Inter-annual variation in responses of water chemistry and epilithon to Pacific salmon spawners in an Alaskan stream

1. Pacific salmon (Oncorhynchus spp.) deliver salmon‐derived nutrients (SDN) to the streams in which they spawn. However, many stream parameters, such as discharge and spawner abundance, can vary from year to year, which could alter the quantity and flux of SDN. 2. Over six consecutive years, we studied responses in streamwater chemistry and epilithon (i.e. the microbial community on submerged rocks) to salmon spawners in Fish Creek, southeastern Alaska, U.S.A. The lower reach of Fish Creek receives spawners of several salmon species, while the upper reach does not receive spawners because of an intervening waterfall. 3. We estimated salmon spawner biomass, analysed water chemistry [ammonium, nitrate, soluble reactive phosphorus (SRP) and dissolved organic carbon (DOC)], and measured epilithon abundance [as chlorophyll a (chl a) and ash‐free dry mass (AFDM)] in Fish Creek. Measurements were made in both the upper and lower reaches, before, during and after the major salmon runs. 4. Absolute values and relative differences indicated that the presence of salmon spawners consistently increased dissolved ammonium (by 58 μg L⁻¹ on average, 41× over background), SRP (by 6 μg L⁻¹, 14×), epilithon chl a (by 35 mg m⁻², 16×), and epilithon AFDM (by 3 g m⁻², 8×). Salmon spawners did not increase nitrate or DOC in either absolute or relative amounts. The persistence and magnitude of spawner effects varied among years and appeared to reflect weather‐driven hydrology as well as spawner biomass. 5. Salmon‐derived nutrients can stimulate the growth of primary producers by increasing streamwater nutrient concentrations, but this positive influence may be modulated by other factors, such as water temperature and discharge. To better assess the ecological influence of SDN on stream biota, future studies should explicitly consider the role of key environmental factors and their temporal and spatial dynamics in stream ecosystems.     

Climate‐induced shift in hydrological regime alters basal resource dynamics in a wilderness river ecosystem

1. We integrated a 20‐year ecological data set from a sparsely inhabited, snowmelt‐dominated catchment with hydrologic models to predict the effects of hydrologic shifts on stream biofilm. 2. We used a stepwise multiple regression to assess the relationship between hydrology and biofilm ash‐free dry mass (AFDM) and chlorophyll‐a (chl‐a) under recent climate conditions. Biofilm AFDM was significantly related to the timing of peak streamflow, and chl‐a was significantly related to the timing of median streamflow. We applied these results to output from the variable infiltration capacity hydrologic model, which predicted hydrology under a baseline scenario (+0 °C) and a range of warming scenarios expected with climate change (+1, +2 or +3 °C). 3. When compared to the baseline, the results indicated that earlier peakflows predicted under warming scenarios may lead to earlier initiation of biofilm growth. This may increase biofilm AFDM during the summer by up to 103% (±29) in the +3 °C scenario. Moreover, interannual variability of AFDM was predicted to increase up to 300%. Average chl‐a during the summer increased by up to 90% (±15) in the +3 °C scenario; however, its response was not significantly different from baseline in most years. 4. Because hydrologic change may alter the temporal dynamics of biofilm growth, it may affect the seasonal dynamics of biofilm quality (i.e. chl‐a‐to‐AFDM ratio). The results indicated that hydrologic shifts may increase biofilm quality during the spring, but may decrease it during the summer. Thus, we provide evidence that predicted hydrologic shifts in snowmelt‐dominated streams may alter the quantity and quality of an important basal resource. However, the magnitudes of these predictions are likely to be affected by other environmental changes that are occurring with climate change (e.g. increased wildfire activity and stream warming).     

