Traditional approaches to the study of food webs emphasize the transfer of local primary productivity in the form of living plant organic matter across trophic levels. However, dead organic matter, or detritus, a common feature of most ecosystems plays a frequently overlooked role as a dynamic heterogeneous resource and habitat for many species. We develop an integrative framework for understanding the impact of detritus that emphasizes the ontogeny and heterogeneity of detritus and the various ways that explicit inclusion of detrital dynamics alters generalizations about the structure and functioning of food webs. Through its influences on food web composition and dynamics, detritus often increases system stability and persistence, having substantial effects on trophic structure and biodiversity. Inclusion of detrital heterogeneity in models of food web dynamics is an essential new direction for ecological research.

Stoichiometric relationships between consumers and resources in detritus‐based ecosystems have received little attention, despite the importance of detritus in most food webs. We analysed carbon (C), nitrogen (N), and phosphorus (P) content of invertebrate consumers, and basal food resources in two forested headwater streams (one reference and the other nutrient‐enriched). We found large elemental imbalances between consumers and food resources compared with living plant‐based systems, particularly in regard to P content, which were reduced with enrichment. Enrichment significantly increased nutrient content of food resources (consistent with uptake of N and P by detritus‐associated microbes). P content of some invertebrates also increased in the enriched vs. reference stream, suggesting deviation from strict homeostasis. Nutrient content varied significantly among invertebrate functional feeding groups, orders and, to some extent, size classes. Future application of stoichiometric theory to detritus‐based systems should consider the potential for relatively large consumer‐resource elemental imbalances and P storage by insect consumers.

SUMMARY 1. The effects of catchment urbanisation on water quality were examined for 30 streams (stratified into 15, 50 and 100 km2 ± 25% catchments) in the Etowah River basin, Georgia, U.S.A. We examined relationships between land cover (implying cover and use) in these catchments (e.g. urban, forest and agriculture) and macroinvertebrate assemblage attributes using several previously published indices to summarise macroinvertebrate response. Based on a priori predictions as to mechanisms of biotic impairment under changing land cover, additional measurements were made to assess geomorphology, hydrology and chemistry in each stream. 2. We found strong relationships between catchment land cover and stream biota. Taxon richness and other biotic indices that reflected good water quality were negatively related to urban land cover and positively related to forest land cover. Urban land cover alone explained 29–38% of the variation in some macroinvertebrate indices. Reduced water quality was detectable at c. >15% urban land cover. 3. Urban land cover correlated with a number of geomorphic variables such as stream bed sediment size (–) and total suspended solids (+) as well as a number of water chemistry variables including nitrogen and phosphorus concentrations (+), specific conductance (+) and turbidity (+). Biotic indices were better predicted by these reach scale variables than single, catchment scale land cover variables. Multiple regression models explained 69% of variation in total taxon richness and 78% of the variation in the Invertebrate Community Index (ICI) using phi variability, specific conductance and depth, and riffle phi, specific conductance and phi variability, respectively. 4. Indirect ordination analysis was used to describe assemblage and functional group changes among sites and corroborate which environmental variables were most important in driving differences in macroinvertebrate assemblages. The first axis in a non‐metric multidimensional scaling ordination was highly related to environmental variables (slope, specific conductance, phi variability; adj. R2=0.83) that were also important in our multiple regression models. 5. Catchment urbanisation resulted in less diverse and more tolerant stream macroinvertebrate assemblages via increased sediment transport, reduced stream bed sediment size and increased solutes. The biotic indices that were most sensitive to environmental variation were taxon richness, EPT richness and the ICI. Our results were largely consistent over the range in basin size we tested.

We conducted two experiments to determine the relative effects of herbivory and nutrients on an algal community in Walker Branch, a stream having effectively two trophic levels: primary producers and herbivorous snails. The first study (1989), performed in streamside channels, tested the effects of three factors: (l) stream water nitrogen (N), (2) phosphorus (P), and (3) snail grazing, on periphyton biomass, productivity, and community composition. The second study ( 1990), conducted in situ, tested the effects of snail grazing and nutrients (N + P). In the 1989 study, nutrients had positive effects, and herbivores had negative effects, on algal biomass (chlorophyll a, ash-free dry mass, total algal biovolume) and primary productivity (areaand chlorophyll-specific). Likewise, both nutrients and snail grazing exerted effects ( + and -, respectively) on biomass measured in the 1990 study (chlorophyll a, algal biovolume). Grazed communities were dominated by chlorophytes and cyanophytes, which were overgrown by diatoms when herbivores were removed. Algal species that were reduced most by herbivores were increased most by nutrient addition, and vice versa, suggesting a trade-off between resistance to herbivory and nutrientsaturated growth rates. Increases in algal biomass and productivity were slight with the addition of either N or P compared to responses observed when both nutrients were added together, suggesting that both nutrients were at growth-limiting levels. The greatest changes in periphyton structure or function were observed when both nutrients were added and simultaneously, grazers were removed, in contrast to lesser effects when nutrients were added under grazed conditions or grazers were removed at low nutrient levels, indicating dual control by both factors. Nutrient addition also positively affected snail growth in both experiments, indicating tight coupling between herbivore and algal growth (top-down effects) and that bottom-up factors that directly affected plant growth could also indirectly affect consumers belonging to higher trophic levels. Indices quantifying the direct effects of top-down factors relative to bottom-up factors (top-down index, TDI) and the importance of interactions between these factors (interaction coefficient, IC) were computed. These indices showed that the relative strength of top-down and bottom-up factors varied among biomass and productivity parameters and that top-down and bottom-up effects, alone, were less important than their combined effects.

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