Stoichiometry, growth, and fecundity responses to nutrient enrichment by invertebrate grazers in sub-tropical turtle grass (Thalassia testudinum) meadows

Although the effectiveness of herbivores in mitigating the effects of nutrient enrichment is well documented, few studies have examined the effects of nutrient enrichment on components of consumer fitness. Enclosures were deployed in shallow turtle grass (Thalassia testudinum) beds in Florida Bay, Florida in fall 2003, spring 2004, and fall 2004 to measure the effects of nitrogen and phosphorous enrichment on the growth, fecundity, and stoichiometry of three invertebrate epiphyte grazers commonly associated with T. testudinum. The gastropod Turbo castanea exhibited significantly greater wet weight gain and lower C:P and N:P in enriched than in ambient treatments. Although nutrient enrichment did not have any significant effects on the growth of caridean shrimp (treatment consisted of several different caridean shrimp species), their C:N was significantly lower in enriched treatments. The final size and stoichiometry of the hermit crab Paguristes tortugae was not significantly affected by nutrient enrichment, nor did nutrient enrichment significantly affect the fecundity of P. tortugae, the only grazer in which gravid individuals or egg masses were present. Our study demonstrated that nutrient enrichment of primary producers can positively affect the growth of marine invertebrate grazers and alter their stoichiometry; however, these effects were species-specific and may be dependent upon the life stage, specific diets, and/or compensatory feeding habits of the grazers.

A Suite of Prey Traits Determine Predator and Nutrient Enrichment Effects in a Tri-Trophic Food Chain

Predation, predation risk, and resource quality affect suites of prey traits that collectively impact individual fitness, population dynamics, and community structure. However, studies of multi-trophic level effects generally focus on a single prey trait, failing to capture trade-offs among suites of covarying traits that govern population responses and emergent community patterns. We used structural equation models (SEM) to summarize the non-lethal and lethal effects of crayfish, Procambarus fallax, and phosphorus (P) addition, which affected prey food quality (periphyton), on the interactive effects of behavioral, morphological, developmental, and reproductive traits of snails, Planorbella duryi. Univariate and multivariate analyses suggested trade-offs between production (growth, reproduction) and defense (foraging behavior, shell shape) traits of snails in response to non-lethal crayfish and P addition, but few lethal effects. SEM revealed that non-lethal crayfish effects indirectly limited per capita offspring standing stock by increasing refuge use, slowing individual growth, and inducing snails to produce thicker, compressed shells. The negative effects of non-lethal crayfish on snails were strongest with P addition; snails increased allocation to shell defense rather than growth or reproduction. However, compared to ambient conditions, P addition with non-lethal crayfish still yielded greater per capita offspring standing stock by speeding individual snail growth enabling them to produce more offspring that also grew faster. Increased refuge use in response to non-lethal crayfish led to a non-lethal trophic cascade that altered the spatial distribution of periphyton. Independent of crayfish effects, snails stimulated periphyton growth through nutrient regeneration. These findings illustrate the importance of studying suites of traits that reveal costs associated with inducing different traits and how expressing those traits impacts population and community level processes.

Phenotypical responses of Fucus vesiculosus germlings to warming, acidification, nutrient enrichment and local upwelling under natural fluctuations (July - September 2014)

During summer 2014 (mid-July - mid-September 2014), early life-stage Fucus vesiculosus were exposed to combined ocean acidification and warming (OAW) in the presence and absence of enhanced nutrient levels (OAW x N experiment). Subsequently, F. vesiculosus germlings were exposed to a final upwelling disturbance during 3 days (mid-September 2014). Experiments were performed in the near-natural scenario "Kiel Outdoor Benthocosms" including natural fluctuations in the southwestern Baltic Sea, Kiel Fjord, Germany (54°27 'N, 10°11 'W). Genetically different sibling groups and different levels of genetic diversity were employed to test to which extent genetic variation would result in response variation. The data presented here show the phenotypical response (growth and survival) of the different experimental populations of F. vesiculosus under OAW, nutrient enrichment and the upwelling event. Log effect ratios demonstrate the responses to enhanced OAW and nutrient concentrations relative to the ambient conditons. Carbon, nitrogen content (% DW) and C:N ratios were measured after the exposure of ambient and high nutrient levels. Abiotic conditions the OAW x nutrient experiment and the upwelling event, are shown.

