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A large number and wide variety of lake ecosystem models have been developed and published during the past four decades. We identify two challenges for making further progress in this field. One such challenge is to avoid developing more models largely following the concept of others ('reinventing the wheel'). The other challenge is to avoid focusing on only one type of model, while ignoring new and diverse approaches that have become available ('having tunnel vision'). In this paper, we aim at improving the awareness of existing models and knowledge of concurrent approaches in lake ecosystem modelling, without covering all possible model tools and avenues. First, we present a broad variety of modelling approaches. To illustrate these approaches, we give brief descriptions of rather arbitrarily selected sets of specific models. We deal with static models (steady state and regression models), complex dynamic models (CAEDYM, CE-QUAL-W2, Delft 3D-ECO, LakeMab, LakeWeb, MyLake, PCLake, PROTECH, SALMO), structurally dynamic models and minimal dynamic models. We also discuss a group of approaches that could all be classified as individual based: super-individual models (Piscator, Charisma), physiologically structured models, stage-structured models and traitbased models. We briefly mention genetic algorithms, neural networks, Kalman filters and fuzzy logic. Thereafter, we zoom in, as an in-depth example, on the multi-decadal development and application of the lake ecosystem model PCLake and related models (PCLake Metamodel, Lake Shira Model, IPH-TRIM3D-PCLake). In the discussion, we argue that while the historical development of each approach and model is understandable given its 'leading principle', there are many opportunities for combining approaches. We take the point of view that a single 'right' approach does not exist and should not be strived for. Instead, multiple modelling approaches, applied concurrently to a given problem, can help develop an integrative view on the functioning of lake ecosystems. We end with a set of specific recommendations that may be of help in the further development of lake ecosystem models.
Human-induced alterations of the environment are causing biotic changes worldwide, including the extinction of species and a mixing of once disparate floras and faunas. One type of biological communities that is expected to be particularly affected by environmental alterations are herb layer plant communities of fragmented forests such as those in the west European lowlands. However, our knowledge about current changes in species diversity and composition in these communities is limited due to a lack of adequate long-term studies. In this thesis, I resurveyed the herb layer communities of ancient forest patches in the Weser-Elbe region (NW Germany) after two decades using 175 semi-permanent plots. The general objectives were (i) to quantify changes in plant species diversity considering also between-community (β) and functional diversity, (ii) to determine shifts in species composition in terms of species’ niche breadth and functional traits and (iii) to find indications on the most likely environmental drivers for the observed changes. These objectives were pursued with four independent research papers (Chapters 1-4) whose results were brought together in a General Discussion. Alpha diversity (species richness) increased by almost four species on average, whereas β diversity tended to decrease (Chapter 1). The latter is interpreted as a beginning floristic homogenization. The observed changes were primarily the result of a spread of native habitat generalists that are able to tolerate broad pH and moisture ranges. The changes in α and β diversity were only significant when species abundances were neglected (Chapters 1 and 2), demonstrating that the diversity changes resulted mainly from gains and losses of low-abundance species. This study is one of the first studies in temperate Europe that demonstrates floristic homogenization of forest plant communities at a larger than local scale. The diversity changes found at the taxonomic level did not result in similar changes at the functional level (Chapter 2). The likely reason is that these communities are functionally “buffered”. Single communities involve most of the functional diversity of the regional pool, i.e., they are already functionally rich, while they are functionally redundant among each other, i.e., they are already homogeneous. Independent of taxonomic homogenization, the abundance of 30 species decreased significantly (Chapter 4). These species included 12 ancient forest species (i.e., species closely tied to forest patches with a habitat continuity > 200 years) and seven species listed on the Red List of endangered plant species in NW Germany. If these decreases continue over the next decades, local extinctions may result. This biotic impoverishment would seriously conflict with regional conservation goals. Community assembly mechanisms changed at the local level particularly at sites that experienced disturbance by forest management activities between the sampling periods (Chapter 3). Disturbance altered community assembly mechanisms in two ways: (i) it relaxed environmental filters and allowed the coexistence of different reproduction strategies, as reflected by a higher diversity of reproductive traits at the time of the resurvey, and (ii) it enhanced light availability and tightened competitive filters. These limited the functional diversity with respect to canopy height and selected for taller species. Thirty-one winner and 30 loser species, which had significantly increased or decreased in abundance, respectively, were characterized by various functional traits and ecological performances to find indications on the most likely environmental drivers for the observed floristic changes (Chapter 4). Winner species had higher seed longevity, flowered later in the season and had more often an oceanic distribution compared to loser species. Loser species tended to have a higher specific leaf area, to be more susceptible to deer browsing and to have a performance optimum at higher soil pH values compared to winner species. Multiple logistic regression analyses indicated that disturbances due to forest management interventions were the primary cause of the species shifts. As one of the first European resurvey studies, this study provides indications that an enhanced browsing pressure due to increased deer densities and increasingly warmer winters are important drivers. The study failed to demonstrate that eutrophication and acidification due to atmospheric deposition substantially drive herb layer changes. The restriction of the sample to the most base-rich sites in the region is discussed as a likely reason. Furthermore, the decline of several ancient forest species is discussed as an indication that the forest patches are still paying off their “extinction debt”, i.e., exhibit a delayed response to forest fragmentation.
