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Brose, U.

Publications and source records attributed to Brose, U..

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The biggest losers: Habitat isolation deconstructs complex food webs from top to bottom

Habitat fragmentation is threatening global biodiversity. To date, there is only limited understanding of how habitat fragmentation or any alteration to the spatial structure of a landscape in general, affects species diversity within complex ecological networks such as food webs. Here, we present a dynamic and spatially-explicit food web model which integrates complex food web dynamics at the local scale and species-specific dispersal dynamics at the landscape scale, allowing us to study the interplay of local and spatial processes in metacommunities. We explore how habitat fragmentation, defined as a decrease of habitat availability and an increase of habitat isolation, affects the species diversity patterns of complex food webs (-, {beta}-,{gamma} -diversity), and specifically test whether there is a trophic dependency in the effect of habitat fragmentation on species diversity. In our model, habitat isolation is the main driver causing species loss and diversity decline. Our results emphasise that large-bodied consumer species at high trophic positions go extinct faster than smaller species at lower trophic levels, despite being superior dispersers that connect fragmented landscapes better. We attribute the loss of top species to a combined effect of higher biomass loss during dispersal with increasing habitat isolation in general, and the associated energy limitation in highly fragmented landscapes, preventing higher trophic levels to persist. To maintain trophic-complex and species-rich communities calls for effective conservation planning which considers the interdependence of trophic and spatial dynamics as well as the spatial context of a landscape and its energy availability.

ecology

The intrinsic predictability of ecological time series and its potential to guide forecasting

Successfully predicting the future states of systems that are complex, stochastic and potentially chaotic is a major challenge. Model forecasting error (FE) is the usual measure of success; however model predictions provide no insights into the potential for improvement. In short, the realized predictability of a specific model is uninformative about whether the system is inherently predictable or whether the chosen model is a poor match for the system and our observations thereof. Ideally, model proficiency would be judged with respect to the systems intrinsic predictability - the highest achievable predictability given the degree to which system dynamics are the result of deterministic v. stochastic processes. Intrinsic predictability may be quantified with permutation entropy (PE), a model-free, information-theoretic measure of the complexity of a time series. By means of simulations we show that a correlation exists between estimated PE and FE and show how stochasticity, process error, and chaotic dynamics affect the relationship. This relationship is verified for a dataset of 461 empirical ecological time series. We show how deviations from the expected PE-FE relationship are related to covariates of data quality and the nonlinearity of ecological dynamics.\n\nThese results demonstrate a theoretically-grounded basis for a model-free evaluation of a systems intrinsic predictability. Identifying the gap between the intrinsic and realized predictability of time series will enable researchers to understand whether forecasting proficiency is limited by the quality and quantity of their data or the ability of the chosen forecasting model to explain the data. Intrinsic predictability also provides a model-free baseline of forecasting proficiency against which modeling efforts can be evaluated.\n\nGlossaryActive information: The amount of information that is available to forecasting models (redundant information minus lost information; Fig. 1).\n\nO_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=45 SRC=\"FIGDIR/small/350017_fig1a.gif\" ALT=\"Figure 1A\">\nView larger version (9K):\norg.highwire.dtl.DTLVardef@934d9eorg.highwire.dtl.DTLVardef@ccdc10org.highwire.dtl.DTLVardef@1839ed9org.highwire.dtl.DTLVardef@31bd70_HPS_FORMAT_FIGEXP M_FIG O_FLOATNOFigure 1A.C_FLOATNO The total information content of an observation of a system at a given state in time, St, is depicted by filled circles with past states (St-1 and St-2) represented by shades of grey, i) lack of overlap between past and present states illustrating a case where no information is transmitted from past states (i.e. a purely stochastic system), with low redundancy and high Shannon entropy rate, ii) intermediate overlap indicating a case when some information is transferred from past to present (i.e. a deterministic system strongly driven by stochastic forcing), with intermediate redundancy and Shannon entropy rate, iii) large overlap indicating a case when the current state is mostly determined by the previous state (i.e. a highly deterministic system), with high redundancy and low Shannon entropy rate. Note that both the redundancy and Shannon entropy rate of a system are intrinsic properties of the system and will only change if the system itself changes.\n\nC_FIG Forecasting error (FE): A measure of the discrepancy between a models forecasts and the observed dynamics of a system. Common measures of forecast error are root mean squared error and mean absolute error.\n\nEntropy: Measures the average amount of information in the outcome of a stochastic process.\n\nInformation: Any entity that provides answers and resolves uncertainty about a process. When information is calculated using logarithms to the base two (i.e. information in bits), it is the minimum number of yes/no questions required, on average, to determine the identity of the symbol (Jost 2006). The information in an observation consists of information inherited from the past (redundant information), and of new information.\n\nIntrinsic predictability: the maximum achievable predictability of a system (Beckage et al. 2011).\n\nLost information: The part of the redundant information lost due to measurement or sampling error, or transformations of the data (Fig. 1).\n\nNew information, Shannon entropy rate: The Shannon entropy rate quantifies the average amount of information per observation in a time series that is unrelated to the past, i.e., the new information (Fig. 1).\n\nNonlinearity: When the deterministic processes governing system dynamics depend on the state of the system.\n\nPermutation entropy (PE): permutation entropy is a measure of the complexity of a time series (Bandt & Pompe, 2002) that is negatively correlated with a systems predictability (Garland et al. 2015). Permutation entropy quantifies the combined new and lost information. PE is scaled to range between a minimum of 0 and a maximum of 1.\n\nRealized predictability: the achieved predictability of a system from a given forecasting model.\n\nRedundant information: The information inherited from the past, and thus the maximum amount of information available for use in forecasting (Fig. 1).\n\nSymbols, words, permutations: symbols are simply the smallest unit in a formal language such as the letters in the English alphabet i.e., {\"A\", \"B\",..., \"Z\"}. In information theory the alphabet is more abstract, such as elements in the set {\"up\", \"down\"} or {\"1\", \"2\", \"3\"}. Words, of length m refer to concatenations of the symbols (e.g., up-down-down) in a set. Permutations are the possible orderings of symbols in a set. In this manuscript, the words are the permutations that arise from the numerical ordering of m data points in a time series.\n\nWeighted permutation entropy (WPE): a modification of permutation entropy (Fadlallah et al., 2013) that distinguishes between small-scale, noise-driven variation and large-scale, system-driven variation by considering the magnitudes of changes in addition to the rank-order patterns of PE.