The response of surf-zone phytoplankton to nutrient enrichment (Cassino Beach, Brazil)

To understand the mechanisms that trigger changes in chlorophyll a and species composition in the phytoplankton of the surf-zone at Cassino Beach (RS), we performed two short nutrient-enrichment experiments (4–5 days each) during the summer and winter of 2010. Seawater was incubated under controlled conditions of temperature (summer 25± 3 °C, winter 18±1 °C), salinity (summer 28, winter 26) and irradiance (100 μmol m−2 s−1 ). Dissolved inorganic nutrients were added in various concentrations in the summer (silicate, Si; nitrate, N; phosphate, P) and winter (N, P) experiments. Samples were taken daily for cell counts and chlorophyll a analysis. In both experiments, chlorophyll a values and cell density showed a significant increase (mainly diatoms) in the treatments with nitrate addition, regardless of the proportion added. In the summer experiment, the largest chlorophyll a increase, approximately threefold (31.5 to 89.5 μg L−1 ), was observed in the NP treatment due to the growth of Asterionellopsis glacialis (Castracane) Round, Skeletonema tropicum Cleve, Thalassiosira sp. Cleve and Pseudo-nitzschia spp. Peragallo. The maximum growth was obtained in the SiNP treatment for S. tropicum (μ=0.7), Thalassiosira (μ= 1.9) and Pseudo-nitzschia (μ= 1.3) and in the SiN treatment for A. glacialis (μ= 1.0). In the winter experiment, the chlorophyll a content increased 4.2 and 5.5 times, respectively, in the N and NP treatments (maxima 38.8 μg L−1 and 31.5 μg L−1 ), where A. glacialis (μ= 1.7–1.9) and Cylindrotheca closterium (Ehrenberg) Reimann & J.C. Lewin (μ= 1.0–1.96) showed the highest amount of growth. These results indicate that nitrate is the most important nutrient controlling phytoplankton chlorophyll a at sandy Cassino Beach. However, the responses of different species to enrichment during the summer and winter indicated that other factors also played a role. A. glacialis, present during both seasons, presented the highest growth rate during the winter, whereas during the summer it was independent of nutrient enrichment but coincided with the lowest growth of S. tropicum. This finding suggested the occurrence of allelopathic interactions between these species. During the summer, multi-enrichment (SiNP) favoured the best growth of S. tropicum, Pseudo-nitzschia spp. and Thalassiosira sp. These results indicated that the phytoplankton composition and diversity in the surf zone of Cassino Beach are shaped by the availability of silicate and phosphorus as well as by the availability of nitrate.

Limiting factors for phytoplankton growth in subtropical reservoirs: the effect of light and nutrient availability in different longitudinal compartments

Limited information is available on the interactions between environmental factors and algal growth in tropical and subtropical aquatic systems. We investigated the relationships between algal biomass (measured as chlorophyll, Chl-a) and light, total phosphorus (TP) and total nitrogen (TN) in longitudinal zones of subtropical reservoirs. We studied the seasonal variation of water variables in Itupararanga Reservoir (Brazil) and compared the results with 16 other subtropical lakes and reservoirs. The longitudinal zones in Itupararanga Reservoir were considered statistically different (p 0.05, MANOVA). From the riverine zone to the dam region of the reservoir, Spearman Correlation Test suggested that light limitation and TP limitation tended to decrease and increase, respectively. Although nitrate concentrations were high (400 g/L), the Spearman coefficients between Chl-a and TN and the TN:TP ratios (11:1 TN:TP 35:1) indicated that nitrogen may be co-limiting algal growth in the studied water body. Putting Itupararanga in a regional context allowed assessment of potential influences of land use on trophic state. Within the subtropical dataset, TP explained a greater percentage of variance in Chl-a (R2 = 0.70) than TN (R2 = 0.17). The main land use type within the reservoirs drainage area significantly influenced the concentrations of TP, TN, and Chl-a (p 0.05, MANOVA), with different relationships between nutrients and chlorophyll in forested (R2 = 0.12-0.33), agricultural (R2 = 0.50-0.68) and urban (R2 = 0.09-0.64) watersheds. Comparisons with literature values and those from reservoirs with less altered watersheds indicated that Itupararanga Reservoir is reaching the mesotrophic-eutrophic boundary, and further nutrient enrichment could cause water quality degradation.

