Plasticity, Intraspecific Variability and Local Adaptation to Climatic Extreme Events of Ecotypes/Provenances of Key Plant Species
- Climate change, and especially an increase of magnitude and frequency of climatic extreme events such as drought periods or heatwaves, will alter growing conditions for plants in the future. Persistent ecosystems, with long-living organisms, such as forest or permanent grassland will be particularly impacted by this development. The velocity of these changes is likely to occur at a pace, which species may not be able to keep track with by natural dispersal or genetic adaptation. Agriculture, forestry and ecosystem management must develop counteracting practices to secure the persistence and functioning of these ecosystems and thus their provision of goods and services. Therefore it is important to develop a better understanding how species and ecosystems may respond to future climatic stressors. Impact assessments, e.g. via climatic envelope modelling are prone to misinterpretations of the adaptive capacity of species, as they do not incorporate the intraspecific genetic and phenotypic differences that exist within the populations accross the distribution range of a species.
Yet, intraspecific variation may exhibit potential tools for the development of climate change adaptation strategies. Here, I focus on key ecosystems in Central Europe. In particular the selective use of plant provenances or ecotypes may help to make ecosystems climate-resilient without a potentially more problematic introduction of exotic species. Especially provenances from warmer, drought-prone regions, with a current climate similar to the projected one for Central Europe recently came into focus as potential substitutes for local provenances, as they might have developed local adaptations to climate conditions at their location of origin. Insights about the response of these provenances to changing averages and extreme event regimes are crucial for a reasonable use of within-species diversity in climate change adaptation.
First, the concept of assisted colonization or migration of species or ecotypes and the role it can play as an adaptation strategy in agriculture, forestry or nature conservation is introduced (Manuscript1). It is suggested that a focus should be laid on keystone species that ensure ecosystem persistence and functioning as they govern the habitat structure and microclimate of a site. The assisted colonization of pre-adapted ecotypes of keystone species from climates similar to future projections for the target site is proposed.
Furthermore, provenances of selected grassland and forest key-species were exposed to drought and warming in two experiments in Bayreuth and Landau, and their ecological responses were analysed. Results suggest that local adaptations to climatic stressors exist. However, the magnitude and direction of responses strongly depend on species and climatic variables. For grassland species, e.g. differences in drought sensitivity could be demonstrated in some cases (Manuscript 4). Fagus sylvatica exhibited differences between the provenances in response to drought conditions, as well (Manuscript 3). It seems that marginal provenances, from the dry margins of the distribution range, show less increment reduction due to the drought treatment. Yet, under more favourable conditions of water supply these provenances did not yield the same high increment rates than more central provenances, indicating a trade-off between stability under stress and yield under non-stress conditions. A pine species that is generally considered to be rather drought-resistant, Pinus nigra, which is a potential substitute for climate-threatened conifers on dry sites in Central Europe, did not show any differences in response to drought and warming (Manuscript 2), maybe due to a weak selective pressure as a result of high drought-resistance across the whole distribution range. The impacts of drought on increment became not visible before the second year after the treatment, stressing the need for more long-time experiments in climate impact research.
Even in a generally warmer environment, cold extremes in winter or spring are expected still to prevail in the future. Therefore, the provenances of the selected species were tested for their cold-hardiness and late frost resistance (Manuscripts 5-7). Growth of the grassland species and F. sylvatica were negatively impacted by a late frost event and differences in late-frost sensitivity between provenances or ecotypes were identified. The (sub-) mediterranean species P. nigra showed differences between provenances in their winter cold hardiness. Correlations between performance under cold stress and winter conditions or late frost proneness of the places of origin could be established for almost all species. However, preceding climate experience, such as the warming or drought treatment of the plants altered their reaction to cold extremes compared to the control treatment, indicating the complexity of the interactive impacts of climate factors on ecosystem and plant performance.
The uncertainty of climate projections and the multitude of changing climatic stressors, though, make the prospect of an easy and rapid success in the search for single “best-adapted” provenances very questionable. In economics the portfolio effect shows that a diversification of investments decreases the risk of a total loss of profits. Hence, in a modelling procedure based on the increment data from the above mentioned experiment it was tested if a “portfolio investment” in several provenances in one stand decreases the risk of yield losses (Manuscript 8). Results indicate that the higher the number of provenances the higher the chance for a “best-performer” to be included in the set. So the likelihood of higher yields, under different climatic conditions increases, yet the risk of low yields stays stable.
Generally, it seems that the selective use of plant species and ecotypes in climate change adaptation can be a feasible tool to maintain ecosystem functionality and productivity. However, the uncertain projections, the multitude of climatic stressors and their interplay with other environmental factors and the potential impacts of assisted colonization of ecotypes on the genetic diversity within species and populations require further research.
Beyond productivity- Effects of extreme weather events on ecosystem processes and biotic interactions
- Under global climate change, extreme weather events, such as heat waves, drought or heavy rain spells, are projected to increase in magnitude and frequency. As these may affect vegetation and ecosystems more than gradual shifts in mean climatic parameters, investigating the consequences of extreme weather events recently became an important issue in climate change research. The main focus of most experiments investigating effects of extreme weather events on vegetation is on primary productivity. In our experiment in artificially planted communities, even an extreme drought of 1000-year recurrence did not have effects on above- or below-ground biomass production from 2005-2010.
