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Mankind has dramatically influenced the nitrogen (N) fluxes between soil, vegetation, water and atmosphere " the global N cycle. Increasing intensification of agricultural land use, caused by the growing demand for agricultural products, has had major impacts on ecosystems worldwide. Particularly nitrogenous gases such as ammonia (NH3) have increased mainly due to industrial livestock farming. Countries with high N deposition rates require a variety of deposition measurements and effective N monitoring networks to assess N loads. Due to high costs, current "conventional"-deposition measurement stations are not widespread and therefore provide only a patchy picture of the real extent of the prevailing N deposition status over large areas. One tool that allows quantification of the exposure and the effects of atmospheric N impacts on an ecosystem is the use of bioindicators. Due to their specific physiology and ecology, especially lichens and mosses are suitable to reflect the atmospheric N input at ecosystem level. The present doctoral project began by investigating the general ability of epiphytic lichens to qualify and quantify N deposition by analysing both lichens and total N and δ15N along a gradient of different N emission sources and severity. The results showed that this was a viable monitoring method, and a grid-based monitoring system with nitrophytic lichens was set up in the western part of Germany. Finally, a critical appraisal of three different monitoring techniques (lichens, mosses and tree bark) was carried out to compare them with national relevant N deposition assessment programmes. In total 1057 lichen samples, 348 tree bark samples, 153 moss samples and 24 deposition water samples, were analysed in this dissertation at different investigation scales in Germany.The study identified species-specific ability and tolerance of various epiphytic lichens to accumulate N. Samples of tree bark were also collected and N accumulation ability was detected in connection with the increased intensity of agriculture, and according to the presence of reduced N compounds (NHx) in the atmosphere. Nitrophytic lichens (Xanthoria parietina, Physcia spp.) have the strongest correlations with high agriculture-related N deposition. In addition, the main N sources were revealed with the help of δ15N values along a gradient of altitude and areas affected by different types of land use (NH3 density classes, livestock units and various deposition types). Furthermore, in the first nationwide survey of Germany to compare lichens, mosses and tree bark samples as biomonitors for N deposition, it was revealed that lichens are clearly the most meaningful monitor organisms in highly N affected regions. Additionally, the study shows that dealing with different biomonitors is a difficult task due to their variety of N responses. The specific receptor surfaces of the indicators and therefore their different strategies of N uptake are responsible for the tissue N concentration of each organism group. It was also shown that the δ15N values depend on their N origin and the specific N transformations in each organism system, so that a direct comparison between atmosphere and ecosystems is not possible.In conclusion, biomonitors, and especially epiphytic lichens may serve as possible alternatives to get a spatially representative picture of the N deposition conditions. Furthermore, bioindication with lichens is a cost-efficient alternative to physico-chemical measurements to comprehensively assess different prevailing N doses and sources of N pools on a regional scale. They can at least support on-site deposition instruments by qualification and quantification of N deposition.
Time series archives of remotely sensed data offer many possibilities to observe and analyse dynamic environmental processes at the Earth- surface. Based on these hypertemporal archives, which offer continuous observations of vegetation indices, typically at repetition rates from one to two weeks, sets of phenological parameters or metrics can be derived. Examples of such parameters are the beginning and end of the annual growing period, as well as its length. Even though these parameters do not correspond exactly to conventional observations of phenological events, they nevertheless provide indications of the dynamic processes occurring in the biosphere. The development of robust algorithms for the derivation of phenological metrics can be challenging. Currently, such algorithms are most commonly based on digital filters or the Fourier analysis of time series. Polynomial spline models offer a useful alternative to existing methods. The possibilities of using spline models in the analytical description of time series are numerous, and their specific mathematical properties may help to avoid known problems occurring with the more common methods for deriving phenological metrics. Based on a selection of different polynomial spline models suitable for the analysis of remotely sensed time series of vegetation indices, a method to derive various phenological parameters from such time series was developed and implemented in this work. Using an example data set from an intensively used agricultural area showing highly dynamic variations in vegetation phenology, the newly developed method was verified by a comparison of the results of the spline based approach to the results of two alternative, well established methods.
Two areas were selected to represent major process regimes of Mediterranean rangelands. In the County of Lagads (Greece), situated east of the city of Thessaloniki, livestock grazing with sheep and goats is a major factor of the rural economy. In suitable areas, it is complemented by agricultural use. The region of Ayora (Spain) is located west of the city of Valencia. It is one of regions most affected by fires in Spain. First of all, long time series of satellite data were compiled for both regions on the basis of Landsat sensors, which cover the time until 1976 (Ayora) and 1984 (Lagadas) with one image per year. Using a rigorous processing scheme, the data were geometrically and radiometrically corrected Specific attention was given to an exact sensor calibration, the radiometric intercalibration of Landsat-TM and "MSS. Proportional cover of photosynthetically active vegetation was identified as a suitable quantitative indicator for assessing the state of rangelands. Using Spectral Mixture Analysis (SMA) it was inferred for all data sets. The extensive data base procured this way enabled to map fire events in the Ayora area based on sequential diachronic sets and provide fire dates, perimeter as well as fire recurrence for each pixel. The increasing fire frequency in the past decades is in large parts attributed to the accelerated abandonment of the area that leads to an encroachment of shrublands and the accumulation of combustible biomass. On the basis of the fire mapping results, a spatial and temporal stratification of the data set allowed to asses plant recovery dynamics on the landscape level through linear trend analysis. The long history of fire events in the Mediterranean frequently leads to processes of auto-succession. Following an initial dominance of herbaceous vegetation this commonly leads to similar plant communities as the ones present before the fire. On a temporal axis, this results in typical exponential post-fire trajectories which could also be shown in this study. The analysis of driving factors for post-fire dynamics confirmed the importance of aspect and slope. Locations with lower amounts of solar irradiation and favourable water supply yielded faster recovery rates and higher post-fire vegetation cover levels. In most cases, the vegetation cover levels observed before the fire were not reached within the post-fire observation period. In the area of Lagadas, linear trend analysis and additional statistical parameters were used to infer a degradation index. This could be used to illustrate a complex pattern of stability, regeneration and degradation of vegetation cover. These different processes and states are found in close proximity and are clearly determined by topography and elevation. Following a sequence of analyses, it was found that in particular steep, narrow valleys show positive trends, while negative trends are more abundant on plain or gently undulating areas. Considering the local grazing regime, this spatial differentiation was related to the accessibility of specific locations. Subsequently, animal numbers on community level were used to calculate efficient stocking rates and assess the temporal development of their relation with vegetation cover. This calculation of temporal trajectories illustrated that only some communities show the expected negative relation. To the contrary, a positive relation or even changing relation patterns are observed. This signifies recent concentration and intensification processes in the grazing scheme, as a result of which animals are kept in sheds, where additional feedstuffs are provided. In these cases, free roaming of livestock animals is often confined to some hours every day, which explains the spatial preference of easily accessible areas by the shepherds. Beyond these temporal trends, it was analysed whether the grazing pattern is equally reflected in a spatial trend. Making use of available geospatial information layers, the efforts required to reach each location was expressed as a cost. Then, cost zones could be defined and woody vegetation cover as a grazing indicator could be inferred for the different zones. Animal sheds were employed as starting features for this piospheric analysis, which could be mapped from very high spatial resolution Quickbird image data. The result was a clearly structured gradient showing increasing woody vegetation cover with increasing cost distance. On the basis of these two pilot studies, the elements of a monitoring and interpretation framework identified at the beginning of the work were evaluated and a formal interpretation scheme was presented.