Multitrophic effects of nutrient addition in upland grassland

Although the effects of nutrient enhancement on aquatic systems are well documented, the consequences of nutritional supplements on soil food webs are poorly understood, and results of past research examining bottom-up effects are often conflicting. In addition, many studies have failed to separate the effects of nutrient enrichment and the physical effects of adding organic matter. In this field study, we hypothesised that the addition of nitrogen to soil would result in a trophic cascade, through detritivores (Collembola) to predators (spiders), increasing invertebrate numbers and diversity. Nitrogen and lime were added to plots in an upland grassland in a randomised block design. Populations of Collembola and spiders were sampled by means of pitfall traps and identified to species. Seventeen species of Collembola were identified from the nitrogen plus lime (N + L) and control plots. Species assemblage, diversity, richness, evenness and total number were not affected by nutrient additions. However, there was an increase in the number of Isotomidae juveniles and Parisotoma anglicana trapped in the N + L plots. Of the 44 spider species identified, over 80% were Linyphiidae. An effect on species assemblage from the addition of N + L to the plots was observed on two of the four sampling dates (July 2002 and June 2003). The linyphiid, Oedothorax retusus, was the only species significantly affected by the treatments and was more likely to be trapped in the control plots. The increased number of juvenile Collembola, and change in community composition of spiders, were consequences of the bottom-up effect caused by nutrient inputs. However, despite efforts to eliminate the indirect effects of nutrient inputs, a reduction in soil moisture in the N + L plots cannot be eliminated as a cause of the invertebrate population changes observed. Even so, this experiment was not confounded by the physical effects of habitat structure reported in most previous studies. It provides evidence of moderate bottom-up influences of epigeic soil invertebrate food webs and distinguishes between nutrient addition and plant physical structure effects. It also emphasises the importance Of understanding the effects of soil management practices on soil biodiversity, which is under increasing pressure from land development and food production.

Macroalgal responses to ocean acidification depend on nutrient and light levels

Ocean acidification may benefit algae that are able to capitalize on increased carbon availability for photosynthesis, but it is expected to have adverse effects on calcified algae through dissolution. Shifts in dominance between primary producers will have knock-on effects on marine ecosystems and will likely vary regionally, depending on factors such as irradiance (light vs. shade) and nutrient levels (oligotrophic vs. eutrophic). Thus experiments are needed to evaluate interactive effects of combined stressors in the field. In this study, we investigated the physiological responses of macroalgae near a CO2 seep in oligotrophic waters off Vulcano (Italy). The algae were incubated in situ at 0.2 m depth using a combination of three mean CO2 levels (500, 700-800 and 1200 µatm CO2), two light levels (100 and 70% of surface irradiance) and two nutrient levels of N, P, and K (enriched vs. non-enriched treatments) in the non-calcified macroalga Cystoseira compressa (Phaeophyceae, Fucales) and calcified Padina pavonica (Phaeophyceae, Dictyotales). A suite of biochemical assays and in vivo chlorophyll a fluorescence parameters showed that elevated CO2 levels benefitted both of these algae, although their responses varied depending on light and nutrient availability. In C. compressa, elevated CO2 treatments resulted in higher carbon content and antioxidant activity in shaded conditions both with and without nutrient enrichment-they had more Chla, phenols and fucoxanthin with nutrient enrichment and higher quantum yield (Fv/Fm) and photosynthetic efficiency (alpha ETR) without nutrient enrichment. In P. pavonica, elevated CO2 treatments had higher carbon content, Fv/Fm, alpha ETR, and Chla regardless of nutrient levels-they had higher concentrations of phenolic compounds in nutrient enriched, fully-lit conditions and more antioxidants in shaded, nutrient enriched conditions. Nitrogen content increased significantly in fertilized treatments, confirming that these algae were nutrient limited in this oligotrophic part of the Mediterranean. Our findings strengthen evidence that brown algae can be expected to proliferate as the oceans acidify where physicochemical conditions, such as nutrient levels and light, permit.

Ocean acidification outweighs nutrient effects in structuring seagrass epiphyte communities

1. Developing a framework for assessing interactions between multiple anthropogenic stressors remains an important goal in environmental research. In coastal ecosystems, the relative effects of aspects of global climate change (e.g. CO2 concentrations) and localized stressors (e.g. eutrophication), in combination, have received limited attention. 2. Using a long-term (11 month) field experiment, we examine how epiphyte assemblages in a tropical seagrass meadow respond to factorial manipulations of dissolved carbon dioxide (CO2(aq)) and nutrient enrichment. In situ CO2(aq) manipulations were conducted using clear, open-top chambers, which replicated carbonate parameter forecasts for the year 2100. Nutrient enrichment consisted of monthly additions of slow-release fertilizer, nitrogen (N) and phosphorus (P), to the sediments at rates equivalent to theoretical maximum rates of anthropogenic loading within the region (1.54 g N/m**2/d and 0.24 g P m**2/d). 3. Epiphyte community structure was assessed on a seasonal basis and revealed declines in the abundance of coralline algae, along with increases in filamentous algae under elevated CO2(aq). Surprisingly, nutrient enrichment had no effect on epiphyte community structure or overall epiphyte loading. Interactions between CO2(aq) and nutrient enrichment were not detected. Furthermore, CO2(aq)-mediated responses in the epiphyte community displayed strong seasonality, suggesting that climate change studies in variable environments should be conducted over extended time-scales. 4. Synthesis. The observed responses indicate that for certain locations, global stressors such as ocean acidification may take precedence over local eutrophication in altering the community structure of seagrass epiphyte assemblages. Given that nutrient-driven algal overgrowth is commonly cited as a widespread cause of seagrass decline, our findings highlight that alternate climate change forces may exert proximate control over epiphyte community structure.