Thus, the main objectives of this thesis were (1) to investigate if extreme weather events have an effect on ecosystem functions beyond productivity, (2) to test if such a high resistance or resilience in response to drought regarding productivity also exists in more naturally grown plant communities and (3) to further elucidate possible mechanisms of the surprisingly large stability of the plant communities.
To investigate these objectives, several experimental studies were conducted in artificially planted, as well as in naturally grown grassland communities and consequences of extreme weather events for ecosystem processes, such as decomposition and herbivory were investigated. In a pot experiment, it was studied, if grass plants react improved towards repeated drought when compared to a first drought and thus reveal a kind of drought memory. Such a memory might be one possible, but up until now widely neglected mechanism of resilience.
Even though biomass production remained stable in our experiment in artificially planted communities, biomass quality was severely affected by extreme drought, thereby strongly affecting the development of a herbivore caterpillar feeding on drought-exposed leaves. Further, plant compounds of the host plant depended on the composition of the plant community it was grown in. This in turn resulted in strong effects on the larval mortality of herbivores feeding on such plants.
In contrast to the study in artificially planted communities, aboveground net primary productivity (ANPP) was reduced in naturally composed grassland in response to extreme rainfall variability, including an extreme drought followed by heavy rainfall. Forage quality was altered by drought. Furthermore, mowing frequency strongly altered forage quality and biomass production, but did not interact with rainfall variability and thus did neither buffer, nor amplify effects of extreme rainfall variability. Despite effects of rainfall variability on ANPP, grassland showed high resilience after drought followed by heavy rain, as effects were large shortly after the extreme event, but did not persist until a second harvest later in the year.
In natural grassland, rainfall variability and drought also affected ecosystem processes, here litter decomposition, beyond productivity. Drought followed by heavy rain pulses decreased decomposition rates. Decomposition in more frequently mown meadows was more vulnerable towards drought exposure. Winter warming and additional winter rain had no long-term effect on decomposition. To conclude, projected increases in drought frequency under climate change may inhibit decomposition and alter nutrient and carbon cycling along with soil quality in temperate grassland, whereas a reduction of snow cover leading to more variable soil surface temperatures may counteract increased decomposition under winter warming.
In this thesis, an ecological stress memory as one possible mechanism of resilience is defined as any response of a single plant after a stress experience that improves the reaction of the plant towards future stress experience and which is assessed on a whole plant level. This thesis further provides evidence of a drought memory in grass plants: Plants repeatedly subjected to drought showed improved photo-protection and a higher rate of living biomass when compared to plants faced with their first drought. Similarly, tree seedlings exposed to drought in summer revealed higher frost resistance during winter, providing evidence of a long-lasting “cross-stress-memory” .
To sum up, the thesis shows that extreme weather events, even though neither severely affecting biomass production in artificially composed, nor in naturally growing communities in the long-term, exert strong influence on physiological or biogeochemical parameters, such as plant compounds or soil biotic activity. These changes in turn modify ecosystem functions beyond productivity, for example herbivory or decomposition, possibly altering biotic interactions and nutrient cycling. Furthermore, the findings imply that plants exhibit a stress memory after stress exposure, which may be one mechanisms leading to a high stability and resilience upon frequent stress.
Climate change impacts on habitats and biodiversity :
From environmental envelope modelling to nature conservation strategies
- Climate change will pose entirely new challenges for nature conservation. A literature study of 852 publications (between 2003 and 2010) illuminates this topic, examines driving research forces as well as focal points and shows recent research gaps. Here could be shown that changes in species distribution, diverse consequences for habitats, changing communities as well as biotic interactions and general aspects of diversity are the major challenges. The potential climatic modifications can alter deeply the distribution of animals and plants. Range changes due to recent climate change already exist and are traceable. In order to quantify such changes, environmental envelope modelling can be used. In addition to individual species, changes in distribution of more complex units are also conceivable. The present work mainly focuses on habitat types listed in the Annex I of the European Habitats Directive. To reveal the potential range changes of habitat types, two principally different modelling approaches have been developed. An indirect approach modelling the distribution of a habitat type using the distribution of its characteristic plant species and a direct approach, using the distribution of the habitat type itself. These two approaches were tested by modelling five grassland habitat types. Looking at the modelled results all habitats are projected to lose between 22% and 93% of their range in the ‘no dispersal’ scenario. Both approaches produce reasonable results. However, modelling an extensive set of habitat types using the indirect approach is currently not possible, because of the required but actually lacking amount of plant distribution