productivity whilst maintaining diversity of grasslands, with a view to informing ecological restoration. The overall approach was to understand interactions between plants, to study diversity effects on productivity, and mainly investigate how plant functional groups that arrive first in the system can create priority effects that influence community productivity both above- and belowground. A grassland field experiment was established and monitored for four years, in order to verify the effects of manipulating the order of arrival of different plant functional groups, as well as the sown diversity level on productivity and methane yield. The overall findings were: a) sowing legumes first created priority effects aboveground (higher biomass) and belowground (lower root length), plants invested less in roots and more in shoots, b) priority effects were more consistent below than aboveground, c) sown diversity did not affect aboveground biomass, d) the order of arrival treatments indirectly affected methane yield by affecting the relative dominance of plant functional groups. Since the researchers lack information on how legumes and non-legumes interact spatially belowground, (particularly related to root foraging) a controlled experiment was performed, using two grass species and one legume. The identity and location of the neighbours played a role in interactions, and the order plants arrived modulated it. When the focal species (grass) was growing with a legume it generally equated to the same outcome as not having a neighbour. Roots from the focal species grew more toward the legume than the grass neighbour, indicating a spatial component of facilitation. Since these studies involved root measurements, a method study was also conducted to verify how comparable and accurate are root length estimates obtained from different techniques. Results showed that the use of different methods can lead to different results, the studied methods did not have the same accuracy, and the automated methods can underestimate the root length. Overall, the results allow to conclude that different groups of plants arriving before others affected above and belowground biomass, roots may be key drivers during the creation of these priority effects, and interaction outcomes between plants depended on neighbour identity and location, modulated by the order they arrive in. The results suggest that priority effects can be used by sowing different species or plant functional groups at different time to steer a community to a desired trajectory depending on the restoration goal. However, there is a need to test contingency, potential, and long term impacts of such possible tools for restoration.