Aeolian dust in Colorado Plateau soils: Nutrient inputs and recent change in source

Aeolian dust (windblown silt and clay) is an important component in arid-land ecosystems because it may contribute to soil formation and furnish essential nutrients. Few geologic surfaces, however, have been characterized with respect to dust-accumulation history and resultant nutrient enrichment. We have developed a combination of methods to identify the presence of aeolian dust in arid regions and to evaluate the roles of this dust in ecosystem processes. Unconsolidated sandy sediment on isolated surfaces in the Canyonlands region of the Colorado Plateau differs greatly in mineralogical and chemical composition from associated bedrock, mainly aeolian sandstone. Detrital magnetite in the surficial deposits produces moderately high values of magnetic susceptibility, but magnetite is absent in nearby bedrock. A component of the surficial deposits must be aeolian to account for the abundance of magnetite, which formed originally in far-distant igneous rocks. Particle-size analysis suggests that the aeolian dust component is typically as much as 20–30%. Dust inputs have enriched the sediments in many elements, including P, Mg, Na, K, and Mo, as well as Ca, at sites where bedrock lacks calcite cement. Soil-surface biologic crusts are effective dust traps that apparently record a change in dust sources over the past several decades. Some of the recently fallen dust may result from human disturbance of land surfaces that are far from the Canyonlands, such as the Mojave Desert. Some land-use practices in the study area have the potential to deplete soil fertility by means of wind-erosion removal of aeolian silt.

Stable isotopes reveal complex changes in trophic relationships following nutrient addition in a coastal marine ecosystem

Complex links between the top-down and bottomup forces that structure communities can be disrupted by anthropogenic alterations of natural habitats.We used relative abundance and stable isotopes to examine changes in epifaunal food webs in seagrass (Thalassia testudinum) beds following 6 months of experimental nutrient addition at two sites in Florida Bay (USA) with different ambient fertility. At a eutrophic site, nutrient addition did not strongly affect food web structure, but at a nutrient-poor site, enrichment increased the abundances of crustacean epiphyte grazers, and the diets of these grazers became more varied. Benthic grazers did not change in abundance but shifted their diet away from green macroalgae + associated epiphytes and towards an opportunistic seagrass (Halodule wrightii) that occurred only in nutrient addition treatments. Benthic predators did not change in abundance, but their diets were more varied in enriched plots. Food chain length was short and unaffected by site or nutrient treatment, but increased food web complexity in enriched plots was suggested by increasingly mixed diets. Strong bottom-up modifications of food web structure in the nutrient-limited site and the limited top-down influences of grazers on seagrass epiphyte biomass suggest that, in this system, the bottom-up role of nutrient enrichment can have substantial impacts on community structure, trophic relationships, and, ultimately, the productivity values of the ecosystem.

Nutrient impacts on epifaunal density and species composition in a subtropical seagrass bed

The capacity of epifauna to control algal proliferation following nutrient input depends on responses of both grazers and upper trophic level consumers to enrichment. We examined the responses of Thalassia testudinum (turtle grass) epifaunal assemblages to nutrient enrichment at two sites in Florida Bay with varying levels of phosphorus limitation. We compared epifaunal density, biomass, and species diversity in 2 m2 plots that had either ambient nutrient concentrations or had been enriched with nitrogen and phosphorus for 6 months. At the severely P-limited site, total epifaunal density and biomass were two times higher in enriched than in unenriched plots. Caridean shrimp, grazing isopods, and gammarid amphipods accounted for much of the increase in density; brachyuran crabs, primary predatory fish, and detritivorous sea cucumbers accounted for most of the increase in biomass. At the less P-limited site, total epifaunal density and biomass were not affected by nutrient addition, although there were more caridean shrimp and higher brachyuran crab and pink shrimp biomass in enriched plots. At both sites, some variation in epifaunal density and biomass was explained by features of the macrophyte canopy, such as T. testudinum and Halodule wrightii percent cover, suggesting that enrichment may change the refuge value of the macrophyte canopy for epifauna. Additional variation in epifaunal density and biomass was explained by epiphyte pigment concentrations, suggesting that enrichment may change the microalgal food resources that support grazing epifauna. Increased epifaunal density in enriched plots suggests that grazers may be able to control epiphytic algal proliferation following moderate nutrient input to Florida Bay.