Water-deficit stress, usually shortened to water- or drought stress, is one of the most critical abiotic stressors limiting plant growth, crop yield and quality concerning food production. Today, agriculture consumes about 80-90% of the global freshwater used by humans and about two thirds are used for crop irrigation. An increasing world population and a predicted rise of 1.0-2.5-°C in the annual mean global temperature as a result of climate change will further increase the demand of water in agriculture. Therefore, one of the most challenging tasks of our generation is to reduce the amount water used per unit yield to satisfy the second UN Sustainable Development Goal and to ensure global food security. Precision agriculture offers new farming methods with the goal to improve the efficiency of crop production by a sustainable use of resources. Plant responses to water stress are complex and co-occur with other environmental stresses under natural conditions. In general, water stress causes plant physiological and biochemical changes that depend on the severity and the duration of the actual plant water deficit. Stomatal closure is one of the first responses to plant water stress causing a decrease in plant transpiration and thus an increase in plant temperature. Prolonged or severe water stress leads to irreversible damage to the photosynthetic machinery and is associated with decreasing chlorophyll content and leaf structural changes (e.g., leaf rolling). Since a crop can already be irreversibly damaged by only mild water deficit, a pre-visual detection of water stress symptoms is essential to avoid yield loss. Remote sensing offers a non-destructive and spatio-temporal method for measuring numerous physiological, biochemical and structural crop characteristics at different scales and thus is one of the key technologies used in precision agriculture. With respect to the detection of plant responses to water stress, the current state-of-the-art hyperspectral remote sensing imaging techniques are based on measurements of thermal infrared emission (TIR; 8-14 -µm), visible, near- and shortwave infrared reflectance (VNIR/SWIR; 0.4-2.5 -µm), and sun-induced fluorescence (SIF; 0.69 and 0.76 -µm). It is, however, still unclear how sensitive these techniques are with respect to water stress detection. Therefore, the overall aim of this dissertation was to provide a comparative assessment of remotely sensed measures from the TIR, SIF, and VNIR/SWIR domains for their ability to detect plant responses to water stress at ground- and airborne level. The main findings of this thesis are: (i) temperature-based indices (e.g., CWSI) were most sensitive for the detection of plant water stress in comparison to reflectance-based VNIR/SWIR indices (e.g., PRI) and SIF at both, ground- and airborne level, (ii) for the first time, spectral emissivity as measured by the new hyperspectral TIR instrument could be used to detect plant water stress at ground level. Based on these findings it can be stated that hyperspectral TIR remote sensing offers great potential for the detection of plant responses to water stress at ground- and airborne level based on both TIR key variables, surface temperature and spectral emissivity. However, the large-scale application of water stress detection based on hyperspectral TIR measures in precision agriculture will be challenged by several problems: (i) missing thresholds of temperature-based indices (e.g., CWSI) for the application in irrigation scheduling, (ii) lack of current TIR satellite missions with suitable spectral and spatial resolution, (iii) lack of appropriate data processing schemes (including atmosphere correction and temperature emissivity separation) for hyperspectral TIR remote sensing at airborne- and satellite level.
A sustainable development of forests and their ecosystem services requires the monitoring of the forests" state and changes as well as the prediction of their future development. To achieve the latter, eco-physiological forest growth models are usually applied. These models require calibration and validation with forestry reference data. This data includes forest structural parameters such as tree height or stem diameter which are easy to measure and can be used to estimate the core model parameters, i.e. the tree- biomass pools. The methods traditionally applied to derive the structural parameters are mainly manual and time-consuming. Hence, the in situ data acquisition is inefficient and limited in its ability to capture the vertical and horizontal variability in stand structure. Ground-based remote sensing bears the potential to overcome the limitations of the traditional methods. As they can be automated, ground-based remote sensing methods allow a much more efficient data acquisition and a larger spatial coverage. They are also able to capture forest structure in its three dimensions. Nevertheless, at present further research is required, in particular with respect to the practical integration of ground-based remote sensing data into forest growth models as well as regarding factors influencing the structural parameter retrieval from this data. Therefore, the goal of this PhD thesis was to investigate the influencing factors of two ground-based remote sensing methods (terrestrial laser scanning and hemispherical photography), which have not or only scarcely been studied to date. In addition, the use of forest structural parameters derived from these methods for the calibration of a forest growth model was assessed. Both goals were achieved. The results of this thesis could contribute significantly to a comprehensive assessment of ground-based remote sensing and its potential to derive the forest structural parameters. However, the use of these methods to calibrate forest growth models proved to be limited. An optimized data sampling design is expected to eliminate the major limitations, though. Furthermore, the combination of ground-based, airborne, and satellite remote sensing sensors was suggested to provide an optimized framework for the general integration of remotely sensed data into forest growth models. This combination of remote sensing observations at different scales will contribute greatly to a modern forest management with the purpose of warranting a sustainable forest development even under growing economic and ecological pressures.
Dry tropical forests are facing massive conversion and degradation processes and they are the most endangered forest type worldwide. One of the largest dry forest types are Miombo forests that stretch across the Southern African subcontinent and the proportionally largest part of this type can be found in Angola. The study site of this thesis is located in south-central Angola. The country still suffers from the consequences of the 27 years of civil war (1975-2002) that provides a unique socio-economic setting. The natural characteristics are a representative cross section which proved ideal to study underlying drivers as well as current and retrospective land use change dynamics. The major land change dynamic of the study area is the conversion of Miombo forests to cultivation areas as well as modification of forest areas, i.e. degradation, due to the extraction of natural resources. With future predictions of population growth, climate change and large scale investments, land pressure is expected to further increase. To fully understand the impacts of these dynamics, both, conversion and modification of forest areas were assessed. By using the conceptual framework of ecosystem services, the predominant trade-off between food and timber in the study area was analyzed, including retrospective dynamics and impacts. This approach accounts for products that contribute directly or indirectly to human well-being. For this purpose, data from the Landsat archive since 1989 until 2013 was applied in different study area adapted approaches. The objectives of these approaches were (I) to detect underlying drivers and their temporal and spatial extent of impact, (II) to describe modification and conversion processes that reach from times of armed conflicts over the ceasefire and the post-war period and (III) to provide an assessment of drivers and impacts in a comparative setting. It could be shown that major underlying drivers for the conversion processes are resettlement dynamics as well as the location and quality of streets and settlements. Furthermore, forests that are selectively used for resource extraction have a higher chance of being converted to a field. Drivers of forest degradation are on one hand also strongly connected to settlement and infrastructural structures. But also to a large extent to fire dynamics that occur mostly in more remote and presumably undisturbed forest areas. The loss of woody biomass as well as its slow recovery after the abandonment of fields could be quantified and stands in large contrast to the amount of potentially cultivated food that is necessarily needed. The results of the thesis support the fundamental understanding of drivers and impacts in the study area and can thus contribute to a sustainable resource management.