Separating physical disturbance and nutrient enrichment caused by Pacific salmon in stream ecosystems

1. Pacific salmon (Oncorhynchus spp.) deliver marine‐derived nutrients to the streams in which they spawn and die, and these resource subsidies can increase the abundance of stream biota. In strong contrast, physical disturbance from salmon spawning activity can reduce the abundance of benthic organisms. Previous experimental designs have not established the relative effects of these two contrasting processes on stream organisms during a salmon run. 2. We combined manipulative and observational field studies to assess the degree of nutrient enrichment, physical disturbance, and the net effect of salmon on the abundance of benthic periphyton. Related salmon‐mediated processes were also evaluated for benthic macroinvertebrates. Mesh exclosures (2 × 2 m plots) prevented salmon from disturbing areas of the stream channel, which were compared with areas to which salmon had access. Sampling was conducted both before and during the late‐summer spawning run of pink (O. gorbushca) and chum (O. keta) salmon. 3. Streamwater nitrogen and phosphorus concentrations increased sharply with the onset of the salmon run, and highly significant positive relationships were observed between the numbers of salmon present in the stream and these dissolved nutrients. Before the salmon run, periphyton biomass (as chlorophyll a) and total macroinvertebrate abundance were very similar between control and exclosure plots. During the salmon run, exclosures departed substantially from controls, suggesting significant disturbance imparted on benthic biota. 4. Comparing exclosures before and during the salmon run enabled us to estimate the effects of salmon in the absence of direct salmon disturbance. This ‘nutrient enrichment potential’ was significant for periphyton biomass, as was a related index for macroinvertebrate abundance (although enhanced invertebrate drift into exclosures during the salmon run could also have been important). Interestingly, however, the net effect of salmon, evaluated by comparing control plots before and during the salmon run, was relatively modest for both periphyton and macroinvertebrates, suggesting that nutrient enrichment effects were largely offset by disturbance. 5. Our results illustrate the importance of isolating the specific mechanisms via which organisms affect ecosystems, and indicate that the relative magnitude of salmon nutrient enrichment and benthic disturbance determines the net effect that these ecologically important fish have on stream ecosystems.     

Pacific salmon effects on stream ecosystems: a quantitative synthesis

Pacific salmon (Oncorhynchus spp.) disturb sediments and fertilize streams with marine-derived nutrients during their annual spawning runs, leading researchers to classify these fish as ecosystem engineers and providers of resource subsidies. While these processes strongly influence the structure and function of salmon streams, the magnitude of salmon influence varies widely across studies. Here, we use meta-analysis to evaluate potential sources of variability among studies in stream ecosystem responses to salmon. Results obtained from 37 publications that collectively included 79 streams revealed positive, but highly inconsistent, overall effects of salmon on dissolved nutrients, sediment biofilm, macroinvertebrates, resident fish, and isotopic enrichment. Variation in these response variables was commonly influenced by salmon biomass, stream discharge, sediment size, and whether studies used artificial carcass treatments or observed a natural spawning run. Dissolved nutrients were positively related to salmon biomass per unit discharge, and the slope of the relationship for natural runs was five to ten times higher than for carcass additions. Mean effects on ammonium and phosphorus were also greater for natural runs than carcass additions, an effect attributable to excretion by live salmon. In contrast, we observed larger positive effects on benthic macroinvertebrates for carcass additions than for natural runs, likely because disturbance by live salmon was absent. Furthermore, benthic macroinvertebrates and biofilm associated with small sediments (<32 mm) displayed a negative response to salmon while those associated with large sediments (>32 mm) showed a positive response. This comprehensive analysis is the first to quantitatively identify environmental and methodological variables that influence the observed effects of salmon. Identifying sources of variation in salmon–stream interactions is a critical step toward understanding why engineering and subsidy effects vary so dramatically over space and time, and toward developing management strategies that will preserve the ecological integrity of salmon streams. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Variation in responses to spawning Pacific salmon among three south-eastern Alaska streams

