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Understanding how variance in environmental factors affects physiological performance, population growth, and persistence is central in ecology. Despite recent interest in the effects of variance in single biological drivers, such as temperature, we have lacked a comprehensive framework for predicting how the variances and covariances between multiple environmental factors will affect physiological rates. Here, we integrate current theory on variance effects with co-limitation theory into a single unified conceptual framework that has general applicability. We show how the framework can be applied (1) to generate mathematically tractable predictions of the physiological effects of multiple fluctuating co-limiting factors, (2) to understand how each co-limiting factor contributes to these effects, and (3) to detect mechanisms such as acclimation or physiological stress when they are at play. We show that the statistical covariance of co-limiting factors, which has not been considered before, can be a strong driver of physiological performance in various ecological contexts. Our framework can provide powerful insights on how the global change-induced shifts in multiple environmental factors affect the physiological performance of organisms.
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 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.
Global change has complex eco-evolutionary consequences for organisms and ecosystems, but related concepts (e.g., novel ecosystems) do not cover their full range. Here we propose an umbrella concept of "ecological novelty" comprising (1) a site-specific and (2) an organism-centered, eco-evolutionary perspective. Under this umbrella, complementary options for studying and communicating effects of global change on organisms, ecosystems, and landscapes can be included in a toolbox. This allows researchers to address ecological novelty from different perspectives, e.g., by defining it based on (a) categorical or continuous measures, (b) reference conditions related to sites or organisms, and (c) types of human activities. We suggest striving for a descriptive, non-normative usage of the term "ecological novelty" in science. Normative evaluations and decisions about conservation policies or management are important, but require additional societal processes and engagement with multiple stakeholders.
This thesis aims to quantify the human impact on the natural resource water at the landscape scale. The drivers in the federal state of Brandenburg (Germany), the area under investigation, are land-use changes induced by policy decisions at European and federal state level. The water resources of the federal state are particularly sensitive to changes in land-use due to low precipitation rates in the summer combined with sandy soils and high evapotranspiration rates. Key elements in landscape hydrology are forests because of their unique capacity to transport water from the soil to the atmosphere. Given these circumstances, decisions made at any level of administration that may have effects on the forest sector in the state are critical in relation to the water cycle. It is therefore essential to evaluate any decision that may change forest area and structure in such a sensitive region. Thus, as a first step, it was necessary to develop and implement a model able to simulate possible interactions and feedbacks between forested surfaces and the hydrological cycle at the landscape scale. The result is a model for simulating the hydrological properties of forest stands based on a robust computation of the temporal and spatial LAI (leaf area index) dynamics. The approach allows the simulation of all relevant hydrological processes with a low parameter demand. It includes the interception of precipitation and transpiration of forest stands with and without groundwater in the rooting zone. The model also considers phenology, biomass allocation, as well as mortality and simple management practices. It has been implemented as a module in the eco-hydrological model SWIM (Soil and Water Integrated Model). This model has been tested in two pre-studies to verify the applicability of its hydrological process description for the hydrological conditions typical for the state. The newly implemented forest module has been tested for Scots Pine (Pinus sylvestris) and in parts for Common Oak (Quercus robur and Q. petraea) in Brandenburg. For Scots Pine the results demonstrate a good simulation of annual biomass increase and LAI in addition to the satisfactory simulation of litter production. A comparison of the simulated and measured data of the May sprout for Scots pine and leaf unfolding for Oak, as well as the evaluation against daily transpiration measurements for Scots Pine, does support the applicability of the approach. The interception of precipitation has also been simulated and compared with weekly observed data for a Scots Pine stand which displays satisfactory results in both the vegetation periods and annual sums. After the development and testing phase, the model is used to analyse the effects of two scenarios. The first scenario is an increase in forest area on abandoned agricultural land that is triggered by a decrease in European agricultural production support. The second one is a shift in species composition from predominant Scots Pine to Common Oak that is based on decisions of the regional forestry authority to support a more natural species composition. The scenario effects are modelled for the federal state of Brandenburg on a 50m grid utilising spatially explicit land-use patterns. The results, for the first scenario, suggest a negative impact of an increase in forest area (9.4% total state area) on the regional water balance, causing an increase in mean long-term annual evapotranspiration of 3.7% at 100% afforestation when compared to no afforestation. The relatively small annual change conceals a much more pronounced seasonal effect of a mean long-term evapotranspiration increase by 25.1% in the spring causing a pronounced reduction in groundwater recharge and runoff. The reduction causes a lag effect that aggravates the scarcity of water resources in the summer. In contrast, in the second scenario, a change in species composition in existing forests (29.2% total state area) from predominantly Scots Pine to Common Oak decreases the long-term annual mean evapotranspiration by 3.4%, accompanied by a much weaker, but apparent, seasonal pattern. Both scenarios exhibit a high spatial heterogeneity because of the distinct natural conditions in the different regions of the state. Areas with groundwater levels near the surface are particularly sensitive to changes in forest area and regions with relatively high proportion of forest respond strongly to the change in species composition. In both cases this regional response is masked by a smaller linear mean effect for the total state area. Two critical sources of uncertainty in the model results have been investigated. The first one originates from the model calibration parameters estimated in the pre-study for lowland regions, such as the federal state. The combined effect of the parameters, when changed within their physical meaningful limits, unveils an overestimation of the mean water balance by 1.6%. However, the distribution has a wide spread with 14.7% for the 90th percentile and -9.9% for the 10th percentile. The second source of uncertainty emerges from the parameterisation of the forest module. The analysis exhibits a standard deviation of 0.6 % over a ten year period in the mean of the simulated evapotranspiration as a result of variance in the key forest parameters. The analysis suggests that the combined uncertainty in the model results is dominated by the uncertainties of calibration parameters. Therefore, the effect of the first scenario might be underestimated because the calculated increase in evapotranspiration is too small. This may lead to an overestimation of the water balance towards runoff and groundwater recharge. The opposite can be assumed for the second scenario in which the decrease in evapotranspiration might be overestimated.
