Nitrate depletion in the riparian zone and stream channel of a small headwater catchment

A mass balance procedure was used to determine rates of nitrate depletion in the riparian zone and stream channel of a small New Zealand headwater stream. In all 12 surveys the majority of nitrate loss (56–100%) occurred in riparian organic soils, despite these soils occupying only 12% of the stream's border. This disproportionate role of the organic soils in depleting nitrate was due to two factors. Firstly, they were located at the base of hollows and consequently a disproportionately high percentage (37–81%) of the groundwater flowed through them in its passage to the stream. Secondly, they were anoxic and high in both denitrifying enzyme concentration and available carbon. Direct estimates ofin situ denitrification rate for organic soils near the upslope edge (338 mg N m^–2 h^–1) were much higher than average values estimated for the organic soils as a whole (0.3–2.1 mg N m^–2 h^–1) and suggested that areas of these soils were limited in their denitrification activity by the supply of nitrate. The capacity of these soils to regulate nitrate flux was therefore under-utilized. The majority of stream channel nitrate depletion was apparently due to plant uptake, with estimates of thein situ denitrification rate of stream sediments being less than 15% of the stream channel nitrate depletion rate estimated by mass balance.This study has shown that catchment hydrology can interact in a variety of ways with the biological processes responsible for nitrate depletion in riparian and stream ecosystems thereby having a strong influence on nitrate flux. This reinforces the view that those seeking to understand the functioning of these ecosystems need to consider hydrological phenomena.     

Nitrate depletion in the riparian zone of a small woodland stream

Field enrichments with nitrate in two spring-fed drainage lines within the riparian zone of a small woodland stream near Toronto, Ontario showed an absence of nitrate depletion. Laboratory experiments with riparian substrates overlain with nitrate enriched solutions revealed a loss of only 5–8% of the nitrate during 48 h incubation at 12°C. However, 22–24% of the initial nitrate was depleted between 24 and 48 h when a second set of substrate cores was incubated at 20°C. Short-term (3 h) incubations of fresh substrates amended with acetylene were used to estimate in situ denitrification potentials which varied from 0.05–3.19 g N g^–1 d^–1 for organic and sandy sediments. Denitrification potentials were highly correlated with initial nitrate content of substrate samples implying that low nitrate levels in ground water and riparian substrates may be an important factor in controlling denitrification rates. The efficiency of nitrate removal in spring-fed drainage lines is also limited by short water residence times of < 1 h within the riparian zone. These data suggest that routes of ground water movement and substrate characteristics are important in determining nitrate depletion within stream riparian areas.     

Nitrogen mass balances and denitrification rates in central Ontario Lakes

Nitrogen mass balances for seven unproductive lakes and 20 forested catchments in central Ontario were measured between 1977 and 1989. Average annual lake denitrification rates calculated with the N/P ratio method were strongly correlated with summer anoxic factor (extent of surficial sediment anoxia) whereas denitrification rates calculated with a²¹⁰Pb sediment N accumulation method were poorly correlated with the anoxic factor suggesting that the N/P method is superior. Substantial denitrification occurred in all lakes — an average of 36% of TN inputs or 75% of the net gain. On a regional area-weighted basis, 67% of bulk atmospheric TN deposition was stored or denitrified terrestrially, 12% was denitrified in lakes, 4% was stored in lake sediments, and 17% was exported from lakes. N/P ratios were generally less in streams than in precipitation suggesting preferential N retention in catchments, whereas the N/P ratios in lake outputs were slightly higher than lake input ratios, suggesting preferential P retention in lakes. This is consistent with the notion that P-limited lakes can exist adjacent to N-limited forests.     

