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1. Single-station diel oxygen curves were used to monitor the oxygen metabolism of an intermittent, forested third-order stream (Fuirosos) in the Mediterranean area, over a period of 22 months. Ecosystem respiration (ER) and gross primary production (GPP) were estimated and related to organic matter inputs and photosynthetically active radiation (PAR) in order to understand the effect of the riparian forest on stream metabolism. 2. Annual ER was 1690 g O 2 m )2 year )1 and annual GPP was 275 g O 2 m )2 year )1 . Fuirosos was therefore a heterotrophic stream, with P : R ratios averaging 0.16. 3. GPP rates were relatively low, ranging from 0.05 to 1.9 g O 2 m )2 day )1 . The maximum values of GPP occurred during a few weeks in spring, and ended when the riparian canopy was fully closed. The phenology of the riparian vegetation was an important determinant of light availability, and consequently, of GPP. 4. On a daily scale, light and temperature were the most important factors governing the shape of photosynthesis-irradiance (P-I) curves. Several patterns could be generalised in the P-I relationships. Hysteresis-type curves were characteristic of late autumn and winter. Light saturation responses (that occurred at irradiances higher than 90 lE m )2 s )1 ) were characteristic of early spring. Linear responses occurred during late spring, summer and early autumn when there was no evidence of light saturation. 5. Rates of ER were high when compared with analogous streams, ranging from 0.4 to 32 g O 2 m )2 day )1 . ER was highest in autumn 2001, when organic matter accumulations on the streambed were extremely high. By contrast, the higher discharge in autumn 2002 prevented these accumulations and caused lower ER. The Mediterranean climate, and in its effect the hydrological regime, were mainly responsible for the temporal variation in benthic organic matter, and consequently of ER.

Low current velocities, high nutrient levels, the lack of riparian forest vegetation, and the development of dense and rich macrophyte communities characterize Pampean streams. The objective of this study was to describe the main physical, chemical, and biological characteristics of a headwater Pampean stream as well as to analyze the role of macrophytes and phytobenthos. The study was conducted in a stream considered to be not much disturbed by human activities. Samples of water and organisms (macrophytes, benthic algae and invertebrates) were taken monthly for 14 months in two sampling stations, in fast flow and slow flow sites. Macrophyte biomass and diversity increased in spring and summer, and they decreased in autumn, when the plant community was greatly affected by an important flood. Phytobenthos biomass was lower in late summer, possibly due to the establishment of a dense cover of the floating macrophyte Lemna gibba L. Density of amphipods and gastropods greatly increases in spring and summer, jointly with the macrophyte development. Analysis of correlation showed that current velocity is the most important factor influencing macrophyte biomass and phytobenthos structure, while depth, nutrients, and herbivores are linked factors. Pampean streams could be considered systems dynamically fragile, because habitat heterogeneity is generated by aquatic vegetation, a substratum that varies along time.

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Environmental heterogeneity in natural ecosystems influences several parameters at the population and community levels. In freshwater ecosystems, habitat heterogeneity can be provided by macrophyte species with different structural shapes. Previous studies suggest that aquatic plants with more complex architectures will support higher number, biomass, and taxon richness of macroinvertebrates than plants with simpler shape. We investigated the influence of macrophyte structural heterogeneity (quantified by fractal dimension) and food availability (represented by epiphytic biomass) on several parameters (number of individuals, biomass, body size distribution, taxon richness, and diversity) of the macroinvertebrate community in a Pampean stream. Four submerged macrophyte species (Egeria densa, Elodea ernstae, Ceratophyllum demersum, and Stuckenia striata) and associated macroinvertebrates were sampled in late spring, summer, and autumn.

We investigated the effect of benthic substratum type (sand and rocks) and nutrient supply (N and P) on biofilm structure and heterotrophic metabolism in a field experiment in a forested Mediterranean stream (Fuirosos). Rock and sand colonization and biofilm formation was intensively studied for 44 d at two stream reaches: control and experimental (continuous addition of phosphate, ammonia, and nitrate). Structural (C, N, and polysaccharide content and bacterial and chlorophyll density) and metabolic biofilm parameters (␤-glucosidase, peptidase, and phosphatase enzyme activities) were analyzed throughout the colonization process. The epilithic biofilm (grown on rocks) had a higher peptidase activity at the impacted reach, together with a higher algal and bacterial biomass. The positive relationship between the peptidase activity per cell and the N content of the epilithic biofilm suggested that heterotrophic utilization of proteinaceous compounds from within the biofilm was occurring. In contrast, nutrient addition caused the epipsammic biofilm (grown on sand) to exhibit lower ␤-glucosidase and phosphatase activities, without a significant increase in bacterial and algal biomass. The differential response to nutrient addition was related to different structural characteristics within each biofilm. The epipsammic biofilm had a constant and high C : N ratio (22.7) throughout the colonization. The epilithic biofilm had a higher C : N ratio at the beginning of the colonization (43.2) and evolved toward a more complex structure (high polysaccharide content and low C : N ratio) during later stages. The epipsammic biofilm was a site for the accumulation and degradation of organic matter: polysaccharides and organic phosphorus compounds had higher degradation activities.Microorganisms play a key role in the degradation of organic matter and the associated release of energy in stream ecosystems (Meyer 1994). Microbial communities feed on allochthonous (dissolved and particulate plant and animal materials) and/or autochthonous organic matter (algal detritus and exudates) and are the main factor responsible for river C cycling (Allan 1995). In low-order streams, the benthic community is the most important site for organic matter cycling. In most cases, rock, cobbles, sand, and wood coexist in a stream reach, and all of these substrata host biofilms with differing structural characteristics (Lock 1993). In association with these structural differences, there are also significant differences in photosynthetic activity, biofilm thick- AcknowledgmentsWe thank two anonymous reviewers for their helpful suggestions about the manuscript. Comments by E. Garcia Berthou and C. Freeman are very much appreciated.

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Stream metabolism at both ecosystem and functional-compartment scales was measured in a low-order Pampean stream (La Choza) over a 3-wk period to characterize metabolic rates and discern the contribution of each functional compartment (submerged macrophytes, benthos, floating macroalgae, water column, and hyporheic zone) to ecosystem metabolism. La Choza stream is an autotrophic ecosystem during low flows and has gross primary production rates of up to 22 g O 2 m 22 d 21 , which are among the highest reported in the literature and set an upper bound on how productive streams can be in the absence of light and nutrient limitations. Floating macroalgae provided most of the primary production (30-90%), whereas the hyporheic zone provided most of the ecosystem respiration (40-80%). The differential effects of high flows on the different functional compartments depressed the production:respiration ratio, suggesting a strong relationship between flow and metabolism. Thus, low flows enhanced primary production and led to diel dissolved O 2 concentration oscillations between 0 and 25 g O 2 /m 3 . In contrast, high flow depressed primary production by an order of magnitude and increased ecosystem respiration. High production rates during the low-flow period and extreme physicochemical conditions (anoxia for 7-8 h on a daily basis) may be typical in this type of ecosystem during extended low-flow periods.