Methods of assessing the structure and function of food webs are needed to provide a basis for assessing large-scale direct (e.g. fisheries) and indirect (e.g. climate change) effects of human activities on marine ecosystems. We present a simple synthesis of the complex structure and function of a real marine food web, based on analyses of body size distributions, production-body size relationships and trophic level-body size relationships. We show how size-based estimates of production, species richness and trophic level (from nitrogen stable isotope analysis) can be used to quantify trophic transfer efficiency, mean predator-prey body-mass ratios and the mean ratio of the number of predator to prey species in marine food webs. We applied these methods to the central North Sea, and estimated transfer efficiencies of 3.7 to 12.4%, a mean predator-prey body-mass ratio of 109:1 and a mean ratio of the number of predator to prey species of 0.34. We conducted sensitivity analyses to show how differences in the fractionation of δ 15 N and changes in the slope of the relationship between production and δ 15 N affected our predictions. Our estimates of transfer efficiency and mean predator-prey body-mass ratios are similar to those obtained by costly and labour-intensive diet-and ecosystem-modelling studies. Coupled analyses of size and trophic structure may provide a method for validating ecosystem models and assessing human impacts on marine ecosystems. KEY WORDS: Body size · Food web · Transfer efficiency · Stable isotope analysis · Production · DiversityResale or republication not permitted without written consent of the publisher

Summary 1.Trawling disturbance has wide-ranging impacts on the marine environment and is well known to modify benthic habitat and community structure. This has led to speculation about the positive and negative impacts of trawling on ecosystem processes such as production. 2. Existing theory suggests that frequent trawling disturbance may lead to the proliferation of smaller benthic species, with faster life histories, because they can withstand the mortality imposed by trawling and benefit from reduced competition or predation as populations of larger species are depleted. Since smaller species are more productive, trawling disturbance may 'farm the sea', with knock-on benefits for consumers, including fish populations. 3. We conducted the first large-scale studies of trawling effects on benthic production across quantified gradients of trawling disturbance on real fishing grounds in two regions (Silver Pit and Hills) of the North Sea. There were 27-and 10-fold differences in levels of beam trawl disturbance among the Silver Pit and Hills sites, respectively. 4. Size structure was described using normalized size-spectra, and the slopes and intercepts of these spectra were related to levels of trawling disturbance. Production was estimated from the size spectra, using a new allometric relationship between body mass and the production to biomass (P:B) ratio of marine invertebrates. The general validity of the relationship was confirmed using a phylogenetic comparative approach. 5. In the Silver Pit region, trawling led to significant decreases in infaunal biomass and production. The abundance of larger individuals was depleted more than smaller ones, as reflected by the positive relationship between the slope of the normalized size spectra and trawling disturbance. The effects of trawling disturbance were not significant in the epifaunal community. In the Hills region, where the range of trawling disturbance was lower, trawling disturbance did not have significant effects on biomass or production. 6. In the Silver Pit, relative infaunal production (production per unit biomass) rose with increased trawling disturbance. This was attributable largely to the dominance of smaller animals in the disturbed communities. The increase in relative production did not compensate for the loss of total production that resulted from the depletion of large individuals. There was some evidence for the proliferation of small polychaetes at moderate levels of disturbance, but at higher levels of disturbance their biomass and production fell. 7. We conclude that reported increases in the biomass and production of small infaunal invertebrates in the North Sea are attributable largely to recent increases in primary production that were driven by climate change, and not to the effects of trawling disturbance.

Models of biomass size spectra assume that organisms with higher body mass feed at higher trophic levels, but explicit empirical tests of this pattern are rare. We used nitrogen stable isotopes (δ 15 N) as an index of the trophic level in a benthic fish and invertebrate size-spectrum, and demonstrated that body mass was positively and significantly related to trophic level. This pattern was consistent with the expectation that predator-prey relationships led to powerful size-based trophic structuring in marine communities and ecosystems. Further analysis of intra-and interspecific relationships between body mass and trophic level in the community showed that increases in trophic level across the size spectrum were predominantly a consequence of intra-specific increases in trophic level with body mass and not a consequence of larger species (species with greater maximum body mass) feeding at higher trophic levels. We confirmed the absence of strong inter-specific relationships between maximum body mass and trophic level with cross-species and phylogenetic comparative approaches. Size-based models are easier and cheaper to parameterise than most food-web models. Subject to the persistence of relationships between body mass and trophic level in space and time, our results suggest that size spectra could be parameterised with body mass-trophic level relationships and used to describe the trophic structure of some marine communities and ecosystems.KEY WORDS: Size-spectra · Trophic level · Food web · Community structure · Stable isotopes · PhylogenyResale or republication not permitted without written consent of the publisher