Coastal eutrophication as a driver of salt marsh loss.

Salt marshes are highly productive coastal wetlands that provide important ecosystem services such as storm protection for coastal cities, nutrient removal and carbon sequestration. Despite protective measures, however, worldwide losses of these ecosystems have accelerated in recent decades. Here we present data from a nine-year whole-ecosystem nutrient-enrichment experiment. Our study demonstrates that nutrient enrichment, a global problem for coastal ecosystems, can be a driver of salt marsh loss. We show that nutrient levels commonly associated with coastal eutrophication increased above-ground leaf biomass, decreased the dense, below-ground biomass of bank-stabilizing roots, and increased microbial decomposition of organic matter. Alterations in these key ecosystem properties reduced geomorphic stability, resulting in creek-bank collapse with significant areas of creek-bank marsh converted to unvegetated mud. This pattern of marsh loss parallels observations for anthropogenically nutrient-enriched marshes worldwide, with creek-edge and bay-edge marsh evolving into mudflats and wider creeks. Our work suggests that current nutrient loading rates to many coastal ecosystems have overwhelmed the capacity of marshes to remove nitrogen without deleterious effects. Projected increases in nitrogen flux to the coast, related to increased fertilizer use required to feed an expanding human population, may rapidly result in a coastal landscape with less marsh, which would reduce the capacity of coastal regions to provide important ecological and economic services.

Asynchrony among local communities stabilises ecosystem function of metacommunities.

Temporal stability of ecosystem functioning increases the predictability and reliability of ecosystem services, and understanding the drivers of stability across spatial scales is important for land management and policy decisions. We used species-level abundance data from 62 plant communities across five continents to assess mechanisms of temporal stability across spatial scales. We assessed how asynchrony (i.e. different units responding dissimilarly through time) of species and local communities stabilised metacommunity ecosystem function. Asynchrony of species increased stability of local communities, and asynchrony among local communities enhanced metacommunity stability by a wide range of magnitudes (1-315%); this range was positively correlated with the size of the metacommunity. Additionally, asynchronous responses among local communities were linked with species' populations fluctuating asynchronously across space, perhaps stemming from physical and/or competitive differences among local communities. Accordingly, we suggest spatial heterogeneity should be a major focus for maintaining the stability of ecosystem services at larger spatial scales.

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 m2 ) 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 µM NO3- ), the most common form of coastal nutrient enrichment, via tidal water at the ecosystem scale (~60,000 m2 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.

A grazing crab drives saltmarsh carbon storage and recovery.

Consumers can directly (e.g., consumption) and indirectly (e.g., trophic cascades) influence carbon cycling in blue carbon ecosystems. Previous work found that large grazers have nuanced effects on carbon stocks, yet, small, bioturbating-grazers, which remove plant biomass and alter sediment properties, remain an understudied driver of carbon cycling. We used field-derived and remote sensing data to quantify how the purple marsh crab, Sesarma reticulatum, influenced carbon stocks, flux, and recovery in salt marshes. Sesarma caused a 40%-70% loss in carbon stocks as fronts propagated inland (i.e., ungrazed to recovered transition), with front migration rates accelerating over time. Despite latitudinal differences, front migration rate had no effect on carbon stocks, flux, or time to replacement. When we included Sesarma disturbance in carbon flux calculations, we found it may take 5-100 years for marshes to replace lost carbon, if at all. Combined, we show that small grazers cause a net loss in carbon stocks as they move through the landscape, and irrespective of migration rate, these grazer-driven impacts persist for decades. This work showcases the significant role of consumers in carbon storage and flux, challenging the classic paradigm of plant-sediment feedbacks as the primary ecogeomorphic driver of carbon cycling in blue carbon ecosystems.

The fiddler crab, Minuca pugnax, follows Bergmann's rule.

Bergmann's rule predicts that organisms at higher latitudes are larger than ones at lower latitudes. Here, we examine the body size pattern of the Atlantic marsh fiddler crab, Minuca pugnax (formerly Uca pugnax), from salt marshes on the east coast of the United States across 12 degrees of latitude. We found that M. pugnax followed Bergmann's rule and that, on average, crab carapace width increased by 0.5 mm per degree of latitude. Minuca pugnax body size also followed the temperature-size rule with body size inversely related to mean water temperature. Because an organism's size influences its impact on an ecosystem, and M. pugnax is an ecosystem engineer that affects marsh functioning, the larger crabs at higher latitudes may have greater per-capita impacts on salt marshes than the smaller crabs at lower latitudes.

Pattern and scale: evaluating generalities in crab distributions and marsh dynamics from small plots to a national scale.