Nitrogen cycling and dynamics in a macrophyte‐rich stream as determined by a release

1. A tracer release study was conducted in a macrophyte‐rich stream, the River Lilleaa in Denmark. The objectives of the study were to compare uptake rates per unit area of by primary producers and consumers in macrophyte and non‐macrophyte habitats, estimate whole‐stream uptake rates of and compare this to other stream types, and identify the pathways and estimate the rate at which enters the food web in macrophyte and non‐macrophyte habitats. 2. Macrophyte habitats had four times higher primary uptake rates and an equal uptake rate by primary consumers per unit habitat area as compared to non‐macrophyte habitats. These rates represent the lower limit of potential macrophyte effects because the rates will be highly dependent on macrophyte bed height and mean bed height in the River Lilleaa was low compared to typical bed heights in many lowland streams. Epiphytes accounted for 30% of primary uptake in macrophyte habitats, illustrating a strong indirect effect of macrophytes as habitat for epiphytes. N flux per unit habitat area from primary uptake compartments to primary consumers was four times lower in macrophyte habitats compared to non‐macrophyte habitats, reflecting much greater biomass accrual in macrophyte habitats. Thus, we did not find higher N flux from macrophyte habitats to primary consumers compared to non‐macrophyte habitats. 3. Whole‐stream uptake rate was 447 mgN m^?2 day^?1. On a habitat‐weighted basis, fine benthic organic matter (FBOM) accounted for 72% of the whole‐stream uptake rate, and macrophytes and epiphytes accounted for 19 and 8%, respectively. 4. We had expected a priori relatively high whole‐stream N uptake in our study stream compared to other stream types mainly due to generally high biomass and the macrophyte’s role as habitat for autotrophic and heterotrophic organisms, but our results did not confirm this. In comparison with other release study streams, we conclude that nutrient concentration is the overall controlling factor for N uptake rates across streams, mostly as a result of high biomass of primary uptake compartments in streams with high nutrient concentrations in general and not in macrophyte streams in particular. 5. Our results indicate that macrophytes play an important role in the longer‐term retention of N and thus a decrease in net downstream transport during the growing season compared to streams without macrophytes, through direct and indirect effects on the stream reach. Direct effects are high uptake efficiency, low turnover rate (partly due to no direct feeding on macrophytes) and high longevity. An indirect effect is increased sedimentation of FBOM in macrophytes compared to non‐macrophyte habitats and streams which possibly also increase denitrification. Increased retention with macrophyte presence would decrease downstream transport during the growing season and thus the N loading on downstream ecosystems.     

Benthic organic carbon influences denitrification in streams with high nitrate concentration

1. Anthropogenic activities have increased reactive nitrogen availability, and now many streams carry large nitrate loads to coastal ecosystems. Denitrification is potentially an important nitrogen sink, but few studies have investigated the influence of benthic organic carbon on denitrification in nitrate‐rich streams. 2. Using the acetylene‐block assay, we measured denitrification rates associated with benthic substrata having different proportions of organic matter in agricultural streams in two states in the mid‐west of the U.S.A., Illinois and Michigan. 3. In Illinois, benthic organic matter varied little between seasons (5.9–7.0% of stream sediment), but nitrate concentrations were high in summer (>10 mg N L⁻¹) and low (<0.5 mg N L⁻¹) in autumn. Across all seasons and streams, the rate of denitrification ranged from 0.01 to 4.77 μg N g⁻¹ DM h⁻¹ and was positively related to stream‐water nitrate concentration. Within each stream, denitrification was positively related to benthic organic matter only when nitrate concentration exceeded published half‐saturation constants. 4. In Michigan, streams had high nitrate concentrations and diverse benthic substrata which varied from 0.7 to 72.7% organic matter. Denitrification rate ranged from 0.12 to 11.06 μg N g⁻¹ DM h⁻¹ and was positively related to the proportion of organic matter in each substratum. 5. Taken together, these results indicate that benthic organic carbon may play an important role in stream nitrogen cycling by stimulating denitrification when nitrate concentrations are high.     