Until today the effects of many chlorinated hydrocarbons (e.g. DDT, PCBs) against the specific organisms are still a subject of controversial discussions. It was also the case for potential endocrine effects to influence the spermatogenesis correlated with possible changes of the population's vitality. To clear this situation, three questions could be at the centre of attention: 1) Do the chemicals cause a special harmful effect on the male reproductive tract? 2) Could some particular chemical mixtures act to bind and activate the human estrogen receptor (hER)? 3) Are the life stages of an organism specially sensitive to the effects of chemicals and therefore be established as Screening-Test-System? the connected effects of DDT and Arochlor 1254 as single substance and in 1:1 mixture according to their estrogenic effectiveness on zebrafish (Brachydanio rerio) were therefore investigated. the concentrations of the pesticides and their mixture ranged between 0.05-µg/l and 500-µg/l and separated by a factor of 10. It was turned out that the test concentrations of 500-µg/l were too toxic to zebrafish in all the cases. The experiment was followed up with four concentrations of DDT, A54 as well as their 1:1 mixture anew each separated by a factor of 10 and ranging between 0.05-µg/l and 50-µg/l. The bioaccumulation test within 8 days showed that the zebrafish accumulated the chemicals, but no equilibrum was reached and the concentration 0.05-µg/l was established as No Observed Effect Concentration (NOEC). Putting up on these analyses, the investigation of the life cycle (LC) starting with fertilized eggs demonstrated a reduction in the rate of hatchability, reproduction and length of fish emerged. These reductions involved the duration of the life cycle stages (LCS) which consequently lasted longer than expected. Exposure time and level of the tested chemicals accelerated the occurrence of these effects which were more significant when the chemical mixtures were used too. To establish whether the parameter assessed were correlated to the male reproductive tract, the quality, quantity and life span of sperm were assessed using the methods of Leong (1988) and Shapiro et al (1994). The sperm degeneration observed, led us to investigate the spermatogenesis and the ultrastructure of the testes. This last experiment showed a significant reduction of the late stage of spermatogenesis and the heterophagic vacuoles which play an important role in the spermatid maturation. It could therefore be concluded that, DDT and A54 could act synergically and cause disorders of the male reproductive tract of male zebrafish and influence also their growth.
This study investigates the endemic centres of Indonesian animals and the biodiversity across geographical gradients. At the same time, it also evaluated different lines suggested for separating the Oriental and Australian faunal region in the Indonesian region. The analyses have mainly used the present-day distribution of terrestrial vertebrates, especially the smallest ranges of species and subspecies. The results show that faunal migration of Oriental and Australian lineages to the Indonesian Archipelago may have been happening since the Palaeocene period and more importantly, island drifts might have facilitated such migration. These events caused major reorganisation of island positions and island forms, which in turn resulted in faunal extinction around the mid-Pliocene. Some islands, especially in the Wallacea region, emerged very late and as a result nowadays they are lacking endemic forms. There are currently at least seven endemic centres, which can be recognised, i.e. Borneo, Java, Sumatra, Sulawesi, North Moluccas, New Guinea and the Lesser Sundas/Banda Arcs. The affinities between these endemic centres revealed that there are two clusters of islands in the Indonesian Archipelago. These different clusters suggest in turn the shifts of biogeographical lines in the Indonesian Archipelago. Furthermore, oscillation in climate, eustatic sea level changes and fluctuations in vegetation in the Quaternary period had much affected the distribution pattern of animals. There was a phase of expansion for montane oak forests, grasslands and woodlands during the period 18,000-14,000 years ago in East Indonesia and 16,500-12,000 years ago in West Indonesia. Such an expansion led to the increased isolation of rainforests and of the faunas adapted to them. These periods are also indicated by the lowering of the tree line which facilitated montane fauna to disperse across lower elevations. At 8,000-9,000 years ago, the climate became warmer and slightly wetter. The mid- to upper montane forests expanded to their full altitudinal range, while montane oak forest, grassland, and woodland areas had contracted. The oscillation in climate, eustatic sea level changes and fluctuations in vegetation in turn determines much the formation of numerous sub endemic centres, which today can be found within the mainland. Recently, there are 14 sub endemic centres on Borneo, 8 on Java, 16 on Sumatra, 14 on Sulawesi and 14 on New Guinea. From the conservation management point of view, the identification of such sub endemic centres would generate valuable information for the protection effort.
Besides well-known positive aspects of conservation tillage combined with mulching, a drawback may be the survival of phytopathogenic fungi like Fusarium species on plant residues. This may endanger the health of the following crop by increasing the infection risk for specific plant diseases. In infected plant organs, these pathogens are able to produce mycotoxins like deoxynivalenol (DON). Mycotoxins like DON persist during storage, are heat resistant and of major concern for human and animal health after consumption of contaminated food and feed, respectively. Among fungivorous soil organisms, there are representatives of the soil fauna which are obviously antagonistic to a Fusarium infection and the contamination with mycotoxins. Earthworms (Lumbricus terrestris), collembolans (Folsomia candida) and nematodes (Aphelenchoides saprophilus) provide a wide range of ecosystem services including the stimulation of decomposition processes which may result in the regulation of plant pathogens and the degradation of environmental contaminants. Several investigations under laboratory conditions and in the field were conducted to test the following hypotheses: (1) Fusarium-infected and DON-contaminated wheat straw provides a more attractive food substrate than non-infected control straw (2) the introduced soil fauna reduce the biomass of F. culmorum and the content of DON in infected wheat straw under laboratory and field conditions (3) the species interaction of the introduced soil fauna enhances the degradation of Fusarium biomass and DON concentration in wheat straw; (4) the degradation efficiency of soil fauna is affected by soil texture. The results of the present thesis pointed out that the degradation performance of the introduced soil fauna must be considered as an important contribution to the biological control of plant diseases and environmental pollutants. As in particular L. terrestris revealed to be the driver of the degradation process, earthworms contribute to a sustainable control of fungal pathogens like Fusarium and its mycotoxins in wheat straw, thus reducing the risk of plant diseases and environmental pollution as ecosystem services.
