Food-web dynamics under climate change.

Climate change affects ecological communities through its impact on the physiological performance of individuals. However, the population dynamic of species well inside their thermal niche is also determined by competitors, prey and predators, in addition to being influenced by temperature changes. We use a trait-based food-web model to examine how the interplay between the direct physiological effects from temperature and the indirect effects due to changing interactions between populations shapes the ecological consequences of climate change for populations and for entire communities. Our simulations illustrate how isolated communities deteriorate as populations go extinct when the environment moves outside the species' thermal niches. High-trophic-level species are most vulnerable, while the ecosystem function of lower trophic levels is less impacted. Open communities can compensate for the loss of ecosystem function by invasions of new species. Individual populations show complex responses largely uncorrelated with the direct impact of temperature change on physiology. Such complex responses are particularly evident during extinction and invasion events of other species, where climatically well-adapted species may be brought to extinction by the changed food-web topology. Our results highlight that the impact of climate change on specific populations is largely unpredictable, and apparently well-adapted species may be severely impacted.

Predicting the consequences of species loss using size-structured biodiversity approaches.

Understanding the consequences of species loss in complex ecological communities is one of the great challenges in current biodiversity research. For a long time, this topic has been addressed by traditional biodiversity experiments. Most of these approaches treat species as trait-free, taxonomic units characterizing communities only by species number without accounting for species traits. However, extinctions do not occur at random as there is a clear correlation between extinction risk and species traits. In this review, we assume that large species will be most threatened by extinction and use novel allometric and size-spectrum concepts that include body mass as a primary species trait at the levels of populations and individuals, respectively, to re-assess three classic debates on the relationships between biodiversity and (i) food-web structural complexity, (ii) community dynamic stability, and (iii) ecosystem functioning. Contrasting current expectations, size-structured approaches suggest that the loss of large species, that typically exploit most resource species, may lead to future food webs that are less interwoven and more structured by chains of interactions and compartments. The disruption of natural body-mass distributions maintaining food-web stability may trigger avalanches of secondary extinctions and strong trophic cascades with expected knock-on effects on the functionality of the ecosystems. Therefore, we argue that it is crucial to take into account body size as a species trait when analysing the consequences of biodiversity loss for natural ecosystems. Applying size-structured approaches provides an integrative ecological concept that enables a better understanding of each species' unique role across communities and the causes and consequences of biodiversity loss.

Adult and offspring size in the ocean: a database of size metrics and conversion factors.

The purpose of this dataset was to compile adult and offspring size estimates for marine organisms. Adult and offspring size estimates of 408 species were compiled from the literature covering >17 orders of magnitude in body mass and including Cephalopoda (ink fish), Cnidaria ("jelly" fish), Crustaceans, Ctenophora (comb jellies), Elasmobranchii (cartilaginous fish), Mammalia (mammals), Sagittoidea (arrow worms) and Teleost (i.e., Actinopterygii, bony fish). Individual size estimates were converted to standardized size estimates (carbon weight, g) to allow for among-group comparisons. This required a number of size estimates to be converted and a compilation of conversion factors obtained from the literature are also presented.

Coexistence of structured populations with size-based prey selection.

Species with a large adult-offspring size ratio and a preferred predator-prey mass ratio undergo ontogenetic trophic niche shift(s) throughout life. Trophic interactions between such species vary throughout life, resulting in different species-level interaction motifs depending on the maximum adult sizes and population size distributions. We explore the assembly and potential for coexistence of small communities where all species experience ontogenetic trophic niche shifts. The life-history of each species is described by a physiologically structured model and species identity is characterised by the trait: size at maturation. We show that a single species can exist in two different states: a 'resource driven state' and a 'cannibalistic state' with a large scope for emergent Allee effects and bistable states. Two species can coexist in two different configurations: in a 'competitive coexistence' state when the ratio between sizes at maturation of the two species is less than a predator-prey mass ratio and the resource level is low to intermediate, or in a 'trophic ladder' state if the ratio of sizes at maturation is larger than the predator-prey mass ratio at all resource levels. While there is a large scope for coexistence of two species, the scope for coexistence of three species is limited and we conclude that further trait differentiation is required for coexistence of more species-rich size-structured communities.

Universal temperature and body-mass scaling of feeding rates.

Knowledge of feeding rates is the basis to understand interaction strength and subsequently the stability of ecosystems and biodiversity. Feeding rates, as all biological rates, depend on consumer and resource body masses and environmental temperature. Despite five decades of research on functional responses as quantitative models of feeding rates, a unifying framework of how they scale with body masses and temperature is still lacking. This is perplexing, considering that the strength of functional responses (i.e. interaction strengths) is crucially important for the stability of simple consumer-resource systems and the persistence, sustainability and biodiversity of complex communities. Here, we present the largest currently available database on functional response parameters and their scaling with body mass and temperature. Moreover, these data are integrated across ecosystems and metabolic types of species. Surprisingly, we found general temperature dependencies that differed from the Arrhenius terms predicted by metabolic models. Additionally, the body-mass-scaling relationships were more complex than expected and differed across ecosystems and metabolic types. At local scales (taxonomically narrow groups of consumer-resource pairs), we found hump-shaped deviations from the temperature and body-mass-scaling relationships. Despite the complexity of our results, these body-mass- and temperature-scaling models remain useful as a mechanistic basis for predicting the consequences of warming for interaction strengths, population dynamics and network stability across communities differing in their size structure.

Food web framework for size-structured populations.

We synthesise traditional unstructured food webs, allometric body size scaling, trait-based modelling, and physiologically structured modelling to provide a novel and ecologically relevant tool for size-structured food webs. The framework allows food web models to include ontogenetic growth and life-history omnivory at the individual level by resolving the population structure of each species as a size-spectrum. Each species is characterised by the trait 'size at maturation', and all model parameters are made species independent through scaling with individual body size and size at maturation. Parameter values are determined from cross-species analysis of fish communities as life-history omnivory is widespread in aquatic systems, but may be reparameterised for other systems. An ensemble of food webs is generated and the resulting communities are analysed at four levels of organisation: community level, species level, trait level, and individual level. The model may be solved analytically by assuming that the community spectrum follows a power law. The analytical solution provides a baseline expectation of the results of complex food web simulations, and agrees well with the predictions of the full model on biomass distribution as a function of individual size, biomass distribution as a function of size at maturation, and relation between predator-prey mass ratio of preferred and eaten food. The full model additionally predicts the diversity distribution as a function of size at maturation.