data. Therefore, the direct approach is an appropriate instrument for modelling habitat types. Here, all 127 widespread terrestrial habitat types defined in the Annex I of the Habitats Directive were modelled and, resulting from this, a map of terrestrial habitat type diversity was calculated. Several habitat types are projected to lose many of their actually suitable areas, in particular bogs, rocky habitats, grassland and in part forests. Due to their developmental time or rather due to their special abiotic requirements, bogs and rocky habitats even lose under the assumption of a full dispersal scenario. However, most heath and grassland habitats are also projected to lose in the full dispersal scenario. Pooling all modelled results together, terrestrial habitat type diversity is shifting partly to mountain regions and the atlantic biogeographical region is projected to decrease in habitat type diversity. According to the Habitats Directive habitat types listed in Annex I are protected in ‘sites of community interest’ aiming to maintain or restore them at a favourable conservation status. Due to the projected changes a static nature conservation concept could face problems which particularly concern the coherence of the protected area network. This could lead to a loss of protective goods in spite of protected areas. To illustrate the potential problems and difficulties emerging with respect to spatial coherence of habitat types between protected areas, an analysis of spatial coherence under future conditions for a variety of habitat types in Germany was conducted. Here, a combination of environmental envelope modelling and graph theory is presented to assess the coherence of nature conservation networks. The possible incapacity of species to reach all climatically suitable areas is currently debated. Therefore, spatial scales are not only crucial for the coherence of protected areas but also for the question if future projected suitable areas could be colonized. Dispersal movements of species are only infrequently possible in our highly fragmented landscape. To answer this raising question, Odonata listed in the Habitats Directive with known dispersal distances were contemplated. The species Coenagrion ornatum, C. mercuriale and Ophiogomphus cecilia are projected to lose range when incorporating specific dispersal distances, while they are projected to extend their range in the unrestricted dispersal scenario. Furthermore, suitable climatic conditions tend to decline for Leucorrhinia albifrons and L. caudalis, whereas L. pectoralis is projected to gain distribution area assuming either species-specific or unrestricted dispersal. The nature conservation measure of translocation is an at least 100 years old methodology with pros and cons. In this thesis, the emerging problems and opportunities of this species preservation strategy are presented. Further, a new question about the ‘focal unit’ is pointed out as well as the problem of genetic variability and the aspect of pre-adopted subspecies. Moreover, a selective assisted colonisation not by moving species but ecotypes is referred.
Applying regional climate change projections for spatio-temporal risk analyses of vector-borne diseases
- Bei vorliegender Dissertation handelt es sich um eine Abhandlung zu vektor-assoziierten Krankheiten in Zeiten des Klimawandels. Bei vektor-assoziierten Krankheiten wird ein Pathogen durch einen Vektor (Überträger), auf ein Wirtstier übertragen. Als solche Vektoren agieren meist Arthropoden. Klimatische Veränderungen beeinflussen vektor-assoziierte Krankheiten insbesondere dadurch, dass Arthropoden ihre Körpertemperatur nicht selbst regeln können und zudem bestimmte Temperaturansprüche zur Pathogenentwicklung im Vektor erfüllt sein müssen. Das Klimaänderungssignal des 21. Jahrhunderts wird von Klimamodellen in verschiedenen räumlichen und zeitlichen Auflösungen wiedergegeben. Die Projektionen beruhen auf Emissionsszenarien klimawirksamer Treibhausgase. In der Arbeit werden die Einsatzmöglichkeiten von regionalen Klimamodellen zur Gefährdungsabschätzung anhand verschiedener Fallbeispiele aufgezeigt. Deren Nutzen und Einsatzmöglichkeiten werden einführend aufgeführt. Für die Risikoanalysen werden regionalen Klimamodelle REMO und COSMO-CLM angewandt, die durch dynamisches „Downscaling“ globaler Modelle generiert wurden. Beide sind in ihrem neuesten Prozesslauf in das globale Modell ECHAM5 eingebettet. Der direkte Übertrag bekannter Temperaturansprüche von Vektor und/oder Pathogen auf künftig zu erwartende Bedingungen stellt den ersten methodologischen Schwerpunkt dieser Arbeit dar. Eine Amplifikation des Dengue-Virus im Überträger der Stechmücke Aedes aegypti könnte demnach zunächst in Südeuropa, im weiteren Verlauf des 21. Jhd. aber auch in weiteren europäischen Regionen möglich sein. Weiterhin verdeutlichen die Ergebnisse, dass sich auch das Zeitfenster einer potentiellen Übertragung des Dengue-Virus verlängern kann. Durch das Überlagern der bekannten Temperaturansprüchen von Sandmücken (Gattung Phlebotomus) und der von ihnen übertragbaren Erreger - Leishmania infantum Komplex - können potentielle Regionen Deutschlands identifiziert werden, in denen einer autochthone Übertragung der Leishmaniose möglich ist. Es ist zu erwarten, dass ein solches Risiko zunächst in südwestlichen und westlichen Regionen Deutschlands, im späteren Verlaufe des des 21. Jhd. jedoch auch für eher nördlich und östlich gelegene Regionen bestehen wird. Der zweite innerhalb dieser Arbeit gewählte methodologische Ansatz zeigt die Einsatzmöglichkeiten regionaler Klimaprojektion für die bioklimatische Nischenmodellierung von Krankheitsüberträgern auf. Die anhand statistischer Verfahren ermittelte bioklimatische Nische der jeweiligen Art wird hierbei auf zukünftig zu erwartende klimatische Bedingungen übertragen. Anhand dieser Analyse kann aufgezeigt werden, dass sich die klimatische Eignung für die invasive Stechmücke Aedes albopictus (Überträger mehrere human-pathogener