1. Benthic algal communities are shaped by the availability of nutrients and light and by herbivore consumption. Many studies have examined how one of these factors affects algal communities, but studies simultaneously addressing all three are rare. 2. We investigated the effects of nutrients, light and a herbivore (the snail Potamopyrgus antipodarum) on benthic stream algae in a fully factorial experiment in 128 circular streamside channels. Four nutrient levels (none added to highly enriched), four snail grazing pressures (no snails to 777 individuals m )2 ) and two light levels (ambient and 65% reduced) were applied. Colonising algae were dominated by diatoms (Bacillariophyta), which were determined to species using acid-cleaned samples and assigned to functional groups according to their physiognomic growth form. 3. Diatom community structure changed considerably in response to our manipulations. Light had the strongest influence (as indicated by M AN O V A M AN O V A effect size), whereas nutrients had an intermediate effect and grazing was fairly weak. Relative abundances of six common diatom taxa decreased under reduced light, whereas five others became more prevalent. Eight taxa benefitted from nutrient enrichment, while three became rarer. Grazing affected the relative density of only one common taxon, which increased at higher grazing pressure. 4. Diatom functional groups also responded strongly. 'Low profile' taxa dominated at low resource levels (nutrients and especially light), whereas 'high profile' and 'motile' taxa became markedly more prevalent at higher resource levels. 5. Two-way interactions between experimental factors were quite common. For example, Planothidium lanceolatum and Rossithidium petersenii responded more strongly to nutrient enrichment at reduced than at ambient light, whereas Cocconeis placentula responded more strongly at ambient light. For diatom functional groups, the benefit of nutrient enrichment for 'motile' diatoms was greater at ambient than at reduced light. 6. Our results imply that multifactor experiments are required to determine the main forces driving the composition of benthic algal communities. Further, our findings highlight the considerable potential of using functional algal groups as indicators of changing environmental conditions to complement the traditional taxonomic approach.

2005. Toward a stoichiometric framework for evolutionary biology. Á/ Oikos 109: 6 Á/17.Ecological stoichiometry, the study of the balance of energy and materials in living systems, may serve as a useful synthetic framework for evolutionary biology. Here, we review recent work that illustrates the power of a stoichiometric approach to evolution across multiple scales, and then point to important open questions that may chart the way forward in this new field. At the molecular level, stoichiometry links hereditary changes in the molecular composition of organisms to key phenotypic functions. At the level of evolutionary ecology, a simultaneous focus on the energetic and material underpinnings of evolutionary tradeoffs and transactions highlights the relationship between the cost of resource acquisition and the functional consequences of biochemical composition. At the macroevolutionary level, a stoichiometric perspective can better operationalize models of adaptive radiation and escalation, and elucidate links between evolutionary innovation and the development of global biogeochemical cycles. Because ecological stoichiometry focuses on the interaction of energetic and multiple material currencies, it should provide new opportunities for coupling evolutionary dynamics across scales from genomes to the biosphere.

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Summary 1.Productivity and grazing pressure interact in determining autotroph diversity, because high productivity increases the capability of a plant community to compensate for grazing losses. However, further factors may play a role in shaping diversity, including primary producer nutrient stoichiometry and grazer activity. 2. Our study focuses on the interactions between light, nutrients and grazing in determining species richness and evenness of stream diatoms. By measuring primary producer productivity and nutrient content as well as grazer activity, we attempt to disentangle the different pathways by which the three factors affect diatom species richness and evenness. 3. We hypothesized that high light intensities and nutrient addition would increase species richness by increasing primary productivity and that higher levels of light and nutrients would compensate for negative grazer effects on species richness of primary producers. We also hypothesized that high light intensities would decrease the nutrient content of primary producers, especially when nutrients are limiting, whereas nutrient addition would increase primary producer nutrient content. Last, in addition to changing primary producer nutrient content, light and nutrients would also change grazer activity, thus modifying the interactions between light, nutrients and grazing. 4. We used periphyton and gastropod grazers in an experiment with circular stream channels with four nutrient, two light and four grazing levels to determine individual and combined effects on benthic diatom richness and evenness. After 3 weeks, we determined algal biomass, periphyton nutrient content, diatom species richness and evenness as well as grazer activity. 5. Our results showed that light and nutrients increased species richness and primary producer productivity and nutrient content. Grazing decreased species richness but only at low light levels, possibly because high light levels reduced grazer activity. Evenness was not affected by any single factor alone, but was influenced by nutrient-light and grazing-light interactions. 6. Synthesis. Light, nutrients and grazing interacted in determining primary producer species richness. Their effects were mainly mediated through changes in productivity but primary producer nutrient content and grazer activity also played important roles.