1. Pacific salmon are thought to stimulate the productivity of the fresh waters in which they spawn by fertilising them with marine‐derived nutrients (MDN). We compared the influence of salmon spawners on surface streamwater chemistry and benthic biota among three south‐eastern Alaska streams. Within each stream, reaches up‐ and downstream of barriers to salmon migration were sampled during or soon after spawners entered the streams. 2. Within streams, concentrations of dissolved ammonium and soluble reactive phosphorus (SRP), abundance of epilithon (chlorophyll a and ash‐free dry mass) and biomass of chironomids were significantly higher in reaches with salmon spawners. In contrast, biomass of the mayflies Epeorus spp. and Rhithrogena spp. was significantly higher in reaches lacking spawners. 3. Among streams, significant differences were found in concentrations of dissolved ammonium, dissolved organic carbon, nitrate and SRP, abundance of epilithon, and the biomass of chironomids and Rhithrogena. These differences did not appear to reflect differences among streams in spawner density, nor the changes in water chemistry resulting from salmon spawners. 4. Our results suggest that the ‘enrichment’ effect of salmon spawners (e.g. increased streamwater nutrient concentrations) was balanced by other concurrent effects of spawners on streams (e.g. sediment disturbance). Furthermore, the collective effect of spawners on lotic ecosystems is likely to be constrained by conditions unique to individual streams, such as temperature, background water chemistry and light attenuation.     

Impacts of Riparian Forest Removal on Palauan Streams

The 2006 completion of the circum‐island Compact Road on the island of Babeldaob in the Republic of Palau resulted in several deforested stream reaches with modified stream channels. To determine the impacts of deforestation and road construction, various ecosystem parameters were compared between road‐impacted reaches, reforested savanna reaches, and forested reaches. Compared to adjacent forested reaches, road‐impacted reaches received significantly more light (0.4 ± 0.1 vs. 87.8 ± 4.1 % light transmittance, respectively), were significantly warmer (25.7 ± 0.1 vs. 26.1 ± 0.1°C, respectively), and received higher nutrient and sediment loads, all of which were attributed to the removal of riparian vegetation and increased surface runoff from the road. These differences were believed to have shifted the benthic algal community in road‐impacted reaches from diatoms to filamentous algae with significantly greater chl a biomass (10×) and benthic algal ash free dry mass AFDM (3×) compared to adjacent forested reaches. Savanna‐impacted and forested reaches had similar chl a, algal AFDM, and received similar amounts of light. Nutrient and sediment concentrations varied between the two reach types. Results from this study emphasize the need for the maintenance of riparian forests especially with predicted increases in population, development, and deforestation. Future studies are needed to determine effective riparian widths and riparian forest community structure to help resource managers and land owners protect and preserve the many ecosystem services that Palauan streams and watersheds provide.     

Effects of spawning Pacific salmon on the isotopic composition of biota differ among southeast Alaska streams

1. Adult Pacific salmon (Oncorhynchus spp.) transport marine nutrients to fresh waters and disturb sediments during spawning. The relative importance of nutrient fertilisation and benthic disturbance by salmon spawners can be modulated by environmental conditions (e.g. biological, chemical and physical conditions in the catchment, including human land use). 2. To determine the importance of the environmental context in modifying the uptake and incorporation of salmon‐derived material into stream biota, we measured the nitrogen (δ¹⁵N) and carbon (δ¹³C) isotopic composition of benthic algae (i.e. epilithon) and juvenile coho salmon (Oncorhynchus kisutch) in seven streams across a timber‐harvest gradient (8–69% catchment area harvested), both before and during the salmon run. Conditional bootstrap modelling simulations were used to assess variability in the response of epilithon and juvenile coho salmon to spawning salmon. 3. In response to spawning salmon, epilithon exhibited enrichment in both δ¹⁵N (mean: 1.5‰) and δ¹³C (2.3‰). Juvenile coho were also enriched in both δ¹⁵N (0.7‰) and δ¹³C (1.4‰). Conditional bootstrap models indicate decreased variation in data as spatial replication increases, suggesting that the number of study sites can influence the results of Pacific salmon isotope studies. 4. Epilithon isotopic enrichment was predicted by environmental conditions, with δ¹⁵N enrichment predicted by stream temperature and timber harvest (R² = 0.87) and δ¹³C enrichment by discharge, sediment size, timber harvest and spawner density (R² = 0.96). Furthermore, we found evidence for a legacy effect of salmon spawners, with pre‐spawner δ¹⁵N and δ¹³C of both epilithon and juvenile coho predicted by salmon run size in the previous year. 5. Our results show that the degree of incorporation of salmon‐derived nitrogen and carbon differs among streams. Furthermore, the environmental context, including putative legacy effects of spawning salmon, can influence background isotopic concentrations and utilisation of salmon‐derived materials in southeast Alaska salmon streams. Future studies should consider the variation in isotopic composition of stream biota when deciding on the number of study sites and samples needed to generate meaningful results.     