Viren) ausgehend von westlichen Regionen Europas über Mitteleuropa und schließlich Osteuropas erhöhen wird. Der Transfer der ermittelten spezifischen klimatischen Nische ausgewählter Sandmücken-Arten (u.a. Überträger der zum Leishmania-Komplex zählenenden Pathogenen) auf künftige Bedingungen lässt vermuten, dass deren klimatische Eignung in Mitteleuropa - abgesehen von alpinen Regionen - zunehmen wird. Künftige potenzielle Ausbreitungswege der Sandmücken in einer sich verändernden Umwelt, werden via “least-cost analysis“ ermittelt. Die Ergebnisse deuten darauf hin, dass aufgrund der eingeschränkten natürlichen Ausbreitungsfähigkeit, einige der künftig potenziell geeigneten Lebensräume nicht erreicht werden. In den verschiedenen Fallstudien kann gezeigt werden, dass die zu erwartenden klimatischen Veränderungen im 21. Jhd. eine mögliche Ausbreitung der in dieser Arbeit adressierten Vektoren und vektor-assoziierter Krankheiten in Europa begünstigen werden. Als einheitliche Tendenz kann speziell für Mitteleuropa festgehalten werden, dass sich die Gefährdung, Ende des 21.Jhd. erhöhen wird. Dies begründet sich höchstwahrscheinlich durch die projizierte raschere Erwärmung in der zweiten Jahrhunderthälfte. Abschließend bleibt jedoch festzuhalten, dass es neben klimatischen Veränderungen weitere Faktoren für die Ausbreitung bzw. Neuetablierung von Vektoren und den damit verbundenen übertragbaren Infektionskrankheiten ausschlaggebend sind. Der Einfluss einzelner Faktoren auf die Etablierung bzw. Ausbreitung vektor-assoziierte Krankheiten variiert auf raum-zeitlichen Skalen. Für die ermittelten klimatisch-abgeleiteten Risikogebiete sollten in Folgestudien auf kleineren Skalen wirksam werdenden Faktoren integriert werden. Diese Ergebnisse können wiederum die Entwicklung von Surveillance- und Monitoringprogramme unterstützen, um somit Maßnahmen gegen die Ausbreitung von vektor-assoziierten Krankheiten initiieren zu können.
Turnover and fluxes of carbon and nitrogen in a spruce forest under natural and extreme meteorological conditions
- Climate models predict an increase in the intensity and frequency of extreme meteorological climate events like extended summer droughts, heavy rainfall or intensive frost periods with largely unknown effects on microbial activity and pysico-chemical soil properties and their impact on availability of soil organic matter. The influence of drying/rewetting (A/W) and freezing/thawing (G/A) events on solution chemistry and leaching losses of soils is barely known. This thesis aimed to study the effects of A/W and G/A events on soil solution chemistry and solute fluxes, in particular, of dissolved organic carbon (DOC) and inorganic nitrogen (NH4+, NO3-) in a podzol soil under a Norway spruce forest. A field experiment was designed to study the effects of (i) summer drought by exclusion of natural throughfall and subsequent rewetting and of (ii) soil frost by removal of natural snow cover. In complementary laboratory experiments with undisturbed soil columns, (i) drying/rewetting cycles were simulated with different rewetting intensities and (ii) freezing/thawing cycles were induced using different freezing temperatures. In the second part of this work, total C and N stocks as well as radiocarbon signatures of soil organic carbon (SOC) from different soil horizons and density fractions were investigated. A/W increased the DOC concentrations in the organic layer and upper mineral soil. More DOC was released from the organic layer to the mineral soil. However, the effects on total DOC leaching were smaller due to reduced water fluxes. Specific UV absorbance and emission fluorescence detected a switch in the release of easily decomposable DOC to hardly decomposable DOC during the wetting phase. Prolonged summer drought and incomplete rewetting due to hydrophobicity of SOM in the organic layer and upper mineral horizon reduced net N mineralisation as well as concentrations and fluxes of the NH4+ and NO3-. The net nitrification rate in the organic layer was more negatively influenced than net ammonification, indicating that nitrifiers are more sensitive to drought stress than ammonifiers. The effect of soil frost strongly depended on soil freezing temperature. Only soil frost at temperature below -8°C led to short periods of additional DOC production in the organic layer. Spectroscopic properties and ∆14C signatures of DOC implied a disruption of soil aggregates and desorption of older DOC from the mineral associated organic matter fraction of the Oa horizons by G/A events. Severe soil frost below -8°C inhibited the activity of nitrifiers and ammonifiers with decreased NH4+ and NO3- concentrations and fluxes in the mesocosm experiment. A delayed (by 4 months) increase in NO3- concentration in the upper soil horizon by moderate soil frost (-5°C) was attributed to reduced Immobilisation by heterotrophic microorganisms. Summarised, drying and the effect of hydrophobicity led to long-term, severe soil frost to short-term reduction in N mineralisation and N leaching. The effect of increased NO3- concentrations as delayed response to G/A needs further research in case of potentially changes in the N balance. Drying as well as freezing induced changes in the soil structure and properties and led to increased DOC concentrations. Moderate soil temperature had much less effects on C and N in this temperate forest soil. The results of this thesis demonstrated the potential of extreme meteorological events on the quality and availability of dissolved C and N. Both, A/W and G/A cycles decreased C and N mineralisation, increased the sink strength of the soil by the accumulation of SOC and N, considering constant C and N litter input. However, optimal temperature and moisture conditions in other seasons could compensate the sink strength of soils. This work underpins the need for holistic and long-term investigations to understand and model the impact of extreme meteorological conditions on the dynamics of dissolved C and N.