Aim The study and prediction of speciesenvironment relationships is currently mainly based on species distribution models. These purely correlative models neglect spatial population dynamics and assume that species distributions are in equilibrium with their environment. This causes biased estimates of species niches and handicaps forecasts of range dynamics under environmental change. Here we aim to develop an approach that statistically estimates process-based models of range dynamics from data on species distributions and permits a more comprehensive quantification of forecast uncertainties.
Innovation We present an approach for the statistical estimation of process-based dynamic range models (DRMs) that integrate Hutchinson's niche concept with spatial population dynamics. In a hierarchical Bayesian framework the environmental response of demographic rates, local population dynamics and dispersal are estimated conditional upon each other while accounting for various sources of uncertainty. The method thus: (1) jointly infers species niches and spatiotemporal population dynamics from occurrence and abundance data, and (2) provides fully probabilistic forecasts of future range dynamics under environmental change. In a simulation study, we investigate the performance of DRMs for a variety of scenarios that differ in both ecological dynamics and the data used for model estimation.
Main conclusions Our results demonstrate the importance of considering dynamic aspects in the collection and analysis of biodiversity data. In combination with informative data, the presented framework has the potential to markedly improve the quantification of ecological niches, the process-based understanding of range dynamics and the forecasting of species responses to environmental change. It thereby strengthens links between biogeography, population biology and theoretical and applied ecology.
Specialisation and diversity of multiple trophic groups are promoted by different forest features
(2019)
While forest management strongly influences biodiversity, it remains unclear how the structural and compositional changes caused by management affect different community dimensions (e.g. richness, specialisation, abundance or completeness) and how this differs between taxa. We assessed the effects of nine forest features (representing stand structure, heterogeneity and tree composition) on thirteen above- and belowground trophic groups of plants, animals, fungi and bacteria in 150 temperate forest plots differing in their management type. Canopy cover decreased light resources, which increased community specialisation but reduced overall diversity and abundance. Features increasing resource types and diversifying microhabitats (admixing of oaks and conifers) were important and mostly affected richness. Belowground groups responded differently to those aboveground and had weaker responses to most forest features. Our results show that we need to consider forest features rather than broad management types and highlight the importance of considering several groups and community dimensions to better inform conservation.
Anthropogenic activities have transformed the Earth's environment, not only on local level, but on the planetary-scale causing global change. Besides industrialization, agriculture is a major driver of global change. This change in turn impairs the agriculture sector, reducing crop yields namely due to soil degradation, water scarcity, and climate change. However, this is a more complex issue than it appears. Crop yields can be increased by use of agrochemicals and fertilizers which are mainly produced by fossil energy. This is important to meet the increasing food demand driven by global demographic change, which is further accelerated by changes in regional lifestyles. In this dissertation, we attempt to address this complex problem exploring agricultural potential globally but on a local scale. For this, we considered the influence of lifestyle changes (dietary patterns) as well as technological progress and their effects on climate change, mainly greenhouse gas (GHG) emissions. Furthermore, we examined options for optimizing crop yields in the current cultivated land with the current cropping patterns by closing yield gaps. Using this, we investigated in a five-minute resolution the extent to which food demand can be met locally, and/or by regional and/or global trade. Globally, food consumption habits are shifting towards calorie rich diets. Due to dietary shifts combined with population growth, the global food demand is expected to increase by 60-110% between 2005 and 2050. Hence, one of the challenges to global sustainability is to meet the growing food demand, while at the same time, reducing agricultural inputs and environmental consequences. In order to address the above problem, we used several freely available datasets and applied multiple interconnected analytical approaches that include artificial neural network, scenario analysis, data aggregation and harmonization, downscaling algorithm, and cross-scale analysis.