Stream geomorphology regulates the effects on periphyton of ecosystem engineering and nutrient enrichment by Pacific salmon

1. Pacific salmon (Oncorhynchus spp.) returning to streams deliver substantial quantities of nutrients (nitrogen and phosphorus) that may stimulate primary production. Salmon can also affect the phytobenthos negatively via physical disturbance during nest excavation, a process that may counteract the positive effects of salmon‐derived nutrients on benthic algae. The ability of salmon to disturb benthic habitats may be a function of substratum particle size, and therefore, the geomorphology of streams could determine the net effect of salmon on benthic communities. 2. Based on surveys of 17 streams in southwest Alaska before the salmon run and during peak salmon density, we identified size thresholds for the disturbance of substratum particles by salmon and classified particles as vulnerable (<60 mm B‐axis), invulnerable (>110 mm) or transitional (61–110 mm). At the scale of individual rocks, algal biomass on vulnerable substrata decreased at peak spawning (relative to values before the run) as a power function of salmon density; transitional and invulnerable substrata showed no quantifiable pattern. However, invulnerable substrata in streams with more than 0.11 salmon m^?2 showed net algal accrual, or relatively smaller declines in algal biomass, than vulnerable substrata, indicating that large rocks provide refuge for benthic algae from salmon disturbance. 3. We expected that streams with proportionally larger rocks would respond positively to salmon at the whole‐stream scale, after accounting for the relative abundance of rocks of different sizes within streams. Invulnerable rocks made up only 0–12% of the total substratum particle size distribution in salmon‐bearing streams, however, and algal accrual on invulnerable substrata did not outweigh the strong disturbance effects on the more spatially extensive vulnerable substrata. The change in whole‐stream benthic algal biomass among streams was negatively related to salmon density. 4. Stable isotopes of nitrogen (δ¹⁵N) were used to track nutrients from salmon into benthic biota. Periphyton δ¹⁵N on rocks of all size classes was higher at peak salmon spawning than before the salmon run, indicating the uptake of salmon‐derived nitrogen. Peak δ¹⁵N values were positively related to salmon abundance and followed a two‐isotope mixing relationship. The per cent of N from salmon in periphyton was also related to salmon density and was best explained by a saturating relationship. Spring δ¹⁵N was unrelated to salmon returns in the previous year, suggesting little annual carryover of salmon nutrients.     

Increases in benthic community production and metabolism in response to marine‐derived nutrients from spawning Atlantic salmon (Salmo salar)

相似文献   

Whole-Stream Metabolism Responds to Spawning Pacific Salmon in Their Native and Introduced Ranges

Pacific salmon (Oncorhynchus spp.) perform important ecological roles in stream ecosystems by provisioning nutrients as resource subsidies and modifying their physical habitat as ecosystem engineers. These contrasting roles result in concurrent nutrient enrichment and benthic disturbance, where local environmental characteristics potentially determine which effect predominates. Whole-stream metabolism quantifies the functional response to salmon and may identify patterns in enrichment and disturbance not apparent from structural measurements alone. We measured ecosystem respiration (ER) and gross primary production (GPP), along with chemical and physical characteristics, in seven Southeast Alaska streams and two Michigan streams, before and during the salmon run. These streams in the native and introduced ranges of salmon differed in environmental characteristics, from geomorphology at the reach scale to climate at the biome scale. Salmon consistently increased ER across streams and biomes, from an average (±SE) of 1.92 ± 0.23 g O₂ m^?2 d^?1 before salmon to 6.30 ± 1.08 g O₂ m^?2 d^?1 during the run. In the cobble-bottom streams of Southeast Alaska, GPP doubled from 0.29 ± 0.05 g O₂ m^?2 d^?1 before salmon to 0.66 ± 0.16 g O₂ m^?2 d^?1 during the run. In contrast, GPP responded inconsistently to salmon in the sand-bottom Michigan streams, increasing in one and decreasing in the other. Patterns in ER and GPP among streams and time periods were predicted by stream water nutrients (for example, ammonium, soluble reactive phosphorus) rather than by physical characteristics (for example, light, sediment size, and so on). This study demonstrates that salmon can periodically override physical controls on ER and GPP and enhance whole-stream metabolism via their dual ecological roles as both resource subsidies and ecosystem engineers.     

