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Die vorliegende Dissertation behandelt die Ökologie von Cnidium dubium (Schkuhr) Thell. (Sumpf-Brenndolde), Gratiola officinalis L. (Gottes-Gnadenkraut) und Juncus atratus Krocker (Schwarze Binse), drei gefährdeten Arten, die als sogenannte Stromtalpflanzen in Mitteleuropa in ihrem Vorkommen eng an die Flussauen gebunden sind. Die Arbeit basiert auf verschiedenen Simulationsexperimenten und Feldstudien in der Unteren Havelniederung, einem „Feuchtgebiet von internationaler Bedeutung“. Sie behandelt Themenkomplexe wie das Samenbankverhalten, die Samenkeimung, die Stickstofflimitierung, die Konkurrenzkraft, das Verhalten der Pflanzen nach einer Sommertrockenheit und nach einer Winter/Frühjahrsüberflutung. Ferner widmet sie sich der Populationsbiologie der Arten und dem Verhalten der Pflanzen nach besonderen Störungsereignissen wie Mahd, Herbivorie und der Sommerflut 2002. Der Leser erfährt, wie die Pflanzen in verschiedenen Lebensphasen auf die auentypische Umwelt reagieren und erhält umfassende Einblicke in physiologische Mechanismen, die der Anpassung an die typischen Bedingungen einer mitteleuropäischen Flussaue dienen. Eine Interpretation der Ergebnisse zeigt auf, welche der spezifischen Eigenschaften zur Gefährdung der drei Stromtalarten beitragen. Die Arbeit ist für den Arten-, Biotop- und Landschaftsschutz interessant. Darüber hinaus bietet sie zahlreiche Anknüpfungspunkte zur ökophysiologischen Grundlagenforschung. Die verstärkte Nutzung physiologischer Methoden bei der Klärung ökologischer Fragestellungen wird angeregt.
Introduction: Many semiarid regions around the world are presently experiencing significant changes in both climatic conditions and vegetation. This includes a disturbed coexistence between grasses and bushes also known as bush encroachment, and altered precipitation patterns with larger rain events. Fewer, more intense precipitation events might promote groundwater recharge, but depending on the structure of the vegetation also encourage further woody encroachment.
Materials and Methods: In this study, we investigated how patterns and sources of water uptake of Acacia mellifera (blackthorn), an important encroaching woody plant in southern African savannas, are associated with the intensity of rain events and the size of individual shrubs. The study was conducted at a commercial cattle farm in the semiarid Kalahari in Namibia (MAP 250 mm/a). We used soil moisture dynamics in different depths and natural stable isotopes as markers of water sources. Xylem water of fifteen differently sized individuals during eight rain events was extracted using a Scholander pressure bomb.
Results and Discussion: Results suggest the main rooting activity zone of A. mellifera in 50 and 75 cm soil depth but a reasonable water uptake from 10 and 25 cm. Any apparent uptake pattern seems to be driven by water availability, not time in the season. Bushes prefer the deeper soil layers after heavier rain events, indicating some evidence for the classical Walter’s two-layer hypothesis. However, rain events up to a threshold of 6 mm/day cause shallower depths of use and suggest several phases of intense competition with perennial grasses. The temporal uptake pattern does not depend on shrub size, suggesting a fast upwards water flow inside. d2H and d18O values in xylem water indicate that larger shrubs rely less on upper and very deep soil water than smaller shrubs. It supports the hypothesis that in environments where soil moisture is highly variable in the upper soil layers, the early investment in a deep tap-root to exploit deeper, more reliable water sources could reduce the probability of mortality during the establishment phase. Nevertheless, independent of size and time in the season, bushes do not compete with potential groundwater recharge. In a savanna encroached by A. mellifera, groundwater will most likely be affected indirectly.
Major drivers of savanna shrub encroachment are climatic conditions, CO2 and unsustainable grazing management including fire prevention. Although all drivers affect ecohydrological processes, and given that water is a seasonally scarce resource in savannas, it remains largely unclear how shrub encroachment itself affects hydrological conditions that feed back into water use and community assembly of the remaining plant community. Hence, understanding direct ecohydrological effects of shrubs that may limit the recovery of the perennial herbaceous vegetation in grazed areas and promote the establishment of shrub seedlings facilitates the identification of areas that are most sensitive to further encroachment. In our trait-based approach, we determined relationships among shrub cover, soil and plant trait characteristics sensitive to water limitation in 120 plots along a shrub cover gradient. We focused on two functional response traits indicating immediate drought stress and subsequent water use for drought stress recovery with associated competition for water (midday leaf/xylem water potential and diurnally recovery rate of leaf water potential), and three functional response traits indicating long-term stress adaptation and related resource use strategies (SLA, plant height and seed release height). To understand species assembly and the associated mechanisms of resource use, we calculated community weighted mean traits, intraspecific trait variability as a proxy for the mechanism of coexistence, and mean traits at plant functional type level including 2-year-old Acacia mellifera-saplings. We found a low intraspecific trait variability in drought stress recovery rate and height suggesting that competitive exclusion via active resource acquisition (i.e. water exploitation) played a minor role for community assembly in a shrub encroaching savanna. The dominant community assembly process was passive stress avoidance via resource conservation up to stress tolerance indicated by the high variability in SLA and midday leaf water potential. Correlations of traits with soil moisture suggest a rooting niche differentiation between annual and perennial grasses and that Acacia-shrub saplings within the first 50 cm of soil already escaped the highest drought stress. Interestingly, immediate drought stress for the herbaceous community was lowest on moderately shrub encroached sites and not on grass dominated sites. Since passive stress avoidance accompanied by a distinct stress tolerance in semi-arid savannas is more important than active competition, and assuming that the low drought stress of the herbaceous community at intermediate levels of shrub cover also applies to newly emerging shrub seedlings, these areas are likely to be most sensitive to further encroachment. As such, they should be considered as focal areas for prevention management.
