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Mathematical models of species dispersal and the resilience of metapopulations against habitat loss
(2021)
Habitat loss and fragmentation due to climate and land-use change are among the biggest threats to biodiversity, as the survival of species relies on suitable habitat area and the possibility to disperse between different patches of habitat. To predict and mitigate the effects of habitat loss, a better understanding of species dispersal is needed. Graph theory provides powerful tools to model metapopulations in changing landscapes with the help of habitat networks, where nodes represent habitat patches and links indicate the possible dispersal pathways between patches.
This thesis adapts tools from graph theory and optimisation to study species dispersal on habitat networks as well as the structure of habitat networks and the effects of habitat loss. In chapter 1, I will give an introduction to the thesis and the different topics presented in this thesis. Chapter 2 will then give a brief summary of tools used in the thesis.
In chapter 3, I present our model on possible range shifts for a generic species. Based on a graph-based dispersal model for a generic aquatic invertebrate with a terrestrial life stage, we developed an optimisation model that models dispersal directed to predefined habitat patches and yields a minimum time until these patches are colonised with respect to the given landscape structure and species dispersal capabilities. We created a time-expanded network based on the original habitat network and solved a mixed integer program to obtain the minimum colonisation time. The results provide maximum possible range shifts, and can be used to estimate how fast newly formed habitat patches can be colonised. Although being specific for this simulation model, the general idea of deriving a surrogate can in principle be adapted to other simulation models.
Next, in chapter 4, I present our model to evaluate the robustness of metapopulations. Based on a variety of habitat networks and different generic species characterised by their dispersal traits and habitat demands, we modeled the permanent loss of habitat patches and subsequent metapopulation dynamics. The results show that species with short dispersal ranges and high local-extinction risks are particularly vulnerable to the loss of habitat across all types of networks. On this basis, we then investigated how well different graph-theoretic metrics of habitat networks can serve as indicators of metapopulation robustness against habitat loss. We identified the clustering coefficient of a network as the only good proxy for metapopulation robustness across all types of species, networks, and habitat loss scenarios.
Finally, in chapter 5, I utilise the results obtained in chapter 4 to identify the areas in a network that should be improved in terms of restoration to maximise the metapopulation robustness under limited resources. More specifically, we exploit our findings that a network’s clustering coefficient is a good indicator for metapopulation robustness and develop two heuristics, a Greedy algorithm and a deducted Lazy Greedy algorithm, that aim at maximising the clustering coefficient of a network. Both algorithms can be applied to any network and are not specific to habitat networks only.
In chapter 6, I will summarize the main findings of this thesis, discuss their limitations and give an outlook of future research topics.
Overall this thesis develops frameworks to study the behaviour of habitat networks and introduces mathematical tools to ecology and thus narrows the gap between mathematics and ecology. While all models in this thesis were developed with a focus on aquatic invertebrates, they can easily be adapted to other metapopulations.
World’s ecosystems are under great pressure satisfying anthropogenic demands, with freshwaters being of central importance. The Millennium Ecosystem Assessment has identified anthropogenic land use and associated stressors as main drivers in jeopardizing stream ecosystem functions and the
biodiversity supported by freshwaters. Adverse effects on the biodiversity of freshwater organisms, such as macroinvertebrates, may propagate to fundamental ecosystem functions, such as organic matter breakdown (OMB) with potentially severe consequences for ecosystem services. In order to adequately protect and preserve freshwater ecosystems, investigations regarding potential and observed as well as direct and indirect effects of anthropogenic land use and associated stressors (e.g. nutrients, pesticides or heavy metals) on ecosystem functioning and stream biodiversity are needed. While greater species diversity most likely benefits ecosystem functions, the direction and magnitude of changes in ecosystem functioning depends primarily on species functional traits. In this context, the functional diversity of stream organisms has been suggested to be a more suitable predictor of changes in ecosystem functions than taxonomic diversity.
The thesis aims at investigating effects of anthropogenic land use on (i) three ecosystem functions by anthropogenic toxicants to identify effect thresholds (chapter 2), (ii) the organic matter breakdown by three land use categories to identify effects on the functional level (chapter 3) and (iii)on the stream community along an established land-use gradient to identify effects on the community level.
