Phosphatase activity as an early warning indicator of wetland eutrophication: problems and prospects

A phosphorus (P) loading experiment conducted in the oligotrophic P-limited Everglades was used to assess the utility of phosphatase activity (PA) of periphyton as an early warning (EW) indicator of wetland eutrophication. Phosphorus loads of 0, 0.4, 0.8, 1.6, 3.2, 6.4 and 12.8 g P m^–2 yr^–1 were applied to mesocosms placed in a slough community consisting of Cladium jamaicense Crantz, Eleocharis spp. and calcareous periphyton mats. Phosphatase activity, expressed on a biomass-specific basis, was not a sensitive indicator of P enrichment for epiphytic periphyton growing on acrylic dowels or floating mat periphyton. However, surface-area-specific PA was a sensitive indicator of P enrichment, responding within 2–3 weeks of the initiation of dosing. Surface-area-specific PA of unenriched periphyton ranged from 0.42 to 0.7 nmol cm^–2 min^–1, while PA of periphyton growing in the highest load (12.8 g P m^–2 yr^–1), ranged from 0.11 to 0.29 nmol cm^–2 min^–1. Conclusions drawn from PA analyses were consistent with those obtained from periphyton primary productivity and P content. Phosphatase activity is a potentially valuable EW indicator when used in conjunction with other complementary indicators.     

Decomposition responses to phosphorus enrichment in an Everglades (USA) slough

The effects of phosphorus (P) enrichment ondecomposition rates were measured in a Ploading experiment conducted in an oligotrophicmarsh in the northern Everglades, USA. In thisstudy, eighteen 2.5 m² enclosures(mesocosms) were placed in a pristineopen-water (slough) wetland and subjectedweekly to 6 inorganic P loads; 0, 0.2, 0.4,0.8, 1.6 and 3.2 g·m^–2g·yr^–1. Phosphorus accumulated rapidly in the benthicperiphyton and unconsolidated detrital (benthicfloc) layer and significantly higher Pconcentrations were recorded after 1 yr of Paddition. In contrast, a significant increasein surface soil (0–3 cm) TP concentrations wasmeasured in the surface soil layer only after 3yr of loading at the highest dose. Plantlitter and benthic floc/soil decompositionrates were measured using litter bags,containing sawgrass (Cladium jamaicenseCrantz) leaves, and cotton (cellulose) strips,respectively. Litter bag weight losses weresimilar among treatments and averaged 30% atthe end of the 3 yr study period. Litter Nconcentrations increased over time by anaverage of 80% at P loads < 1.6g·m^–2·yr^–1, and by > 120% at Ploads 1.6 g·m^–2·yr^–1.In contrast,litter P concentrations declined up to 50% inthe first 6 months in all P loads and onlysubsequently increased in the two highestP-loaded mesocosms. Cotton strip decaydemonstrated that benthic floc and soilmicrobial activity increased within 5 mo of Paddition with more significant treatmenteffects in the benthic than the soil layer. The influence of soil microbial transformationswas shown in porewater chemistry changes. While porewater P levels remained close tobackground concentrations throughout the study,porewater NH₄ ⁺ and Ca²⁺increased in response to P enrichment,suggesting that one significant effect of Penrichment in this oligotrophic peat system isenhanced nutrient regeneration.     

Periphyton nitrogenase activity as an indicator of wetland eutrophication: spatial patterns and response to phosphorus dosing in a northern Everglades ecosystem

The use of periphyton nitrogenase activity (biological N₂ fixation) as an indicator of wetland P impact was assessed using patterns of nutrient content (C, N, P, Ca, Mg, K, Fe, and Mn) and acetylene reduction (AR) in floating cyanobacterial periphyton mat (metaphyton) communities of a P-enriched portion of the Florida Everglades, USA (Water Conservation Area-2A, WCA-2A). Spatial patterns of nutrients indicate the enrichment of floating mat periphyton N, P, Fe, and K, and the reduction of Mn and TN:TP in enriched marsh areas. In highly enriched areas, floating mat periphyton AR was approximately threefold greater than that in less enriched, interior marsh zones. Multiple regression models indicated AR dependence on P in eutrophic WCA-2A areas while the AR of more interior marsh periphyton mats was more closely related to tissue levels of Ca and Fe. Nitrogenase activity of floating mat periphyton from P-loaded mesocosms revealed a significant enhancement of N₂ fixation in samples receiving approximately 2–3 mg P m⁻² of cumulative P dosing or with biomass TP content of 100–300 mg kg⁻¹. At P contents above the optimum, mat periphyton AR was suppressed possibly as a result of changes in species composition or increased levels of NH₄⁺. After 3 years of dosing, consistently high AR occurred only at low rates of P enrichment (0.4–0.8 g P m⁻² yr⁻¹), and the patterns appeared to be seasonal. These findings agree with the hypothesis that P availability is a key determinant of nitrogenase activity in aquatic systems, and thus, may support the use of periphyton nitrogenase to indicate P impacts in P-limited systems. These results also demonstrate the potential existence of a P threshhold for biogeochemical alteration of periphyton mat function in the Everglades, and that cumulative loading of limiting nutrients (i.e., P), rather than instantaneous concentrations, should be considered when evaluating nutrient criteria.     