Encroachment of shrubs into the unique pastoral grassland ecosystems of the Tibetan Plateau has significant impact on ecosystem services, especially forage production. We developed a process-based ecohydrological model to identify the relative importance of the main drivers of shrub encroachment for the alpine meadows within the Qinghai province. Specifically, we explored the effects of summer livestock grazing (intensity and type of livestock) together with the effects of climate warming, including interactions between herbaceous and woody vegetation and feedback loops between soil, water and vegetation. Under current climatic conditions and a traditional herd composition, an increasing grazing intensity above a threshold value of 0.32 +/- 0.10 large stock units (LSU) ha(-1) day(-1) changes the vegetation composition from herbaceous towards a woody and bare soil dominated system. Very high grazing intensity (above 0.8 LSU ha(-1) day(-1)) leads to a complete loss of any vegetation. Under warmer conditions, the vegetation showed a higher resilience against livestock farming. This resilience is enhanced when the herd has a higher browser : grazer ratio. A cooler climate has a shrub encroaching effect, whereas warmer conditions increase the cover of the herbaceous vegetation. This effect was primarily due to season length and an accompanied competitive loss of slower growing shrubs, rather than evaporative water loss leading to less soil water in deeper soil layers for deeper rooting shrubs. If climate warming is driving current shrub encroachment, we conclude it is only indirectly so. It would be manifest by an advancing shrubline and could be regarded as a climatic escape of specific shrub species such as Potentilla fruticosa. Under the recent high intensity of grazing, only herding by more browsing animals can potentially prevent both shrub encroachment and the complete loss of herbaceous vegetation.
Fire and grazing shape biodiversity in savannah landscapes. In land use management, knowing the effects of fire and grazing on biodiversity are important in order to ensure environmental sustainability. Beetles specifically are indicators of the biodiversity response to fire and grazing. A grazing exclusion and burning experiment in a split-plot design was used in order to investigate the interacting effects of fire and wildlife grazing on biomass, diversity, and species composition of darkling beetles (Coleoptera, Tenebrionidae) over time after fire. Darkling beetle species richness and diversity were responding in a three-way-interaction to fire, grazing, and time after fire, whereby biomass of darkling beetles remained unaffected and species compositional changes were attributed to seasonal changes of time only. Fire on ungrazed plots had a negative effect on species diversity and richness 2 weeks and 6 months post fire, whereas fire on grazed plots had no impact on species diversity and richness. Grazing only lowered species diversity and richness 6 months after fire treatments. Results suggest that grazing overrides the effects of fire and that the similar effects caused by fire and grazing are due to niche and assemblage simplification of the habitat.
As a contribution to conservation, we investigated germination requirements of three perennial, endangered river corridor plants of Central European lowlands coexisting in subcontinental flood meadows, but preferring particular zones of decreasing flooding frequency and duration along the elevational gradient of the banks. It was hypothesized that the species have specific germination requirements to respond successfully to open patch creation depending on their occurrence along the gradient of spring flooding in the field. This study involved controlled experiments and phenological studies. Juncus atratus and Gratiola officinalis, which frequently occupy flooded, naturally disturbed sites, have an absolute light requirement for germination, typical of pioneer species. Summer-dispersed, non-dormant seeds off. atratus did hardly germinate at high temperatures and lacked a gap sensitivity based on temperature fluctuation. Since the temperature amplitude decreases beneath an insulating cover of vegetation or water, seeds seem to be prepared for rapid germination at open, wet, maybe even inundated sites. Late-summer-dispersed seeds of G. officinalis were in a state of conditional primary dormancy. Dormancy could be completely broken by cold-wet stratification, indicating spring germination. Similar to J. atratus, daily temperature fluctuations did not control germination at suitable microsites. In Cnidium dubium that occurs at higher elevated sites, the level of primary dormancy of seeds was sufficient to prevent germination following dispersal, but the level was dependent on the year of harvest. Buried seeds showed an annual dormancy/conditional dormancy cycle. Dormancy was only partially broken by cold- wet stratification. It was completely broken by application of a high concentration of gibberellic acid. C. dubium had no absolute light requirement for germination, but it was stimulated by high light levels and in contrast to the other two species, seeds were stimulated by daily temperature fluctuations. Germination would therefore be maximized by zaps in early spring when the flooding water has receded. Re-entering dormancy in the late spring fails to support that germination occurs immediately after early-summer mowing - an important factor at subcontinental flood meadows.