N-acetylation by N-acetyltransferase 1 (NAT1) is an important biotransformation pathway of the human skin and it is involved in the deactivation of the arylamine and well-known contact allergen para-phenylenediamine (PPD). Here, NAT1 expression and activity were analyzed in antigen presenting cells (monocyte-derived dendritic cells, MoDCs, a model for epidermal Langerhans cells) and human keratinocytes. The latter were used to study exogenous and endogenous NAT1 activity modulations. Within this thesis, MoDCs were found to express metabolically active NAT1. Activities were between 23.4 and 26.6 nmol/mg/min and thus comparable to peripheral blood mononuclear cells. These data suggest that epidermal Langerhans cells contribute to the cutaneous N-acetylation capacity. Keratinocytes, which are known for their efficient N-acetylation, were analyzed in a comparative study using primary keratinocytes (NHEK) and different shipments of the immortalized keratinocyte cell line HaCaT, in order to investigate the ability of the cell line to model epidermal biotransformation. N-acetylation of the substrate para-aminobenzoic acid (PABA) was 3.4-fold higher in HaCaT compared to NHEK and varied between the HaCaT shipments (range 12.0"44.5 nmol/mg/min). Since B[a]P induced cytochrome p450 1 (CYP1) activities were also higher in HaCaT compared to NHEK, the cell line can be considered as an in vitro tool to qualitatively model epidermal metabolism, regarding NAT1 and CYP1. The HaCaT shipment with the highest NAT1 activity showed only minimal reduction of cell viability after treatment with PPD and was subsequently used to study interactions between NAT1 and PPD in keratinocytes. Treatment with PPD induced expression of cyclooxygenases (COX) in HaCaT, but in parallel, PPD N-acetylation was found to saturate with increasing PPD concentration. This saturation explains the presence of the PPD induced COX induction despite the high N-acetylation capacities. A detailed analysis of the effect of PPD on NAT1 revealed that the saturation of PPD N-acetylation was caused by a PPD-induced decrease of NAT1 activity. This inhibition was found in HaCaT as well as in primary keratinocytes after treatment with PPD and PABA. Regarding the mechanism, reduced NAT1 protein level and unaffected NAT1 mRNA expression after PPD treatment adduced clear evidences for substrate-dependent NAT1 downregulation. These results expand the existing knowledge about substrate-dependent NAT1 downregulation to human epithelial skin cells and demonstrate that NAT1 activity in keratinocytes can be modulated by exogenous factors. Further analysis of HaCaT cells from different shipments revealed an accelerated progression through the cell cycle in HaCaT cells with high NAT1 activities. These findings suggest an association between NAT1 and proliferation in keratinocytes as it has been proposed earlier for tumor cells. In conclusion, N-acetylation capacity of MoDCs as well as keratinocytes contribute to the overall N-acetylation capacity of human skin. NAT1 activity of keratinocytes and consequently the detoxification capacities of human skin can be modulated by the presence of exogenous NAT1 substrates and endogenous by the cell proliferation status of keratinocytes.
Chemical communication in the reproductive behaviour of Neotropical poison frogs (Dendrobatidae)
(2013)
Chemical communication is the evolutionary oldest communication system in the animal kingdom that triggers intra- and interspecific interactions. It is initiated by the emitter releasing either a signal or a cue that causes a reaction of the receiving individual. Compared to other animals there are relatively few studies regarding chemical communication in anurans. In this thesis the impact of chemical communication on the behaviour of the poison frog Ranitomeya variabilis (Dendrobatidae) and its parental care performance was investigated. This species uses phytotelmata (small water bodies in plants) for both clutch and tadpole depositions. Since tadpoles are cannibalistic, adult frogs do not only avoid conspecifics when depositing their eggs but also transport their tadpoles individually into separated phytotelmata. The recognition of already occupied phytotelmata was shown to be due to chemical substances released by the conspecific tadpoles. In order to gain a deeper comprehension about the ability of adult R. variabilis to generally recognize and avoid tadpoles, in-situ pool choice experiments were conducted, offering chemical substances of tadpole of different species to the frogs (Chapter I). It turned out that they were able to recognize all species and avoid their chemical substances for clutch depositions. However, for tadpole depositions only dendrobatid tadpoles occurring in phytotelmata were avoided, while those species living in rivers were not. Additionally, the chemical substances of a treefrog tadpole (Hylidae) were recognized by R. variabilis. Yet, they were not avoided but preferred for tadpole depositions; thus these tadpoles might be recognized as a potential prey for the predatory poison frog larvae. One of the poison frog species which was avoided for both tadpole and clutch depositions, was the phytotelmata breeding Hyloxalus azureiventris. The chemical substances released by its tadpoles were analysed together with those of the R. variabilis tadpoles (Chapter II). After finding a suitable solid-phase extraction sorbent (DSC-18), the active chemical compounds from the water of both tadpole species were extracted and fractionated. In order to determine which fractions triggered the avoidance behaviour of the frogs, in-situ bioassays were conducted. It was found that the biologically active compounds differed between both species. Since the avoidance of the conspecific tadpoles is not advantageous to the releaser tadpoles (losing a potential food resource) the chemicals released by them might be defined as chemical cues. However, as it turned out that the avoidance of the heterospecific tadpoles was not triggered by a mere byproduct based on the close evolutionary relationship between the two species, the chemical compounds released by H. azureiventris tadpoles might be defined as chemical signals (being advantageous to the releasing tadpoles) or, more specifically as synomones, interspecificly acting chemicals that are advantageous for both emitter and receiver (since R. variabilis avoids a competition situation for its offspring, too). Another interspecific communication system investigated in this thesis was the avoidance of predator kairomones (Chapter III). Using chemical substances from damselfly larvae, it could be shown that R. variabilis was unable to recognize and avoid kairomones of these tadpole predators. However, when physically present, damselfly larvae were avoided by the frogs. For the recognition of conspecific tadpoles in contrast, chemical substances were necessary, since purely visible artificial tadpole models were not avoided. If R. variabilis is also capable to chemically communicate with adult conspecifics was investigated by presenting chemical cues/signals of same-sex or opposite-sex conspecifics to the frogs (Chapter IV). It was suggested that males would be attracted to chemical substances of females and repelled by those of conspecific males. But instead all individuals showed avoidance behaviour towards the conspecific chemicals. This was suggested to be an artefact due to confinement stress of the releaser animals, emitting disturbance cues that triggered avoidance behaviour in their conspecifics. The knowledge gained about chemical communication in parental care thus far, was used to further investigate a possible provisioning behaviour in R. variabilis. In-situ pool-choice experiments with chemical cues of conspecific tadpoles were carried out throughout the change from rainy to dry season (Chapter V). With a changepoint analysis, the exact seasonal change was defined and differences between frogs" choices were analysed. It turned out that R. variabilis does not avoid but prefer conspecific cues during the dry season for tadpole depositions, what might be interpreted as a way to provide their tadpoles with food (i.e. younger tadpoles) in order to accelerate their development when facing desiccation risk. That tadpoles were also occasionally fed with fertilized eggs could be shown in a comparative study, where phytotelmata that contained a tadpole deposited by the frogs themselves received more clutch depositions than freshly erected artificial phytotelmata containing unfamiliar tadpoles (i.e. their chemical cues; Chapter VI). Conducting home range calculations with ArcGIS, it turned out that R. variabilis males showed unexpectedly strong site fidelity, leading to the suggestion that they recognize their offspring by phytotelmata location. However, in order to test if R. variabilis is furthermore able to perform chemical offspring recognition, frogs were confronted in in-situ pool-choice experiments with chemical cues of single tadpoles that were found in their home ranges (Chapter VII). Genetic kinship analyses were conducted between those tadpoles emitting the chemical cues and those deposited together with or next to them. The results, however, indicated that frogs did not choose to deposit their offspring with or without another tadpole due to relatedness, i.e. kin recognition by chemical cues could not be confirmed in R. variabilis.