Soil organic matter dynamics in a temperate forest influenced by extreme weather events
- Climate models predict an increase in surface temperature and a change in intensity and kind of precipitation in the future for Europe depending on the region with effects on C cycling and soil organic matter (SOM). We investigated the influence of extreme weather events (frost/drought) on the quality and quantity of SOM in a Haplic Podzol under a 140 years old Norway spruce forest in the Fichtelgebirge mountains (Bavaria, German) within two laboratory and two field studies. In one laboratory study, we investigated the effect of frost intensity and repeated freeze/thaw cycles. Undisturbed soil columns comprising organic layer and top mineral soil were treated as followed: Control (+5 °C), frost at –3 °C, –8 °C and –13 °C. After a two-week freezing period, frozen soils were thawed at +5 °C and irrigated with 80 mm water at a rate of 4 mm per day. After the third cycle, SOM pools of the treatments were compared with those of non-dried control columns. Under field conditions from late December 2005 until middle of February 2006 we removed the natural snow cover during winter on three replicate plots. Hence we induced soil frost to 15 cm depth (in a depth of 5 cm below surface up to -5°C) from January to April 2006, while the snow-covered control plots never reached temperatures below 0 °C. In the second laboratory experiment after air-drying for five weeks, undisturbed soil columns were re-wetted at different intensities (8, 20 and 50 mm per day) and time intervals, so that all treatments received the same amount of water per cycle (100 mm). After the third cycle, SOM pools of the treatments were compared with those of non-dried control columns. Under field conditions, a throughfall exclusion (TE) experiment was conducted in the summers 2006 and 2007 using a roof installation followed by re-wetting compared to non-manipulated control plots. On 18th January 2007, the heavy low pressure system Kyrill caused large damages at our control plots whereas the TE sites were less influenced. Therefore, for this study, only data were used from the control plots before Kyrill and from the soil structure undisturbed TE plots. SOM quantity and quality was followed by biomarker analysis: lignin, neutral sugars and phospholipid fatty acids (PLFA) as measure for microbial biomass. Amounts of lignin contents were not significantly affected by repeated freeze/thaw cycles. However, intensive frost slightly enhanced lignin mobilization in the O layer and the translocation into the B horizon. While soil frost did not influence lignin concentrations, the decomposition rate of vanillyl monomers (Ac/Ad)v decreased at the end of the frost period, these results confirm reduced mineralisation under frost. In contrast, lignin phenols were not systematically affected by the drying/rewetting-experiment and the moisture regime. The sum of PLFA (soil microbial biomass) was not affected by the frost respectively drying event, suggesting that most soil microorganisms were well adapted or recovered more quickly than the accumulation of microbial residues such as microbial sugars directly after the experiment. However, PLFA patterns indicate that fungi are more susceptible to soil frost than bacteria. The ratio of fungi to bacteria were generally not altered through drying, however, at least in the L horizon, warmer and drier weather led to a dominance of fungi while a cooler and moister regime favoured bacteria. Increasing water stress was indicated by a higher PLFA (cy17:0+cy19:0)/ (16:1w7c+18:1w7c) ratio suggesting that the microbes suffered from water stress in the organic layer and uppermost mineral soil. While soil microbial biomass was not affected by the moisture regime, the structure of soil microbial community changed. Gram-positive bacteria and actinomycetes were reduced whereas gram-negative bacteria, fungi and protozoa were stimulated by the reduced moisture regime. In the subsequent summer after the freezing experience, soil microbial biomass was significantly higher at the snow-removal plots (SM) compared to the control despite lower CO2 respiration and increasing water stress indicator. These results suggest that soil microbial respiration and therefore the activity was not closely related to soil microbial biomass but more strongly controlled by substrate availability and quality. Both freezing/thawing and drying/re-wetting reduced the amount of microbial sugars due to reduced mineralisation. However, also the hydrolysable plant sugars decreased in all soil horizons. We postulated that the only possible explanation for the disappearance of plant and microbial sugars upon soil freezing or drying are chemical alterations of sugar molecules leading to SOM stabilization, also known as SOM aging. Further studies are required to quantify the effect of temperature or moisture regime to the observed changes in soil sugar concentrations.
