Scaling of the risk landscape drives optimal life history strategies and the evolution of grazing

Consumers face numerous risks that can be minimized by incorporating different life-history strategies. How much and when a consumer adds to its energetic reserves or invests in reproduction are key behavioral and physiological adaptations that structure much of how organisms interact. Here we develop a theoretical framework that explicitly accounts for stochastic fluctuations of an individual consumer's energetic reserves while foraging and reproducing on a landscape with resources that range from uniformly distributed to highly clustered. First, we show that optimal life-history strategies vary in response to changes in the mean productivity of the resource landscape, where depleted environments promote reproduction at lower energetic states, greater investment in each reproduction event, and smaller litter sizes. We then show that if resource variance scales with body size due to landscape clustering, consumers that forage for clustered foods are susceptible to strong Allee effects, increasing extinction risk. Finally, we show that the proposed relationship between consumer body size, resource clustering, and Allee effect-induced population instability offers key ecological insights into the evolution of large-bodied grazing herbivores from small-bodied browsing ancestors.Comment: 9 pages, 5 figures, 3 Supplementary Appendices, 2 Supplementary Figure

On the dynamics of mortality and the ephemeral nature of mammalian megafauna

Energy flow through consumer-resource interactions is largely determined by body size. Allometric relationships govern the dynamics of populations by impacting rates of reproduction, as well as alternative sources of mortality, which have differential impacts on smaller to larger organisms. Here we derive and investigate the timescales associated with four alternative sources of mortality for terrestrial mammals: mortality from starvation, mortality associated with aging, mortality from consumption by predators, and mortality introduced by anthropogenic subsidized harvest. The incorporation of these allometric relationships into a minimal consumer-resource model illuminates central constraints that may contribute to the structure of mammalian communities. Our framework reveals that while starvation largely impacts smaller-bodied species, the allometry of senescence is expected to be more difficult to observe. In contrast, external predation and subsidized harvest have greater impacts on the populations of larger-bodied species. Moreover, the inclusion of predation mortality reveals mass thresholds for mammalian herbivores, where dynamic instabilities may limit the feasibility of megafaunal populations. We show how these thresholds vary with alternative predator-prey mass relationships, which are not well understood within terrestrial systems. Finally, we use our framework to predict the harvest pressure required to induce mass-specific extinctions, which closely align with previous estimates of anthropogenic megafaunal exploitation in both paleontological and historical contexts. Together our results underscore the tenuous nature of megafaunal populations, and how different sources of mortality may contribute to their ephemeral nature over evolutionary time.Comment: 10 pages, 5 figures, 1 table, 4 appendices, 8 supplementary figure

Collapse of an ecological network in Ancient Egypt

The dynamics of ecosystem collapse are fundamental to determining how and why biological communities change through time, as well as the potential effects of extinctions on ecosystems. Here we integrate depictions of mammals from Egyptian antiquity with direct lines of paleontological and archeological evidence to infer local extinctions and community dynamics over a 6000-year span. The unprecedented temporal resolution of this data set enables examination of how the tandem effects of human population growth and climate change can disrupt mammalian communities. We show that the extinctions of mammals in Egypt were nonrandom, and that destabilizing changes in community composition coincided with abrupt aridification events and the attendant collapses of some complex societies. We also show that the roles of species in a community can change over time, and that persistence is predicted by measures of species sensitivity, a function of local dynamic stability. Our study is the first high-resolution analysis of the ecological impacts of environmental change on predator-prey networks over millennial timescales, and sheds light on the historical events that have shaped modern animal communities

Eco-Evolutionary Origins of Diverse Abundance, Biomass, and Trophic Structures in Food Webs

Organismal traits and their evolution can strongly influence food web structure and dynamics. To what extent the evolution of such traits impacts food web structure, however, is poorly understood. Here, we investigate a simple three-species omnivory food web module where the attack rates of all predators evolve as ecological dynamics unfold, such that predator trophic levels are themselves dynamic. We assume a timescale where other vital rates that govern population dynamics are constant and incorporate a well-known tradeoff between attack rates and the conversion of prey into predator biomass. We show that this eco-evolutionary model yields a surprisingly rich array of dynamics. Moreover, even small amounts of selection lead to important differences in the abundance, trophic, and biomass structure of the food web. Systems in which intermediate predators are strongly constrained by tradeoffs lead to hourglass-shaped food webs, where basal resources and top predators have large abundances, but intermediate predators are rare, like those observed in some marine ecosystems. Such food webs are also characterized by a relatively low maximum trophic level. Systems in which intermediate predators have weaker tradeoffs lead to pyramid-shaped food webs, where basal resources are more abundant than intermediate and top predators, such as those observed in some terrestrial system. These food webs also supported a relatively higher maximum trophic level. Overall, our results suggest that eco-evolutionary dynamics can strongly influence the abundance-, trophic-, and biomass-structure of food webs, even in the presence of small levels of selection, thus stressing the importance of taking traits and trait evolution into account to further understand community-level patterns and processes