Algal responses to experimental nutrient addition in the littoral community of a subtropical lake

1. An in situ experiment was performed in the littoral zone of a large, subtropical lake to quantify effects of phosphorus (P) and nitrogen (N) on algal biomass, productivity, nutrient content and phosphate uptake kinetics. 2. We hypothesized that resident periphyton rapidly sequester added nutrients from the water column, but once a certain threshold is reached, nutrients remain in the water and permit a shift to a phytoplankton-dominated community. 3. Three duplicate sets of 1.2-m diameter mesocosms were treated with 10, 20 or 50 μg P L^??1 in combination with 100, 200 or 500 μg N L^??1, respectively. The nutrients were added thrice weekly for 14 days, after which the treatment doses were doubled for an additional 9 days. The cumulative amounts of P and N added over the course of the study were 700 and 7000 μg L^??1, respectively. Two untreated mesocosms and two open reference sites were used as controls. 4. The total P concentration in the water column of nutrient-treated mesocosms remained low, even after prolonged high dosing. However, there was a two-fold increase in the P content of surface algal mats and epiphyton. This indicates that some of the added P was sequestered by those components of the community. In contrast, metaphyton and epipelon displayed little or no increase in their P content. Large quantities of added P could not be accounted for in the periphyton community, and may reflect unmeasured losses to the sediments or other pools. 5. Nitrogen also was depleted from the water column, but there were no significant increases in periphyton N content. Much of the added N could not be accounted for in mass balances, and may have been lost from the mesocosms through volatilization or other biochemical processes. 6. Chlorophyll-a in epiphyton increased significantly after 14 days in the highest nutrient treatment, where there also was a proliferation of Spirogyra on day 28. 7. On day 28, water column samples from the highest nutrient treatment also displayed a significantly higher rate of carbon uptake, and a significantly higher concentration of midday dissolved oxygen. 8. The hypothesis that phytoplankton become dominant at high nutrient loading rates was not supported. However, there were dramatic changes in community structure (increased dominance by epiphytic Spirogyra) and function (increased productivity and dissolved oxygen) in response to nutrient additions.     

Effects of increased phosphorus loading on dissolved oxygen in a subtropical wetland, the Florida Everglades

The Florida Everglades is an oligotrophic, phosphorus (P)-limited wetland that is experiencing eutrophication as a result of P-enriched agricultural runoff. Effects of P enrichment on diel water-column dissolved oxygen concentration (DO) profiles were measured along nutrient gradients downstream of agricultural discharges in two northern Everglades marshes and in field enclosures (mesocosms) exposed to different P loading rates. Reference (i.e., water-column TP < 10 g/L) areas in the marsh interior were characterized by strong diel fluctuations in DO, and aerobic conditions generally were maintained throughout the diel cycle. Enriched stations (water-column TP elevated to between 12 and 131 g/L) were characterized by dampened diel fluctuations and reduced DO, and the extent of these changes was correlated strongly with marsh P concentrations. Mean DO declined from between 1.81 and 7.52 mg/L at reference stations to between 0.04 and 3.18 mg/L in highly enriched areas. Similarly, minimum DO declined from between 0.33 and 5.86 mg/L to between 0 and 0.84 mg/L with increasing enrichment, and the frequency of extremely low DO (< 1 mg/L) increased from between 0 and 20% to as high as 100% in the most enriched areas. Diel oxygen profiles in P-enriched mesocosms declined progressively with time; all loading treatments exhibited similar DO during the 1^st year of P loading, but concentrations declined significantly at higher loads by year 3. Reductions in water-column DO with increased P enrichment were associated with reduced oxygen production by submersed periphyton and macrophytes and increased sediment oxygen demand. Increased emergent macrophyte cover in enriched areas likely contributed to these changes by shading the water-column, which inhibited submerged productivity, and by providing inputs of nutrient-rich detritus, which increased oxygen demand. Declines in marsh DO are associated with other ecological changes such as increased anaerobic metabolism and an increase in invertebrate taxa that tolerate low DO. While background oxygen concentrations in wetlands can be lower than those in lakes and rivers, declines in water-column DO caused by eutrophication can result in biological impacts similar to those in other aquatic ecosystems.     