ecology

Applying generalised allometric regressions to predict live body mass of tropical and temperate arthropods

1. The ecological implications of body size extend from the biology of individual organisms to ecosystem-level processes. Measuring body mass for high numbers of invertebrates can be logistically challenging, making length-mass regressions useful for predicting body mass with minimal effort. However, standardised sets of scaling relationships covering a large range in body length, taxonomic groups, and multiple geographical regions are scarce.\n\n2. We collected 6293 arthropods from 19 higher-level taxa in both temperate and tropical locations to compile a comprehensive set of linear models relating live body mass to a range of predictor variables. For each individual, we measured live weight (hereafter, body mass), body length and width, and conducted linear regressions to predict body mass using body length, body width, taxonomic group and geographic region. Additionally, we quantified prediction error when using parameters from arthropods of a different geographic region.\n\n3. Incorporating body width into taxon- and region-specific length-mass regressions yielded the highest prediction accuracy for body mass. Using regression parameters from a different geographic location increased prediction error, causing over- or underestimation of body mass depending on geographical origin and whether body width was included.\n\n4. We present a comprehensive range of parameters for predicting arthropod body mass and provide guidance for selecting optimal scaling relationships. Given the importance of body mass for functional invertebrate ecology and a paucity of adequate regressions to predict arthropod body mass from different geographical regions, our study provides a long-needed resource for quantifying live body mass in invertebrate ecology research.

zoology

Metabolic adjustment enhances food web stability

Understanding ecosystem stability is one of the greatest challenges of ecology. Over several decades, it has been shown that allometric scaling of biological rates and feeding interactions provide stability to complex food web models. Moreover, introducing adaptive responses of organisms to environmental changes (e.g. like adaptive foraging that enables organisms to adapt their diets depending on resources abundance) improved species per-sistence in food webs. Here, we introduce the concept of metabolic adjustment, i.e. the ability of species to slow down their metabolic rates when facing starvation and to increase it in time of plenty. We study the reactions of such a model to nutrient enrichment and the adjustment speed of metabolic rates. We found that increasing nutrient enrichment leads to a paradox of enrichment (increase in biomasses and oscillation amplitudes and ultimately extinction of species) but metabolic adjustment stabilises the system by dampening the oscillations. Metabolic adjustment also increases the average biomass of the top predator in a tri-trophic food chain. In complex food webs, metabolic adjustment has a stabilising effect as it promotes species survival by creating a large diversity of metabolic rates. However, this stabilising effect is mitigated in enriched ecosystems. Phenotypic plasticity of organisms must be considered in food web models to better understand the response of organisms to their environment. As metabolic rate is central in describing biological rates, we must pay attention to its variations to fully understand the population dynamics of natural communities.