Plant Species and Functional Diversity along Altitudinal Gradients, Southwest Ethiopian Highlands
Desalegn Wana Dalacho
- Understanding how biodiversity is organized across space and time has long been a central focus of ecologists and biogeographers. Altitudinal patterns of richness gradients are one of such striking patterns in the landscape. Despite its historical and ecological importance as a heuristic natural experimental site for development of ecological theories, the emergent patterns and mechanisms that structure them are poorly understood. This is partly because of the complex relationships of species to the environment and the choice of the response variable itself, i.e. using taxonomic richness as a metrics of diversity. This thesis, therefore, applies plant functional types (hereafter PFTs) approach to study the response of vegetation to environmental factors in the southwest Ethiopian highlands. It focuses on the classification of the vegetation into a few main plant functional response categories and relate them to environmental variables. For pattern identification and mechanistic explanations, a deconstructive approach of the taxonomic richness into its constituent components was used. Furthermore, the potential effects of land use/land cover change and global warming on the biodiversity of the study area was investigated. The results reveal that the application of plant functional types is a promising tool to understand vegetation-environment relationships. Local topographic attributes (altitude and slope) and soil properties found to structure the variance in the relative abundance of PFTs along environmental gradients. Moreover, specific response to drought favours the abundance of species with thorns/spines and tussocks in the lowlands as opposed to chilling which favours rosettes and rhizomes PFTs in the highlands. Concerning patterns of richness along altitudinal gradients, various structures of richness appear for total vascular plant species and growth forms. Woody plants, graminoids and climbers showed a uni-modal structure while ferns and herbs revealed an increasing pattern of richness along the altitudinal gradient. By contrast, total vascular plants species richness did not show any strong response to altitudinal gradients. Climate related water-energy dynamics, species area relationships due to the physical shape of the mountain, local topographic and soil conditions were found to be predominant factors structuring the observed richness in the study area. The threats to biodiversity loss due to land use/land cover change and global warming is eminent in the study area. Land conversion for agricultural purposes was a pervasive process that had a deleterious effect on the biodiversity of the study area. Population growth, socio-economic challenges (poverty) and government policy regimes drive land cover change processes. In addition, recent climate change poses a serious challenge to the biodiversity of the study area. The results of model predictions indicated that biodiversity of the study area will suffer severe consequences of lowland biotic attrition (i.e. the net loss of species richness in the tropical lowlands caused by altitudinal range shifts in the absence of new species arriving), range gap shifts and contraction, and extinction due to expected warming at the end of this century. The model also predicted that endangered and endemic species with restricted elevational ranges will disproportionately suffer from range contraction and extinction due to warming. In conclusion, the plant functional types approach was found to be an essential tool to reduce complexity of the vegetation of the study system and to elucidate vegetation-environment relationships. Moreover, the identification of emergent patterns and attributing them to mechanistic explanations are pre-requisites for conservation planning to save biodiversity of the study area. The study also evidenced that land use/land cover change and global warming will present strong threats to the loss of biodiversity in the study area. Salvaging biodiversity in the future requires the consideration of the effect of land use and climate change on vegetation responses. Consequently, nature conservation strategies and future reserve designs should take into account options of human assisted migration across fragmented landscapes and creating dispersal routes for species to track to new thermal niches.
Dynamics and underlying processes of N2O and NO soil-atmosphere exchange under extreme meteorological boundary conditions
- Climate models predict an increasing frequency and intensity of summer drought periods with subsequent heavy rainfall or soil frost and thaw events in mountain regions of Central Europe. These indirect effects of global warming may considerably influence soil microbial processes and in consequence emissions of climate-relevant trace gases. Regarding the nitrogen cycle, N2O and NO emissions are of concern, since they are involved in climate warming and soils represent a main source for these two gases. In spite of a growing number of studies on this subject, knowledge on effects of climate change on soil N2O and NO emissions is still scarce. This is mainly due to a hitherto poor mechanistic understanding of underlying processes within soil. In this thesis, the impact of extreme meteorological boundary conditions on N2O and NO fluxes in a Norway spruce forest and an acidic fen in the Fichtelgebirge area was investigated. The summer drought period and precipitation were experimentally increased in the forest and the fen over a 2-year span. Soil frost was induced in the forest by removal of the natural snow cover. The experiments were run in three replicates each and non-manipulated plots served as controls. Throughout the experiments, N2O and NO fluxes were recorded in weekly to monthly intervals. In addition, N2O concentrations and isotope signatures in soil air were measured along soil profiles to identify and localise the underlying biogenic production and consumption processes. Prolonged drought continuously reduced the N2O emission from the forest soil and even turned the soil temporarily into a sink for atmospheric N2O. Soil freezing and thawing caused a burst of N2O release contributing 84 % of the annual emission. Soil air N2O concentration and stable isotope profiles provide a new mechanistic explanation tool for all of these findings. N2O concentration in the soil air decreased in most cases exponentially from the subsoil to the soil surface. This observation identifies microbial activity in the subsoil (at >= 70 cm soil depth) as an additional source for N2O and diffusion to the soil surface along a concentration gradient. Furthermore, isotope abundance analysis identified simultaneous microbial N2O consumption (reduction to N2). Drought reduced the source strength of the organic layers for N2O while simultaneously the sink function of the mineral soil for N2O remained active. This resulted in a net N2O sink function of the forest soil under severe drought. Frost in the topsoil was the only exception for these trends in N2O concentration and isotope