Response of Periphytic Algae to Gradients in Nitrogen and Phosphorus in Streamside Mesocosms

In this study we manipulated both nitrogen and phosphorus concentrations in stream mesocosms to develop quantitative relationships between periphytic algal growth rates and peak biomass with inorganic N and P concentrations. Stream water from Harts Run, a 2nd order stream in a pristine catchment, was constantly added to 36 stream-side stream mesocosms in low volumes and then recirculated to reduce nutrient concentrations. Clay tiles were colonized with periphyton in the mesocosms. Nutrients were added to create P and N concentrations ranging from less than Harts Run concentrations to 128 μg SRP l⁻¹ and 1024 μg NO₃-N l⁻¹. Algae and water were sampled every 3 days during colonization until periphyton communities reached peak biomass and then sloughed. Nutrient depletion was substantial in the mesocosms. Algae accumulated in all streams, even streams in which no nutrients were added. Nutrient limitation of algal growth and peak biomass accrual was observed in both low P and low N conditions. The Monod model best explained relationships between P and N concentrations and algal growth and peak biomass. Algal growth was 90% of maximum rates or higher in nutrient concentrations 16 μg SRP l⁻¹ and 86 μg DIN l⁻¹. These saturating concentrations for growth rates were 3–5 times lower than concentrations needed to produce maximum biomass. Modified Monod models using both DIN and SRP were developed to explain algal growth rates and peak biomass, which respectively explained 44 and 70% of the variance in algal response.     

Effects of food‐web structure on periphyton stoichiometry in eutrophic lakes: a mesocosm study

1. Aquatic herbivores typically have much higher concentrations of nutrients (e.g. N and P) in their tissues than there is in the food they eat. These stoichiometric differences can cause herbivores to be limited by the elemental quality of their food, which could affect, in turn, the structure of consumer communities and even alter key ecosystem processes. 2. In streams and in the littoral zone of shallow lakes, periphyton is an important food resource for benthic animals. Studying the elemental composition of periphyton may help us to understand food‐web structure, and any reciprocal effect of this structure on periphyton stoichiometry. 3. To understand how alterations in the food‐web structure affect the elemental composition of periphyton in a eutrophic lake, we carried out a long‐term experiment (14 months) in large‐scale mesocosms (40 m³), in which we manipulated food‐web structure, and which were dominated either by planktivorous fish (Rutilus rutilus) or herbivorous invertebrates (without fish). Periphyton was sampled monthly at three depths (0.5, 1.5 and 2.5 m) to determine its biomass and elemental composition (C/N/P ratio). Food‐web structure, physical and chemical parameters were monitored throughout the experiment. 4. Fish had indirect positive effect on periphyton biomass, leading to twofold higher levels than in herbivore‐dominated mesocosms. This result was probably due to control of benthic consumers by fish, suggesting a strong top–down control on periphyton by their consumers in fishless enclosures. 5. The elemental ratios C/P and C/N were lower in deep water in both treatments, mainly mediated by light availability, in accordance with the light/nutrient ratio hypothesis. These ratios were also lower in fishless treatments, probably due to increases in inorganic nutrient availability and grazing pressure in herbivore‐dominated systems. During winter, periphyton elemental composition was similar in both treatments, and was unrelated to inorganic nutrient availability. 6. These results indicate that any alteration of food‐web structure in lakes, such as in biomanipulation experiments, is likely to modify both the biomass and elemental quality of periphyton. Resultant effects on the consumers of periphyton and macrophytes could play a key role in the success of biomanipulations and should be taken into account in further studies.     

