Hydrologic Drivers and Controls of Stream Ecological Processes

The temporal variability of streamflows is a key feature structuring and controlling ecological communities and ecosystem processes. The magnitude, frequency and predictability of streamflows, and thus of velocity and near-bed shear stress fields, control ecological structure and function, particularly of benthic organisms. River ecosystem dynamics is indeed deeply connected to streamflow variability, which chiefly relies on rainfall, climate, land use, and geomorphologic properties. Although alterations of streamflow regime due to climate change, habitat fragmentation or other anthropogenic factors are ubiquitous, their ecological implications remain poorly understood. The present Thesis therefore addresses a quantitative analysis of the implications of hydrological fluctuations on river ecosystems positing that they have major ecological importance. From a food-web perspective, the response of benthic biota to hydrologic change is analyzed through experimental and theoretical approaches. Starting from the theoretical characterization of the probability distribution function of streamflows, flow velocities, water depths and near-bed shear stresses at a site, where a comparison between measured and analytical streamflow statistics across several catchments is outlined, a flume experimental campaign has been carried out in order to analyze how flow variability affects biofilm growth and invertebrate grazing activity. Here, two contrasting flow regimes (a constant and a time-varying discharge sequence) and four different light conditions (from 90% to 27% of incoming light radiation), as a key control on biofilm algal productivity and grazer activity, have been performed. Average grazing rates were significantly enhanced under time-varying flow conditions and highest at intermediate light availability. This result suggests that the stochastic flow regime offers increased opportunities for grazing under more favorable shear stress conditions, with implications for stream ecology and trophic carbon transfer in stream food webs. A spatial generalization of the above results, upscaling them to whole river basins, is made possible by employing scaling relationships that are known to characterize the geometry and topology of natural landforms and riverine patterns. The proposed generalization accounts for hydrological characterizations of the driving streamflow variability where the explicit dependence of invertebrate suitability on geomorphic controls (e.g. river depth, bed shear stress and flow velocity) and on hydrologic conditions (e.g. embedded in the probability of streamflow, and the ensuing temporal correlations that define the persistence of the hydrologic signal in time) has been accounted for. Different invertebrate grazing species have been compared. Suitable dynamic models, describing benthic biota dynamics, in particular stream biofilm growth under both ungrazed and grazed conditions (that is, via the absence and presence of invertebrates, respectively), have been explored in order to assess the relevant processes that controlled biofilm-invertebrate temporal pattern. The present Thesis therefore aims to further our understanding of the linkages between hydrology and ecology, possibly leading to a comprehensive theory of hydrologic drivers and controls of biotic processes in stream benthic environments.