Environmental DNA for the enumeration and management of Pacific salmon

Pacific salmon are a keystone resource in Alaska, generating annual revenues of well over ~US$500 million/year. Due to their anadromous life history, adult spawners distribute amongst thousands of streams, posing a huge management challenge. Currently, spawners are enumerated at just a few streams because of reliance on human counters and, rarely, sonar. The ability to detect organisms by shed tissue (environmental DNA, eDNA) promises a more efficient counting method. However, although eDNA correlates generally with local fish abundances, we do not know if eDNA can accurately enumerate salmon. Here we show that daily, and near‐daily, flow‐corrected eDNA rate closely tracks daily numbers of returning sockeye and coho spawners and outmigrating sockeye smolts. eDNA thus promises accurate and efficient enumeration, but to deliver the most robust numbers will need higher‐resolution stream‐flow data, at‐least‐daily sampling, and a focus on species with simple life histories, since shedding rate varies amongst jacks, juveniles, and adults.     

Reproductive patterns and secondary production of <Emphasis Type="Italic">Gammaropsis japonicus</Emphasis> (Crustacea,Amphipoda) on the seagrass <Emphasis Type="Italic">Zostera marina</Emphasis> of Korea

It is essential to know the nutrient limitation status of biofilms to understand how they may buffer uptake and export of nutrients from polluted watersheds. We tested the effects of nutrient additions on biofilm biomass (chlorophyll a, ash free dry mass (AFDM), and autotrophic index (AI, AFDM/chl a)) and metabolism via nutrient-diffusing substrate bioassays (control, nitrogen (N), phosphorus (P), and N + P treatments) at 11 sites in the Upper Snake River basin (southeast Idaho, USA) that differed in the magnitude and extent of human-caused impacts. Water temperature, turbidity, and dissolved inorganic N concentrations all changed seasonally at the study sites, while turbidity and dissolved inorganic N and P also varied with impact level. Chl a and AI on control treatments suggested that the most heavily impacted sites supported more autotrophic biofilms than less-impacted sites, and that across all sites biofilms were more heterotrophic in autumn than in summer. Nutrient stimulation or suppression of biofilm biomass was observed for chl a in 59% of the experiments and for AFDM in 33%, and the most frequent response noted across all study sites was N limitation. P suppression of chl a was observed only at the most-impacted sites, while AFDM was never suppressed by nutrients. When nutrient additions did have significant effects on metabolism, they were driven by differences in biomass rather than by changes in metabolic rates. Our study demonstrated that biofilms in southeast Idaho rivers were primarily limited by N, but nutrient limitation was more frequent at sites with good water quality than at those with poor water quality. Additionally, heterotrophic and autotrophic biofilm components may respond differently to nutrient enrichment, and nutrient limitation of biofilm biomass should not be considered a surrogate for metabolism in these rivers. Handling editor: D. Ryder     

Persistent nitrogen limitation of stream biofilm communities along climate gradients in the Arctic

Climate change is rapidly reshaping Arctic landscapes through shifts in vegetation cover and productivity, soil resource mobilization, and hydrological regimes. The implications of these changes for stream ecosystems and food webs is unclear and will depend largely on microbial biofilm responses to concurrent shifts in temperature, light, and resource supply from land. To study those responses, we used nutrient diffusing substrates to manipulate resource supply to biofilm communities along regional gradients in stream temperature, riparian shading, and dissolved organic carbon (DOC) loading in Arctic Sweden. We found strong nitrogen (N) limitation across this gradient for gross primary production, community respiration and chlorophyll‐a accumulation. For unamended biofilms, activity and biomass accrual were not closely related to any single physical or chemical driver across this region. However, the magnitude of biofilm response to N addition was: in tundra streams, biofilm response was constrained by thermal regimes, whereas variation in light availability regulated this response in birch and coniferous forest streams. Furthermore, heterotrophic responses to experimental N addition increased across the region with greater stream water concentrations of DOC relative to inorganic N. Thus, future shifts in resource supply to these ecosystems are likely to interact with other concurrent environmental changes to regulate stream productivity. Indeed, our results suggest that in the absence of increased nutrient inputs, Arctic streams will be less sensitive to future changes in other habitat variables such as temperature and DOC loading.     