Nitrogen transformations in a small mountain stream

Ammonium, urea, and nitrate were added to Bear Brook, a second and third order stream in the Hubbard Brook Experimental Forest, New Hampshire. Removal of ammonium and urea during downstream transport coincided with the release of nitrate. Nitrate removal did not occur when it was added alone or with dissolved organic carbon. Laboratory experiments showed that coarse particulate organic material (detritus) and bryophytes taken from the streambed were active in the removal of ammonium from enriched stream water, and in the release of nitrate upon the addition of ammonium.The patterns of removal and release observed in these experiments suggest a biologically mediated, oxidation process. Budgetary calculations show that the in-stream transformation of nitrogen inputs during summer and autumn could represent 12 to 25 percent of the nitrogen exported as nitrate during winter and spring from heterotrophic streams like Bear Brook. This type of internal cycling affects the timing and form of nitrogen export from small streams draining forested watersheds in the northeastern United States.     

Nitrate transformation and water movement in a wetland area

The NO₃ ^– transformation capacity of a riparian zone at Rabis stream, Denmark, was investigated for a period of 2 years. The riparian zone of 15–25 m received NO₃ ^–-containing groundwater from the adjoining agricultural areas. The water flows as surface runoff along the surface of the wetland and in the root zone towards the stream. Changes in water chemistry, water balance and mass transport were investigated. The riparian zone acted as a buffer zone for NO₃ ^–, PO₄ ^3– and dissolved Fe²⁺. The NO₃ ^–-transformation capacity of the wetland was about 400 kg N ha^–1 y^–1, but varied seasonally. A simple rearrangement of drain systems in wetland areas can probably reduce the NO₃ ^– content of Danish surface waters by 20 000–50 000 t N y^–1.     

Size-mass relationships of stream invertebrates in a northern Spain stream

We show the regression equations that relate the head width of 10 freshwater insect species or the length of the first thoracic segment of 2 amphipods with their dry mass in 3 reaches along a small stream in northern Spain. Spatial differences in size–mass relationships were found. Organisms stored in alcohol showed lower dry mass than those specimens that were dried immediately after collection. The use of both size measures for dry mass estimation in field samples is discussed. It is recommended that the size–mass relationships be constructed using non-preserved organisms. Furthermore, great care is required when such allometric equations are taken from the literature.     

Longitudinal distribution of macroinvertebrate assemblages in a glacially influenced stream system in the Italian Alps

1. The longitudinal distribution of macroinvertebrates was investigated in June, August and September 1996 and 1997 in the Conca glacial stream and its tributary (Italian Alps; 46°N, 10°E). The principal aim was to test the 22 model that predicts the succession of faunal groups downstream of the glacial snout in relation to water temperature and channel stability. The effect of a non‐glacial tributary on the taxonomic richness and density patterns occurring in the glacial stream was also considered. 2. Channel stability showed an atypical longitudinal trend in the Conca glacial stream, being high in the upper part with Pfankuch Index values between 30 and 33. Water temperature exceeded 6 °C at all stations, with average values below 2 °C occurring only within 700 m from the glacial snout. 3. Taxonomic richness and diversity increased downstream. Taxonomic richness in the glacial stream (at about 1.5 km from the glacier) was comparable with the tributary and the reach after the confluence. Abundance also increased downstream in the glacial stream, but not as greatly as the number of taxa. 4. At higher taxonomic levels, the community structure in the tributary stations appeared to be similar to the two stations in the glacial stream just upstream of the confluence. The effect of the tributary was evident mainly at the genus or species level of the Chironomidae community. Some taxa found in the non‐glacial stream (e.g. Cricotopus fuscus, Eukiefferiella coerulescens, Metriocnemus sp., Paratrichocladius rufiventris, P. skirwitensis, Rheocricotopus effusus and Smittia sp.) were found also in the Conca stream but only after the confluence. 5. The upper glacial reach (within 700 m from the glacier snout) was dominated by the chironomid Diamesa spp. Less than 400 m from the glacier other Diamesinae (Pseudokiefferiella parva) and a few Orthocladiinae, especially Orthocladius (Euorthocladius) rivicola gr., colonized the stream. Some Diamesinae maintained relatively dense populations at mean water temperature around 5 °C, while some Orthocladiinae colonized reaches with mean water temperature <3 °C. 6. Contrary to the 22 model, Dipteran families such as Empididae and Limoniidae were more abundant in the upper stations than Simuliidae; non‐insects such as Nematoda and Oligochaeta were also numerous at some sites. Leuctridae, Taeniopterygidae and Nemouridae were the first Plecoptera to appear upstream, while Chloroperlidae were restricted to the lower reaches. Among Ephemeroptera, Heptageniidae were more abundant than Baetidae in the glacial sites. 7. In this glacial system channel stability and maximum temperature did not show the expected longitudinal trend and thus a typical kryal community was confined within 700 m from the glacier snout where summer mean water temperature was below 4 °C.     

