Covariations in ecological scaling laws fostered by community dynamics

Scaling laws in ecology, intended both as functional relationships among ecologically relevant quantities and the probability distributions that characterize their occurrence, have long attracted the interest of empiricists and theoreticians. Empirical evidence exists of power laws associated with the number of species inhabiting an ecosystem, their abundances, and traits. Although their functional form appears to be ubiquitous, empirical scaling exponents vary with ecosystem type and resource supply rate. The idea that ecological scaling laws are linked has been entertained before, but the full extent of macroecological pattern covariations, the role of the constraints imposed by finite resource supply, and a comprehensive empirical verification are still unexplored. Here, we propose a theoretical scaling framework that predicts the linkages of several macroecological patterns related to species’ abundances and body sizes. We show that such a framework is consistent with the stationary-state statistics of a broad class of resource-limited community dynamics models, regardless of parameterization and model assumptions. We verify predicted theoretical covariations by contrasting empirical data and provide testable hypotheses for yet unexplored patterns. We thus place the observed variability of ecological scaling exponents into a coherent statistical framework where patterns in ecology embed constrained fluctuations.

Scaling as a key conceptual tool for the interpretation of empirical and experimental patterns in ecology and biology

Silvia Zaoli

This thesis is concerned with regular patterns found in ecology and biology, their linkages and the statistical description of their fluctuations around average trends. Among such patterns, often conforming to power laws, well-known examples include the Species-Area relationship (SAR), quantifying the increase of the number of species S inhabiting an ecosystem with ecosystem area, and the scale-invariant body size spectrum, routinely observed, e.g., in aquatic ecosystems. In biology, Kleiber's law is an allometric relationship describing how metabolic rates scale with an organism's body size. While ecological laws have often been studied independently, simple heuristic reasonings show that they are linked. The need for a unifying effort in ecology, coherently synthesizing the vast and diverse set of empirical observations across scales, has been often voiced. However, a theoretical framework answering this need was still lacking. Furthermore, ecological variables are the result of the interplay between several stochastic ecological processes, and are therefore stochastic variables fluctuating around average values. Ecological and biological scaling laws typically make predictions for such averages, but the issue of fluctuations received scarce attention in the literature. Similarly, biological fluctuations have been typically neglected in the study of the size-scaling of metabolic rates, even though body sizes and metabolic rates may have a significant variability within a species. Fluctuations may be relevant to interpret empirical observations, judge the reliability of predictions and understand ecosystem dynamics. An hypothesis for the distribution of abundances and body sizes of species inhabiting an ecosystem of finite area is proposed here. The hypothesis is inspired by finite-size scaling and is used to derive macroecological patterns and their linkages within a coherent theoretical framework. Stochastic models of community dynamics are used to support the hypothesis, and the derived linkages are tested on empirical datasets. Several stochastic models of community dynamics are also used here to study the fluctuations of S and how they scale with the average S. The intra-specific variability of metabolic rates and body sizes is investigated experimentally using freshwater phytoplankton species by nanoscale secondary ion mass spectrometry (NanoSIMS). The linkages among ecological scaling laws predicted by the theoretical framework are verified in several empirical datasets. Theoretical generalizations including deviations from pure power-law behavior and heavy-tailed intra-specific size distributions are also addressed. The theoretical study of the relative scaling of the fluctuations of S with the mean

\langle S \rangle

in various community dynamics models shows that different ecological processes predict radically different fluctuations scalings, highlighting the need of empirical investigations to sort out which scenario applies to real ecosystems. Experiments on phytoplankton metabolic rate scaling with body size suggest that intra-specific metabolic rate distributions are described by a universal scaling form across different taxa and over three orders of magnitude in body size. This thesis, along with previous works, suggests that scaling concepts derived for inanimate systems can provide new insights into the dynamics of ecosystems and help unraveling regularities across scales of biological complexity.

EPFL2018

Ecohydrological Footprints

Samir Suweis

Ecohydrological footprints are defined as the response of ecosystem functions or services to changes in their hydrologic drivers. In this thesis, several diverse footprints are addressed: noise-driven effects on storage-discharge relations and catchment streamflow distributions, that are important drivers of biodiversity; soil salinization and its ecohydrological implications; topological effects of the ecological interaction networks on living communities (e.g. on their species persistence); and form and function of the global virtual water trade network. The coherence of the conceptual framework is provided by the study of drivers and controls of ecohydrological variability using methodological approaches based on statistical mechanics. In fact, this thesis work outlines a significant portion of environmental statistical mechanics, an overarching discipline that is emerging in recent years, which applies mathematical tools from statistical mechanics to model several ecohydrological processes. The proposed relevance of this thesis lies in the major effects of hydrologic drivers on ecological process. The view that emerges from current research in ecohydrology, that this thesis supports, is that there exists a definite need for an integrated understanding of ecological and hydrological processes. Because stochasticity is intrinsic to environmental and ecohydrological variability, noise plays an important and constructive role in ecohydrological processes. In this thesis, a stochastic approach is applied to analyze different ecohydrological processes, ranging from green and blue water flows in river basins (part I), ecosystem dynamics affected by the directional dispersal provided by river networks (part II) to water footprints of human society (part III).Methods range from novel exact solutions to stochastic differential equations to random graph theory applications, and imply the analysis of suitable field data. An analytical framework for quantitative analysis is laid out to tackle complex problems and to estimate the effects of environmental change on the interaction of the hydrologic processes with the biota. The main results of this thesis are: i) the achievement of exact solutions for the probability distribution of catchment streamflow, that takes in account stochastic fluctuations in the storage-discharge relation and for the condition of a noise induced phenomena to the streamflows regimes; ii) the stationary solutions of soil salinity under stochastic hydrologic forcing; iii) a novel solution of the Ito-Stratonovich problem in multiplicative Poisson processes; iv) the proper framework for species' persistence time distributions, as a function of topological constraints on the ecosystem, and its connection with other important macroecological laws. A related length-bias sampling problem is also solved. v) A statistical analysis of the global virtual trade network and a semi-analytical model that is able to describe most of the observed properties.

