Food webs are more than the sum of their tritrophic parts.

Many studies have aimed to understand food webs by investigating components such as trophic links (one consumer taxon eats one resource taxon), tritrophic interactions (one consumer eats an intermediate taxon, which eats a resource), or longer chains of links. We show here that none of these components (links, tritrophic interactions, and longer chains), individually or as an ensemble, accounts fully for the properties of the next higher level of organization. As a cell is more than its molecules, as an organ is more than its cells, and as an organism is more than its organs, in a food web, new structure emerges at every organizational level up to and including the whole web. We demonstrate the emergence of properties at progressively higher levels of structure by using all of the directly observed, appropriately organized, publicly available food web datasets with relatively complete trophic link data and with average body mass and population density data for each taxon. There are only three such webs, those of Tuesday Lake, Michigan, in 1984 and 1986, and Ythan Estuary, Scotland. We make the data freely available online with this report. Differences in web patterns between Tuesday Lake and Ythan Estuary, and similarities of Tuesday Lake in 1984 and 1986 despite 50% turnover of species, suggest that the patterns we describe respond to major differences between ecosystem types.

Influence of macrofaunal assemblages and environmental heterogeneity on microphytobenthic production in experimental systems.

Despite the complexity of natural systems, heterogeneity caused by the fragmentation of habitats has seldom been considered when investigating ecosystem processes. Empirical approaches that have included the influence of heterogeneity tend to be biased towards terrestrial habitats; yet marine systems offer opportunities by virtue of their relative ease of manipulation, rapid response times and the well-understood effects of macrofauna on sediment processes. Here, the influence of heterogeneity on microphytobenthic production in synthetic estuarine assemblages is examined. Heterogeneity was created by enriching patches of sediment with detrital algae (Enteromorpha intestinalis) to provide a source of allochthonous organic matter. A gradient of species density for four numerically dominant intertidal macrofauna (Hediste diversicolor, Hydrobia ulvae, Corophium volutator, Macoma balthica) was constructed, and microphytobenthic biomass at the sediment surface was measured. Statistical analysis using generalized least squares regression indicated that heterogeneity within our system was a significant driving factor that interacted with macrofaunal density and species identity. Microphytobenthic biomass was highest in enriched patches, suggesting that nutrients were obtained locally from the sediment-water interface and not from the water column. Our findings demonstrate that organic enrichment can cause the development of heterogeneity which influences infaunal bioturbation and consequent nutrient generation, a driver of microphytobenthic production.