Saltmarsh plant responses to eutrophication

In saltmarsh plant communities, bottom-up pressure from nutrient enrichment is predicted to increase productivity, alter community structure, decrease biodiversity, and alter ecosystem functioning. Previous work supporting these predictions has been based largely on short-term, plot-level (e.g., 1-300 m(2)) studies, which may miss landscape-level phenomena that drive ecosystem-level responses. We implemented an ecosystem-scale, nine-year nutrient experiment to examine how saltmarsh plants respond to simulated conditions of coastal eutrophication. Our study differed from previous saltmarsh enrichment studies in that we applied realistic concentrations of nitrate (70-100 mu M NO3-), the most common form of coastal nutrient enrichment, via tidal water at the ecosystem scale (similar to 60,000 m(2) creeksheds). Our enrichments added a total of 1,700 kg N.creek(-1).yr(-1), which increased N loading 10-fold vs. reference creeks (low-marsh, 171 g N.m(-2).yr(-1); high-marsh, 19 g N.m(-2).yr(-1)). Nutrients increased the shoot mass and height of low marsh, tall Spartina alterniflora; however, declines in stem density resulted in no consistent increase in aboveground biomass. High-marsh plants S. patens and stunted S. alterniflora did not respond consistently to enrichment. Nutrient enrichment did not shift community structure, contrary to the prediction of nutrient-driven dominance of S. alterniflora and Distichlis spicata over S. patens. Our mild responses may differ from the results of previous studies for a number of reasons. First, the limited response of the high marsh may be explained by loading rates orders of magnitude lower than previous work. Low loading rates in the high marsh reflect infrequent inundation, arguing that inundation patterns must be considered when predicting responses to estuarine eutrophication. Additionally, we applied nitrate instead of the typically used ammonium, which is energetically favored over nitrate for plant uptake. Thus, the form of nitrogen enrichment used, not just N-load, may be important in predicting plant responses. Overall, our results suggest that when coastal eutrophication is dominated by nitrate and delivered via flooding tidal water, aboveground saltmarsh plant responses may be limited despite moderate-to-high water-column N concentrations. Furthermore, we argue that the methodological limitations of nutrient studies must be considered when using results to inform management decisions about wetlands

Global change accelerates carbon assimilation by a wetland ecosystem engineer

The primary productivity of coastal wetlands is changing dramatically in response to rising atmospheric carbon dioxide (CO2) concentrations, nitrogen (N) enrichment, and invasions by novel species, potentially altering their ecosystem services and resilience to sea level rise. In order to determine how these interacting global change factors will affect coastal wetland productivity, we quantified growing-season carbon assimilation (≈gross primary productivity, or GPP) and carbon retained in living plant biomass (≈net primary productivity, or NPP) of North American mid-Atlantic saltmarshes invaded by Phragmites australis (common reed) under four treatment conditions: two levels of CO2 (ambient and +300 ppm) crossed with two levels of N (0 and 25 g N added m−2 yr−1). For GPP, we combined descriptions of canopy structure and leaf-level photosynthesis in a simulation model, using empirical data from an open-top chamber field study. Under ambient CO2 and low N loading (i.e., the Control), we determined GPP to be 1.66 ± 0.05 kg C m−2 yr−1 at a typical Phragmites stand density. Individually, elevated CO2 and N enrichment increased GPP by 44 and 60%, respectively. Changes under N enrichment came largely from stimulation to carbon assimilation early and late in the growing season, while changes from CO2 came from stimulation during the early and mid-growing season. In combination, elevated CO2 and N enrichment increased GPP by 95% over the Control, yielding 3.24 ± 0.08 kg C m−2 yr−1. We used biomass data to calculate NPP, and determined that it represented 44%–60% of GPP, with global change conditions decreasing carbon retention compared to the Control. Our results indicate that Phragmites invasions in eutrophied saltmarshes are driven, in part, by extended phenology yielding 3.1× greater NPP than native marsh. Further, we can expect elevated CO2 to amplify Phragmites productivity throughout the growing season, with potential implications including accelerated spread and greater carbon storage belowground

Responses of stomatal features and photosynthesis to porewater N enrichment and elevated atmospheric CO2 in Phragmites australis, the common reed

PREMISE Biological invasions increasingly threaten native biodiversity and ecosystem services. One notable example is the common reed, Phragmites australis, which aggressively invades North American salt marshes. Elevated atmospheric CO2 and nitrogen pollution enhance its growth and facilitate invasion because P. australis responds more strongly to these enrichments than do native species. We investigated how modifications to stomatal features contribute to strong photosynthetic responses to CO2 and nitrogen enrichment in P. australis by evaluating stomatal shifts under experimental conditions and relating them to maximal stomatal conductance (g(wmax)) and photosynthetic rates.METHODS Plants were grown in situ in open-top chambers under ambient and elevated atmospheric CO2 (eCO(2)) and porewater nitrogen (N-enr) in a Chesapeake Bay tidal marsh. We measured light-saturated carbon assimilation rates (A(sat)) and stomatal characteristics, from which we calculated g(wmax) and determined whether CO2 and N-enr altered the relationship between g(wmax) and A(sat).RESULTS eCO(2) and N-enr enhanced both g(wmax) and A(sat), but to differing degrees; g(wmax) was more strongly influenced by N-enr through increases in stomatal density while A(sat) was more strongly stimulated by eCO(2). There was a positive relationship between g(wmax) and A(sat) that was not modified by eCO(2) or N-enr, individually or in combination.CONCLUSIONS Changes in stomatal features co-occur with previously described responses of P. australis to eCO(2) and N-enr. Complementary responses of stomatal length and density to these global change factors may facilitate greater stomatal conductance and carbon gain, contributing to the invasiveness of the introduced lineage