Phosphorus Biogeochemistry and the Impact of Phosphorus Enrichment: Why Is the Everglades so Unique?

The Florida Everglades is extremely oligotrophic and sensitive to small increases in phosphorus (P) concentrations. P enrichment is one of the dominant anthropogenic impacts on the ecosystem and is therefore a main focus of restoration efforts. In this review, we synthesize research on P biogeochemistry and the impact of P enrichment on ecosystem structure and function in the Florida Everglades. There are clear patterns of increased P concentrations and altered structure and processes along nutrient-enrichment gradients in the water, periphyton, soils, macrophytes, and consumers. Periphyton, an assemblage of algae, bacteria, and associated microfauna, is abundant and has a large influence on phosphorus cycling in the Everglades. The oligotrophic Everglades is P-starved, has lower P concentrations and higher nitrogen–phosphorus (N:P) ratios, and has oxidized to only slightly reduced soil profiles compared to other freshwater wetland ecosystems. Possible general causes and indications of P limitation in the Everglades and other wetlands include geology, hydrology, and dominance of oxidative microbial nutrient cycling. The Everglades may be unique with respect to P biogeochemistry because of the multiple causes of P limitation and the resulting high degree of limitation.

Ocean Acidification Outweighs Nutrient Effects in Structuring Seagrass Epiphyte Communities

Developing a framework for assessing interactions between multiple anthropogenic stressors remains an important goal in environmental research. In coastal ecosystems, the relative effects of aspects of global climate change (e.g. CO2 concentrations) and localized stressors (e.g. eutrophication), in combination, have received limited attention. Using a long-term (11 month) field experiment, we examine how epiphyte assemblages in a tropical seagrass meadow respond to factorial manipulations of dissolved carbon dioxide (CO2(aq)) and nutrient enrichment. In situ CO2(aq) manipulations were conducted using clear, open-top chambers, which replicated carbonate parameter forecasts for the year 2100. Nutrient enrichment consisted of monthly additions of slow-release fertilizer, nitrogen (N) and phosphorus (P), to the sediments at rates equivalent to theoretical maximum rates of anthropogenic loading within the region (1.54 g N m−2 d−1 and 0.24 g P m−2 d−1). Epiphyte community structure was assessed on a seasonal basis and revealed declines in the abundance of coralline algae, along with increases in filamentous algae under elevated CO2(aq). Surprisingly, nutrient enrichment had no effect on epiphyte community structure or overall epiphyte loading. Interactions between CO2(aq) and nutrient enrichment were not detected. Furthermore, CO2(aq)-mediated responses in the epiphyte community displayed strong seasonality, suggesting that climate change studies in variable environments should be conducted over extended time-scales. Synthesis. The observed responses indicate that for certain locations, global stressors such as ocean acidification may take precedence over local eutrophication in altering the community structure of seagrass epiphyte assemblages. Given that nutrient-driven algal overgrowth is commonly cited as a widespread cause of seagrass decline, our findings highlight that alternate climate change forces may exert proximate control over epiphyte community structure.

Boundary Effects on Benthic Microbial Phosphorus Concentrations and Diatom Beta Diversity in a Hydrologically-modified, Nutrient-limited Wetland

Water flow and flooding duration in wetlands influence the structure and productivity of microbial communities partly through their influence on nutrient loading. The effect of flow-regulated nutrient loads is especially relevant for microbial communities in nutrient-poor settings, where delivery controls nutrient uptake rates and the intensity of microbial interactions. We examined the effect of hydrologic history and proximity to water sources on nutrient enrichment of benthic microbial assemblages (periphyton) and on their diatom species composition, along the artificial boundaries of Taylor Slough, a historically phosphorus-depleted drainage of the Florida Everglades. Concentrations of phosphorus in periphyton declined from the wetland boundary near inflow structures to 100-m interior, with spatial and temporal variability in rates dependent on proximity to and magnitude of water flow. Phosphorus availability influenced the beta diversity of diatom assemblages, with higher values near inflow structures where resources were greatest, while interior sites and reference transects contained assemblages with constant composition of taxa considered endemic to the Everglades. This research shows how hydrologic restoration may have unintended consequences when incoming water quality is not regulated, including a replacement of distinctive microbial assemblages by ubiquitous, cosmopolitan ones.