Theoretical and Experimental Studies of Dendritic Metacommunities

Francesco Carrara

The present thesis deals with the understanding of the origins and the mechanisms of maintenance of biodiversity in natural landscapes, in particular by identifying key processes that define large-scale patterns of abundance and diversity. Biological communities often occur in spatially structured habitats where connectivity directly affects dispersal and metacommunity processes. Recent theoretical work suggests that dispersal constrained by the connectivity of specific habitat structures affects diversity patterns and species interactions. This is particularly relevant in dendritic networks epitomized by fluvial ecological corridors. This thesis addresses whether connectivity alone can explain observed features of biodiversity and selectively promote different components of community composition in river-like landscapes, such as local species richness or the among-community similarity. The relevance of this thesis lies in the major ecological challenges posed by the topic, and its fundamental importance to conservation biology. The studies pursued herein are also deemed relevant because of the influence of the spatial connectivity and dispersal on population dynamics and of the relevance of biodiversity to ecosystem functioning. Mechanisms of species coexistence were investigated with a blend of theoretical tools (broadly related to statistical mechanics and the theory of stochastic processes) and experimental work using laboratory microbial communities. The research tools ranged from aspects of modern coexistence theory in a local perspective to the recent concept of the metacommunity in spatial ecology, within a unified framework. The study of biodiversity in riverine ecosystems guided by observational data has been addressed by combining theoretical metacommunity models with laboratory experiments. The results are diverse. First, they show experimentally that connectivity per se shapes key components of biodiversity in metacommunities. Local dispersal in isotropic lattice landscapes homogenizes local species richness and leads to pronounced spatial persistence. By contrast, dispersal along dendritic landscapes leads to higher variability in local diversity and among-community composition. Although headwaters exhibit relatively lower species richness, they are crucial for the maintenance of regional biodiversity. By suitably arranging patch sizes within river-like networks the effect of local habitat capacity (i.e., the patch size) and dendritic connectivity on biodiversity can be experimentally disentangled in aquatic microcosm metacommunities. It is shown in this thesis that species coexistence and community assembly depend on intricate, non-trivial interactions of local community capacity and network positioning. Furthermore, an interaction of spatial arrangement of habitat capacity and dispersal along river-like networks also affects a key ecosystem descriptor, namely regional evenness. High regional evenness in community composition is found only in landscapes preserving geomorphological scaling properties of patch sizes. In riverine environments some of the rarer species sustained regionally more abundant populations and were better able to track their own niche requirements compared to landscapes with homogeneous patch size or landscapes with spatially uncorrelated patch size. All the experimental results were supported and extended by a theoretical analysis where the above mechanisms have been generalized. This thesis provides the first direct experimental evidence that spatially constrained dendritic connectivity is a key factor for community composition and population persistence in riverine landscapes. As such, this thesis assesses a longstanding issue in spatial community ecology. It offers unique insights into the ecological forces structuring natural communities in a key ecosytem, and demonstrates principles that can be further tested in theoretical metacommunity models possibly to be extended to real riverine ecosystems. Taken together, the analyses show how the structure of ecological networks interacts with the spatial environmental matrix in determining biodiversity patterns and the functioning of biological communities. The analyses also suggest that altering the natural linkage between dendritic connectivity and patch size strongly affects community properties at multiple scales. The first part of this thesis (chapters 2 and 3) addresses key aspects of biodiversity-ecosystem functioning research where the combination of theory-guided experiments and theoretical investigations shows how a stochastic implementation of population dynamics proves fundamental for key community properties such as species persistence and community stability. The diversity-productivity and diversity-stability relationships are explored. Both experimental findings and the results of a stochastic model fitted to the experimental interaction matrix, suggest the emergence of strong stabilizing forces when species from different functional groups interact in the same environment, increasing species coexistence and community biomass production. The last part (chapter 6) provides a synthesis of this thesis work, in that it aims at unifying aspects from niche-theory, usually adopted in spatially implicit models, with those characteristic of a spatially explicit context from a typical real-life mountainous regions. It is dedicated to the possible explanation for a macroecological pattern routinely observed from organisms in different domains of life, that is, the mid-elevational peak in local species richness. Guided by empirical observations on diversity of macroinvertebrates in Swiss river basins, a theoretical ansatz is provided which is deemed to capture the essential geomorphological drivers and controls relating species-fitness to altitude. A set of overarching conclusions and perspectives for future research are discussed in the concluding chapter.

EPFL2013

Glacier shrinkage will accelerate downstream decomposition of organic matter and alters microbiome structure and function

Tom Ian Battin, Massimo Bourquin, Jade Brandani, Susheel Bhanu Busi, Vincent Henri De Staercke, Nicola Deluigi, Leïla Ezzat, Stylianos Fodelianakis, Tyler Joe Kohler, Grégoire Marie Octave Edouard Michoud, Hannes Markus Peter, Paraskevi Pramateftaki, Martina Andrea Schön, Michail Styllas, Matteo Tolosano