In chapter 2, I reviewed the literature regarding pesticide and heavy metal effects on OMB, primary production and community respiration. From each reviewed study that met inclusion criteria, the toxicant concentration resulting in a reduction of at least 20% in an ecosystem function was standardized based on laboratory toxicity data. Effect thresholds were based on the relationship between ecosystem functions and standardized concentration-effect relationships. The analysis revealed that more than one third of pesticide observations indicated reductions in ecosystem functions at concentrations that are assumed being protective in regulation. However, high variation within and between studies hampered the derivation of a concentration-effect relationship and thus effect thresholds.
In chapter 3, I conducted a field study to determine the microbial and invertebrate-mediated OMB by deploying fine and coarse mesh leaf bags in streams with forested, agricultural, vinicultural
and urban riparian land use. Additionally, physicochemical, geographical and habitat parameters were monitored to explain potential differences in OMB among land use types and sites. Regarding results, only microbial OMB differed between land use types. The microbial OMB showed a negative relationship with pH while the invertebrate-mediated OMB was positively related to tree cover. OMB responded to stressor gradients rather than directly to land use.
In chapter 4, macroinvertebrates were sampled in concert with leaf bag deployment and after species identification (i) the taxonomic diversity in terms of Simpson diversity and total taxonomic
richness (TTR) and (ii) the functional diversity in terms of bio-ecological traits and Rao’s quadratic entropy was determined for each community. Additionally, a land-use gradient was established and the response of the taxonomic and functional diversity of invertebrate communities along this gradient was investigated to examine whether these two metrics of biodiversity are predictive for the rate of OMB. Neither bio-ecological traits nor the functional diversity showed a significant relationship with
OMB. Although, TTR decreased with increasing anthropogenic stress and also the community structure and 26 % of bio-ecological traits were significantly related to the stress gradient, any of these shifts propagated to OMB.
Our results show that the complexity of real-world situations in freshwater ecosystems impedes the effect assessment of chemicals and land use for functional endpoints, and consequently our potential to predict changes. We conclude that current safety factors used in chemical risk assessment may not be sufficient for pesticides to protect functional endpoints. Furthermore, simplifying real-world stressor gradients into few land use categories was unsuitable to predict and quantify losses in OMB. Thus, the monitoring of specific stressors may be more relevant than crude land use categories to detect effects on ecosystem functions. This may, however, limit the large scale assessment of the status of OMB. Finally, despite several functional changes in the communities the functional diversity over several trait modalities remained similar. Neither taxonomic nor functional diversity were suitable predictors of OMB. Thus, when understanding anthropogenic impacts on the linkage between biodiversity and ecosystem functioning is of main interest, focusing on diversity metrics that are clearly linked to the stressor in question (Jackson et al. 2016) or integrating taxonomic and functional metrics (Mondy et al., 2012) might enhance our predictive capacity.
Change of ecosystems and the associated loss of biodiversity is among the most important environmental issues. Climate change, pollution, and impoundments are considered as major drivers of biodiversity loss. Organism traits are an appealing tool for the assessment of these three stressors, due to their ability to provide mechanistic links between organism responses and stressors, and consistency over wide geographical areas.
Additionally, traits such as feeding habits influence organismal performance and ecosystem processes. Although the response of traits of specific taxonomic groups to stressors is known, little is known about the response of traits of different taxonomic groups to stressors. Additionally, little is known about the effects of small impoundments on stream ecosystem processes, such as leaf litter decomposition, and food webs.
After briefly introducing the theoretical background and objectives of the studies, this thesis begins by synthesizing the responses of traits of different taxonomic groups to climate change and pollution. Based on 558 peer-reviewed studies, the uniformity (i.e., convergence) in trait response across taxonomic groups was evaluated through meta-analysis (Chapter 2). Convergence was primarily limited to traits related to tolerance.
In Chapter 3, the hypothesis that small impoundments would modify leaf litter decomposition rates at the sites located within the vicinity of impoundments, by altering habitat variables and invertebrate functional feeding groups (FFGs) (i.e., shredders), was tested. Leaf litter decomposition rates were significantly reduced at the study sites located immediately upstream (IU) of impoundments, and were significantly related to the abundance of invertebrate shredders.
In Chapter 4, the invertebrate FFGs were used to evaluate the effect of small impoundments on stream ecosystem attributes. The results showed that heterotrophic production was significantly reduced at the sites IU. With regard to food webs, the contribution of methane gas derived carbon to the biomass of chironomid larvae was evaluated through correlation of stable carbon isotope values of chironomid larvae and methane gas concentrations.