Globally, we identified sixteen dietary patterns between 1961 and 2007 with food intakes ranging from 1,870 to 3,400 kcal/cap/day. These dietary patterns also reflected changing dietary habits to meat rich diets worldwide. Due to the large share of animal products, very high calorie diets that are common in the developed world, exhibit high total per capita emissions of 3.7-6.1 kg CO2eq./day. This is higher than total per capita emissions of 1.4-4.5 kg CO2eq./day associated with low and moderate calorie diets that are common in developing countries. Currently, 40% of the global crop calories are fed to livestock and the feed calorie use is four times the produced animal calories. However, these values vary from less than 1 kcal to greater 10 kcal around the world. On the local and national scale, we found that the local and national food production could meet demand of 1.9 and 4.4 billion people in 2000, respectively. However, 1 billion people from Asia and Africa require intercontinental agricultural trade to meet their food demand. Nevertheless, these regions can become food self-sufficient by closing yield gaps that require location specific inputs and agricultural management strategies. Such strategies include: fertilizers, pesticides, soil and land improvement, management targeted on mitigating climate induced yield variability, and improving market accessibility. However, closing yield gaps in particular requires global N-fertilizer application to increase by 45-73%, P2O5 by 22-46%, and K2O by 2-3 times compare to 2010. Considering population growth, we found that the global agricultural GHG emissions will approach 7 Gt CO2eq./yr by 2050, while the global livestock feed demand will remain similar to 2000. This changes tremendously when diet shifts are also taken into account, resulting in GHG emissions of 20 Gt CO2eq./yr and an increase of 1.3 times in the crop-based feed demand between 2000 and 2050. However, when population growth, diet shifts, and technological progress by 2050 were considered, GHG emissions can be reduced to 14 Gt CO2eq./yr and the feed demand to nearly 1.8 times compare to that in 2000. Additionally, our findings shows that based on the progress made in closing yield gaps, the number of people depending on international trade can vary between 1.5 and 6 billion by 2050. In medium term, this requires additional fossil energy. Furthermore, climate change, affecting crop yields, will increase the need for international agricultural trade by 4% to 16%.
In summary, three general conclusions are drawn from this dissertation. First, changing dietary patterns will significantly increase crop demand, agricultural GHG emissions, and international food trade in the future when compared to population growth only. Second, such increments can be reduced by technology transfer and technological progress that will enhance crop yields, decrease agricultural emission intensities, and increase livestock feed conversion efficiencies. Moreover, international trade dependency can be lowered by consuming local and regional food products, by producing diverse types of food, and by closing yield gaps. Third, location specific inputs and management options are required to close yield gaps. Sustainability of such inputs and management largely depends on which options are chosen and how they are implemented. However, while every cultivated land may not need to attain its potential yields to enable food security, closing yield gaps only may not be enough to achieve food self-sufficiency in some regions. Hence, a combination of sustainable implementations of agricultural intensification, expansion, and trade as well as shifting dietary habits towards a lower share of animal products is required to feed the growing population.
Biodiversity maintains soil multifunctionality and soil organic carbon in novel urban ecosystems
(2022)
Biodiversity in urban ecosystems has the potential to increase ecosystem functions and support a suite of services valued by society, including services provided by soils. Specifically, the sequestration of carbon in soils has often been advocated as a solution to mitigate the steady increase in CO2 concentration in the atmosphere as a key driver of climate change. However, urban ecosystems are also characterized by an often high level of ecological novelty due to profound human-mediated changes, such as the presence of high numbers of non-native species, impervious surfaces or other disturbances. Yet it is poorly understood whether and how biodiversity affects ecosystem functioning and services of urban soils under these novel conditions. In this study, we assessed the influence of above- and below-ground diversity, as well as urbanization and plant invasions, on multifunctionality and organic carbon stocks of soils in non-manipulated grasslands along an urbanization gradient in Berlin, Germany. We focused on plant diversity (measured as species richness and functional trait diversity) and, in addition, on soil organism diversity as a potential mediator for the relationship of plant species diversity and ecosystem functioning. Our results showed positive effects of plant diversity on soil multifunctionality and soil organic carbon stocks along the entire gradient. Structural equation models revealed that plant diversity enhanced soil multifunctionality and soil organic carbon by increasing the diversity of below-ground organisms. These positive effects of plant diversity on soil multifunctionality and soil fauna were not restricted to native plant species only, but were also exerted by non-native species, although to a lesser degree. Synthesis. We conclude that enhancing diversity in plants and soil fauna of urban grasslands can increase the multifunctionality of urban soils and also add to their often underestimated but very valuable role in mitigating effects of climate change.
