Tomas Jonsson

Ecological community description using the food web, species abundance, and body size

Measuring the numerical abundance and average body size of individuals of each species in an ecological communitys food web reveals new patterns and illuminates old ones. This approach is illustrated using data from the pelagic community of a small lake: Tuesday Lake, Michigan, United States. Body mass varies almost 12 orders of magnitude. Numerical abundance varies almost 10 orders of magnitude. Biomass abundance (average body mass times numerical abundance) varies only 5 orders of magnitude. A new food web graph, which plots species and trophic links in the plane spanned by body mass and numerical abundance, illustrates the nearly inverse relationship between body mass and numerical abundance, as well as the pattern of energy flow in the community. Species with small average body mass occur low in the food web of Tuesday Lake and are numerically abundant. Larger-bodied species occur higher in the food web and are numerically rarer. Average body size explains more of the variation in numerical abundance than does trophic height. The trivariate description of an ecological community by using the food web, average body sizes, and numerical abundance includes many well studied bivariate and univariate relationships based on subsets of these three variables. We are not aware of any single community for which all of these relationships have been analyzed simultaneously. Our approach demonstrates the connectedness of ecological patterns traditionally treated as independent. Moreover, knowing the food web gives new insight into the disputed form of the allometric relationship between body mass and abundance.

Food Webs, Body Size, and Species Abundance in Ecological Community Description

I. Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 A. Trivariate Relationships. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 B. Bivariate Relationships . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 C. Univariate Relationships . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 D. EVect of Food Web Perturbation . . . . . . . . . . . . . . . . . . . . . . . . 4 I

Functional identity and diversity of animals predict ecosystem functioning better than species-based indices

Drastic biodiversity declines have raised concerns about the deterioration of ecosystem functions and have motivated much recent research on the relationship between species diversity and ecosystem functioning. A functional trait framework has been proposed to improve the mechanistic understanding of this relationship, but this has rarely been tested for organisms other than plants. We analysed eight datasets, including five animal groups, to examine how well a trait-based approach, compared with a more traditional taxonomic approach, predicts seven ecosystem functions below- and above-ground. Trait-based indices consistently provided greater explanatory power than species richness or abundance. The frequency distributions of single or multiple traits in the community were the best predictors of ecosystem functioning. This implies that the ecosystem functions we investigated were underpinned by the combination of trait identities (i.e. single-trait indices) and trait complementarity (i.e. multi-trait indices) in the communities. Our study provides new insights into the general mechanisms that link biodiversity to ecosystem functioning in natural animal communities and suggests that the observed responses were due to the identity and dominance patterns of the trait composition rather than the number or abundance of species per se.

Stepping in Elton’s footprints: a general scaling model for body masses and trophic levels across ecosystems

Despite growing awareness of the significance of body-size and predator-prey body-mass ratios for the stability of ecological networks, our understanding of their distribution within ecosystems is incomplete. Here, we study the relationships between predator and prey size, body-mass ratios and predator trophic levels using body-mass estimates of 1313 predators (invertebrates, ectotherm and endotherm vertebrates) from 35 food-webs (marine, stream, lake and terrestrial). Across all ecosystem and predator types, except for streams (which appear to have a different size structure in their predator-prey interactions), we find that (1) geometric mean prey mass increases with predator mass with a power-law exponent greater than unity and (2) predator size increases with trophic level. Consistent with our theoretical derivations, we show that the quantitative nature of these relationships implies systematic decreases in predator-prey body-mass ratios with the trophic level of the predator. Thus, predators are, on an average, more similar in size to their prey at the top of food-webs than that closer to the base. These findings contradict the traditional Eltonian paradigm and have implications for our understanding of body-mass constraints on food-web topology, community dynamics and stability.

The Role of Body Size in Complex Food Webs: A Cold Case

Human-induced habitat destruction, overexploitation, introduction of alien species and climate change are causing species to go extinct at unprecedented rates, from local to global scales. There are growing concerns that these kinds of disturbances alter important functions of ecosystems. Our current understanding is that key parameters of a community (e.g. its functional diversity, species composition, and presence/absence of vulnerable species) reflect an ecological network’s ability to resist or rebound from change in response to pressures and disturbances, such as species loss. If the food web structure is relatively simple, we can analyse the roles of different species interactions in determining how environmental impacts translate into species loss. However, when ecosystems harbour species-rich communities, as is the case in most natural systems, then the complex network of ecological interactions makes it a far more challenging task to perceive how species’ functional roles influence the consequences of species loss. One approach to deal with such complexity is to focus on the functional traits of species in order to identify their respective roles: for instance, large species seem to be more susceptible to extinction than smaller species. Here, we introduce and analyse the marine food web from the high Antarctic Weddell Sea Shelf to illustrate the role of species traits in relation to network robustness of this complex food web. Our approach was threefold: firstly, we applied a new classification system to all species, grouping them by traits other than body size; secondly, we tested the relationship between body size and food web parameters within and across these groups and finally, we calculated food web robustness. We addressed questions regarding (i) patterns of species functional/trophic roles, (ii) relationships between species functional roles and body size and (iii) the role of species body size in terms of network robustness. Our results show that when analyzing relationships between trophic structure, body size and network structure, the diversity of predatory species types needs to be considered in future studies.