High-resolution projections of the future climate are required to assess climate change realistically at a regional scale. This is in particular important for climate change impact studies since global projections are much too coarse to represent local conditions adequately. A major concern is thereby the change of extreme values in a warming climate due to their severe impact on the natural environment, socio-economical systems and the human health. Regional climate models (RCMs) are, however, able to reproduce much of those local features. Current horizontal resolutions are about 18-25km, which is still too coarse to directly resolve small-scale processes such as deep-convection. For this reason, projections of a possible future climate were simulated in this study with the regional climate model COSMO-CLM at horizontal resolutions of 4.5km and 1.3km for the region of Saarland-Lorraine-Luxemburg and Rhineland-Palatinate for the first time. At a horizontal scale of about 1km deep-convection is treated explicitly, which is expected to improve particularly the simulation of convective summer precipitation and a better resolved orography is expected to improve near surface fields such as 2m temperature. These simulations were performed as 10-year long time-slice experiments for the present climate (1991"2000), the near future (2041"2050) and the end of the century (2091"2100). The climate change signals of the annual and seasonal means and the change of extremes are analysed with respect to precipitation and 2m temperature and a possible added value due to the increased resolution is investigated. To assess changes in extremes, extreme indices have been applied and 10- and 20-year return levels were estimated by "peak-over-threshold" models. Since it is generally known that model output of RCMs should not directly be used for climate change impact studies, the precipitation and temperature fields were bias-corrected with several quantile-matching methods. Among them is a new developed parametric method which includes an extension for extreme values and is hence expected to improve the correction. In addition, the impact of the bias-correction on the climate change signals and on the extreme value statistics was investigated. The results reveal a significant warming of the annual mean by about +1.7 -°C until 2041"2050 and +3.7 -°C until 2091"2100, but considerably stronger signals of up to +5 -°C in summer in the Rhine Valley. Furthermore, the daily variability increases by about +0.8 -°C in summer but decreases by about -0.8 -°C in winter. Consequently, hot extremes increase moderately until the mid of the century but strongly thereafter, in particular in the Rhine Valley. Cold extremes warm continuously in the complete domain in the next 100 years but strongest in mountainous areas. The change signals with regard to annual precipitation are of the order -±10% but not significant. Significant, however, are a predicted increase of +32% of the seasonal precipitation in autumn until 2041"2050 and a decrease of -28% in summer until 2091-2100. No significant changes were found for days with intensities > 20 mm/day, but the results indicate that extremes with return periods ≤2 years increase as well as the frequency and duration of dry periods. The bias-corrections amplified positive signals but dampened negative signals and considerably reduced the power of detection. Moreover, absolute values and frequencies of extremes were altered by the correction but change signals remained approximately constant. The new method outperformed other parametric methods, in particular with regard to extreme value correction and related extreme indices and return levels. Although the bias correction removed systematic errors, it should be treated as an additional layer of uncertainty in climate change studies. Finally, the increased resolution of 1.3km improved predominantly the representation of temperature fields and extremes in terms of spatial heterogeneity. The benefits for summer precipitation were not as clear due to a severe dry-bias in summer, but it could be shown that in principle the onset and intensity of convection improves. This work demonstrates that climate change will have severe impacts in this investigation area and that in particular extremes may change considerably. An increased resolution provides thereby an added value to the results. These findings encourage further investigations, for other variables as for example near-surface wind, which will be more feasible with growing computing resources. These analyses should, however, be repeated with longer time series, different RCMs and anthropogenic scenarios to determine the robustness and uncertainty of these results more extensively.
Climate change and habitat fragmentation modify the natural habitat of many wetland biota and lead to new compositions of biodiversity in these ecosystems. While the direct effects of climate are often well known, indirect effects due to biotic interactions remain poorly understood. The water meadow grasshopper, Chorthippus montanus, is a univoltine habitat specialist, which is adapted to permanently moist habitats. Land use change and drainage led to highly fragmented populations of this generally flightless species. In large parts of the Palaearctic Ch. montanus occurs sympatrically with its widespread congener, the meadow grasshopper Chorthippus parallelus. Due to their close relationship and their similar songs, hybridization is likely to occur in syntopic populations. Such a species pair of a habitat specialist and a habitat generalist represents an ideal model system to examine the role of ongoing climate change and an accumulation of extreme climatic events on the life history strategies, population dynamics and inter-specific interactions. In Chapter I a laboratory experiment was conducted to identify the impact of environmental factors on intra-specific life-history traits of Ch. montanus. Like other Orthoptera species, Ch. montanus follows a converse temperature size rule. In line with the dimorphic niche hypothesis, which states that sexual size dimorphism evolved in response to the different sexual reproductive roles, both sexes showed different responses to increasing density at lower temperatures. Males attained smaller body sizes at high densities, whereas females had a prolonged development time. This is the first evidence for a sex-specific phenotypic plasticity in Ch. montanus. Females benefit from the prolonged development as their reproductive success depends on the size and number of egg clutches they may produce. By contrast, the reproductive success of males depends on the chance to fertilize virgin females, which increases with faster development. This may become a disadvantage for Ch. montanus as an intraspecific phenology shift may increase hybridization risk with the sibling species. Despite the widespread assumption that hybridization between two sympatric species is rare due to complete reproductive barriers, the genetic analyses of 16 populations (Chapter II) provided evidence for wide prevalence of hybridization between both species in the wild. As no complete admixture was found in the examined population, it is assumed that hybridization only occurs in ecotones between wetlands and drier parts. Reproductive barriers (habitat isolation, behavior, phenology) seem to prevent the genetic swamping of Ch. montanus populations. Although a behavioral experiment showed that mate choice presents an important reproductive barrier between both species, the experiment also revealed that reproductive barriers could be altered by environmental change (e.g. increasing heterospecific frequency). Chapter III analyzes the impact of extreme climatic events on population dynamics and interspecific hybridization. A mark-recapture analysis combined with weather records over five years provides evidence that the embryonic development in Ch. montanus is vulnerable to extreme climatic events. Strong population declines in Ch. montanus lead to a disequilibrium between Ch. montanus and Ch. parallelus populations and increases the risk of hybridization. The highest hybridization risk was found in the first weeks of a season, when both species had an overlapping phenology. Furthermore, hybrids were generally localized at the edge of the Ch. montanus distribution with higher heterospecific encounter probabilities. The hybridization rate reached up to 19.6%. The genetic analyses in Chapter II and III show that hybridization differentially affects specialists and generalists. While generalists may benefit from hybridization by an increasing genetic diversity, such a positive correlation was not found for Ch. montanus. The results underline the importance of reproductive barriers for the co-existence of these sympatric species. However, climate change and other anthropogenic disturbances alter reproductive barriers and promote hybridization, which may threaten small populations by genetic displacement. As anthropogenic hybridization is recognized as a major threat to biodiversity, it should be considered in environmental law and policy. In Chapter IV the role of hybrids and hybridization in three levels of law and the historical backgrounds of hybrids becoming a part of legal instruments is analyzed. Due to legal uncertainties and the complexity of this topic a legal assessment of hybrids is challenging and argues for species-specific approaches. Nonetheless, existing legal norms provide a suitable basis, but need to be specified. Finally, this chapter discusses different opportunities for the management of hybrids and hybridization in a conservation perspective and their necessity.