Effects of nutrient loading,temperature regime and grazing pressure on nutrient limitation of periphyton in experimental ponds

相似文献   

THE ROLE OF PERIPHYTON IN PHOSPHORUS RETENTION IN SHALLOW FRESHWATER AQUATIC SYSTEMS

Eutrophication caused by phosphorus (P) leads to water quality problems in aquatic systems, particularly freshwaters, worldwide. Processing of nutrients in shallow habitats removes P from water naturally and periphyton influences P removal from the water column in flowing waters and wetlands. Periphyton plays several roles in removing P from the water column, including P uptake and deposition, filtering particulate P from the water, and attenuating flow, which decreases advective transport of particulate and dissolved P from sediments. Furthermore, periphyton photosynthesis locally increases pH by up to 1 unit, which can lead to increased precipitation of calcium phosphate, concurrent deposition of carbonate-phosphate complexes, and long-term burial of P. Actively photosynthesizing periphyton can cause super-saturated O₂ concentrations near the sediment surface encouraging deposition of metal phosphates. However, anoxia associated with periphyton respiration at night may offset this effect. Linking the small-scale functional role of periphyton to ecosystem-level P retention will require more detailed studies in a variety of ecosystems or large mesocosms. A case study from the Everglades illustrates the importance of considering the role of periphyton in P removal from wetlands. In general, periphyton tends to increase P retention and deposition. In pilot-scale constructed periphyton-dominated wetlands in South Florida, about half of the inflowing total P was removed.     

Feedbacks of consumer nutrient recycling on producer biomass and stoichiometry: separating direct and indirect effects

Herbivores can have both direct (consumptive) and indirect (nutrient‐mediated) effects on primary producer biomass and nutrient stoichiometry. Ecological stoichiometry theory predicts that herbivores of contrasting body stoichiometry will differentially remineralize nutrients, resulting in feedbacks on producer stoichiometry. We experimentally separated direct and indirect effects of aquatic vertebrate grazers on periphyton by manipulating grazer abundance and identity in mesocosms, and using grazer exclusion cages to expose periphyton to recycled nutrients in the absence of direct grazing. In experiment 1, we used a catfish with high body phosphorus (low body N:P), Ancistrus triradiatus, to assess consumptive versus nutrient‐mediated effects of grazer density on periphyton. In experiment 2, we compared the nutrient‐mediated effects of grazing by Ancistrus triradiatus and Rana palmipes, a tadpole with low body phosphorus and high body N:P. In experiment 1, we found that increasing catfish density led to lower biomass and particulate nutrients in periphyton through direct consumptive effects, but that nutrient‐mediated indirect effects enhanced periphyton biomass when grazers were experimentally separated from direct contact with periphyton. As predicted by stoichiometry theory, nutrient recycling by this P‐rich grazer tended to increase algal C:P and N:P (although effects were not statistically significant), while their consumptive effects reduced algal C:P and N:P. In experiment 2, grazer identity had strong effects on dissolved water nutrient concentrations, N recycling (measured with a ¹⁵N tracer), and periphyton stoichiometry. In accordance with stoichiometry theory, catfish increased N concentrations and recycling rates leading to higher periphyton N:P, while tadpoles had greater effects on P availability leading to lower periphyton N:P. Our experiments elucidate the importance of both the density and identity of grazers in controlling periphyton biomass and stoichiometry through consumptive and nutrient‐mediated effects, and support the power of ecological stoichiometry theory to predict feedbacks on producer stroichiometry arising from consumer stoichiometry through nutrient recycling.     

Periphyton nutrient status in a temperate stream with mixed land-uses: implications for watershed nitrogen storage

We sampled periphyton communities in a highly productive stream to characterize how longitudinal changes in watershed geology and land use affect periphyton nutrient status and elemental composition. Nutrient status was evaluated from measures of periphyton nutrient composition (carbon, nitrogen, and phosphorus), stable isotope signatures (δ¹⁵N and δ¹³C), and the response of periphyton to experimental enrichment with nitrogen. Biomass and nutrient content increased dramatically from the headwaters to downstream, while tissue nutrient ratios (C:P and C:N) were more consistent and did not indicate strong N- or P-limitation. Nitrogen enrichment experiments did not exhibit a consistent response upstream or downstream, and periphyton C:N:P stoichiometry showed no significant response to N-enrichment. Absolute densities of periphyton N were 5- to 90-fold greater than the overlying N concentrations in stream water (159- to 353-fold greater for P), and the δ¹⁵N signal indicates downstream enrichment from likely watershed sources (urban and agriculture land-use). These results suggest that periphyton in Spring Creek are not N-limited and store large quantities of both N and P, which in turn can be transported downstream during high flow events. Handling editor: David Hamilton     