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.
Models are useful tools for understanding and predicting ecological patterns and processes. Under ongoing climate and biodiversity change, they can greatly facilitate decision-making in conservation and restoration and help designing adequate management strategies for an uncertain future. Here, we review the use of spatially explicit models for decision support and to identify key gaps in current modelling in conservation and restoration. Of 650 reviewed publications, 217 publications had a clear management application and were included in our quantitative analyses. Overall, modelling studies were biased towards static models (79%), towards the species and population level (80%) and towards conservation (rather than restoration) applications (71%). Correlative niche models were the most widely used model type. Dynamic models as well as the gene-to-individual level and the community-to-ecosystem level were underrepresented, and explicit cost optimisation approaches were only used in 10% of the studies. We present a new model typology for selecting models for animal conservation and restoration, characterising model types according to organisational levels, biological processes of interest and desired management applications. This typology will help to more closely link models to management goals. Additionally, future efforts need to overcome important challenges related to data integration, model integration and decision-making. We conclude with five key recommendations, suggesting that wider usage of spatially explicit models for decision support can be achieved by 1) developing a toolbox with multiple, easier-to-use methods, 2) improving calibration and validation of dynamic modelling approaches and 3) developing best-practise guidelines for applying these models. Further, more robust decision-making can be achieved by 4) combining multiple modelling approaches to assess uncertainty, and 5) placing models at the core of adaptive management. These efforts must be accompanied by long-term funding for modelling and monitoring, and improved communication between research and practise to ensure optimal conservation and restoration outcomes.
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.
Aim: Across the planet, grass-dominated biomes are experiencing shrub encroachment driven by atmospheric CO2 enrichment and land-use change. By altering resource structure and availability, shrub encroachment may have important impacts on vertebrate communities. We sought to determine the magnitude and variability of these effects across climatic gradients, continents, and taxa, and to learn whether shrub thinning restores the structure of vertebrate communities. Location: Worldwide. Time period: Contemporary. Major taxa studied: Terrestrial vertebrates. Methods: We estimated relationships between percentage shrub cover and the structure of terrestrial vertebrate communities (species richness, Shannon diversity and community abundance) in experimentally thinned and unmanipulated shrub-encroached grass-dominated biomes using systematic review and meta-analyses of 43 studies published from 1978 to 2016. We modelled the effects of continent, biome, mean annual precipitation, net primary productivity and the normalized difference vegetation index (NDVI) on the relationship between shrub cover and vertebrate community structure. Results: Species richness, Shannon diversity and total abundance had no consistent relationship with shrub encroachment and experimental thinning did not reverse encroachment effects on vertebrate communities. However, some effects of shrub encroachment on vertebrate communities differed with net primary productivity, amongst vertebrate groups, and across continents. Encroachment had negative effects on vertebrate diversity at low net primary productivity. Mammalian and herpetofaunal diversity decreased with shrub encroachment. Shrub encroachment also had negative effects on species richness and total abundance in Africa but positive effects in North America. Main conclusions: Biodiversity conservation and mitigation efforts responding to shrub encroachment should focus on low-productivity locations, on mammals and herpetofauna, and in Africa. However, targeted research in neglected regions such as central Asia and India will be needed to fill important gaps in our knowledge of shrub encroachment effects on vertebrates. Additionally, our findings provide an impetus for determining the mechanisms associated with changes in vertebrate diversity and abundance in shrub-encroached grass-dominated biomes.
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.