River health monitoring traditionally has made use of structural measurements (water quality or taxonomic composition of aquatic organisms). We argue that a more complete assessment of river health should include functional metrics, such as rates of organic matter decomposition and ecosystem metabolism. Leaf breakdown links the characteristics of riparian vegetation with the activity of both aquatic invertebrates and microbial organisms and is affected by natural and human-induced variation in a wide range of environmental factors. Measurement of leaf breakdown is relatively simple and has modest equipment requirements. River metabolism (gross primary productivity and ecosystem respiration) measures the rates of production and use of organic C in river ecosystems as a whole, providing a direct estimate of the food base that determines life-supporting capacity. Metabolism measurements require more sophisticated equipment than do measurements of leaf breakdown, but improvements in technology have made metabolism measurements relatively easy. We review the factors that influence leaf breakdown and river metabolism and pay particular attention to the effects of human-induced environmental stressors. We also describe how measurements can be standardized and suggest criteria for interpreting functional measures in terms of river ecosystem health. Last, we consider the strengths and weaknesses of both methods as functional measures and provide recommendations for their use as biomonitoring tools.

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Organic matter breakdown and ecosystem metabolism: functional indicators for assessing river ecosystem health

SUMMARY 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

Ecological stoichiometry in freshwater benthic systems: recent progress and perspectives

The decomposition of plant litter is one of the most important ecosystem processes in the biosphere and is particularly sensitive to climate warming. Aquatic ecosystems are well suited to studying warming effects on decomposition because the otherwise confounding influence of moisture is constant. By using a latitudinal temperature gradient in an unprecedented global experiment in streams, we found that climate warming will likely hasten microbial litter decomposition and produce an equivalent decline in detritivore-mediated decomposition rates. As a result, overall decomposition rates should remain unchanged. Nevertheless, the process would be profoundly altered, because the shift in importance from detritivores to microbes in warm climates would likely increase CO2 production and decrease the generation and sequestration of recalcitrant organic particles. In view of recent estimates showing that inland waters are a significant component of the global carbon cycle, this implies consequences for global biogeochemistry and a possible positive climate feedback.

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A global experiment suggests climate warming will not accelerate litter decomposition in streams but might reduce carbon sequestration

Feeding strategies are typical traits reflecting the adaptation of species to environmental conditions. This concept is currently developed in some water quality systems (e.g. Index of Trophic completeness) and the structure of functional feeding groups (FFGs) could form part of a unified measure across communities differing in taxonomic composition. However, in South America, information about the FFG classification of invertebrates in streams is almost absent and existing studies using FFG structure follows classification from North America. But even taxonomically related species may have different diets in tropical and temperate areas and therefore, studies about FFG structure in neotropics could be biased. For this reason, we determined diet composition, trophic level and FFGs, using gut contents analysis and mouthpart observations of 49 macroinvertebrate taxa (mostly at genus level) from neotropical streams. We observed that practically all macroinvertebrates fed upon fine detritus which indicates the importance of this food resource in neotropical streams. As the assignment to a single FFG does not accurately reflect the functional profile of taxa, we transcribed the affinity of taxa to each FFG using fuzzy codes. Finally, we published the coding of diet composition and FFG of the taxa examined, which could be used in future community analyses of lotic ecosystems in the Neotropical zone.

Trophic levels and functional feeding groups of macroinvertebrates in neotropical streams

Food traceability: New trends and recent advances. A review

1. The importance of leaf quality to the nutritional ecology of lotic shredders is well established for temperate species but virtually unknown for tropical taxa. In the present study, we compared the feeding behaviour and performance of two tropical and two temperate shredders in a series of pair-wise experiments.



2. Specifically, we tested whether leaf conditioning status (stream-conditioned versus unconditioned leaves) and geographical origin (temperate Alnus glutinosa versus tropical Hura crepitans leaves) affect the food preference, survivorship, and growth of selected shredders from low and high latitudes in a consistent manner. The animals used in experiments were the caddis-flies Nectopsyche argentata and Phylloicus priapulus from Venezuela, Sericostoma vittatum from Central Portugal, and the amphipod Gammarus pulex from Northern Germany.



3. In general, all shredders exhibited the same high preference for conditioned over unconditioned leaves, irrespective of the geographical origin of the leaf or shredder species.



4. A corresponding tendency for higher growth was found for sets of animals offered conditioned leaves, with the differences in growth being clearer in the two tropical shredders. Survivorship of the two temperate species was consistently high (> 83%) regardless of the diet offered, whereas the tropical shredders survived better on conditioned (77–90%) as compared with unconditioned (54–87%) leaves, although not significantly so.



5. With the exception of the temperate S. vittatum, shredders did not select or perform better on leaves to which they had previously been exposed, indicating a potential adaptation to native leaf species is over-ridden by intrinsic leaf properties.



6. Taken together, the results of this study suggest that tropical shredders may exhibit the same basic patterns of food exploitation as their temperate counterparts. Consequently, current concepts relating to the role of shredders in stream detritus dynamics may well be applicable to tropical streams, although essentially derived from temperate systems.

Barrios

Papers

Food quality, feeding preferences, survival and growth of shredders from temperate and tropical streams