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In der vorliegenden Dissertation wird die Bedeutung von Brachen für Artenvielfalt und Stabilität von Blüte-Bestäuber-Nahrungsnetzen in agrarisch genutzten Landschaften anhand ausgewählter blütenbesuchender Insektengruppen (Syrphidae, Lepidoptera) untersucht. Die Freilandarbeiten fanden von 1998-2000 im Raum der Feldberger Seenlandschaft, Mecklenburg-Vorpommern, statt. Es werden die beiden Hauptnahrungsquellen Nektar und Pollen betrachtet, dabei fanden Untersuchungen zur Intensität der Blüte-Bestäuber-Interaktion auf Stilllegungsflächen, zum flächenbezogenen quantitativen Nektarangebot im Jahresverlauf, zur individuellen Pollennutzung bei Syrphiden und zur Breite und Überlappung der Nahrungsnischen bei den dominanten Arten Episyrphus balteatus, Metasyrphus corollae, Syritta pipiens und Sphaerophoria scripta statt. Im Ergebnis zeigt sich eine hohe Bedeutung der Brachflächen für die Stabilität des Blüte-Bestäuber-Netzes, während die Diversität von anderen, eher landschaftsbezogenen Faktoren abhängig ist.
Spatial and temporal temperature and moisture patterns across the Tibetan Plateau are very complex. The onset and magnitude of the Holocene climate optimum in the Asian monsoon realm, in particular, is a subject of considerable debate as this time period is often used as an analogue for recent global warming. In the light of contradictory inferences regarding past climate and environmental change on the Tibetan Plateau, I have attempted to explain mismatches in the timing and magnitude of change. Therefore, I analysed the temporal variation of fossil pollen and diatom spectra and the geochemical record from palaeo-ecological records covering different time scales (late Quaternary and the last 200 years) from two core regions in the NE and SE Tibetan Plateau. For interpretation purposes I combined my data with other available palaeo-ecological data to set up corresponding aquatic and terrestrial proxy data sets of two lake pairs and two sets of sites. I focused on the direct comparison of proxies representing lacustrine response to climate signals (e.g., diatoms, ostracods, geochemical record) and proxies representing changes in the terrestrial environment (i.e., terrestrial pollen), in order to asses whether the lake and its catchments respond at similar times and magnitudes to environmental changes. Therefore, I introduced the established numerical technique procrustes rotation as a new approach in palaeoecology to quantitatively compare raw data of any two sedimentary records of interest in order to assess their degree of concordance. Focusing on the late Quaternary, sediment cores from two lakes (Kuhai Lake 35.3°N; 99.2°E; 4150 m asl; and Koucha Lake 34.0°N; 97.2°E; 4540 m asl) on the semi-arid northeastern Tibetan Plateau were analysed to identify post-glacial vegetation and environmental changes, and to investigate the responses of lake ecosystems to such changes. Based on the pollen record, five major vegetation and climate changes could be identified: (1) A shift from alpine desert to alpine steppe indicates a change from cold, dry conditions to warmer and more moist conditions at 14.8 cal. ka BP, (2) alpine steppe with tundra elements points to conditions of higher effective moisture and a stepwise warming climate at 13.6 cal. ka BP, (3) the appearance of high-alpine meadow vegetation indicates a further change towards increased moisture, but with colder temperatures, at 7.0 cal. ka BP, (4) the reoccurrence of alpine steppe with desert elements suggests a return to a significantly colder and drier phase at 6.3 cal. ka BP, and (5) the establishment of alpine steppe-meadow vegetation indicates a change back to relatively moist conditions at 2.2 cal. ka BP. To place the reconstructed climate inferences from the NE Tibetan Plateau into the context of Holocene moisture evolution across the Tibetan Plateau, I applied a five-scale moisture index and average link clustering to all available continuous pollen and non-pollen palaeoclimate records from the Tibetan Plateau, in an attempt to detect coherent regional and temporal patterns of moisture evolution on the Plateau. However, no common temporal or spatial pattern of moisture evolution during the Holocene could be detected, which can be assigned to the complex responses of different proxies to environmental changes in an already very heterogeneous mountain landscape, where minor differences in elevation can result in marked variations in microenvironments. Focusing on the past 200 years, I analysed the sedimentary records (LC6 Lake 29.5°N, 94.3°E, 4132 m asl; and