The subsurface riparian zone was examined as an ecotone with two interfaces. Inland is a terrestrial boundary, where transport of water and dissolved solutes is toward the channel and controlled by watershed hydrology. Streamside is an aquatic boundary, where exchange of surface water and dissolved solutes is bi-directional and flux is strongly influenced by channel hydraulics. Streamside, bi-directional exchange of water was qualitatively defined using biologically conservative tracers in a third order stream.In several experiments, penetration of surface water extended 18 m inland. Travel time of water from the channel to bankside sediments was highly variable. Subsurface chemical gradients were indirectly related to the travel time. Sites with long travel times tended to be low in nitrate and DO (dissolved oxygen) but high in ammonium and DOC (dissolved organic carbon). Sites with short travel times tended to be high in nitrate and DO but low in ammonium and DOC. Ammonium concentration of interstitial water also was influenced by sorption-desorption processes that involved clay minerals in hyporheic sediments. Denitrification potential in subsurface sediments increased with distance from the channel, and was limited by nitrate at inland sites and by DO in the channel sediments. Conversely, nitrification potential decreased with distance from the channel, and was limited by DO at inland sites and by ammonium at channel locations. Advection of water and dissolved oxygen away from the channel resulted in an oxidized subsurface habitat equivalent to that previously defined as the hyporheic zone. The hyporheic zone is viewed as stream habitat because of its high proportion of surface water and the occurrence of channel organisms. Beyond the channel's hydrologic exchange zone, interstitial water is often chemically reduced. Interstitial water that has not previously entered the channel, groundwater, is viewed as a terrestrial component of the riparian ecotone. Thus, surface water habitats may extend under riparian vegetation, and terrestrial groundwater habitats may be found beneath the stream channel.

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The influence of hydrogeologic setting on the susceptibility of streams to legacy nitrate was examined at seven study sites having a wide range of base flow index (BFI) values. BFI is the ratio of base flow to total streamflow volume. The portion of annual stream nitrate loads from base flow was strongly correlated with BFI. Furthermore, dissolved oxygen concentrations in streambed pore water were significantly higher in high BFI watersheds than in low BFI watersheds suggesting that geochemical conditions favor nitrate transport through the bed when BFI is high. Results from a groundwater-surface water interaction study at a high BFI watershed indicate that decades old nitrate-laden water is discharging to this stream. These findings indicate that high nitrate levels in this stream may be sustained for decades to come regardless of current practices. It is hypothesized that a first approximation of stream vulnerability to legacy nutrients may be made by geospatial analysis of watersheds with high nitrogen inputs and a strong connection to groundwater (e.g., high BFI).

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In extreme environments, retention of nutrients within stream ecosystems contributes to the persistence of aquatic biota and continuity of ecosystem function. In the McMurdo Dry Valleys, Antarctica, many glacial meltwater streams flow for only 5-12 weeks a year and yet support extensive benthic microbial communities. We investigated NO 3 Ϫ uptake and denitrification in Green Creek by analyzing small-scale microbial mat dynamics in mesocosms and reach-scale nutrient cycling in two whole-stream NO 3 Ϫ enrichment experiments. Nitrate uptake results indicated that microbial mats were nitrogen (N)-limited, with NO 3 Ϫ uptake rates as high as 16 nmol N cm Ϫ2 h Ϫ1. Denitrification potentials associated with microbial mats were also as high as 16 nmol N cm Ϫ2 h Ϫ1. During two whole-stream NO 3 Ϫ -enrichment experiments, a simultaneous pulse of NO 2 Ϫ was observed in the stream water. The one-dimensional solute transport model with inflow and storage was modified to simulate two storage zones: one to account for short time scale hydrologic exchange of stream water into and out of the benthic microbial mat, the other to account for longer time scale hydrologic exchange with the hyporheic zone. Simulations indicate that injected NO 3 Ϫ was removed both in the microbial mat and in the hyporheic zone and that as much as 20% of the NO 3 Ϫ that entered the microbial mat and hyporheic zone was transformed to NO 2 Ϫ by dissimilatory reduction. Because of the rapid hydrologic exchange in microbial mats, it is likely that denitrification is limited either by biotic assimilation, reductase limitation, or transport limitation (reduced NO 2 Ϫ is transported away from reducing microbes).Dry valley glacial meltwater streams are simple physical systems, ideal for studying stream-hyporheic biogeochemical interactions. There is no contribution to stream flow from precipitation (either directly or from snowmelt), and there are no hillslope additions of water or solutes. Further, the hyporheic zones are well defined, bounded at depths of 0.4 to 0.7 m by permafrost, extending several meters from stream edges . In these streams, hyporheic zone exchange and biotic assimilation are important controls on biogeochemical cycling. At the mineral grain scale, Maurice et al. (2002) recently documented weathering processes occurring in the hyporheic zone of a dry valley stream at rates higher than typically found in temperate watersheds, as well as the presence of denitrifying bacteria. Similarly, at the reach scale, Gooseff et al. (2002) have

Nutrient recycling by animals is a potentially important biogeochemical process in both terrestrial and aquatic ecosystems. Stoichiometric traits of individual species may result in some taxa playing disproportionately important roles in the recycling of nutrients relative to their biomass, acting as keystone nutrient recyclers. We examined factors controlling the relative contribution of 12 Neotropical fish species to nutrient recycling in four streams spanning a range of phosphorus (P) levels. In high-P conditions (135 microg/L soluble reactive phosphorus, SRP), most species fed on P-enriched diets and P excretion rates were high across species. In low-P conditions (3 microg/L SRP), aquatic food resources were depleted in P, and species with higher body P content showed low rates of P recycling. However, fishes that were subsidized by terrestrial inputs were decoupled from aquatic P availability and therefore excreted P at disproportionately high rates. One of these species, Astyanax aeneus (Characidae), represented 12% of the total population and 18% of the total biomass of the fish assemblage in our focal low-P study stream but had P excretion rates > 10-fold higher than other abundant fishes. As a result, we estimated that P excretion by A. aeneus accounted for 90% of the P recycled by this fish assemblage and also supplied approximately 90% of the stream P demand in this P-limited ecosystem. Nitrogen excretion rates showed little variation among species, and the contribution of a given species to ecosystem N recycling was largely dependent upon the total biomass of that species. Because of the high variability in P excretion rates among fish species, ecosystem-level P recycling could be particularly sensitive to changes in fish community structure in P-limited systems.

Denitrification was assayed by the acetylene blockage technique in hyporheic sediments. Samples were obtained along transects perpendicular to the stream at two sites: (1) the base of a slope dominated by old-growth redwood and (2) the base of a slope dominated by alder regenerating from a clearcut in 1965. Denitrification was evident at in situ nitrate concentrations at all locations tested. Activity was stimulated by nitrate but nitrate plus glucose had no additional effect. Denitrifying potentials increased with increasing distance from the stream channel. Dissolved oxygen was 100% of the concentration expected in equilibrium with the atmosphere in water obtained from monitoring wells immediately adjacent to the stream but was as low as 7% of the expected value in water 11.4 m inland. Both nitrate and dissolved organic carbon decreased over summer in wells at the base of the alder-forested slope. A 48-h injection of nitrate-amended stream water into hyporheic water 8.4 m inland stimulated nitrous oxide production in the presence of acetylene. Nitrous oxide was generated as nitrate and acetylene were co-transported to a well 13 m down-gradient. The acetylene-block experiments coupled with the chemistry data suggest that denitrification can modify the chemistry of water during passage through the hyporheic zone.