ecology

fluxweb: a package to easily estimate energy fluxes in food webs

O_LIUnderstanding how changes in biodiversity will impact the stability and functioning of ecosystems is a central challenge in ecology. Food-web approaches have been advocated to link community composition with ecosystem functioning by describing the fluxes of energy among species or trophic groups. However, estimating such fluxes remains problematic because current methods become unmanageable as network complexity increases.\nC_LIO_LIWe developed a generalisation of previous indirect estimation methods assuming a steady state system [1, 2, 3]: the model estimates energy fluxes in a top-down manner assuming system equilibrium; each nodes losses (consumption and physiological) balances its consumptive gains. Jointly, we provide theoretical and practical guidelines to use the fluxweb R package (available on CRAN at https://bit.ly/2OC0uKF).\nWe also present how the framework can merge with the allometric theory of ecology [4] to calculate fluxes based on easily obtainable organism-level data (i.e. body masses and species groups -eg, plants animals), opening its use to food webs of all complexities. Physiological losses (metabolic losses or losses due to death other than from predation within the food web) may be directly measured or estimated using allometric relationships based on the metabolic theory of ecology, and losses and gains due to predation are a function of ecological efficiencies that describe the proportion of energy that is used for biomass production.\nC_LIO_LIThe primary output is a matrix of fluxes among the nodes of the food web. These fluxes can be used to describe the role of a species, a function of interest (e.g. predation; total fluxes to predators), multiple functions, or total energy flux (system throughflow or multitrophic functioning). Additionally, the package includes functions to calculate network stability based on the Jacobian matrix, providing insight into how resilient the network is to small perturbations at steady state.\nC_LIO_LIOverall, fluxweb provides a flexible set of functions that greatly increase the feasibility of implementing food-web energetic approaches to more complex systems. As such, the package facilitates novel opportunities for mechanistically linking quantitative food webs and ecosystem functioning in real and dynamic natural landscapes.\nC_LI

ecology

Interactive effects of shifting body size and feeding adaptation drive interaction strengths of protist predators under warming

Global change is heating up ecosystems fuelling biodiversity loss and species extinctions. High-trophic-level predators are especially prone to extinction due to an energetic mismatch between increasing feeding rates and metabolism with warming. Different adaptation mechanisms such as decreasing body size to reduce energy requirements (morphological response) as well as direct effects of adaptation to feeding parameters (physiological response) have been proposed to overcome this problem. Here, we use protist-bacteria microcosm experiments to show how those adaptations may have the potential to buffer the impact of warming on predator-prey interactions. After adapting the ciliate predator Tetrahymena pyriformis to three different temperatures (15{degrees}C, 20{degrees}C and 25{degrees}C) for approximately 20 generations we conducted functional response experiments on bacterial prey along an experimental temperature gradient (15{degrees}C, 20{degrees}C and 25{degrees}C). We found an increase of maximum feeding rates and half-saturation densities with rising experimental temperatures. Adaptation temperature had on average slightly negative effects on maximum feeding rates, but maximum feeding rates increased more strongly with rising experimental temperature in warm adapted predators than in cold adapted predators. There was no effect of adaptation temperature on half-saturation densities characterising foraging efficiency. Besides the mixed response in functional response parameters, predators also adapted by decreasing body size. As smaller predators need less energy to fulfil their energetic demands, maximum feeding rates relative to the energetic demands increased slightly with increased adaptation temperature. Accordingly, predators adapted to 25{degrees}C showed the highest feeding rates at 25{degrees}C experimental temperature, while predators adapted to 15{degrees}C showed the highest maximum feeding rate at 15{degrees}C. Therefore, adaptation to different temperatures potentially avoids an energetic mismatch with warming. Especially a shift in body size with warming additionally to an adaptation of physiological parameters potentially helps to maintain a positive energy balance and prevent predator extinction with rising temperatures.

ecology

Universal scaling of maximum speed with body mass - Why the largest animals are not the fastest

Speed is the fundamental constraint on animal movement, yet there is no general consensus on the determinants of maximum speed itself. Here, we provide a universal scaling model of maximum speed with body mass, which holds across locomotion modes, ecosystem types and taxonomic groups. In contrast to traditional power-law scaling, we predict a hump-shaped relationship due to a finite acceleration time for animals. This model is strongly supported by extensive empirical data (470 species with body masses ranging from 5.7x10-8 to 108,000 kg) from terrestrial as well as aquatic ecosystems. Our approach offers a novel concept of what determines the upper limit of animal movement, thus enabling a better understanding of realized movement patterns in nature and their multifold ecological consequences.

ecology