Wuxu Lake 29.9°N, 101.1°E, 3705 m asl) from the southeastern Tibetan Plateau. I found that despite presumed significant temperature increases over that period, pollen and diatom records from the SE Tibetan Plateau reveal only very subtle changes throughout their profiles. The compositional species turnover investigated over the last 200 years appears relatively low in comparison to the species reorganisations during the Holocene. The results indicate that climatically induced ecological thresholds are not yet crossed, but that human activity has an increasing influence, particularly on the terrestrial ecosystem. Forest clearances and reforestation have not caused forest decline in our study area, but a conversion of natural forests to semi-natural secondary forests. The results from the numerical proxy comparison of the two sets of two pairs of Tibetan lakes indicate that the use of different proxies and the work with palaeo-ecological records from different lake types can cause deviant stories of inferred change. Irrespective of the timescale (Holocene or last 200 years) or region (SE or NE Tibetan Plateau) analysed, the agreement in terms of the direction, timing, and magnitude of change between the corresponding terrestrial data sets is generally better than the match between the corresponding lacustrine data sets, suggesting that lacustrine proxies may partly be influenced by in-lake or local catchment processes whereas the terrestrial proxy reflects a more regional climatic signal. The current disaccord on coherent temporal and spatial climate patterns on the Tibetan Plateau can partly be ascribed to the complexity of proxy response and lake systems on the Tibetan Plateau. Therefore, a multi-proxy, multi-site approach is important in order to gain a reliable climate interpretation for the complex mountain landscape of the Tibetan Plateau.
Numerous pollen records across the upper Tibetan Plateau indicate that in the early part of the mid-Holocene, Kobresia-rich high-alpine meadows invaded areas formerly dominated by alpine steppe vegetation rich in Artemisia. We examine climate, land-use, and CO2 concentration changes as potential drivers for this marked vegetation change. The climatic implications of these vegetational shifts are explored by applying a newly developed pollen-based moisture-balance transfer-function to fossil pollen spectra from Koucha Lake on the north-eastern Tibetan Plateau (34.0 degrees N; 97.2 degrees E; 4540 m a.s.l.) and Xuguo Lake on the central Tibetan Plateau (31.97 degrees N; 90.3 degrees E; 4595 m a.s.l.), both located in the meadow-steppe transition zone. Reconstructed moisture-balances were markedly reduced (by similar to 150-180 mm) during the early mid-Holocene compared to the late-Holocene. These findings contradict most other records from the Indian monsoonal realm and also most non-pollen records from the Tibetan Plateau that indicate a rather wet early- and mid-Holocene. The extent and timing of anthropogenic land-use involving grazing by large herbivores on the upper Tibetan Plateau and its possible impacts on high-alpine vegetation are still mostly unknown due to the lack of relevant archaeological evidence. Arguments against a mainly anthropogenic origin of Kobresia high-alpine meadows are the discovery of the widespread expansion of obviously 'natural' Kobresia meadows on the south-eastern Tibetan Plateau during the Lateglacial period indicating the natural origin of this vegetation type and the lack of any concurrence between modern human-driven vegetation shifts and the mid-Holocene compositional changes. Vegetation types are known to respond to atmospheric CO2 concentration changes, at least on glacial-interglacial scales. This assumption is confirmed by our sensitivity study where we model Tibetan vegetation at different CO2 concentrations of 375 (present-day), 260 (early Holocene), and 650 ppm (future scenario) using the BIOME4 global vegetation model. Previous experimental studies confirm that vegetation growing on dry and high sites is particularly sensitive to CO2 changes. Here we propose that the replacement of drought-resistant alpine steppes (that are well adapted to low CO2 concentrations) by mesic Kobresia meadows can, at least, be partly interpreted as a response to the increase of CO2 concentration since 7000 years ago due to fertilization and water-saving effects. Our hypothesis is corroborated by former CO2 fertilization experiments performed on various dry grasslands and by the strong recent expansion of high-alpine meadows documented by remote sensing studies in response to recent CO2 increases.