Movers and shakers: nutrient subsidies and benthic disturbance predict biofilm biomass and stable isotope signatures in coastal streams

相似文献   

Ecoregion and land‐use influence invertebrate and detritus transport from headwater streams

Summary 1. Habitats are often connected by fluxes of energy and nutrients across their boundaries. For example, headwater streams are linked to surrounding riparian vegetation through invertebrate and leaf litter inputs, and there is evidence that consumers in downstream habitats are subsidised by resources flowing from headwater systems. However, the strength of these linkages and the manner in which potential headwater subsidies vary along climatic and disturbance gradients are unknown. 2. We quantified the downstream transport of invertebrates, organic matter and inorganic sediment from 60 fishless headwater streams in the Wenatchee River Basin located on the eastern slope of the Cascade Range in Washington, U.S.A. Streams were classified into four groups (each n = 15) based on their position within two ecological subregions (wet and dry) and the extent of past timber harvest and road development (logged and unlogged). 3. Time and ecoregion were significant for all response variables as transport varied across sampling periods, and dry ecoregion streams displayed significantly higher mean values. Logged sites also generally showed higher mean transport, but only inorganic sediment transport was significantly higher in logged sites. Both ecoregion and land‐use interacted significantly with time depending on the response variable. Differences among stream categories were driven by relatively low levels of transport in unlogged drainages of the wet ecoregion. Interestingly, unlogged dry ecoregion streams showed comparable transport rates to logged sites in the wet ecoregion. Dominance by deciduous riparian vegetation in all but unlogged streams in the wet ecoregion is a primary hypothesised mechanism determining transport dynamics in our study streams. 4. Understanding the quantity and variation of headwater subsidies across climate and disturbance gradients is needed to appreciate the significance of ecological linkages between headwaters and associated downstream habitats. This will enable the accurate assessment of resource management impacts on stream ecosystems. Predicting the consequences of natural and anthropogenic disturbances on headwater stream transport rates will require knowledge of how both local and regional factors influence these potential subsidies. Our results suggest that resources transported from headwater streams reflect both the meso‐scale land‐use surrounding these areas and the constraints imposed by the ecoregion in which they are embedded.     

THE RELATIONSHIP OF LIPIDS WITH LIGHT AND CHLOROPHYLL MEASUREMENTS IN FRESHWATER ALGAE AND PERIPHYTON1

Lipid content and lipid class composition were determined in stream periphyton and the filamentous green algae Cladophora sp. and Spirogyra sp, Sterols and phospholipids were compared to chlorophyll a (chl a) as predictors of biomass for stream periphyton and algae. Chlorophyll a, phospholipids, and sterols were each highly correlated with ash-free dry mass (AFDM) (r² > 0.98). Stream periphyton exposed naturally to high light (HL) and low light (LL) had chl a concentrations (μg chl a-mg^?1AFDM) of 7.9± 0.7 and 12.4 ± 2.9, respectively, while the sterol concentrations of these HL and LL stream periphyton (1.6 ± 0.4) were not significantly different (P > 0.05). Periphyton exposed to an irradiance of 300 μmol photons·m^?2s^?1 in the laboratory for 60 h had 5.6 ± 0.55 μg chl a·mg^?1 AFDM, but the same periphyton exposed to 2% incident light for the same amount of time had 11.0 ± 0.56 μg chl a·mg^?1 AFDM. Sterol concentrations in these periphyton communities remained unchanged (1.5 ± 0.3 μg·mg^?1AFDM), Similar results (i.e. changes in chl a but stability of sterol concentrations in response to irradiance changes) were also found for Cladophora and Spirogyra in laboratory experiments. Sterols can be quantified rapidly from a few milligrams of algae and appear to be a useful predictor of eukaryote biomass, whereas cellular levels of chl a vary substantially with light conditions. Phospholipids (or phospholipid fatty acids) are considered to be a reliable measure of viable microbial biomass. Nevertheless, phospholipid content varied substantially and unpredictably among algae and periphyton under different light regimes. Irradiance also had a significant effect on storage lipids: HL Cladophora and HL periphyton had 2 × and 5 × greater concentrations of triacylglycerols, respectively, compared to their LL forms. HL and LL algae also differed in the concentration of several major fatty acids. These light-induced changes in algal lipids and fatty acids have important implications for grazers.     