Factors influencing periphyton growth in agricultural streams of central Illinois

Factors limiting periphyton accrual in east-central Illinois agricultural streams were investigated. Nutrient-diffusing substrata were used to examine periphyton macronutrient limitation in streams in two agricultural watersheds. Substrata consisted of sand-agar mixtures with one of six experimental treatments. Macronutrients included carbon, nitrate, phosphate and combinations of the three. Substrata were collected after a 5 and 9 day period and analyzed for chlorophyll a. None of the treatments were significantly greater than the controls at any of the seven stations, thus we conclude that periphyton in these streams was not nutrient limited. Highest periphyton colonization/growth rates were associated with the smaller upstream reaches, while lower rates occurred in the larger downstream reaches. Multiple regression showed that most of the variance in the rate of chlorophyll a accrual after five days was explained through water temperature and turbidity (r² = 0.91); whereas, stream nitrate and phosphate concentrations accounted for no significant portion of the variance. We conclude that instream primary production in agricultural streams of central Illinois is limited by temperature and light.     

Groundwater controls on nutrient cycling in a Mojave desert stream

1. Groundwater fluxes of nitrogen and dissolved organic carbon (DOC) were investigated in Grape Vine Canyon Stream in the Mojave Desert focusing on the rate of inputs and the fate of groundwater-derived nutrients in the stream. Discharge rates from different ground waters were measured using an end-member mixing model coupled with injections of a conservative solute tracer into the stream channel.
2. In surface water, nitrate concentration averaged 1.13 mg N L^–1 and DOC concentration averaged 1.82 mg C L^–1.
3. Groundwater discharge into Grape Vine Canyon Stream was derived from three sources. Nitrate concentration varied among the three groundwater sources with mean concentrations of 0.56, 0.94 and 0.08 mg N L^–1. DOC, in contrast, did not vary among ground water sources, with an overall average concentration of 2.96 mg C L^–1.
4. In the surface stream, nitrate concentration was two-fold greater than the concentration predicted from groundwater input, indicating that in-stream processes generated nitrate. Stream DOC concentration was lower than predicted based upon groundwater input rate. The production of nitrate and loss of DOC suggest that DOC is lost through mineralisation of dissolved organic matter, possibly resulting in the mineralisation of dissolved organic nitrogen to ammonium and subsequent transformation to nitrate via nitrification. In further support of this hypothesised linkage, DOC loss explained 80–89% of the variance in nitrate production in Grape Vine Canyon Stream.     

The role of ammonium and nitrate retention in the acidification of lakes and forested catchments