The past climate in central Asia, and especially on the Tibetan Plateau (TP), is of great importance for an understanding of global climate processes and for predicting the future climate. As a major influence on the climate in this region, the Asian Summer Monsoon (ASM) and its evolutionary history are of vital importance for accurate predictions. However, neither the evolutionary pattern of the summer monsoon nor the driving mechanisms behind it are yet clearly understood. For this research, I first synthesized previously published Late Glacial to Holocene climatic records from monsoonal central Asia in order to extract the general climate signals and the associated summer monsoon intensities. New climate and vegetation sequences were then established using improved quantitative methods, focusing on fossil pollen records recovered from Tibetan lakes and also incorporating new modern datasets. The pollen-vegetation and vegetation-climate relationships on the TP were also evaluated in order to achieve a better understanding of fossil pollen records. The synthesis of previously published moisture-related palaeoclimate records in monsoonal central Asia revealed generally different temporal patterns for the two monsoonal subsystems, i.e. the Indian Summer Monsoon (ISM) and East Asian Summer Monsoon (EASM). The ISM appears to have experienced maximum wet conditions during the early Holocene, while many records from the area affected by the EASM indicate relatively dry conditions at that time, particularly in north-central China where the maximum moisture levels occurred during the middle Holocene. A detailed consideration of possible driving factors affecting the summer monsoon, including summer solar insolation and sea surface temperatures, revealed that the ISM was primarily driven by variations in northern hemisphere solar insolation, and that the EASM may have been constrained by the ISM resulting in asynchronous patterns of evolution for these two subsystems. This hypothesis is further supported by modern monsoon indices estimated using the NCEP/NCAR Reanalysis data from the last 50 years, which indicate a significant negative correlation between the two summer monsoon subsystems. By analogy with the early Holocene, intensification of the ISM during coming decades could lead to increased aridification elsewhere as a result of the asynchronous nature of the monsoon subsystems, as can already be observed in the meteorological data from the last 15 years. A quantitative climate reconstruction using fossil pollen records was achieved through analysis of sediment core recovered from Lake Donggi Cona (in the north-eastern part of the TP) which has been dated back to the Last Glacial Maximum (LGM). A new data-set of modern pollen collected from large lakes in arid to semi-arid regions of central Asia is also presented herein. The concept of "pollen source area" was introduced to modern climate calibration based on pollen from large lakes, and was applied to the fossil pollen sequence from Lake Donggi Cona. Extremely dry conditions were found to have dominated the LGM, and a subsequent gradually increasing trend in moisture during the Late Glacial period was terminated by an abrupt reversion to a dry phase that lasted for about 1000 years and coincided with the first Heinrich Event of the northern Atlantic region. Subsequent periods corresponding to the warm Bølling-Allerød period and the Younger Dryas cold event were followed by moist conditions during the early Holocene, with annual precipitation of up to about 400 mm. A slightly drier trend after 9 cal ka BP was then followed by a second wet phase during the middle Holocene that lasted until 4.5 cal ka BP. Relatively steady conditions with only slight fluctuations then dominated the late Holocene, resulting in the present climatic conditions. In order to investigate the relationship between vegetation and climate, temporal variations in the possible driving factors for vegetation change on the northern TP were examined using a high resolution late Holocene pollen record from Lake Kusai. Moving-window Redundancy Analyses (RDAs) were used to evaluate the correlations between pollen assemblages and individual sedimentary proxies. These analyses have revealed frequent fluctuations in the relative abundances of alpine steppe and alpine desert components, and in particular a decrease in the total vegetation cover at around 1500 cal a BP. The climate was found to have had an important influence on vegetation changes when conditions were relatively wet and stable. However, after the 1500 cal a BP threshold in vegetation cover was crossed the vegetation appears to have been affected more by extreme events such as dust storms or fluvial erosion than by the general climatic trends. In addition, pollen spectra over the last 600 years have been revealed by Procrustes analysis to be significantly different from those recovered from older samples, which is attributed to an increased human impact that resulted in unprecedented changes to the composition of the vegetation. Theoretical models that have been developed and widely applied to the European area (i.e. the Extended R-Value (ERV) model and the Regional Estimates of Vegetation Abundance from Large Sites (REVEALS) model) have been applied to the high alpine TP ecosystems in order to investigate the pollen-vegetation relationships, as well as for quantitative reconstructions of vegetation abundance. The modern pollen–vegetation relationships for four common pollen species on the TP have been investigated using Poaceae as the reference taxa. The ERV Submodel 2 yielded relatively high PPEs for the steppe and desert taxa (Artemisia Chenopodiaceae), and low PPEs for the Cyperaceae that are characteristic of the alpine Kobresia meadows. The plant abundances on the central and north-eastern TP were quantified by applying these PPEs to four post-Late Glacial fossil pollen sequences. The reconstructed vegetation assemblages for the four pollen sequences always yielded smaller compositional species turnovers than suggested by the pollen spectra, indicating that the strength of the previously-reported vegetation changes may therefore have been overestimated. In summary, the key findings of this thesis are that (a) the two ASM subsystems show asynchronous patterns during both the Holocene and modern time periods, (b) fossil pollen records from large lakes reflect regional signals for which the pollen source areas need to be taken into account, (c) climate is not always the main driver for vegetation change, and (d) previously reported vegetation changes on the TP may have been overestimated because they ignored inter-species variations in pollen productivity.