signature along soil profiles. Under conditions of soil frost the topsoil served no longer as a sink for N2O, thus leading to the observed burst in N2O emission. NO emissions from the forest soil exceeded the N2O emissions by up to two orders of magnitude. Prolonged drought in- or decreased NO emissions depending on the soil moisture content of the organic layers. Wetting after long-lasting drought periods – which turned out to be of less importance regarding N2O fluxes – strongly increased biogenic NO emissions and contributed 44 % to the annual loss. In contrast to the forest soil, NO fluxes from the fen were always one to two orders of magnitude lower than the N2O fluxes. These results support earlier findings that this highly reactive gas is either only marginally produced in the fen soil or undergoes chemical conversion before escaping from the soil surface. Nevertheless, water table reduction resulted in significantly increased net NO emission. Regarding N2O, this thesis suggests that summer drought periods may drastically increase emissions from minerotrophic fens depending on the reduction of water table height. Furthermore, heavy rainfall following drought periods caused short lived, but strong N2O peaks having significant impact on the annual N2O loss, that have not been reported so far. N-15 and O-18 isotope data provide evidence that these N2O peaks are due to newly produced N2O in the upper soil. This thesis documents the huge impact of extreme weather events on soil N2O and NO emissions and provides so far scarcely considered mechanistic explanations for these observations. A major outcome of this work is the finding of a hitherto unconsidered sink function of forest soils for atmospheric N2O, when soil net N2O production is compensated for by net consumption during long-lasting droughts. This work underlines the importance of investigating the fate of N2O within soil profiles next to flux measurements to improve the current knowledge on the complex interactions between meteorological boundary conditions and soil biogenic processes and thus help further upgrading global N2O balances.
Dynamics of soil processes under extreme meteorological boundary conditions - response of below-ground carbon, sulfur, and iron cycling in fen soils
- Northern peatlands store approximately 30 % of the global soil carbon stocks. On the other hand, peatlands contribute about 3-10 % to the global methane burden into the atmosphere. Climate predictions foresee not only an increase in the global mean temperature, but also a change in precipitation patterns. As peatlands critically depend on hydrological conditions, a change in precipitation distribution is likely to affect the carbon sink and source function of peatlands. Thus, these ecosystems have become the focus of an increasing number of studies over the past decades. Low water table levels, high temperatures, and a higher nutrient availability were mostly found to increase respiratory activity, but to reduce methane production and –emission. Existing studies, however, investigated changes in average environmental conditions in the long term, while the impact of extreme weather on peatland elemental cycles is still uncertain. Moreover, most studies do not provide a mechanistic understanding of the redox processes underlying the response of peatlands to fluctuations of the water table level. Based on laboratory studies, a thermodynamically constrained competition of the different terminal electron accepting processes for common electron donors was postulated. A detailed validation of this concept under natural or near-natural conditions is, however, still lacking to date. Furthermore, the processes that renew alternative electron accepting capacity during drought are still not yet understood. Fens were also identified to be notable sources or sinks for arsenic. The close association of arsenic with the iron- and sulphur-dynamics – and thus redox dynamics during fluctuations of the water table level in general – is already known. Nevertheless, there exist hardly any study investigating arsenic dynamics and solid phase associations for fens. The main objective of this work was therefore to study the effects of more pronounced drying and rewetting events on redox processes of carbon, iron, and sulphur – and concomitantly arsenic – in an electron acceptor rich fen-ecosystem. In contrast to some existing studies, we could not find a notable effect of the drying/wetting treatment on the overall carbon budgets of the peat. There was an obvious effect of drying/wetting on respiration within the soil, increasing drastically during drought, but the net carbon budget was by far dominated by the autotrophic activity of the vegetation (55-65 %) which was hardly affected by the treatment. Due to the drought event, methanogenesis was effectively suppressed in the unsaturated part of the profile and re-established after rewetting only after a notable time lag of some weeks. This suppression of methanogenic activity – in the laboratory and in the field approach – could successfully be explained by a reoxidation of reduced iron and sulphur compounds, providing alternative electron accepting capacity during and after drought. Only after depletion of alternative electron acceptors, methanogenic conditions could re-establish. Locally, however, in micro-environments especially in the uppermost, intensively rooted layers, methanogenesis re-established even before alternative electron acceptors had been depleted. Based on the obtained data, we propose the high availability of easily degradable organic material, a still high water content, and poor aeration of the peat to responsible for this observation. These factors could support a local depletion of alternative electron acceptors and methanogenesis could thus occur in locally distinct micro-environments. The analysis of the isotopic composition of the dissolved CO2 and the methane produced suggested that the methane was formed via the CO2-reduction pathway with H2 as the electron donor. This pattern was not affected by the drying/wetting treatment. Exceptionally high isotope fractionation factors suggested thermodynamic conditions to be quite unfavourable for methanogens. This coincided with the observation that most of the peat was likely structured in small micro-environments of locally distinct redox conditions and the rapid switches between methanogenic and methanotrophic conditions. The arsenic dynamics under variable redox conditions generally followed the dynamics of ferrous iron, especially in the intensively rooted uppermost soil layers. Coincidingly, a major part of the arsenic was found in the reactive iron-hydroxide fraction, readily available for microbial reduction. Although the total arsenic content in the solid phase was comparably low in the fen under study, concentrations of arsenic exceeded common drinking water standards mostly by far. Methylated arsenic species did not play a noteworthy role in this fen and the immobilization of arsenic in sulfidic phases during reducing conditions was also negligible when compared to mobilization from iron-hydroxide reduction.