Mesocosm experiments on nutrient and fish effects on shallow lake food webs in a Mediterranean climate

1. Nutrient and fish manipulations in mesocosms were carried out on food‐web interactions in a Mediterranean shallow lake in south‐east Spain. Nutrients controlled biomass of phytoplankton and periphyton, while zooplankton, regulated by planktivorous fish, influenced the relative percentages of the dominant phytoplankton species. 2. Phytoplankton species diversity decreased with increasing nutrient concentration and planktivorous fish density. Cyanobacteria grew well in both turbid and clear‐water states. 3. Planktivorous fish increased concentrations of soluble reactive phosphorus (SRP). Larger zooplankters (mostly Ceriodaphnia and copepods) were significantly reduced when fish were present, whereas rotifers increased, after fish removal of cyclopoid predators and other filter feeders (cladocerans, nauplii). The greatest biomass and diversity of zooplankton was found at intermediate nutrient levels, in mesocosms without fish and in the presence of macrophytes. 4. Water level decrease improved underwater light conditions and favoured macrophyte persistence. Submerged macrophytes (Chara spp.) outcompeted algae up to an experimental nutrient loading equivalent to added concentrations of 0.06 mg L^?1 PO₄‐P and 0.6 mg L^?1 NO₃‐N, above which an exponential increase in periphyton biomass and algal turbidity caused characean biomass to decline. 5. Declining water levels during summer favoured plant‐associated rotifer species and chroococcal cyanobacteria. High densities of chroococcal cyanobacteria were related to intermediate nutrient enrichment and the presence of small zooplankton taxa, while filamentous cyanobacteria were relatively more abundant in fishless mesocosms, in which Crustacea were more abundant, and favoured by dim underwater light. 6. Benthic macroinvertebrates increased significantly at intermediate nutrient levels but there was no relationship with planktivorous fish density. 7. The thresholds of nutrient loading and in‐lake P required to avoid a turbid state and maintain submerged macrophytes were lower than those reported from temperate shallow lakes. Mediterranean shallow lakes may remain turbid with little control of zooplankton on algal biomass, as observed in tropical and subtropical lakes. Nutrient loading control and macrophyte conservation appear to be especially important in these systems to maintain high water quality.     

Periphyton nutrient content,biomass and algal community on artificial substrate: response to experimental nutrient enrichment and the effect of its interruption in a tropical reservoir

Periphyton plays an important functional role in the retention of nutrients in aquatic ecosystems, especially phosphorus. We evaluated the effects of enrichment with N and P and the effect after 20 days of no additional N and P on periphyton on artificial substratum in open-bottom mesocosms. The aim was to jointly evaluate periphyton, phytoplankton and zooplankton in the presence of macrophytes. Experimental conditions simulated natural conditions and nutrient addition was based on the maximum concentration recorded in mesotrophic reservoir. Our hypothesis is that the periphyton is sensitive to the effects of N and P enrichment and its interruption, despite the positive response of phytoplankton and zooplankton. Two treatments were designed using open-bottom mesocosms (n = 3): control (no nutrient addition); NP+ (combined phosphorus and nitrogen addition). Sampling for the measurement of biotic and abiotic variables was performed, with 10 days of continuous enrichment, on the 3rd, 6th and 11th, and 20 days after enrichment had ended (31st day). Periphyton chlorophyll a, dry mass and algal density increased significantly with the addition of N and P and decreased 20 days after the interruption of the enrichment. The highest periphyton P content was found in the NP+ treatment. The enrichment had a positive effect on Chrysophyceae (Chromulina spp.) and rotifer (Polyarthra spp.) density and the interruption of enrichment favored Bacillariophyceae (Gomphonema sp.) and rotifers (Gastropus stylifer). Phytoplankton responded positively to enrichment. Along with the high macrophyte coverage over the experimental period, we evidenced the positive effect enrichment had on phytoplankton biomass and zooplankton abundance. Therefore, periphyton on artificial substrate was sensitive to effects of N and P enrichment and its interruption, responding promptly to changes in nutrient availability in a scenario of high competition and grazing.     

