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Epigenetic modifications, of which DNA methylation is the best studied one, can convey environmental information through generations via parental germ lines. Past studies have focused on the maternal transmission of epigenetic information to the offspring of isogenic mice and rats in response to external changes, whereas heterogeneous wild mammals as well as paternal epigenetic effects have been widely neglected. In most wild mammal species, males are the dispersing sex and have to cope with differing habitats and thermal changes. As temperature is a major environmental factor we investigated if genetically heterogeneous Wild guinea pig (Cavia aperea) males can adapt epigenetically to an increase in temperature and if that response will be transmitted to the next generation(s). Five adult male guinea pigs (F0) were exposed to an increased ambient temperature for 2 months, i.e. the duration of spermatogenesis. We studied the liver (as the main thermoregulatory organ) of F0 fathers and F1 sons, and testes of F1 sons for paternal transmission of epigenetic modifications across generation(s). Reduced representation bisulphite sequencing revealed shared differentially methylated regions in annotated areas between F0 livers before and after heat treatment, and their sons’ livers and testes, which indicated a general response with ecological relevance. Thus, paternal exposure to a temporally limited increased ambient temperature led to an ‘immediate’ and ‘heritable’ epigenetic response that may even be transmitted to the F2 generation. In the context of globally rising temperatures epigenetic mechanisms may become increasingly relevant for the survival of species.
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.
Global change, especially land-use intensification, affects human well-being by impacting the delivery of multiple ecosystem services (multifunctionality). However, whether biodiversity loss is a major component of global change effects on multifunctionality in real-world ecosystems, as in experimental ones, remains unclear. Therefore, we assessed biodiversity, functional composition and 14 ecosystem services on 150 agricultural grasslands differing in land-use intensity. We also introduce five multifunctionality measures in which ecosystem services were weighted according to realistic land-use objectives. We found that indirect land-use effects, i.e. those mediated by biodiversity loss and by changes to functional composition, were as strong as direct effects on average. Their strength varied with land-use objectives and regional context. Biodiversity loss explained indirect effects in a region of intermediate productivity and was most damaging when land-use objectives favoured supporting and cultural services. In contrast, functional composition shifts, towards fast-growing plant species, strongly increased provisioning services in more inherently unproductive grasslands.
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.
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.
It has been proposed that growth and reproduction of animals is frequently limited by multiple nutrients simultaneously. To improve our understanding of the consequences of multiple nutrient limitations (i.e. co-limitation) for the performance of animals, we conducted standardized population growth experiments using an important aquatic consumer, the rotifer Brachionus calyciflorus. We compared nutrient profiles (sterols, fatty acids and amino acids) of rotifers and their diets to reveal consumerdiet imbalances and thus potentially limiting nutrients. In concomitant growth experiments, we directly supplemented potentially limiting substances (sterols, fatty acids, amino acids) to a nutrient-deficient diet, the cyanobacterium Synechococcus elongatus, and recorded population growth rates. The results from the supplementation experiments corroborated the nutrient limitations predicted by assessing consumerdiet imbalances, but provided more detailed information on co-limiting nutrients. While the fatty acid deficiency of the cyanobacterium appeared to be of minor importance, the addition of both cholesterol and certain amino acids (leucine and isoleucine) improved population growth rates of rotifers, indicating a simultaneous limitation by sterols and amino acids. Our results add to growing evidence that consumers frequently face multiple nutrient limitations and suggest that the concept of co-limitation has to be considered in studies assessing nutrient-limited growth responses of consumers.
The spatial distribution of a species is determined by dynamic processes such as reproduction, mortality and dispersal. Conventional static species distribution models (SDMs) do not incorporate these processes explicitly. This limits their applicability, particularly for non-equilibrium situations such as invasions or climate change. In this paper we show how dynamic SDMs can be formulated and fitted to data within a Bayesian framework. Our focus is on discrete state-space Markov process models which provide a flexible framework to account for stochasticity in key demographic processes, including dispersal, growth and competition. We show how to construct likelihood functions for such models (both discrete and continuous time versions) and how these can be combined with suitable observation models to conduct Bayesian parameter inference using computational techniques such as Markov chain Monte Carlo. We illustrate the current state-of-the-art with three contrasting examples using both simulated and empirical data. The use of simulated data allows the robustness of the methods to be tested with respect to deficiencies in both data and model. These examples show how mechanistic understanding of the processes that determine distribution and abundance can be combined with different sources of information at a range of spatial and temporal scales. Application of such techniques will enable more reliable inference and projections, e.g. under future climate change scenarios than is possible with purely correlative approaches. Conversely, confronting such process-oriented niche models with abundance and distribution data will test current understanding and may ultimately feedback to improve underlying ecological theory.
Aim Biotic interactions within guilds or across trophic levels have widely been ignored in species distribution models (SDMs). This synthesis outlines the development of species interaction distribution models (SIDMs), which aim to incorporate multispecies interactions at large spatial extents using interaction matrices. Location Local to global. Methods We review recent approaches for extending classical SDMs to incorporate biotic interactions, and identify some methodological and conceptual limitations. To illustrate possible directions for conceptual advancement we explore three principal ways of modelling multispecies interactions using interaction matrices: simple qualitative linkages between species, quantitative interaction coefficients reflecting interaction strengths, and interactions mediated by interaction currencies. We explain methodological advancements for static interaction data and multispecies time series, and outline methods to reduce complexity when modelling multispecies interactions. Results Classical SDMs ignore biotic interactions and recent SDM extensions only include the unidirectional influence of one or a few species. However, novel methods using error matrices in multivariate regression models allow interactions between multiple species to be modelled explicitly with spatial co-occurrence data. If time series are available, multivariate versions of population dynamic models can be applied that account for the effects and relative importance of species interactions and environmental drivers. These methods need to be extended by incorporating the non-stationarity in interaction coefficients across space and time, and are challenged by the limited empirical knowledge on spatio-temporal variation in the existence and strength of species interactions. Model complexity may be reduced by: (1) using prior ecological knowledge to set a subset of interaction coefficients to zero, (2) modelling guilds and functional groups rather than individual species, and (3) modelling interaction currencies and species effect and response traits. Main conclusions There is great potential for developing novel approaches that incorporate multispecies interactions into the projection of species distributions and community structure at large spatial extents. Progress can be made by: (1) developing statistical models with interaction matrices for multispecies co-occurrence datasets across large-scale environmental gradients, (2) testing the potential and limitations of methods for complexity reduction, and (3) sampling and monitoring comprehensive spatio-temporal data on biotic interactions in multispecies communities.
The distribution of poikilotherms is determined by the thermal structure of the marine environment that they are exposed to. Recent research has indicated that changes in migration phenology of beluga whales in the Arctic are triggered by changes in the thermal structure of the marine environment in their summering area. If sea temperatures reflect the spatial distribution of food resources, then changes in the thermal regime will affect how homogeneous or clumped food is distributed. We explore, by individual-based modelling, the hypothesis that changes in migration phenology are not necessarily or exclusively triggered by changes in food abundance, but also by changes in the spatial aggregation of food. We found that the level of food aggregation can significantly affect the relationship between the timing of the start of migration to the winter grounds and the total prey capture of individuals. Our approach strongly indicates that changes in the spatial distribution of food resources should be considered for understanding and quantitatively predicting changes in the phenology of animal migration.
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.