Temporally changing drivers for late-Holocene vegetation changes on the northern Tibetan Plateau
(2012)
Fossil pollen records have been widely used as indicators of past changes in vegetation and variations in climate. The driving mechanisms behind these vegetation changes have, however, remained unclear. In order to evaluate vegetation changes that have occurred in the northern part of the Tibetan Plateau and the possible drivers behind these changes, we have applied a moving-window Redundancy Analysis (RDA) to high resolution (10-15 years) pollen and sedimentary data from Lake Kusai covering the last 3770 years. Our analyses reveal frequent fluctuations in the relative abundances of alpine steppe and alpine desert components. The sedimentary proxies (including total organic carbon content, total inorganic carbon content, and "end-member" indices from grain-size analyses) that explain statistically some of the changes in the pollen assemblage vary significantly with time, most probably reflecting multiple underlying driving processes. Climate appears to have had an important influence on vegetation changes when conditions were relatively wet and stable. However, a gradual decrease in vegetation cover was identified after 1500 cal a BP, after which the vegetation appears to have been affected more by extreme events such as dust-storms or fluvial erosion than by general climatic trends. Furthermore, pollen spectra over the last 600 years are shown by Procrustes analysis to be statistically different from those recovered from older samples, which we attribute to increased human impact that resulted in unprecedented changes to the vegetation composition. Overall, changes in vegetation and climate on the northern part of the Tibetan Plateau appear to have roughly followed the evolution of the Asian Summer Monsoon. After taking into account the highly significant millennial (1512 years) periodicity revealed by time-series analysis, the regional vegetation and climate changes also show variations that appear to match variations in the mid-latitude westerlies.
Beringian climate and environmental history are poorly characterized at its easternmost edge. Lake sediments from the northern Yukon Territory have recorded sedimentation, vegetation, summer temperature and precipitation changes since similar to 16 cal ka BP. Herb-dominated tundra persisted until similar to 14.7 cal ka BP with mean July air temperatures <= 5 degrees C colder and annual precipitation 50 to 120 mm lower than today. Temperatures rapidly increased during the Bolling/Allerod interstadial towards modern conditions, favoring establishment of Betula-Salix shrub tundra. Pollen-inferred temperature reconstructions recorded a pronounced Younger Dryas stadial in east Beringia with a temperature drop of similar to 1.5 degrees C (similar to 2.5 to 3.0 degrees C below modern conditions) and low net precipitation (90 to 170 mm) but show little evidence of an early Holocene thermal maximum in the pollen record. Sustained low net precipitation and increased evaporation during early Holocene warming suggest a moisture-limited spread of vegetation and an obscured summer temperature maximum. Northern Yukon Holocene moisture availability increased in response to a retreating Laurentide Ice Sheet, postglacial sea level rise, and decreasing summer insolation that in turn led to establishment of Alnus-Berula shrub tundra from similar to 5 cal ka BP until present, and conversion of a continental climate into a coastal-maritime climate near the Beaufort Sea.
Phytoplankton development affects the community structure and dynamics of freshwater bacteria by changing the availability of nutrients, algal exudates and biological surfaces. To elucidate these effects of phytoplankton development in spring in oligotrophic Lake Stechlin (Germany), we measured limnological and biological parameters, including the bacterial community composition (BCC), at the depth of the highest chlorophyll a concentration. To increase the resolution of BCC measurements, we separated particle-associated (PA) and free-living (FL) bacteria using serial filtration through 5.0 and 0.2 mu m pore size filters, respectively. The BCC of ultramicrobacteria was also determined by collecting the 0.2 mu m filtrate on 0.1 mu m filters. Changes in the community composition of Bacteria and particularly of Actinobacteria, one of the most important bacterial groups in temperate freshwater habitats, were studied via DGGE analysis of PCR-amplified 16S rRNA gene fragments. Patterns in BCC dynamics of FL Bacteria and Actinobacteria remained fairly constant throughout the study period, while patterns of PA Bacteria were more variable over time. At the breakdown of the diatom spring bloom, bacterial production and abundance sharply increased, indicating a close coupling between heterotrophic bacteria and algal detritus. The succession in BCC revealed life-style dependent patterns related to specific environmental variables. Our results indicate independent dynamics of PA and FL Bacteria as well as Actinobacteria during succession of phytoplankton spring blooms. These differences in bacterial lifestyle can only be resolved when the PA and FL fractions of microorganisms are separated.