The relative contribution of HN0₃ to precipitation acidity in eastern Canada has increased in recent years leading to some concern that the relative importance of NO^– ₃ deposition in acidification of terrestrial and aquatic ecosystems may increase. To gauge the extent of this impact, annual mass balances for N0^– ₃ and NH^{+ 4} were calculated for several forested catchments and lakes in Ontario. Retention of NH⁺ ₄ (R _NH4) by forested catchments was consistently high compared to retention of NO₃ ^– (R _NO3) which was highly variable. Retention of inorganic nitrogen was influenced by catchment grade and areal water discharge. In lakes, the reciprocals of retention of N0^– ₃ and NH⁺ ₄ were linearly related to the ratio of lake mean depth to water residence time (z/; equal to areal water discharge), and retention did not appear to be a function of degree of acidification of the lakes. Net N consumption-based acidification of lakes, defined as the ratio of annual NH; mass to N0^– ₃ mass consumption, was negatively correlated with / and N consumption-related acidification was most likely to occur when – was < 1.5 m yr^–1.If retention mechanisms are unaffected by changes in deposition, changes in deposition will still result in changes in surface water concentrations although the changes will be of similar proportions. Therefore, NO^– ₃ saturation should not be defined by concentrations alone, but should be defined as decreasing long-term, average NO^– ₃ retention in streams and lakes in response to long-term increases in NO^– ₃ deposition. Analysis o f survey data will be facilitated by grouping lakes and catchments according to similar characteristics.     

Hierarchy,spatial configuration,and nutrient cycling in a desert stream

Abstract Recent studies of nutrient cycling in Sycamore Creek in Arizona, USA, suggest that a thorough understanding requires a spatially explicit, hierarchical approach. Physical configuration determines the path that water follows as it moves downstream. Water follows flowpaths through surface stream components, the hyporheic zone beneath the surface stream, and the parafluvial (sand bar) zone. Characteristic biogeochemical processes in these subsystems alter nitrogen (N) species in transport, in part as a function of available concentrations of N species. At several hierarchical levels, substrate materials are an important determinant of nitrogen dynamics in desert streams. Sand is present in bars of variable size and shape, each of which can be considered a unit, interacting with the surface stream. Groups of these stream-sandbar units form a higher level, the reach. At the next higher scale, sandy reaches (runs) alternate with riffles. Where flowpaths converge, rates of N transformation are high and, as a result, change in concentration is a non-linear function of flowpath length. Disturbance by flash floods alters sandbar configuration. Between floods, the interaction of subsurface and surface flowpaths shapes configuration in each, thus a self-organizing element of spatial structure exists. Sandy runs are dominated by subsurface processes and are likely to be net nitrifiers while riffles are dominated by surface flow and are nitrogen fixers. Whether a stream ecosystem retains nitrogen, or transports it to downstream recipient systems, or is a net emitter of gaseous forms of N, depends upon the dynamics of a spatial mosaic of interacting elements. An understanding of the net effect of this mosaic requires a spatially explicit, hierarchical, multi-scale approach.     

Macroinvertebrate stream communities along regional and physico-chemical gradients in Western Greenland

1. Macroinvertebrates were collected and physico‐chemical variables measured at 16 stream sites in Western Greenland during July 1999. Eight sites were located on Disko Island in an arctic oceanic climate and eight sites in the Kangerlussuaq area close to the icecap where the climate is arctic continental. The streams had different water sources (glacial, groundwater, snowmelt and lake water). 2. The streams showed pronounced differences in water temperature (2.2–17.3 °C), concentrations of suspended solids (0–2400 mg L^?1), and conductivity (10–109 μS cm^?1). Principal component analysis (PCA) analysis of the physico‐chemical variables separated the Disko Island sites into a distinct group, whereas the sites in the Kangerlussuaq area were more dispersed. 3. A total of 56 macroinvertebrate species were found, including 31 species of Chironomidae, the most abundant of which was Orthocladius thienemanni. Diamesa sp. was only the sixth most abundant chironomid taxon. Species composition varied between sites, and abundance varied from about 20 individuals m^?2 in a glacier fed stream to more than 16 000 m^?2 in a lake outlet. 4. The macroinvertebrate communities of the 16 streams were separated into five TWINSPAN groups reflecting water source, irrespective of region. Lake outlets and ground‐water‐fed streams had the highest species richness and abundance, temperature and bed stability, while glacier‐fed streams were characterized by low species richness, abundance, temperature, bed stability and high concentrations of suspended solids. Macroinvertebrate species richness was positively correlated with water temperature and negatively with bed stability. Conductivity was positively correlated with invertebrate abundance. 5. The results of this study suggest that the source of stream water can be used to predict invertebrate community composition in Greenlandic streams and thus the effects of changes in water balance and flow regime, and to identify sites of special conservation interest.     