Comparing the results of the phylogeographies of the four species included in this thesis, some accordances have been found, even though certain patterns are only represented in one or two species. In all cases, the findings of the studied species strongly support the existence of forests or forest-like ecosystems beyond the classic forest refugia in the Mediterranean areas (Iberian, Apennine and Balkan peninsulas) during glacial times. However, evidence of glacial refugial areas in Southeastern Europe, especially the Balkans, have been found in this study as well. The analysed populations of Aposeris foetida, Melampyrum sylvaticum and Erebia euryale showed high genetic diversity values and mostly higher private fragments in this area, which is a strong indicator for centres of glacial survival during Würm and, regarding the results of M. sylvaticum, even during the Riss ice age. Three of the analysed species (A. foetida, M. sylvaticum and Colias palaeno) supported a second main glacial refuge area located along the Northern Alps. Again, high genetic diversity values and the uniqueness of the populations living in this region today prove the importance of this area as a glacial centre of survival. Those results confirm several recently published studies on forest species and strongly indicate the persistence of forest-like structures or even forests during the ice ages along the foothills of the Northern Alps. Additionally, the persistence of C. palaeno in this area furthermore supports the existence of peatlands north of the Alps, at least during the last glacial. The results of M. sylvaticum and E. euryale further indicate the vicinity of the Tatra Mountains as core areas for glacial survival. However, the genetic patterns found for E. euryale are ambiguous. Due to an intermediate position of two genetic lineages (originating in the Eastern Alps and Southeastern Europe), the Tatras could also reflect a postglacial mixture zone of those lineages. Moreover, the glacial and postglacial importance of this area for woodland species was accentuated, supporting other phylogeographic studies published. Besides the congruities among the results of the study species, some unique patterns and therefore further potential glacial refugia have also been illuminated in this thesis. For instance, the calcicole species, A. foetida, most probably had further survival area at both sides of the Dinaric Alps, supported by high genetic diversity values and a high number of private fragments found in Croatian populations. Furthermore, the surroundings of the German Uplands and the margin of the Southern Alps provided suitable conditions for glacial survival for M. sylvaticum, while the Eastern and Southeastern Alpine region most probably sheltered the Large Ringlet E. euryale during ice ages. Additionally, this butterfly species survived at least the glaciation along the foothills of the Massif Central, whose present populations showed a unique genetic lineage and their genetic diversity values have been measurably higher than in other populations for this species. Finally, a large and continuous Würm distribution is highly likely south of the Fennoscandian glaciers in Central Europe for C. palaeno, which might indicate extended peatland areas during Würm glacial. With all the patterns found in this study, the understanding of glacial persistence of forest, respectively forest-like structures and peatlands during Würm or even Riss glacial in Europe could be advanced. The congruencies among the analysed woodland and bog species illustrate the importance and location of extra-Mediterranean refugia for European mountain forests and the glacial presence of Central European peatlands. Thus, already postulated theories could be supported and further pieces of the overall puzzle could be added. The varieties of the different survival centres once more clarified that further phylogeographic studies on mountain forest of different habitat requirements and especially peatland species have to be implemented to get a clearer picture of the glacial history of these habitats.
Mechanical and Biological Treatment (MBT) generally aims to reduce the amount of solid waste and emissions in landfills and enhance the recoveries. MBT technology has been studied in various countries in Europe and Asia. Techniques of solid waste treatment are distinctly different in the study areas. A better understanding of MBT waste characteristics can lead to an optimization of the MBT technology. For a sustainable waste management, it is essential to determine the characteristics of the final MBT waste, the effectiveness of the treatment system as well as the potential application of the final material regarding future utilization. This study aims to define and compare the characteristics of the final MBT materials in the following countries: Luxembourg (using a high degree technology), Fridhaff in Diekirch/Erpeldange, Germany (using a well regulated technology), Singhofen in Rhein-Lahn district, Thailand (using a low cost technology): Phitsanulok in Phitsanulok province. The three countries were chosen for this comparative study due to their unique performance in the MBT implementation. The samples were taken from the composting heaps of the final treatment process prior to sending them to landfills, using a random sampling standard strategy from August 2008 onwards. The size of the sample was reduced to manageable sizes before characterization. The size reduction was achieved by the quartering method. The samples were first analyzed for the size fraction on the day of collection. They were screened into three fractions by the method of dry sieving: small size with a diameter of <10 mm, medium size with a diameter of 10-40 mm and large size with a diameter of >40 mm. These fractions were further analyzed for their physical and chemical parameters such as particle size distribution (total into 12 size fractions), particle shape, porosity, composition, water content, water retention capacity and respiratory activity. The extracted eluate was analyzed for pH-value, heavy metals (lead, cadmium and arsenic), chemical oxygen demand, ammonium, sulfate and chloride. In order to describe and evaluate the potential application of the small size material as a final cover of landfills, the fraction of small size samples were tested for the geotechnical properties as well. The geotechnical parameters were the compaction test, permeability test and shear strength test. The detailed description of the treatment facilities and methods of the study areas were included in the results. The samples from the three countries are visibly smaller than waste without pretreatment. Maximum particle size is found to be less than 100 mm. The samples are found to consist of dust to coarse fractions. The small size with a diameter of <10 mm was highest in the sample from Germany (average 60% by weight), secondly in the sample from Luxembourg (average 43% by weight) and lowest in the sample from Thailand (average 15% by weight). The content of biodegradable material generally increased with decreasing particle sizes. Primary components are organic, plastics, fibrous materials and inert materials (glass and ceramics). The percentage of each components greatly depends on the MBT process of each country. Other important characteristics are significantly reduced water content, reduced total organic carbon and reduced potential heavy metals. The geotechnical results show that the small fraction is highly compact, has a low permeability and lot of water adsorbed material. The utilization of MBT material in this study shows a good trend as it proved to be a safe material which contained very low amounts of loadings and concentrations of chemical oxygen demand, ammonium, and heavy metals. The organic part can be developed to be a soil conditioner. It is also suitably utilized as a bio-filter layer in the final cover of landfill or as a temporary cover during the MBT process. This study showed how to identify the most appropriate technology for municipal solid waste disposal through the study of waste characterization.