A high resolution multi proxy (pollen, grain size, total organic carbon) record from a small mountain lake (Lake Khuisiin; 46.6 degrees N, 101.8 degrees E; 2270 m a.s.l.) in the south eastern Khangai Mountains of central Mongolia has been used to explore changes in vegetation and climate over the last 1200 years. The pollen data indicates that the vegetation changed from dry steppe dominated by Poaceae and Artemisia (ca AD 760-950), to Larix forest steppe (ca AD 950-1170), Larix Betula forest steppe (ca AD 1170-1380), meadow dominated by Cyperaceae and Poaceae (ca AD 1380-1830), and Larix Betula forest steppe (after similar to AD 1830). The cold-wet period between AD 1380 and 1830 may relate to the Little Ice Age. Environmental changes were generally subtle and climate change seems to have been the major driver of variations in vegetation until at least the early part of the 20th century, suggesting that either the level of human activity was generally low, or the relationship between human activity and vegetation did not alter substantially between AD 760 and 1830. A review of centennial scale moisture records from China and Mongolia revealed that most areas experienced major changes at ca AD 1500 and AD 1900. However, the moisture availability since AD 1500 varied between sites, with no clear regional pattern or relationship to present day conditions. Both the reconstructions and the moisture levels simulation on a millennium scale performed in the MPI Earth System Model indicate that the monsoon-westerlies transition area shows a greater climate variability than those areas influenced by the westerlies, or by the summer monsoon only.
The Tibetan Plateau is a region that is highly sensitive to recent global warming, but the complexity and heterogeneity of its mountainous landscape can result in variable responses. In addition, the scarcity and brevity of regional instrumental and palaeoecological records still hamper our understanding of past and present patterns of environmental change. To investigate how the remote, high-alpine environments of the Nianbaoyeze Mountains, eastern Tibetan Plateau, are affected by climate change and human activity over the last similar to 600 years, we compared regional tree-ring studies with pollen and diatom remains archived in the dated sediments of Dongerwuka Lake (33.22A degrees N, 101.12A degrees E, 4,307 m a.s.l.). In agreement with previous studies from the eastern Tibetan Plateau, a strong coherence between our two juniper-based tree-ring chronologies from the Nianbaoyeze and the Anemaqin Mountains was observed, with pronounced cyclical variations in summer temperature reconstructions. A positive directional trend to warmer summer temperatures in the most recent decades, was, however, not observed in the tree-ring record. Likewise, our pollen and diatom spectra showed minimal change over the investigated time period. Although modest, the most notable change in the diatom relative abundances was a subtle decrease in the dominant planktonic Cyclotella ocellata and a concurrent increase in small, benthic fragilarioid taxa in the similar to 1820s, suggesting higher ecosystem variability. The pollen record subtly indicates three periods of increased cattle grazing activity (similar to 1400-1480 AD, similar to 1630-1760 AD, after 1850 AD), but shows generally no significant vegetation changes during past similar to 600 years. The minimal changes observed in the tree-ring, diatom and pollen records are consistent with the presence of localised cooling centres that are evident in instrumental and tree-ring data within the southeastern and eastern Tibetan Plateau. Given the minor changes in regional temperature records, our complacent palaeoecological profiles suggest that climatically induced ecological thresholds have not yet been crossed in the Nianbaoyeze Mountains region.
Honey bees display a fascinating division of labour among foragers. While some bees solely collect pollen, others only collect nectar. It is assumed that individual differences in sensory response thresholds are at the basis of this division of labour. Biogenic amines and their receptors are important candidates for regulating the division of labour, because they can modulate sensory response thresholds. Here, we investigated the role of the honey bee tyramine receptor AmTYR1 in regulating the division of foraging labour. We report differential splicing of the Amtyr1 gene and show differential gene expression of one isoform in the suboesophageal ganglion of pollen and nectar foragers. This ganglion mediates gustatory inputs. These findings imply a role for the honey bee tyramine receptor in regulating the division of foraging labour, possibly through the suboesophageal ganglion.