Are rocky shore ecosystems affected by nutrient-enriched seawater? Some preliminary results from a mesocosm experiment

The response of rocky shore ecosystems to increased nutrient availability was examined in eight land-based mesocosms designed for hard-bottom littoral communities built at Marine Research Station Solbergstrand (Norway). The average seawater volume in each basin was 9 m³ with an average water residence time of about 2 h. A tidal regime resembling that in the fjord was maintained in the basins, and waves were generated regularly. NH₄NO₃ and H₃PO₄, at a constant molar NP ratio of 16:1, was added into 6 basins at concentrations 1, 2, 4, 8, 16, 32 M DIN above the background DIN concentration during 1 1/2 years. Two mesocosms were kept as control treatment. Marine communities were introduced into the basins two years prior to the start of nutrient dosage. The effects of nutrient enrichment were few and only marginal during the first year of nutrient addition, while some effects became more obvious during the second year. The growth rate of the periphyton and fast-growing macroalgae communities was stimulated by nutrient enrichment, while the response was less evident among the perennial fucoids. The structure of the macroalgal communities, however, did not change during 16 months' measurements. In contrast, growth on artificial rock substrates during the same period of time revealed intensive growth of the fast-growing Ulva lactuca in high-dosed basins compared with low-dosed and control basins, which were dominated by the fucoid Fucus serratus. The fauna communities exhibited only a minor response to nutrient treatment. The common periwinkle Littorina littorea, however, appeared with increased abundance in the high-dosed basins. The total system metabolism tended to increase slightly, but not significantly, with increased nutrient loading.     

Using porewater profiles to assess nutrient availability in seagrass-vegetated carbonate sediments

We measured porewater profiles of inorganic (NH₄ ⁺, NO₃ ^–(+NO₂ ^–), PO₄ ^3– (hereafter referred to as DIP)) and organic (DON, DOP) nutrients in seagrass-vegetated sediments at two sites in a shallow bay in Bermuda within close proximity (200 m) but subject to different nutrient loading. At both sites, total dissolved and inorganic nutrient concentrations were usually 1–2 orders of magnitude higher in the sediments than in the water column, with the exception of NO₃ ^–. Organic N and P were significant components of the total dissolved nutrient pools both in the sediment porewater and in the overlying water column (up to 75% for DON and 40% for DOP), and may be important in meeting plant nutrient demands. We used two approaches to examine how well porewater nutrient concentrations reflected the relative availabilities of N and P for seagrasses: (1) a simple stoichiometric nutrient regeneration model based on the N:P ratio of decomposing organic matter and porewater NH₄ ⁺ concentrations to predict porewater DIP, and (2) fitting of the porewater profiles to estimate rates of net nutrient production (or consumption), which reflects the balance between nutrient sources and sinks in the rhizosphere. The stoichiometric model indicated that sediment porewaters were depleted in P relative to N in the low-nutrient outer bay site, and enriched in P relative to N in the higher-nutrient inner bay site. These results are consistent with the mechanism of carbonate sediments in oligotrophic tropical environments being a strong sink for dissolved inorganic P and our previous work suggesting that nutrient enrichment causes P to become disproportionately more available than N. Net nutrient production rates of porewater P at both sites and N at the inner bay site were low (typically < 2%) relative to the nutrient demands of the seagrasses. The implications of the profile interpretation are two-fold: (1) the low rates of net nutrient production indicate diffusive losses from the root zone were insignificant and that nutrient turnover rates were high, except in the P-limited outer bay where N accumulated in sediment porewaters; and (2) because standing stock nutrient concentrations often represent a small fraction of the total nutrients cycled in the sediments, they are in many cases a poor indicator of nutrient availability. Based on our estimates of losses from the root zone, decomposition, and plant uptake we have constructed a rough budget for the cycling of P and N at our two sites.     

The importance of nutrient co‐limitation in regulating algal community composition,productivity and algal‐derived DOC in an oligotrophic marsh in interior Alaska