Interaction strength, food web topology and the relative importance of species in food webs.

1. We established complex marine communities, consisting of over 100 species, in large subtidal experimental mesocosms. We measured the strength of direct interactions and the net strength of direct and indirect interactions between the species in those communities, using a combination of theoretical and empirical approaches. 2. Theoretical predictions of interaction strength were derived from the interaction coefficient matrix, which was parameterised using allometric predator-prey relationships. Empirical estimates of interaction strength were quantified using the ln-ratio, which measures the change in biomass density of species A in the presence and absence of species B. 3. We observed that highly connected species tend to have weak direct effects and net effects in our experimental food webs, whether we calculate interaction strength theoretically or empirically. 4. We found a significant correlation between our theoretical predictions and empirical estimates of direct effects and net effects. The net effects correlation was much stronger, indicating that our experimental communities were dominated by a mixture of direct and indirect effects. 5. Re-calculation of the theoretical predictions of net effects after randomising predator and prey body masses did not affect the negative relationship with connectance. 6. These results suggest that food web topology, which in this system is constrained by body mass, is overwhelmingly important for the magnitude of direct and indirect interactions and hence species importance in the face of biodiversity declines.

Body size and food web structure : testing the equiprobability assumption of the cascade model

Abstract The cascade model successfuly predicts many patterns in reported food webs. A key assumption of this model is the existence of a predetermined trophic hierarchy; prey are always lower in the hierarchy than their predators. At least three studies have suggested that, in animal food webs, this hierarchy can be explained to a large extent by body size relationships. A second assumption of the standard cascade model is that trophic links not prohibited by the hierarchy occur with equal probability. Using nonparametric contingency table analyses, we tested this ”equiprobability hypothesis” in 16 published animal food webs for which the adult body masses of the species had been estimated. We found that when the hierarchy was based on body size, the equiprobability hypothesis was rejected in favor of an alternative, ”predator-dominance” hypothesis wherein the probability of a trophic link varies with the identity of the predator. Another alternative to equiprobabilty is that the probability of a trophic link depends upon the ratio of the body sizes of the two species. Using nonparametric regression and liklihood ratio tests, we show that a size-ratio based model represents a significant improvement over the cascade model. These results suggest that models with heterogeneous predation probabilities will fit food web data better than the homogeneous cascade model. They also suggest a new way to bridge the gap between static and dynamic food web models.

Temporal Variability in Predator–Prey Relationships of a Forest Floor Food Web

Summary Connectance webs represent the standard data description in food web ecology, but their usefulness is often limited in understanding the patterns and processes within ecosystems. Increasingly, efforts have been made to incorporate additional, biologically meaningful, data into food web descriptions, including the construction of food webs using data describing the body size and abundance of each species. Here, data from a terrestrial forest floor food web, sampled seasonally over a 1-year period, were analysed to investigate (i) how stable the body size–abundance and predator–prey relationships of an ecosystem are through time and (ii) whether there are system-specific differences in body size–abundance and predator–prey relationships between ecosystem types.

EVOLUTION OF STABLE POPULATION DYNAMICS THROUGH NATURAL SELECTION

Analyses of very simple models of population growth indicate that the dynamic behaviour of populations may be very complex — ranging from stable equilibrium points through limit cycles to chaos — even in constant environments. Yet, convincing evidence for chaotic dynamics in natural populations is limited to very few cases. This apparent paradox has received considerable attention among ecologists. We have investigated the population dynamic consequences of density dependent selection in a stage-structured population model by means of an invasibility-analysis. We focus on the situation where juvenile survival and maturation rates are density dependent, which is common in organisms with plastic growth. Based on energetic considerations we assume that there exists a trade-off between the sensitivities of juvenile survival and maturation rate to population density. Given this plausible assumption we show that selection tends to minimize the sensitivity of juvenile survival to population density at the expense of a lower maturation rate and hence a longer time to reach maturity. As a consequence selection will often lead to more stable populations.

Species-rich ecosystems are vulnerable to cascading extinctions in an increasingly variable world

Global warming leads to increased intensity and frequency of weather extremes. Such increased environmental variability might in turn result in increased variation in the demographic rates of interacting species with potentially important consequences for the dynamics of food webs. Using a theoretical approach, we here explore the response of food webs to a highly variable environment. We investigate how species richness and correlation in the responses of species to environmental fluctuations affect the risk of extinction cascades. We find that the risk of extinction cascades increases with increasing species richness, especially when correlation among species is low. Initial extinctions of primary producer species unleash bottom-up extinction cascades, especially in webs with specialist consumers. In this sense, species-rich ecosystems are less robust to increasing levels of environmental variability than species-poor ones. Our study thus suggests that highly species-rich ecosystems such as coral reefs and tropical rainforests might be particularly vulnerable to increased climate variability.