Catchment urbanisation and increased benthic algal biomass in streams: linking mechanisms to management

1. Urbanisation is an important cause of eutrophication in waters draining urban areas. We determined whether benthic algal biomass in small streams draining urban areas was explained primarily by small‐scale factors (benthic light, substratum type and nutrient concentrations) within a stream, or by catchment‐scale variables that incorporate the interacting multiple impacts of urbanisation (i.e. variables that describe urban density and the intensity of drainage or septic tank systems). 2. Benthic algal biomass was assessed as chlorophyll a density (chl a) in 16 streams spanning a rural–urban gradient, with both a wide range of urban density and of piped stormwater infrastructure intensity on the eastern fringe of metropolitan Melbourne, Australia. The gradient of urban density among streams was broadly correlated with catchment imperviousness, drainage connection (proportion of impervious areas connected to streams by stormwater pipes), altitude, longitude and median phosphorus concentration. Catchment area, septic tank density, median nitrogen concentration, benthic light (photosynthetically active radiation) and substratum type were not strongly correlated with the urban gradient. 3. Variation in benthic light and substratum type within streams explained a relatively small amount of variation in log chl a (3–11 and 1–13%, respectively) compared with between‐site variation (39–54%). 4. Median chl a was positively correlated with catchment urbanisation, with a large proportion of variance explained jointly (as determined by hierarchical partitioning) by those variables correlated with urban density. Independent of this correlation, the contributions of drainage connection and altitude to the explained variance in chl a were significant. 5. The direct connection of impervious surfaces to streams by stormwater pipes is hypothesised as the main determinant of algal biomass in these streams through its effect on the supply of phosphorus, possibly in interaction with stormwater‐related impacts on grazing fauna. Management of benthic algal biomass in streams of urbanised catchments is likely to be most effective through the application of stormwater management approaches that reduce drainage connection.     

Stream food web response to a salmon carcass analogue addition in two central Idaho, U.S.A. streams

1. Pacific salmon and steelhead once contributed large amounts of marine‐derived carbon, nitrogen and phosphorus to freshwater ecosystems in the Pacific Northwest of the United States of America (California, Oregon, Washington and Idaho). Declines in historically abundant anadromous salmonid populations represent a significant loss of returning nutrients across a large spatial scale. Recently, a manufactured salmon carcass analogue was developed and tested as a safe and effective method of delivering nutrients to freshwater and linked riparian ecosystems where marine‐derived nutrients have been reduced or eliminated. 2. We compared four streams: two reference and two treatment streams using salmon carcass analogue(s) (SCA) as a treatment. Response variables measured included: surface streamwater chemistry; nutrient limitation status; carbon and nitrogen stable isotopes; periphyton chlorophyll a and ash‐free dry mass (AFDM); macroinvertebrate density and biomass; and leaf litter decomposition rates. Within each stream, upstream reference and downstream treatment reaches were sampled 1 year before, during, and 1 year after the addition of SCA. 3. Periphyton chlorophyll a and AFDM and macroinvertebrate biomass were significantly higher in stream reaches treated with SCA. Enriched stable isotope (δ¹⁵N) signatures were observed in periphyton and macroinvertebrate samples collected from treatment reaches in both treatment streams, indicating trophic transfer from SCA to consumers. Densities of Ephemerellidae, Elmidae and Brachycentridae were significantly higher in treatment reaches. Macroinvertebrate community composition and structure, as measured by taxonomic richness and diversity, did not appear to respond significantly to SCA treatment. Leaf breakdown rates were variable among treatment streams: significantly higher in one stream treatment reach but not the other. Salmon carcass analogue treatments had no detectable effect on measured water chemistry variables. 4. Our results suggest that SCA addition successfully increased periphyton and macroinvertebrate biomass with no detectable response in streamwater nutrient concentrations. Correspondingly, no change in nutrient limitation status was detected based on dissolved inorganic nitrogen to soluble reactive phosphorus ratios (DIN/SRP) and nutrient‐diffusing substrata experiments. Salmon carcass analogues appear to increase freshwater productivity. 5. Salmon carcass analogues represent a pathogen‐free nutrient enhancement tool that mimics natural trophic transfer pathways, can be manufactured using recycled fish products, and is easily transported; however, salmon carcass analogues should not be viewed as a replacement for naturally spawning salmon and the important ecological processes they provide.     