1. Compared to lakes and streams, we know relatively little about the factors that regulate algae in freshwater wetlands. This discrepancy is particularly acute in boreal regions, where wetlands are abundant and processes related to climate change (i.e. increased permafrost collapse and soil weathering) are expected to increase nutrient inputs into aquatic systems. To investigate how accelerated nutrient inputs might affect algal structure and function in northern boreal wetlands, we added nitrogen, phosphorus and silica to mesocosms in an oligotrophic marsh in interior Alaska. 2. We conducted two in situ mesocosm enrichment experiments during consecutive summer growing seasons, each lasting 24 days. In 2007, we investigated the effects of +N, +P, +Si and +N+P+Si enrichment on benthic algal biomass (chlorophyll‐a, ash‐free dry mass, biovolume), chemistry (N : P ratio) and community composition. In 2008, we expanded our first experiment to investigate the effects +N+P, +N+Si, +P+Si and +N+P+Si on the same algal parameters as well as productivity (mg C m^?2 h^?1). 3. In both experiments, we measured water‐column dissolved organic carbon (DOC) inside treatment enclosures and related changes in DOC to standing algal biomass. 4. Benthic algal accrual did not increase following 24 days of enrichment with any nutrient alone or with P and Si together (+P+Si), but increased significantly with the addition of N in any combination with P and Si (+N+P, +N+Si, +N+P+Si). 5. Algal productivity (20 mg C m^?2 h^?1) increased between three‐ and seven‐fold (57–127 mg C m^?2 h^?1) with the addition of N in combination with any other nutrient (+N+P, +N+Si, +N+P+Si). Water‐column DOC concentration was significantly higher inside N‐combination treatments compared to the control during each season, and DOC increased linearly with benthic algal biomass in 2007 (r² = 0.89, P < 0.0001) and 2008 (r² = 0.74, P < 0.0001). 6. Taxonomic composition of the wetland algal community responded most strongly to N‐combination treatments in both seasons. In 2007, there was a significant shift from Euglena and Mougeotia in the control treatment to Chroococcus and Gloeocystis with +N+P+Si enrichment, and in 2008, a Mougeotia‐dominated community was replaced by Gloeocystis in the +N+P treatment and by Nitzschia in +N+Si and +N+P+Si treatments. 7. Together, these data provide several lines of evidence for co‐limitation, and the central importance of N as a co‐limiting nutrient for the wetland algal community. Changes in algal dynamics with increased nutrient concentrations could have important implications for wetland food webs and suggest that algae may provide a functional link between increasing nutrient inputs and altered wetland carbon cycling in this region.     

Nutrient loading and grazing by the minnow Phoxinus erythrogaster shift periphyton abundance and stoichiometry in mesocosms

1. Anthropogenic activities in prairie streams are increasing nutrient inputs and altering stream communities. Understanding the role of large consumers such as fish in regulating periphyton structure and nutritional content is necessary to predict how changing diversity will interact with nutrient enrichment to regulate stream nutrient processing and retention. 2. We characterised the importance of grazing fish on stream nutrient storage and cycling following a simulated flood under different nutrient regimes by crossing six nutrient concentrations with six densities of a grazing minnow (southern redbelly dace, Phoxinus erythrogaster) in large outdoor mesocosms. We measured the biomass and stoichiometry of overstory and understory periphyton layers, the stoichiometry of fish tissue and excretion, and compared fish diet composition with available algal assemblages in pools and riffles to evaluate whether fish were selectively foraging within or among habitats. 3. Model selection indicated nutrient loading and fish density were important to algal composition and periphyton carbon (C): nitrogen (N). Nutrient loading increased algal biomass, favoured diatom growth over green algae and decreased periphyton C : N. Increasing grazer density did not affect biomass and reduced the C : N of overstory, but not understory periphyton. Algal composition of dace diet was correlated with available algae, but there were proportionately more diatoms present in dace guts. We found no correlation between fish egestion/excretion nutrient ratios and nutrient loading or fish density despite varying N content of periphyton. 4. Large grazers and nutrient availability can have a spatially distinct influence at a microhabitat scale on the nutrient status of primary producers in streams.     

Phosphorus budgets in Everglades wetland ecosystems: the effects of hydrology and nutrient enrichment