Coordinated Tree Responses to Drought -Vulnerability and Sustainable Production: Hypotheses on Arid Ecosystem Adjustments to Limitations in Water Resources
- Field and controlled greenhouse experiments were carried out to investigate tree responses to declining soil water content. Field experiments were conducted on naturally growing trees of Acacia tortilis and A. xanthophloea in the savanna region of Kenya and Quercus suber in the Mediterranean region of Portugal. The selected field sites were regions that experience regular drought periods during the year. Greenhouse experiments constituted two watering regimes. Seedlings of A. tortilis and A. xanthophloea grown from seeds initially obtained from the Kenya field site were raised and arranged on a greenhouse bench into two groups per species. The first set of plants were watered every other day (controls) while the second set were watered every seven days (water stress treatments). Field measurements included weather parameters, soil and plant water status, growth, sap flux density, leaf transpiration and stomatal conductance, tissue water relations and isotope labeling. Similar measurements were conducted on plants growing in the greenhouse. Also examined in the greenhouse were root biomass, root structure as well as whole plant biomass accumulation. A second set of experiments was carried out in the greenhouse by subjecting plants initially stressed and non-stressed to severe water stress by withholding water until plants were wilted overnight. The wilted plants were then re-watered regularly and their recovery after stress alleviation was monitored. Declining soil water content significantly affected plant water status in all the trees studied. Lowest psi pd recorded during the study period occurred in the month of June and were –2.0 and –1.1 MPa for A. xanthophloea and A. tortilis respectively. The same species subjected to repeated water stress in the greenhouse attained mean minimum psi pd of –2.4 and –1.2 MPa for A. xanthophloea and A. tortilis respectively at the end of the drying cycle. Mean minimum psi pd recorded for Q. suber during summer was –1.8 MPa and occurred in September. There were however, significant differences among trees. Decline in psi associated with increasing soil drought led to decline in leaf initiation and leaf expansion and both processes ceased at higher water stress levels. For the Acacia species, even leaf shedding occurred at higher stress levels. There was also a decline in stomatal conductance (gs) during water stress, leading to decrease in transpiration rates (E). Maximum stomatal conductance of 340 mmol m-2 s-1 were observed during rainy seasons for the Acacia trees while mean maximum values of 300 mmol m-2 s-1 were recorded for Q. suber when soil moisture conditions were favorable. Stomatal conductance declined by 31%, 67% and 67% in A. tortilis and A. xanthophloea in the savanna and Q. suber in the Mediterranean regions respectively. Daily tree water use (Qtree) as well as leaf transpiration reflected changes in psi and gs. Root to leaf hydraulic conductance equally declined with increasing soil drought. Q. suber trees adjusted osmotically by a magnitude of 0.7 MPa, while bulk modulus of elasticity (epsilon) increased by a magnitude of 17 MPa. Osmotic adjustment of 0.48 MPa was observed in greenhouse plants of A. tortilis while epsilon declined by a magnitude of 7 MPa in A. xanthophloea. A. tortilis plants in the greenhouse showed increased absolute root growth, root depth and root:shoot (r:s) ratio. The dimorphic rooting pattern in Q. suber resulted into hydraulic lift and this could as well occur in A. tortilis because of similarity in their rooting patterns. Most plant responses were reactionary and were aimed at enhancing soil water uptake and reducing transpiration water loss when soil water content was declining. Similar responses were observed for both greenhouse and naturally growing field plants of the same species. Decline in leaf initiation and leaf expansion as well as leaf senescence reduced tree crown size hence potential tree transpiration. This however, had negative impact on plant productivity. Increased root growth as well as osmotic adjustment increased tree water uptake from the soil. The balance between root water uptake and leaf transpiration through growth and stomatal regulation was aimed at protecting xylem integrity. The overall results showed that soil characteristics, root activities and root distribution patterns are the main factors determining tree functioning and productivity in drylands, while the coordinated interaction between the aboveground shoot and belowground root activities ensures survival during drought. Maintained production and survival will ensure distribution and success in the arid environments. Repeated water stress imparted water stress resistance qualities on seedlings enabling them to survive longer during severe stress. The study emphasizes the role of soil resource base as well as species interactions in the functioning and balance of dryland ecosystems.