The role of water exchange between a stream channel and its hyporheic zone in nitrogen cycling at the terrestrial-aquatic interface

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.     

Dispersal of organisms in a patchy stream environment under different settlement scenarios

1.  Previous work in streams, where many organisms disperse downstream by drifting in the water column, has suggested dispersal distances could be related to the overall proportion of the stream within dead water zones (DWZ), places such as backwaters, pools or behind large obstacles, where the water is out of the main flow and is stored temporarily. However, dispersal distances might also be influenced by the spatial organization of DWZ within a reach; this would require us to distinguish between reaches containing many small DWZ compared with those having a few large ones in order to understand fully differences in dispersal patterns among reaches differing in this regard.
2.  We constructed a spatial model of flow in stream reaches, in which we varied the density and size distribution of DWZ distributed through the reach. We defined flow conditions under which stream organisms, modelled as passive particles, would settle or continue to drift downstream. We then examined the horizontal dispersal profiles of such particles through reaches differing in arrays of DWZ.
3.  Our results support those of previous studies, but suggest that the spatial arrangement of DWZ may be just as important as the overall proportion of DWZ within a reach. On a more general note, the importance of patch configuration observed in this study reflects the growing view that a thorough understanding of population and ecosystem processes will require the explicit consideration of spatial pattern in the environment.     

Climate-induced changes in high elevation stream nitrate dynamics

Mountain terrestrial and aquatic ecosystems are responsive to external drivers of change, especially climate change and atmospheric deposition of nitrogen (N). We explored the consequences of a temperature-warming trend on stream nitrate in an alpine and subalpine watershed in the Colorado Front Range that has long been the recipient of elevated atmospheric N deposition. Mean annual stream nitrate concentrations since 2000 are higher by 50% than an earlier monitoring period of 1991–1999. Mean annual N export increased by 28% from 2.03 kg N ha⁻¹ yr⁻¹ before 2000 to 2.84 kg N ha⁻¹ yr⁻¹ in Loch Vale watershed since 2000. The substantial increase in N export comes as a surprise, since mean wet atmospheric N deposition from 1991 to 2006 (3.06 kg N ha⁻¹ yr⁻¹) did not increase. There has been a period of below average precipitation from 2000 to 2006 and a steady increase in summer and fall temperatures of 0.12 °C yr⁻¹ in both seasons since 1991. Nitrate concentrations, as well as the weathering products calcium and sulfate, were higher for the period 2000–2006 in rock glacier meltwater at the top of the watershed above the influence of alpine and subalpine vegetation and soils. We conclude the observed recent N increases in Loch Vale are the result of warmer summer and fall mean temperatures that are melting ice in glaciers and rock glaciers. This, in turn, has exposed sediments from which N produced by nitrification can be flushed. We suggest a water quality threshold may have been crossed around 2000. The phenomenon observed in Loch Vale may be indicative of N release from ice features such as rock glaciers worldwide as mountain glaciers retreat.     

Physico-chemistry and aquatic insects of a glacier-fed and a spring-fed alpine stream

1. Physico-chemical conditions and benthic macroinvertebrates were studied in two adjacent alpine streams in the Tyrolean Alps, Austria, for 2 years, and aquatic insect emergence was recorded for 1 year.
2. In the spring-fed system, maximum discharge and increased concentrations of suspended solids, nitrate and particulate phosphorus occurred during snowmelt in June. In the glacier-fed stream, high discharge and strong diel fluctuations in flow and concentrations of suspended solids created a harsh and unstable environment during summer. Glacial ablation, variation in groundwater inflow, and water inputs from tributaries draining calcareous rocks caused water chemistry to vary both seasonally and longitudinally in glacier-fed Rotmoosache.
3. A total of 126 aquatic or semi-aquatic invertebrate taxa were collected, 94 of which were found in the glacier-fed stream and 120 in the spring-fed stream. Chironomid abundance was 2–8 times and taxa richness 2–3 times lower in the glacier-fed stream than in the spring-fed stream, as was the number of chironomid taxa (72 versus 93 total).
4. These results broadly support the conceptual model by Milner & Petts (1994) concerning glacier-fed stream systems. However, single samples and seasonal means showed relatively high invertebrate abundance and richness, especially during winter, indicating a considerable degree of spatial and temporal variability.
5. We suggest that the seasonal shifts from harsh environmental conditions in summer to less severe conditions in autumn and a rather constant environment in winter are an important factor affecting larval development, life-history patterns and the maintenance of relatively high levels of diversity and productivity in glacier-fed streams.     