The Florida Everglades is a naturally oligotrophic hydroscape that has experienced large changes in ecosystem structure and function as the result of increased anthropogenic phosphorus (P) loading and hydrologic changes. We present whole-ecosystem models of P cycling for Everglades wetlands with differing hydrology and P enrichment with the goal of synthesizing existing information into ecosystem P budgets. Budgets were developed for deeper water oligotrophic wet prairie/slough (‘Slough’), shallower water oligotrophic Cladium jamaicense (‘Cladium’), partially enriched C. jamaicense/Typha spp. mixture (‘Cladium/Typha’), and enriched Typha spp. (‘Typha’) marshes. The majority of ecosystem P was stored in the soil in all four ecosystem types, with the flocculent detrital organic matter (floc) layer at the bottom of the water column storing the next largest proportion of ecosystem P pools. However, most P cycling involved ecosystem components in the water column (periphyton, floc, and consumers) in deeper water, oligotrophic Slough marsh. Fluxes of P associated with macrophytes were more important in the shallower water, oligotrophic Cladium marsh. The two oligotrophic ecosystem types had similar total ecosystem P stocks and cycling rates, and low rates of P cycling associated with soils. Phosphorus flux rates cannot be estimated for ecosystem components residing in the water column in Cladium/Typha or Typha marshes due to insufficient data. Enrichment caused a large increase in the importance of macrophytes to P cycling in Everglades wetlands. The flux of P from soil to the water column, via roots to live aboveground tissues to macrophyte detritus, increased from 0.03 and 0.2 g P m⁻² yr⁻¹ in oligotrophic Slough and Cladium marsh, respectively, to 1.1 g P m⁻² yr⁻¹ in partially enriched Cladium/Typha, and 1.6 g P m⁻² yr⁻¹ in enriched Typha marsh. This macrophyte translocation P flux represents a large source of internal eutrophication to surface waters in P-enriched areas of the Everglades.     

Interactions between nutrient availability and hydroperiod shape macroinvertebrate communities in Florida Everglades marshes

Hydroperiod and nutrient status are known to influence aquatic communities in wetlands, but their joint effects are not well explored. I sampled floating periphyton mat and flocculent detritus (floc) infaunal communities using 6-cm diameter cores at short- and long-hydroperiod and constantly inundated sites across a range of phosphorus (P) availability (total phosphorus in soil, floc and periphyton). Differences in community structure between periphyton and floc microhabitats were greater than any variation attributable to hydroperiod, P availability, or other spatial factors. Multivariate analyses indicated community structure of benthic-floc infauna was driven by hydroperiod, although crowding (no. g⁻¹ AFDM) of individual taxa showed no consistent responses to hydroperiod or P availability. In contrast, community structure of periphyton mat infauna was driven by P availability, while densities of mat infauna (no. m⁻²) were most influenced by hydroperiod (+correlations). Crowding of mat infauna increased significantly with P availability in short-hydroperiod marshes, but was constant across the P gradient in long-hydroperiod marshes. Increased abundance of floating-periphyton mat infauna with P availability at short-hydroperiod sites may result from a release from predation by small fish. Community structure and density were not different between long-hydroperiod and constantly inundated sites. These results have implications for the use of macroinvertebrates as indicators of water quality in wetlands and suggest the substrate sampled can influence interpretation of ecological responses observed in these communities.     

Ecological effects of low-level phosphorus additions on two plant communities in a neotropical freshwater wetland ecosystem

We conducted a low-level phosphorus (P) enrichment study in two oligotrophic freshwater wetland communities (wet prairies [WP] and sawgrass marsh [SAW]) of the neotropical Florida Everglades. The experiment included three P addition levels (0, 3.33, and 33.3 mg P m^–2 month^–1), added over 2 years, and used in situ mesocosms located in northeastern Everglades National Park, Fla., USA. The calcareous periphyton mat in both communities degraded quickly and was replaced by green algae. In the WP community, we observed significant increases in net aboveground primary production (NAPP) and belowground biomass. Aboveground live standing crop (ALSC) did not show a treatment effect, though, because stem turnover rates of Eleocharis spp., the dominant emergent macrophyte in this community, increased significantly. Eleocharis spp. leaf tissue P content decreased with P additions, causing higher C:P and N:P ratios in enriched versus unenriched plots. In the SAW community, NAPP, ALSC, and belowground biomass all increased significantly in response to P additions. Cladium jamaicense leaf turnover rates and tissue nutrient content did not show treatment effects. The two oligotrophic communities responded differentially to P enrichment. Periphyton which was more abundant in the WP community, appeared to act as a P buffer that delayed the response of other ecosystem components until after the periphyton mat had disappeared. Periphyton played a smaller role in controlling ecosystem dynamics and community structure in the SAW community. Our data suggested a reduced reliance on internal stores of P by emergent macrophytes in the WP that were exposed to P enrichment. Eleocharis spp. rapidly recycled P through more rapid aboveground turnover. In contrast, C. jamaicense stored added P by initially investing in belowground biomass, then shifting growth allocation to aboveground tissue without increasing leaf turnover rates. Our results suggest that calcareous wetland systems throughout the Caribbean, and oligotrophic ecosystems in general, respond rapidly to low-level additions of their limiting nutrient.