The generality of ecological patterns depends inextricably on the scale at which they are examined. We investigated patterns of crab distribution and the relationship between crabs and vegetation in salt marshes at multiple scales. By using consistent monitoring protocols across 15 U.S. National Estuarine Research Reserves, we were able to synthesize patterns from the scale of quadrats to the entire marsh landscape to regional and national scales. Some generalities emerged across marshes from our overall models, and these are useful for informing broad coastal management policy. We found that crab burrow distribution within a marsh could be predicted by marsh elevation, distance to creek and soil compressibility. While these physical factors also affected marsh vegetation cover, we did not find a strong or consistent overall effect of crabs at a broad scale in our multivariate model, though regressions conducted separately for each site revealed that crab burrows were negatively correlated with vegetation cover at 4 out of 15 sites. This contrasts with recent smaller-scale studies and meta-analyses synthesizing such studies that detected strong negative effects of crabs on marshes, likely because we sampled across the entire marsh landscape, while targeted studies are typically limited to low-lying areas near creeks, where crab burrow densities are highest. Our results suggest that sea-level rise generally poses a bigger threat to marshes than crabs, but there will likely be interactions between these physical and biological factors. Beyond these generalities across marshes, we detected some regional differences in crab community composition, richness, and abundance. However, we found striking differences among sites within regions, and within sites, in terms of crab abundance and relationships to marsh integrity. Although generalities are broadly useful, our findings indicate that local managers cannot rely on data from other nearby systems, but rather need local information for developing salt marsh management strategies.

It's about time: A synthesis of changing phenology in the Gulf of Maine ecosystem.

The timing of recurring biological and seasonal environmental events is changing on a global scale relative to temperature and other climate drivers. This study considers the Gulf of Maine ecosystem, a region of high social and ecological importance in the Northwest Atlantic Ocean and synthesizes current knowledge of (a) key seasonal processes, patterns, and events; (b) direct evidence for shifts in timing; (c) implications of phenological responses for linked ecological-human systems; and (d) potential phenology-focused adaptation strategies and actions. Twenty studies demonstrated shifts in timing of regional marine organisms and seasonal environmental events. The most common response was earlier timing, observed in spring onset, spring and winter hydrology, zooplankton abundance, occurrence of several larval fishes, and diadromous fish migrations. Later timing was documented for fall onset, reproduction and fledging in Atlantic puffins, spring and fall phytoplankton blooms, and occurrence of additional larval fishes. Changes in event duration generally increased and were detected in zooplankton peak abundance, early life history periods of macro-invertebrates, and lobster fishery landings. Reduced duration was observed in winter-spring ice-affected stream flows. Two studies projected phenological changes, both finding diapause duration would decrease in zooplankton under future climate scenarios. Phenological responses were species-specific and varied depending on the environmental driver, spatial, and temporal scales evaluated. Overall, a wide range of baseline phenology and relevant modeling studies exist, yet surprisingly few document long-term shifts. Results reveal a need for increased emphasis on phenological shifts in the Gulf of Maine and identify opportunities for future research and consideration of phenological changes in adaptation efforts.

Sea level rise may increase extinction risk of a saltmarsh ontogenetic habitat specialist.

Specialist species are more vulnerable to environmental change than generalist species. For species with ontogenetic niche shifts, specialization may occur at a particular life stage making those stages more susceptible to environmental change. In the salt marshes in the northeast U.S., accelerated sea level rise is shifting vegetation patterns from flood-intolerant species such as Spartina patens to the flood-tolerant Spartina alterniflora. We tested the potential impact of this change on the coffee bean snail, Melampus bidentatus, a numerically dominant benthic invertebrate with an ontogenetic niche shift. From a survey of eight marshes throughout the northeast U.S., small snails were found primarily in S. patens habitats, and large snails were found primarily in stunted S. alterniflora habitats. When transplanted into stunted S. alterniflora, small snails suffered significantly higher mortality relative to those in S. patens habitats; adult snail survivorship was similar between habitats. Because other habitats were not interchangeable with S. patens for young snails, these results suggest that Melampus is an ontogenetic specialist where young snails are habitat specialists and adult snails are habitat generalists. Temperature was significantly higher and relative humidity significantly lower in stunted S. alterniflora than in S. patens. These data suggest that thermal and desiccation stress restricted young snails to S. patens habitat, which has high stem density and a layer of thatch that protects snails from environmental stress. Other authors predict that if salt marshes in the northeast U.S. are unable to migrate landward, sea level rise will eliminate S. patens habitats. We suggest that if a salt marsh loses its S. patens habitats, it will also lose its coffee bean snails. Our results demonstrate the need to consider individual life stages when determining a species' vulnerability to global change.