It has been the overall aim of this research work to assess the potential of hyperspectral remote sensing data for the determination of forest attributes relevant to forest ecosystem simulation modeling and forest inventory purposes. A number of approaches for the determination of structural and chemical attributes from hyperspectral remote sensing have been applied to the collected data sets. Many of the methods to be found in the literature were up to now just applied to broadband multispectral data, applied to vegetation canopies other than forests, reported to work on the leaf level or with modelled data, not validated with ground truth data, or not systematically compared to other methods. Attributes that describe the properties of the forest canopy and that are potentially open to remote sensing were identified, appropriate methods for their retrieval were implemented and field, laboratory and image data (HyMap sensor) were acquired over a number of forest plots. The study on structural attributes compared statistical and physical approaches. In the statistical section, linear predictive models between vegetation indices derived from HyMap data and field measurements of structural forest stand attributes were systematically evaluated. The study demonstrates that for hyperspectral image data, linear regression models can be applied to quantify leaf area index and crown volume with good accuracy. For broadband multispectral data, the accuracy was generally lower. The physically-based approach used the invertible forest reflectance model (INFORM), a combination of well established sub-models FLIM, SAIL and LIBERTY. The model was inverted with HyMap data using a neural network approach. In comparison to the statistical approach, it could be shown that the reflectance model inversion works equally well. In opposition to empirically derived prediction functions that are generally limited to the local conditions at a certain point in time and to a specified sensor type, the calibrated reflectance model can be applied more easily to different optical remote sensing data acquired over central European forests. The study on chemical forest attributes evaluated the information content of HyMap data for the estimation of nitrogen, chlorophyll and water concentration. A number of needle samples of Norway spruce were analysed for their total chlorophyll, nitrogen and water concentrations. The chemical data was linked to needle spectra measured in the laboratory and canopy spectra measured by the HyMap sensor. Wavebands selected in statistical models were often located in spectral regions that are known to be important for chlorophyll detection (red edge, green peak). Predictive models were applied on the HyMap image to compute maps of chlorophyll concentration and nitrogen concentration. Results of map overlay operations revealed coherence between total chlorophyll and zones of stand development stage and between total chlorophyll and zones of soil type. Finally, it can be stated that the hyperspectral remote sensing data generally contains more information relevant to the estimation of the forest attributes compared to multispectral data. Structural forest attributes, except biomass, can be determined with good accuracy from a hyperspectral sensor type like HyMap. Among the chemical attributes, chlorophyll concentration can be determined with good accuracy and nitrogen concentration with moderate accuracy. For future research, additional dimensions have to be taken into account, for instance through exploitation of multi-view angle data. Additionally, existing forest canopy reflectance models should be further improved.
A big challenge for agriculture in the 21st century is the provision of food safety to fast growing world- population, which not only demands the well utilisation of the available agricultural resources but also to develop new advancements in the mass production of food crops. Wheat is the third largest food crop of the world and Pakistan is the eighth largest wheat producing country globally. Rice is the second most important staple food of Pakistan after wheat, grown in all provinces of the country. Maize is the world- top ranking food crop followed by wheat and rice. The harvested produts have to be stored in different types of storage structures on small or large scale for food as well as seed purpose. In Pakistan, the harvested grains are stored for the whole year till the introduction of fresh produce in order to ensure the regular food supply throughout the year. However, it is this extended storage period making the commodity more vulnerable to insect attacks. Rhyzopertha dominica (Coleoptera: Bostrychidae), Cryptolestes ferrugineus (Coleoptera: Laemophloeidae), Tribolium castaneum (Coleoptera: Tenebrionidae) and Liposcelis spp. (Psocoptera: Liposcelididae) are the major and most damaging insect pests of stored products all around the world. Various management strategies have been adopted for stored grain insect pests mostly relying upon the use of a broad spectrum of insecticides, but the injudicious use of these chemicals raised various environmental and human health related issues, which necessitate the safe use of the prevailing control measures and evaluation of new and alternative control methods. The application of new chemical insecticides, microbial insecticides (particularly entomopathogenic fungi) and the use of inert dusts (diatomaceous earths) is believed amongst the potential alternatives to generally used insecticides in stored grain insect management system. In the current investigations, laboratory bioassays conducted to evaluate the effects of combining Imidacloprid (new chemistry insecticide) with and without Protect-It (diatomaceous earth formulation) against R. dominica, L. paeta, C. ferrugineus and T. castaneum, on three different grain commodities (i.e. wheat, maize and rice) revealed differences in adult mortality levels among grains and insect species tested. Individually, Imidacloprid was more effective as compared with Protect-It alone and the highest numbers of dead adults were recorded in wheat. The insecticidal efficacy of B. bassiana with Protect-It and DEBBM was also assessed against all test insect species under laboratory conditions. The findings of these studies revealed that the more extended exposure period and the higher combined application rate of B. bassiana and DEs provided the highest mortality of the test insect species. The progeny emergence of each insect species was also greatly suppressed where the highest dose rates of the combined treatments were applied. The residual efficacy of all three control measures Imidacloprid, B. bassiana and DEBBM formulation was also evaluated against all test insect species. The bioassays were carried out after grain treatments and monthly for 6 months. The results indicated that the adult mortality of each test insect species was decreased within the six month storage period, and the integarted application of the test grain protectants enhanced the mortality rates than their alone treatments. The maximum mortality was noted in the combined treatment of DEBBM with Imidacloprid. At the end, the effectiveness of B. bassiana, DEBBM and Imidacloprid applied alone as well as in combinations, against all above mentioned test insect species was also evaluated under field conditions in trials conducted in four districts of Punjab, Pakistan. For each district, a significant difference was observed between treatments, while the combined treatments gave better control of test species as compared with them alone. The least number of surviving adults and minimum percentage of grain damage was observed for the DEBBM and Imidacloprid combination, but DEBBM with B. bassiana provided the best long-term protection as compared with the remaining treatments.