The results indicated that the contribution of methane gas derived carbon into stream benthic food web is low. In conclusion, traits are a useful tool in detecting ecological responses to stressors across taxonomic groups, and the effects of small impoundments on stream ecological integrity and food web are limited.
The adoption of the EU Water Framework Directive (WFD) in 2000 marked the beginning of a new era of European water policy. However, more than a decade later, the majority of European rivers are still failing to meet one of the main objectives of the WFD: the good ecological status. Pesticides are a major stressor for stream ecosystems. This PhD thesis emphasises the need for WFD managers to consider all main agricultural pesticide sources and influencing landscape parameters when setting up River Basin Management Plans and Programmes of Measures. The findings and recommendations of this thesis can help to successfully tackle the risk of pesticide contamination to achieve the WFD objectives.
A total of 663 sites that were situated in the German Federal States of Saxony, Saxony-Anhalt, Thuringia and Hesse were studied (Chapter 3 and 4). In addition to an analysis of the macroinvertebrate data of the governmental WFD monitoring network, a detailed GIS analysis of the main agricultural pesticide sources (arable land and garden allotments as well as wastewater treatment plants (WWTPs)) and landscape elements (riparian buffer strips and forested upstream reaches) was conducted. Based on the results, a screening approach was developed that allows an initial rapid and cost-effective identification of those sites that are potentially affected by pesticide contamination. By using the trait-based bioindicator SPEARpesticides, the insecticidal long-term effects of the WWTP effluents on the structure of the macroinvertebrate community were identified up to at least 1.5 km downstream (in some cases even 3 km) of the WWTPs. The results of the German Saprobic Index revealed that the WWTPs can still be important sources of oxygen-depleting substances. Furthermore, the results indicate that forested upstream reaches and riparian buffer strips at least 5 m in width can be appropriate measures in mitigating the effects and exposure of pesticides.
There are concerns that the future expansion of energy crop cultivation will lead to an increased pesticide contamination of ecosystems in agricultural landscapes. Therefore, the potential of energy crops for pesticide contamination was examined based on an analysis of the development of energy crop cultivation in Germany and a literature search on perennial energy crops (Chapter 5). The results indicate that the future large-scale expansion of energy crop cultivation will not necessarily cause an increase or decrease in the amounts of pesticides that are released into the environment. The potential effects will depend on the future design of the agricultural systems. Instead of creating energy monocultures, annual energy crops should be integrated into the existing food production systems. Financial incentives and further education are needed to encourage the use of sustainable crop rotations, innovative cropping systems and perennial energy crops, which may contribute to crop diversity and generate lower pesticide demands than do intensive farming systems.
The aquatic environment is exposed to multiple environmental pressures and mixtures of chemical substances, among them petroleum and petrochemicals, metals, and pesticides. Aquatic invertebrate communities are used as bioindicators to reflect long-term and integral effects. Information on the presence of species can be supplemented with information on their traits. SPEAR-type bioindicators integrate such trait information on the community level.
This thesis aimed at enhancing specificity of SPEAR-type bioindicators towards particular groups of chemicals, namely to mixtures of oil sands-derived compounds, hydrocarbons, and metals.
For developing a bioindicator for discontinuous contamination with oil-derived organic toxicants, a field study was conducted in the Canadian oil sands development region in Northern Alberta. The traits ‘physiological sensitivity towards organic chemicals’ and ‘generation time’ were integrated to develop the bioindicator SPEARoil, reflecting the community sensitivity towards oil sands derived contamination in relation to fluctuating hydrological conditions.
According to the SPEARorganic approach, a physiological sensitivity ranking of taxa was developed for hydrocarbon contamination originating from crude oil or petroleum distillates. For this purpose, ecotoxicological information from acute laboratory tests was enriched with rapid and mesocosm test results. The developed Shydrocarbons sensitivity values can be used in SPEAR-type bioindicators.
To specifically reflect metal contamination in streams via bioindicators, Australian field studies were re-evaluated with focus on the traits ‘physiological metal sensitivity’ and ‘feeding type’. Metal sensitivity values, however, explained community effects in the field only weakly. Instead, the trait ‘feeding type’ was strongly related to metal exposure. The fraction of predators in a community can, thus, serve as an indicator for metal contamination in the field.
Furthermore, several metrics reflecting exposure to chemical cocktails in the environment were compared using existing pesticide datasets. Exposure metrics based on the 5% fraction of species sensitivity distributions were found to perform best, however, closely followed by Toxic Unit metrics based on the most sensitive species of a community or Daphnia magna.