The shrinking of glaciers is among the most iconic consequences of climate change. Despite this, the downstream consequences for ecosystem processes and related microbiome structure and function remain poorly understood. Here, using a space-for-time substitution approach across 101 glacier-fed streams (GFSs) from six major regions worldwide, we investigated how glacier shrinkage is likely to impact the organic matter (OM) decomposition rates of benthic biofilms. To do this, we measured the activities of five common extracellular enzymes and estimated decomposition rates by using enzyme allocation equations based on stoichiometry. We found decomposition rates to average 0.0129 (% d−1), and that decreases in glacier influence (estimated by percent glacier catchment coverage, turbidity, and a glacier index) accelerates decomposition rates. To explore mechanisms behind these relationships, we further compared decomposition rates with biofilm and stream water characteristics. We found that chlorophyll-a, temperature, and stream water N:P together explained 61% of the variability in decomposition. Algal biomass, which is also increasing with glacier shrinkage, showed a particularly strong relationship with decomposition, likely indicating their importance in contributing labile organic compounds to these carbon-poor habitats. We also found high relative abundances of chytrid fungi in GFS sediments, which putatively parasitize these algae, promoting decomposition through a fungal shunt. Exploring the biofilm microbiome, we then sought to identify bacterial phylogenetic clades significantly associated with decomposition, and found numerous positively (e.g., Saprospiraceae) and negatively (e.g., Nitrospira) related clades. Lastly, using metagenomics, we found evidence of different bacterial classes possessing different proportions of EEA-encoding genes, potentially informing some of the microbial associations with decomposition rates. Our results, therefore, present new mechanistic insights into OM decomposition in GFSs by demonstrating that an algal-based “green food web” is likely to increase in importance in the future and will promote important biogeochemical shifts in these streams as glaciers vanish.

Wiley2022

Improved framework to estimate travel time and derived distributions in hydrological control volumes

Mitra Asadollahi

Crossing properties of soil saturation, defined as the duration and excursion of soil saturation below and above certain thresholds, are key variables to ecosystem functioning and evolution by primarily influencing the plant and soil microbes physiological dynamics. Under climate change, the variability in the precipitation patterns is expected to be more ubiquitous and directly translate to a shift in soil saturation levels. This, in turn, will influence crossing properties of soil saturation and thus be expected to leave a mark on ecosystems. However, the lack of a deep understanding of the links between biological processes and the change in soil saturation patterns hinders foretelling the impact of climate variability on ecohydrological systems even at the smallest scale, that is, the level of a single plant.To that end, this thesis presents experimental results from laboratory soil column experiments and extensive computational studies on the interactions between soil saturation and biological activities, that is, plants and microbes. To this aim, the depth-time soil saturation patterns are mapped by means of modeling the transport processes of relevance and experimental tools at a minimal hydrological control volume in search for upscaling methods. In this context, two modeling approaches can be employed. Physically based models that explicitly represent transport processes with high costs of application at larger scales. Age-based models that rely on generic functions with a focus on time past since arrival, that is, the age of event water in the storage. This thesis work combines models (HYDRUS-1D as physically based and tran-SAS as age-based) with soil column experimental data and Bayesian inference methods to explore the links between age characteristics and transport processes. In this thesis, data from four different experimental designs on monthly to yearly timescale under varied environmental conditions are examined in terms of tracer type, precipitation pattern, vegetation, soil type, resolution and type of collected data. Of the modeled data, two sets of results from specific experiments were carried out within the framework of this thesis.This thesis addresses also the role of the balance between dispersive and convective forces, known as Péclet number, in controlling the shape of age functions in drainage. While sole reliance on drainage flow and tracer data (reflective of Péclet number) suffice to capture age dynamics in drainage, un- certainties may arise for predictions on (evapo)transpiration. This thesis deals also with the role of root distributions as controls in determining the age to root uptake and shows plants with similar uniform-equivalent root distribution render very similar age dynamics. It was shown in this thesis that nitrate consumption by microbes can be linked to age characteristics of a conservative tracer and that the relation between microbial respiration and soil saturation may bear more complexity than is currently integrated in most models. Overall, this thesis lays steps to the development of modeling tools with high predictive power for soil saturation by originating upscaling methods and bridging the gaps between physically based and age-based models with a view on the consequences of (possi- bly changing) variable soil saturation on adaptations of microbial communities and their metabolic product.

EPFL2022