This PhD thesis presents the spatio-temporal distribution of terrestrial carbon fluxes for the time period of 1982 to 2002 simulated by a combination of the process-based dynamic global vegetation model LPJ and a 21-year time series of global AVHRR-fPAR data (fPAR – fraction of photosynthetically active radiation). Assimilation of the satellite data into the model allows improved simulations of carbon fluxes on global as well as on regional scales. As it is based on observed data and includes agricultural regions, the model combined with satellite data produces more realistic carbon fluxes of net primary production (NPP), soil respiration, carbon released by fire and the net land-atmosphere flux than the potential vegetation model. It also produces a good fit to the interannual variability of the CO2 growth rate. Compared to the original model, the model with satellite data constraint produces generally smaller carbon fluxes than the purely climate-based stand-alone simulation of potential natural vegetation, now comparing better to literature estimates. The lower net fluxes are a result of a combination of several effects: reduction in vegetation cover, consideration of human influence and agricultural areas, an improved seasonality, changes in vegetation distribution and species composition. This study presents a way to assess terrestrial carbon fluxes and elucidates the processes contributing to interannual variability of the terrestrial carbon exchange. Process-based terrestrial modelling and satellite-observed vegetation data are successfully combined to improve estimates of vegetation carbon fluxes and stocks. As net ecosystem exchange is the most interesting and most sensitive factor in carbon cycle modelling and highly uncertain, the presented results complementary contribute to the current knowledge, supporting the understanding of the terrestrial carbon budget.
Recent research has shown that many cold-adapted species survived the last glacial maximum (LGM) in northern refugia. Whether this evolutionary history has had consequences for their genetic diversity and adaptive potential remains unknown. We sampled 14 populations of Carex limosa, a sedge specialized to bog ecosystems, along a latitudinal gradient from its Scandinavian core to the southern lowland range-margin in Germany. Using microsatellite and experimental common-garden data, we evaluated the impacts of global climate change along this gradient and assessed the conservation status of the southern marginal populations. Microsatellite data revealed two highly distinct genetic groups and hybrid individuals. In our common-garden experiment, the two groups showed divergent responses to increased nitrogen/phosphorus (N/P) availability, suggesting ecotypic differentiation. Each group formed genetically uniform populations at both northern and southern sampling areas. Mixed populations occurred throughout our sampling area, an area that was entirely glaciated during the LGM. The fragmented distribution implies allopatric divergence at geographically separated refugia that putatively differed in N/P availability. Molecular data and an observed low hybrid fecundity indicate the importance of clonal reproduction for hybrid populations. At the southern range-margin, however, all populations showed effects of clonality, lowered fecundity and low competitiveness, suggesting abiotic and biotic constraints to population persistence.
This is a publication-based dissertation comprising three original research stud-ies (one published, one submitted and one ready for submission; status March 2019). The dissertation introduces a generic computer model as a tool to investigate the behaviour and population dynamics of animals in cyclic environments. The model is further employed for analysing how migratory birds respond to various scenarios of altered food supply under global change. Here, ecological and evolutionary time-scales are considered, as well as the biological constraints and trade-offs the individual faces, which ultimately shape response dynamics at the population level. Further, the effect of fine-scale temporal patterns in re-source supply are studied, which is challenging to achieve experimentally. My findings predict population declines, altered behavioural timing and negative carry-over effects arising in migratory birds under global change. They thus stress the need for intensified research on how ecological mechanisms are affected by global change and for effective conservation measures for migratory birds. The open-source modelling software created for this dissertation can now be used for other taxa and related research questions. Overall, this thesis improves our mechanistic understanding of the impacts of global change on migratory birds as one prerequisite to comprehend ongoing global biodiversity loss. The research results are discussed in a broader ecological and scientific context in a concluding synthesis chapter.
Aim To understand the role and significance of the reindeer, Rangifer tarandus (Linnaeus, 1758), as a specific indicator in terms of late Quaternary biogeography and to determine the effects of global climate change on its range and local extinction dynamics at the end of the Ice Age.
Location Late Pleistocene/early Holocene range of reindeer over all of central and western Europe, including southern Scandinavia and northern Iberia, but excluding Russia, Belarus and the Ukraine.
Methods Radiocarbon-dated subfossil records of R. tarandus from both archaeological and natural deposits younger than 25,000 years were assembled in a database. The distribution area was divided into six representative regions. The C-14 dates were calibrated and plotted chronologically in maps in order to compare presence and absence and regional extinction patterns from one region to another.
Main conclusions The late Quaternary record for reindeer in Europe during the last 25 kyr shows a climate-driven dispersal and retreat in response to climate change, with regional variations. The collapse of the mammoth steppe biome did not lead to the local extinction in Europe, as in the case of other megafaunal species. Rangifer tarandus co-existed for about 3000 years during the Late Glacial and early Holocene with typical temperate species such as red deer and roe deer in non-analogue faunal communities. The regional extinction at the end of the Pleistocene coincides with the transition from light open birch/pine forests to pine/deciduous forests.