Ecological stoichiometry in freshwater benthic systems: recent progress and perspectives

1. Ecological stoichiometry deals with the mass balance of multiple key elements [e.g. carbon (C), nitrogen (N), phosphorus (P)] in ecological systems. This conceptual framework, largely developed in the pelagic zone of lakes, has been successfully applied to topics ranging from population dynamics to biogeochemical cycling. More recently, an explicit stoichiometric approach has also been used in many other environments, including freshwater benthic ecosystems. 2. Description of elemental patterns among benthic resources and consumers provides a useful starting point for understanding causes of variation and stoichiometric imbalance in feeding interactions. Although there is considerable overlap among categories, terrestrially-derived resources, such as wood, leaf litter and green leaves have substantially higher C : nutrient ratios than other resources of both terrestrial and aquatic origin, such as periphyton and fine particulate organic matter. The elemental composition of these resources for benthic consumers is modulated by a range of factors and processes, including nutrient availability and ratios, particle size and microbial colonisation. 3. Among consumers in benthic systems, bacteria are the most nutrient-rich, followed (in descending order) by fishes, invertebrate predators, invertebrate primary consumers, and fungi. Differences in consumer C : nutrient ratios appear to be related to broad-scale phylogenetic differences which determine body size, growth rate and resource allocation to structural body constituents (e.g. P-rich bone). 4. Benthic consumers can influence the stoichiometry of dissolved nutrients and basal resources in multiple ways. Direct consumption alters the stoichiometry of food resources by increasing nutrient availability (e.g. reduced boundary layer thickness on substrata) or through removal of nutrient-rich patches (e.g. selective feeding on fungal patches within leaf litter). In addition, consumers alter the stoichiometry of resources and dissolved nutrient pools through the return of egested or excreted nutrients. In some cases, consumer excretion supplies a large proportion of the nutrients required by algae and heterotrophic microbes and alters elemental ratios of dissolved nutrient pools. 5. Organic matter decomposition in benthic systems is accompanied by significant changes in the elemental composition of organic matter. Microbial colonisation of leaf litter influences C : nutrient ratios, and patterns of microbial succession (e.g. fungi followed by bacteria) may be under some degree of stoichiometric control. Large elemental imbalances