Summary1. High-latitude species (and populations within species) are adapted to short and cold summers. They often have high growth and development rates to fully use the short growing season and mature before the onset of winter. 2. Within the context of ecological stoichiometry theory, this study combines ecology with evolution by relating latitudinal life-history adaptations to their molecular consequences in body nutrient composition in Rana temporaria tadpoles. 3. Temperature and food quality were manipulated during the development of tadpoles from Arctic and Boreal origins. We determined tadpole growth rate, development rate, body size and nutrient content, to test whether (i) Arctic tadpoles could realize higher growth rates and development rates with the help of higher-quality food even when food quantity was unchanged, (ii) Arctic and Boreal tadpoles differed in their stoichiometric (and life history) response to temperature changes, (iii) higher growth rates lead to higher tadpole P content (growth rate hypothesis) and (iv) allometric scaling affects tadpole nutrient allocation. 4. We found that especially Arctic tadpoles grew and developed faster with the help of higher-quality food and that tadpoles differed in their stoichiometric (and life history) response to temperature changes depending on region of origin (probably due to different temperature optima). There was no evidence that higher growth rates mediated the positive effect of temperature on tadpole P content. On the contrary, the covariate growth rate was negatively connected with tadpole P content (refuting the growth rate hypothesis). Lastly, tadpole P content was not related to body size, but tadpole C content was higher in larger tadpoles, probably due to increased fat storage. 5. We conclude that temperature had a strong effect on tadpole life history, nutrient demand and stoichiometry and that this effect depended on the evolved life history.

A. 2004. Effects of macrograzers and light on periphyton stoichiometry. Á/ Oikos 106: 93 Á/104.Ecological stoichiometry describes the biochemical constraints of trophic interactions emerging from the different nutrient content and nutrient demand of producers and consumers, respectively. Most research on this topic originates from well-mixed pelagic food webs, whereas the idea has received far less attention in spatially structured habitats. Here, we test how light as well as grazing and nutrient regeneration by consumers affects growth and biomass of benthic primary producers. In the first laboratory experiment, we manipulated grazer presence (two different snail species plus ungrazed control), in the second experiment we factorially combined manipulation of grazer presence and light intensity. We monitored snail and periphyton biomass as well as dissolved and particulate nutrients (nitrogen and phosphorus) over time. Grazers significantly reduced algal biomass in both experiments. Grazers affected periphyton nutrient content depending on the prevailing nutrient limitation and their own body stoichiometry. In the nitrogen (N-) limited first experiment, grazers increased N both in the periphyton and in the water column. The effect was stronger for grazers with lower N-content. In the phosphorus (P-) limited second experiment, grazers increased the Pcontent of the periphyton, but the grazer with lower N-content had additionally positive effects on algal N. Light reduction did not affect periphyton biomass, but increased chlorophyll-, N-and P-content of the periphyton. These experiments revealed that the indirect effects of grazers on periphyton were bound by stoichiometric constraints of nutrient incorporation and excretion.

The turnover and distribution of energy and nutrients in food webs is influenced by consumer stoichiometry. Although the stoichiometry of heterotrophs is generally considered to vary only little, there may be intraspecific variation due to factors such as habitat, resources, ontogeny and size. We examined intraspecific variation in Eurasian perch Perca fluviatilis stoichiometry, a common species that exhibits habitat and resource specialization, ontogenetic niche shifts and a large size range. This study investigated the elemental stoichiometry of a wide size range of perch from littoral and pelagic habitats. The mean C:N:P stoichiometry of whole perch was 37:9:1 (molar ratios). However, %C, %P, C:N, C:P and N:P varied with size, morphology, habitat and diet category. These factors together explained 24–40% of the variation in C:N:P stoichiometry. In contrast, perch stoichiometry was not related to diet stoichiometry, suggesting that the former is homeostatically regulated. The results suggest that the high P content of perch may result in stoichiometric constraints on the growth of non‐piscivorous perch, and that piscivory is an efficient strategy for acquiring P. Resource polymorphism, individual diet specialization and intraspecific size variation are widespread among animals. Thus changes in stoichiometry with size, habitat, morphology and resource use, and therefore also stoichiometric demands, are probably common.