EPFL2011

Resistance of plant communities to invasive species

Aurélie Thébault

Invasive species are nowadays considered as one of the most important threat to biodiversity. By displacing native species, modifying ecosystem functioning and causing substantial losses to agricultural production, they represent a menace to natural and managed ecosystems. Although ecology of invasions has become an important research topic since the last decades, the mechanisms that determine why a given species may invade a given ecosystem and why some biomes are less resistant to invasion are still not clarified. Ecology of invasions is divided into 2 main topics: invasiveness and invasibility. While invasiveness refers to species ability to invade a community, invasibility focuses on the resistance of a community to invasion. Invasiveness may be the result of ecological processes, such as release from biotic constraints or human alteration of the environment (disturbance, stress…) or the consequence of evolutionary processes, such as hybridization or polyploidization that may increase genetic variation and therefore, enhance niche breadth. Invasibility has been said to be influenced by disturbance and biotic factors such as community diversity, dominant species identity, biotic interactions or community compositional stability. The invasion success is the consequence of the interaction between species invasiveness and community invasibility. Most studies in ecological invasions have focused on either invasiveness or invasibility, but hardly both together. By working at the same time and in the same conditions with native and introduced genotypes and by comparing their ecological performances, this thesis aims at a better understanding of both invasiveness and invasibility mechanisms. Two worldwide invasive species, Centaurea maculosa and Senecio inaequidens, were used in several experiments (pot, microcosm, field) to disentangle the importance of invasiveness and community invasibility in their invasion success. Both species encountered polyploidisation in their native range, leading to the presence of diploid and tetraploid populations, whereas only tetraploid populations have been found in the introduced range. Using native diploid, native tetraploid and introduced tetraploid genotypes of the two model-species, allows assessing the effects of genetic variation (diploid vs. tetraploid genotypes) and environmental variation (genotype from native vs. introduced range) on species phenotypic traits variations and consequently on fitness variation and invasiveness. In the community context, studying response of different genotypes to experimental factors and community change gives information on the interaction between invasiveness and invasibility. Plants were grown in pot, in field or in artificially built communities where (i) the management treatment, (ii) the community diversity and (iii) the spatial organisation of resident species were manipulated. In addition, (iv) community species composition, (v) community competitive ability and (vi) compositional and functional stability of the community were monitored. According to the experiments, data were gathered on survival, morphological traits (vegetative height, lateral spread, shoot and root biomass), leaf traits (specific leaf area, leaf dry matter content) and reproductive traits (probability of flowering, capitulum production) of the genotypes (native diploid, native tetraploid and introduced tetraploid) of both model species. Through the use of statistical models and multivariate approaches, the effects of management and biotic factors on survival, growth and reproduction of native and introduced genotypes of these two worldwide invasive species were assessed. Invasion strategy of the two model species was investigated through a growth experiment in optimal conditions in a pot experiment. High investment in seed production could explain invasive success of S. inaequidens through high propagule pressure, whereas C. maculosa's strategy seemed to be oriented towards interactions with belowground communities, as shown by the shift in rhizosphere bacterial communities between genotypes. For both species, polyploidisation in the native range could be linked to a specialisation towards higher competitive ability, which could have allowed the first step of invasion. Introduction in the new range could be related to a loss of specialisation through selection of traits allowing coping with various or changing environments, improving successful spread. Survival of both model species was highly affected by community spatial pattern, management and neighbouring competition. Responses of growth and reproductive output of native and introduced genotypes to management, community spatial pattern and community diversity were species-specific. Growth and reproductive output of both genotypes of S. inaequidens were affected by experimental factors whereas introduced genotypes of C. maculosa were less affected than native ones. Comparison of response of native and introduced genotypes to experimental factors allowed defining two strategies of invasion based on phenotypic plasticity. Centaurea maculosa was able to maintain fitness in all kinds of environments, either favourable or stressful ("Jack-of-all-trades" invader). Senecio inaequidens was able to deal with all kinds of environments and was also able to increase its fitness in favourable conditions ("Jack-and-Master" invader). The combination of the effects of management, community spatial pattern and community diversity on species genotypes allowed defining, for both invasive species, two invasion phases which were impacted by different factors. The introduction phase corresponds to the survival of seedlings and their ability to deal with neighbouring competition. If seedlings manage to survive despite neighbouring competition, they grow and reproduced in order to spread, which corresponds to the establishment phase. In terms of management perspectives, regular mowing or use of highly covering species could limit invasive success of S. inaequidens and C. maculosa respectively. The synthesis of all the experiments conducted in this thesis with C. maculosa and S. inaequidens highlights (1) the importance of polyploidisation in the invasion process as well as (2) the species-specific invasion strategies and consequently (3) the species-specific response of invasive species to abiotic and biotic factors. It also emphasizes on (4) the temporal evolution of the interaction between invasiveness and invasibility since the community factors that affected invasive species fitness changed according to the invasion stage (introduction vs. establishment phase) of the invader. The provided insights into the importance of the interaction between species invasiveness and community invasibility will contribute to improve knowledge in ecology of invasions, in addition to supply some clues for management efficiency.

EPFL2009