Restoration and management of abandoned, dry continental heathland and sandy grassland communities
(2018)
Land-use changes and long-term abandonment are main drivers that change ecosystem functioning and cause biodiversity loss of many semi-natural habitats, such as heathlands and grasslands. Traditional management measures such as low-intensity grazing created these semi-open landscapes and maintained a high species richness. However, from the middle of the 19th century onwards, traditional management practices began to decline due to socio-economic changes, and large areas of heathlands and grasslands were subjected to succession and reverted to woodland. Nowadays, dry heathlands as well as dry sandy grasslands are recognized as being of high conservation value and classified as ´habitats of community interest´ (´European dry heaths´, habitat code 4030; ´Xeric sand calcareous grasslands´, habitat code *6120). 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, even though they comprise 30 percent of all dry heaths as well as 89 percent of all dry sandy grasslands, respectively, in Europe. 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 (chapter I). Thus, the present thesis provides substantial knowledge about the population dynamics of the key plant species of dry heaths Calluna vulgaris (L.) HULL (henceforth referred to as Calluna) 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 (chapter II). 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 (chapter III) 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) I 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. I 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 my 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 that limit Calluna recruitment and survival in the first year in both degraded heaths as well as in mosaics with sandy grasslands, since seedlings receiving full sunlight die significantly more frequently than slightly or fully shaded seedlings. In the third study (chapter IV), I 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). I 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 my 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.
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.