Climate change effects on upland stream macroinvertebrates over a 25-year period

Climate change effects on some ecosystems are still poorly known, particularly where they interact with other climatic phenomena or stressors. We used data spanning 25 years (1981–2005) from temperate headwaters at Llyn Brianne (UK) to test three hypotheses: (1) stream macroinvertebrates vary with winter climate; (2) ecological effects attributable to directional climate change and the North Atlantic Oscillation (NAO) are distinguishable and (3) climatic effects on macroinvertebrates depend on whether streams are impacted by acidification. Positive (i.e. warmer, wetter) NAO phases were accompanied by reduced interannual stability (=similarity) in macroinvertebrate assemblage in all streams, but associated variations in composition occurred only in acid moorland. The NAO and directional climate change together explained 70% of interannual variation in temperature, but forest and moorland streams warmed respectively by 1.4 and 1.7°C (P<0.001) between 1981 and 2005 after accounting for NAO effects. Significant responses among macroinvertebrates were confined to circumneutral streams, where future thermal projections (+1, +2, +3°C) suggested considerable change. Spring macroinvertebrate abundance might decline by 21% for every 1°C rise. Although many core species could persist if temperature gain reached 3°C, 4–10 mostly scarce taxa (5–12% of the species pool) would risk local extinction. Temperature increase in Wales approaches this magnitude by the 2050s under the Hadley HadCM3 scenarios. These results support all three hypotheses and illustrate how headwater stream ecosystems are sensitive to climate change. Altered composition and abundance could affect conservation and ecological function, with the NAO compounding climate change effects during positive phases. We suggest that acidification, in impacted streams, overrides climatic effects on macroinvertebrates by simplifying assemblages and reducing richness. Climatic processes might, nevertheless, exacerbate acidification or offset biological recovery.     

A ground water nitrogen budget for a headwater swamp in an area of permanent ground water discharge

Ground water inputs and outputs of N were studied for a small ground water discharge swamp situated in a headwater drainage basin in southern Ontario, Canada. Darcy's equation with data for piezometers was used to measure inputs of shallow local ground water at the swamp margin and deep regional ground water beneath the swamp. Ground water flux was also quantified by measuring ground water discharge to the outlet stream draining the swamp in combination with a chemical mixing model to separate shallow and deep ground water components based on chloride differences. Estimates of shallow ground water flux determined by these two approaches agreed closely however, the piezometer data seriously underestimated the deep ground water input to the swamp. An average ground water input-output budget of total N (TN) total organic nitrogen (TON) ammonium (NH₄ ⁺-N) and nitrate (NO₃ ^--N) was estimated for stream base flow periods which occurred on an average of 328 days each year during 1986–1990. Approximately 90% of the annual NO₃ ^--N input was contributed by shallow ground water at the swamp margin. Deep ground water represented about 65% of the total ground water input and a similar proportion of TON and NH₄ ⁺-N inputs. Annual ground water NO₃ ^--N inputs and outputs were similar whereas NH₄ ⁺-N retention was 4 kg ha^-1 representing about 68% of annual ground water input. Annual TON inputs in ground water exceeded outputs by 7.7 kg ha (27%). The capacity of the swamp to regulate ground water N fluxes was influenced by the N chemistry of ground water inputs and the hydrologic pathways of transport within the swamp.