Tropospheric ozone (O3) is known to have various detrimental effects on plants, such as visible leaf injury, reduced growth and premature senescence. Flux models offer the determination of the harmful ozone dose entering the plant through the stomata. This dose can then be related to phytotoxic effects mentioned above to obtain dose-response relationships, which are a helpful tool for the formulation of abatement strategies of ozone precursors. rnOzone flux models are dependant on the correct estimation of stomatal conductance (gs). Based on measurements of gs, an ozone flux model for two white clover clones (Trifolium repens L. cv Regal; NC-S (ozone-sensitive) and NC-R (ozone-resistant)) differing in their sensitivity to ozone was developed with the help of artificial neural networks (ANNs). White clover is an important species of various European grassland communities. The clover plants were exposed to ambient air at three sites in the Trier region (West Germany) during five consecutive growing seasons (1997 to 2001). The response parameters visible leaf injury and biomass ratio of NC-S/NC-R clone were regularly assessed. gs-measurements of both clones functioned as output of the ANN-based gs model, while corresponding climate parameters (i.e. temperature, vapour pressure deficit (VPD) and photosynthetic active radiation (PAR)) and various ozone concentration indices were inputs. The development of the model was documented in detail and various model evaluation techniques (e.g. sensitivity analysis) were applied. The resulting gs model was used as a basis for ozone flux calculations, which were related to above mentioned response parameters. rnThe results showed that the ANNs were capable of revealing and learning the complex relationship between gs and key meteorological parameters and ozone concentration indices. The dose-response relationships between ozone fluxes and visible leaf injury were reasonably strong, while those between ozone fluxes and NC-S/NC-R biomass ratio were fairly weak. The results were discussed in detail with respect to the suitability of the chosen experimental methods and model type.
This dissertation develops a rationale of how to use fossil data in solving biogeographical and ecological problems. It is argued that large amounts of fossil data of high quality can be used to document the evolutionary processes (the origin, development, formation and dynamics) of Arealsystems, which can be divided into six stages in North America: the Refugium Stage (before 15,000 years ago: > 15 ka), the Dispersal Stage (from 8,000 to 15,000 years ago: 8.0 - 15 ka), the Developing Stage (from 3,000 to 8,000 years ago: 3.0 - 8.0 ka), the Transitional Stage (from 1,000 to 3,000 years ago: 1 - 3 ka), the Primitive Stage (from 5,00 to 1,000 years ago: 0.5 - 1 ka) and the Human Disturbing Stage (during the last 500 years: < 0.5 ka). The division into these six stages is based on geostatistical analysis of the FAUNMAP database that contains 43,851 fossil records collected from 1860 to 1994 in North America. Fossil data are one of the best materials to test the glacial refugia theory. Glacial refugia represent areas where flora and fauna were preserved during the glacial period, characterized by richness in species and endemic species at present. This means that these (endemic) species should have distributed purely or primarily in these areas during the glacial period. The refugia can therefore be identified by fossil records of that period. If it is not the case, the richness in (endemic) species may not be the result of the glacial refugia. By exploring where mammals lived during the Refugium Stage (> 15 ka), seven refugia in North America can be identified: the California Refugium, the Mexico Refugium, the Florida Refugium, the Appalachia Refugium, the Great Basin Refugium, the Rocky Mountain Refugium and the Great Lake Refugium. The first five refugia coincide well with De Lattin- dispersal centers recognized by biogeographical methods using data on modern distributions. The individuals of a species are not evenly distributed over its Arealsystem. Brown- Hot Spots Model shows that in most cases there is an enormous variation in abundance within an areal of a species: In a census, zero or only a very few individuals occur at most sample locations, but tens or hundreds are found at a few sample sites. Locations where only a few individuals can be sampled in a survey are called "cool spots", and sites where tens or hundreds of individuals can be observed in a survey are called "hot spots". Many areas within the areal are uninhabited, which are called "holes". This model has direct implications for analyzing fossil data: Hot spots have a much higher local population density than cool spots. The chances to discover fossil individuals of a species are much higher in sediments located in a "hot spot" area than in a "cool spot" area. Therefore much higher MNIs (Minimum Number of Individuals) of the species should be found in fossil localities located in the hot spot than in the cool spot area. There are only a few hot spots but many cool spots within an areal of a single hypothetical species, consequently only a few fossil sites can provide with much high MNIs, whereas most other sites can only provide with very low MNIs. This prediction has been proved to be true by analysis of 70 species in FAUMAP containing over 100 fossil records. The temporal and spatial variation in abundance can be reconstructed from the temporospatial distribution of the MNIs of a species over its Arealsystem. Areas with no fossil records from the last thousands of years may be holes, and sites with much higher MNIs may be hot spots, while locations with low MNIs may be cool spots. Although the hot spots of many species can remain unchanged in an area over thousands of years, our study shows that a large shift of hot spots occurred mainly around 1,500-1,000 years ago. There are three directions of movement: from the west side to the east side of the Rockies, from the East of the USA to the east side of the Rockies and from the west side of the Rockies to the Southwest of the USA. The first two directions of shift are called Lewis and Clark- pattern, which can be verified with the observations mad by Lewis and Clark during their expedition in 1805-1806. The historical process of this pattern may well explain the 200-year-old puzzle why big game then abundant on the east side were rare on the west side of the Rocky Mountains noted by modern ecologists and biogeographers. The third direction of shift is called Bayham- pattern. This pattern can be tested by the model of Late Holocene resource intensification first described by Frank E. Bayham. The historical process creating the Bayham pattern will challenge the classic explanation of the Late Holocene resource intensification. An environmental change model has been proposed to account for the shift of hot spots. Implications of glacial refugia and hot spots areas for wildlife management and effective conservation are discussed. Suggestions for paleontologists and zooarchaeologists regarding how to provide more valuable information in their future excavation and research for other disciplines are given.
This dissertation focuses on e-marketing strategy's effective elements in tourism industry. As case study, research focus is on Airlines, tour operator, chain hotels in Iran and Germany. It aims to show various possibilities to enhance the company- e-marketing strategy and successfully performance e-marketing strategies with recognition effective elements and their important during the strategy designing and implementation process. For the purpose of this research due to the nature of the research, Explanatory -exploratory-applicable; after studying and consulting, Delphi technique has been chosen. In results, we have some effective elements and their important according the Delphi and AHP method. For example between elements "Tourists' Needs, Experience and Expects" with the importance coefficient of %204 is the most remarkable elements and "Customer satisfactions' elements group" with average value 5.54 according the research results have more important than other groups.