Plant species determine tidal wetland methane response to sea level rise

Blue carbon (C) ecosystems are among the most effective C sinks of the biosphere, but methane (CH4) emissions can offset their climate cooling effect. Drivers of CH4 emissions from blue C ecosystems and effects of global change are poorly understood. Here we test for the effects of sea level rise (SLR) and its interactions with elevated atmospheric CO2, eutrophication, and plant community composition on CH4 emissions from an estuarine tidal wetland. Changes in CH4 emissions with SLR are primarily mediated by shifts in plant community composition and associated plant traits that determine both the direction and magnitude of SLR effects on CH4 emissions. We furthermore show strong stimulation of CH4 emissions by elevated atmospheric CO2, whereas effects of eutrophication are not significant. Overall, our findings demonstrate a high sensitivity of CH4 emissions to global change with important implications for modeling greenhouse-gas dynamics of blue C ecosystems

Plant species determine tidal wetland methane response to sea level rise

Blue carbon (C) ecosystems are among the most effective C sinks of the biosphere, but methane (CH4) emissions can offset their climate cooling effect. Drivers of CH4 emissions from blue C ecosystems and effects of global change are poorly understood. Here we test for the effects of sea level rise (SLR) and its interactions with elevated atmospheric CO2, eutrophication, and plant community composition on CH4 emissions from an estuarine tidal wetland. Changes in CH4 emissions with SLR are primarily mediated by shifts in plant community composition and associated plant traits that determine both the direction and magnitude of SLR effects on CH4 emissions. We furthermore show strong stimulation of CH4 emissions by elevated atmospheric CO2, whereas effects of eutrophication are not significant. Overall, our findings demonstrate a high sensitivity of CH4 emissions to global change with important implications for modeling greenhouse-gas dynamics of blue C ecosystems

Livestock as a potential biological control agent for an invasive wetland plant

Invasive species threaten biodiversity and incur costs exceeding billions of US$. Eradication efforts, however, are nearly always unsuccessful. Throughout much of North America, land managers have used expensive, and ultimately ineffective, techniques to combat invasive Phragmites australis in marshes. Here, we reveal that Phragmites may potentially be controlled by employing an affordable measure from its native European range: livestock grazing. Experimental field tests demonstrate that rotational goat grazing (where goats have no choice but to graze Phragmites) can reduce Phragmites cover from 100 to 20% and that cows and horses also readily consume this plant. These results, combined with the fact that Europeans have suppressed Phragmites through seasonal livestock grazing for 6,000 years, suggest Phragmites management can shift to include more economical and effective top-down control strategies. More generally, these findings support an emerging paradigm shift in conservation from high-cost eradication to economically sustainable control of dominant invasive species

Not All Nitrogen Is Created Equal: Differential Effects of Nitrate and Ammonium Enrichment in Coastal Wetlands

Excess reactive nitrogen (N) flows from agricultural, suburban, and urban systems to coasts, where it causes eutrophication. Coastal wetlands take up some of this N, thereby ameliorating the impacts on nearshore waters. Although the consequences of N on coastal wetlands have been extensively studied, the effect of the specific form of N is not often considered. Both oxidized N forms (nitrate, NO3−) and reduced forms (ammonium, NH4+) can relieve nutrient limitation and increase primary production. However, unlike NH4+, NO3− can also be used as an electron acceptor for microbial respiration. We present results demonstrating that, in salt marshes, microbes use NO3− to support organic matter decomposition and primary production is less stimulated than when enriched with reduced N. Understanding how different forms of N mediate the balance between primary production and decomposition is essential for managing coastal wetlands as N enrichment and sea level rise continue to assail our coasts

Cosmopolitan Species As Models for Ecophysiological Responses to Global Change: The Common Reed \u3cem\u3ePhragmites australis\u3c/em\u3e

Phragmites australis is a cosmopolitan grass and often the dominant species in the ecosystems it inhabits. Due to high intraspecific diversity and phenotypic plasticity, P. australis has an extensive ecological amplitude and a great capacity to acclimate to adverse environmental conditions; it can therefore offer valuable insights into plant responses to global change. Here we review the ecology and ecophysiology of prominent P. australis lineages and their responses to multiple forms of global change. Key findings of our review are that: (1) P. australis lineages are well-adapted to regions of their phylogeographic origin and therefore respond differently to changes in climatic conditions such as temperature or atmospheric CO2; (2) each lineage consists of populations that may occur in geographically different habitats and contain multiple genotypes; (3) the phenotypic plasticity of functional and fitness-related traits of a genotype determine the responses to global change factors; (4) genotypes with high plasticity to environmental drivers may acclimate or even vastly expand their ranges, genotypes of medium plasticity must acclimate or experience range-shifts, and those with low plasticity may face local extinction; (5) responses to ancillary types of global change, like shifting levels of soil salinity, flooding, and drought, are not consistent within lineages and depend on adaptation of individual genotypes. These patterns suggest that the diverse lineages of P. australis will undergo intense selective pressure in the face of global change such that the distributions and interactions of co-occurring lineages, as well as those of genotypes within-lineages, are very likely to be altered. We propose that the strong latitudinal clines within and between P. australis lineages can be a useful tool for predicting plant responses to climate change in general and present a conceptual framework for using P. australis lineages to predict plant responses to global change and its consequences