Effects of upstream lakes and nutrient limitation on periphytic biomass and nitrogen fixation in oligotrophic, subalpine streams

1. We conducted bioassays of nutrient limitation to understand how macronutrients and the position of streams relative to lakes control nitrogen (N₂) fixation and periphytic biomass in three oligotrophic Rocky Mountain catchments. We measured periphytic chlorophyll‐a (chl‐a) and nitrogen‐fixation responses to nitrogen (N) and phosphorus (P) additions using nutrient‐diffusing substrata at 19 stream study sites, located above and below lakes within the study catchments. 2. We found that periphytic chl‐a was significantly co‐limited by N and P at 13 of the 19 sites, with sole limitation by P observed at another four sites, and no nutrient response at the final two sites. On average, the addition of N, P and N + P stimulated chl‐a 35%, 114% and 700% above control values respectively. The addition of P alone stimulated nitrogen fixation by 2500% at five of the 19 sites. The addition of N, either with or without simultaneous P addition, suppressed nitrogen fixation by 73% at nine of the 19 sites. 3. Lake outlet streams were warmer and had higher dissolved organic carbon concentrations than inlet streams and those further upstream, but position relative to lakes did not affect chl‐a and nitrogen fixation in the absence of nutrient additions. Chl‐a response to nutrient additions did not change along the length of the study streams, but nitrogen fixation was suppressed more strongly by N, and stimulated more strongly by P, at lower altitude sites. The responses of chl‐a and nitrogen fixation to nutrients were not affected by location relative to lakes. Some variation in responses to nutrients could be explained by nitrate and/or total N concentration. 4. Periphytic chl‐a and nitrogen fixation were affected by nutrient supply, but responses to nutrients were independent of stream position in the landscape relative to lakes. Understanding interactions between nutrient supply, nitrogen fixation and chl‐a may help predict periphytic responses to future perturbations of oligotrophic streams, such as the deposition of atmospheric N.     

The effect of agriculture on the seasonal dynamics and functional diversity of benthic biofilm in tropical headwater streams

Tropical streams are one of the most endangered ecosystems in the world due to the constant pressures from human activities. Among these activities, agriculture represents a land use that is crucial for human development but also a key driver of stream degradation and biodiversity decline in the tropics. Against this background, we investigated indirect effects of agriculture (alterations in stream flow and nutrient availability) and climate characteristics (water temperature) on benthic biofilm communities in tropical streams (São Paulo State, Brazil). Three first‐order streams draining catchments dominated by agricultural land use (sugarcane for bioenergy production, pasture) with some remaining riparian forest were studied for 1 year. We focused on the relationships of benthic biofilm biomass, algal biomass, diatom community, and functional structure with streamflow dynamics, nitrate concentrations, and water temperature. Our results indicate that these biological responses were mainly mediated by flow and water temperature and not by resource availability in the studied headwater streams. This result could be explained by the heavy rains and elevated runoff generation in these tropical catchments under agricultural influence, which might override the known effects of nutrient enrichment on benthic biofilm communities. Considering forecast climate and land‐use changes in tropical streams, our findings may suggest potential shifts in benthic biofilm communities, with functional consequences for aquatic food webs in these environments. Abstract in Brazilian Portuguese is available with online material.