Eastern Mediterranean ecosystems are prone to desertification when under grazing pressure. Therefore, management of grazing intensity plays a crucial role to avoid or to diminish land degradation and to sustain both livelihoods and ecosystem functioning. The dynamic land-use model LandSHIFT was applied to a case study on the country level for Jordan. The impacts of different stocking densities on the environment were assessed through a set of simulation experiments for various combinations of climate input and assumptions about the development of livestock numbers. Indicators used for the analysis include a set of landscape metrics to account for habitat fragmentation and the "Human Appropriation of Net Primary Production" (HANPP), i.e., the difference between the amount of net primary production (NPP) that would be available in a natural ecosystem and the amount of NPP that remains under human management. Additionally, the potential of the economic valuation of ecosystem services, including landscape and grazing services, as an analysis concept was explored. We found that lower management intensities had a positive effect on HANPP but at the same time resulted in a strong increase of grazing area. This effect was even more pronounced under climate change due to a predominantly negative effect on the biomass productivity of grazing land. Also Landscape metrics tend to indicate decreasing habitat fragmentation as a consequence of lower grazing pressure. The valuation of ecosystem services revealed that low grazing intensity can lead to a comparatively higher economic value on the country level average. The results from our study underline the importance of considering grazing management as an important factor to manage dry-land ecosystems in a sustainable manner.
Introduction: Many semiarid regions around the world are presently experiencing significant changes in both climatic conditions and vegetation. This includes a disturbed coexistence between grasses and bushes also known as bush encroachment, and altered precipitation patterns with larger rain events. Fewer, more intense precipitation events might promote groundwater recharge, but depending on the structure of the vegetation also encourage further woody encroachment.
Materials and Methods: In this study, we investigated how patterns and sources of water uptake of Acacia mellifera (blackthorn), an important encroaching woody plant in southern African savannas, are associated with the intensity of rain events and the size of individual shrubs. The study was conducted at a commercial cattle farm in the semiarid Kalahari in Namibia (MAP 250 mm/a). We used soil moisture dynamics in different depths and natural stable isotopes as markers of water sources. Xylem water of fifteen differently sized individuals during eight rain events was extracted using a Scholander pressure bomb.
Results and Discussion: Results suggest the main rooting activity zone of A. mellifera in 50 and 75 cm soil depth but a reasonable water uptake from 10 and 25 cm. Any apparent uptake pattern seems to be driven by water availability, not time in the season. Bushes prefer the deeper soil layers after heavier rain events, indicating some evidence for the classical Walter’s two-layer hypothesis. However, rain events up to a threshold of 6 mm/day cause shallower depths of use and suggest several phases of intense competition with perennial grasses. The temporal uptake pattern does not depend on shrub size, suggesting a fast upwards water flow inside. d2H and d18O values in xylem water indicate that larger shrubs rely less on upper and very deep soil water than smaller shrubs. It supports the hypothesis that in environments where soil moisture is highly variable in the upper soil layers, the early investment in a deep tap-root to exploit deeper, more reliable water sources could reduce the probability of mortality during the establishment phase. Nevertheless, independent of size and time in the season, bushes do not compete with potential groundwater recharge. In a savanna encroached by A. mellifera, groundwater will most likely be affected indirectly.
Shrub encroachment in semi-arid savannas is induced by interacting effects of climate, fire suppression, and unsustainable livestock farming; it carries a severe risk of land degradation and strongly influences natural communities that provide key ecosystem functions. However, species-specific effects of shrub cover on many animal groups that act as indicators of degradation remain largely unknown. We analysed the consequences of shrub encroachment for ground-dwelling beetles in a semi-arid Namibian savanna rangeland, where beetles and vegetation were recorded along a shrub cover gradient (30%). Focusing on species niche breadths and optima, we identified two crucial shrub cover thresholds (2.9% and 10.0%), corresponding to major changes in the beetle communities with implications for savanna ecosystem functioning. Niche optima of most species were between the first and second thresholds; beyond the second threshold, saprophagous, coprophagous, and rare predatory beetles declined in numbers and diversity. This is problematic because beetles provide important ecosystem functions, such as decomposition and nutrient cycling. However, we also found that certain species were adapted to high shrub cover, thus providing examples of niche differentiation. Despite the predominantly negative effects of heavy shrub encroachment on beetle communities, shrubs in their early life stages apparently provide essential structures, which enhance habitat quality for ground-dwelling beetles. Our results demonstrate that shrub encroachment can have mixed effects on ground-dwelling beetle communities and hence on savanna ecosystem functioning. We, therefore, conclude that rangeland management and restoration should consider the complex trade-offs between species-specific effects and the level of encroachment for sustainable land use.