Brian R. Silliman

The value of estuarine and coastal ecosystem services

The global decline in estuarine and coastal ecosystems (ECEs) is affecting a number of critical benefits, or ecosystem services. We review the main ecological services across a variety of ECEs, including marshes, mangroves, nearshore coral reefs, seagrass beds, and sand beaches and dunes. Where possible, we indicate estimates of the key economic values arising from these services, and discuss how the natural variability of ECEs impacts their benefits, the synergistic relationships of ECEs across seascapes, and management implications. Although reliable valuation estimates are beginning to emerge for the key services of some ECEs, such as coral reefs, salt marshes, and mangroves, many of the important benefits of seagrass beds and sand dunes and beaches have not been assessed properly. Even for coral reefs, marshes, and mangroves, important ecological services have yet to be valued reliably, such as cross-ecosystem nutrient transfer (coral reefs), erosion control (marshes), and pollution control (mangroves). An important issue for valuing certain ECE services, such as coastal protection and habitat–fishery linkages, is that the ecological functions underlying these services vary spatially and temporally. Allowing for the connectivity between ECE habitats also may have important implications for assessing the ecological functions underlying key ecosystems services, such coastal protection, control of erosion, and habitat–fishery linkages. Finally, we conclude by suggesting an action plan for protecting and/or enhancing the immediate and longer-term values of ECE services. Because the connectivity of ECEs across land–sea gradients also influences the provision of certain ecosystem services, management of the entire seascape will be necessary to preserve such synergistic effects. Other key elements of an action plan include further ecological and economic collaborative research on valuing ECE services, improving institutional and legal frameworks for management, controlling and regulating destructive economic activities, and developing ecological restoration options.

A blueprint for blue carbon: toward an improved understanding of the role of vegetated coastal habitats in sequestering CO2

Recent research has highlighted the valuable role that coastal and marine ecosystems play in sequestering carbon dioxide (CO(2)). The carbon (C) sequestered in vegetated coastal ecosystems, specifically mangrove forests, seagrass beds, and salt marshes, has been termed blue carbon. Although their global area is one to two orders of magnitude smaller than that of terrestrial forests, the contribution of vegetated coastal habitats per unit area to long-term C sequestration is much greater, in part because of their efficiency in trapping suspended matter and associated organic C during tidal inundation. Despite the value of mangrove forests, seagrass beds, and salt marshes in sequestering C, and the other goods and services they provide, these systems are being lost at critical rates and action is urgently needed to prevent further degradation and loss. Recognition of the C sequestration value of vegetated coastal ecosystems provides a strong argument for their protection and restoration; however, it is necessary to improve scientific understanding of the underlying mechanisms that control C sequestration in these ecosystems. Here, we identify key areas of uncertainty and specific actions needed to address them.

Coastal Ecosystem-Based Management with Nonlinear Ecological Functions and Values

A common assumption is that ecosystem services respond linearly to changes in habitat size. This assumption leads frequently to an “all or none” choice of either preserving coastal habitats or converting them to human use. However, our survey of wave attenuation data from field studies of mangroves, salt marshes, seagrass beds, nearshore coral reefs, and sand dunes reveals that these relationships are rarely linear. By incorporating nonlinear wave attenuation in estimating coastal protection values of mangroves in Thailand, we show that the optimal land use option may instead be the integration of development and conservation consistent with ecosystem-based management goals. This result suggests that reconciling competing demands on coastal habitats should not always result in stark preservation-versus-conversion choices.

Non‐linearity in ecosystem services: temporal and spatial variability in coastal protection

Natural processes tend to vary over time and space, as well as between species. The ecosystem services these natural processes provide are therefore also highly variable. It is often assumed that ecosystem services are provided linearly (unvaryingly, at a steady rate), but natural processes are characterized by thresholds and limiting functions. In this paper, we describe the variability observed in wave attenuation provided by marshes, mangroves, seagrasses, and coral reefs and therefore also in coastal protection. We calculate the economic consequences of assuming coastal protection to be linear. We suggest that, in order to refine ecosystem-based management practices, it is essential that natural variability and cumulative effects be considered in the valuation of ecosystem services.

A trophic cascade regulates salt marsh primary production.

Nutrient supply is widely thought to regulate primary production of many ecosystems including salt marshes. However, experimental manipulation of the dominant marsh grazer (the periwinkle, Littoraria irrorata) and its consumers (e.g., blue crabs, Callinectes sapidus, terrapins, Malaclemys terrapin) demonstrates plant biomass and production are largely controlled by grazers and their predators. Periwinkle grazing can convert one of the most productive grasslands in the world into a barren mudflat within 8 months. Marine predators regulate the abundance of this plant-grazing snail. Thus, top-down control of grazer density is a key regulatory determinant of marsh grass growth. The discovery of this simple trophic cascade implies that over-harvesting of snail predators (e.g., blue crabs) may be an important factor contributing to the massive die-off (tens of km2) of salt marshes across the southeastern United States. In addition, our results contribute to a growing body of evidence indicating widespread, predator regulation of marine macrophyte production via trophic cascades (kelps, seagrasses, intertidal algae).

Anthropogenic modification of New England salt marsh landscapes

Salt marshes play a critical role in the ecology and geology of wave-protected shorelines in the Western Atlantic, but as many as 80% of the marshes that once occurred in New England have already been lost to human development. Here we present data that suggest that the remaining salt marshes in southern New England are being rapidly degraded by shoreline development and eutrophication. On the seaward border of these marshes, nitrogen eutrophication stimulated by local shoreline development is shifting the competitive balance among marsh plants by releasing plants from nutrient competition. This shift is leading to the displacement of natural high marsh plants by low marsh cordgrass. On the terrestrial border of these same marshes, shoreline development is also precipitating the invasion of the common reed, Phragmites, by means of nitrogen eutrophication caused by the removal of the woody vegetation buffer between terrestrial and salt marsh communities. As a consequence of these human impacts, traditional salt marsh plant communities and the plants and animals that are dependent on these habitats are being displaced by monocultures of weedy species.

Drought, Snails, and Large-Scale Die-Off of Southern U.S. Salt Marshes

Salt marshes in the southeastern United States have recently experienced massive die-off, one of many examples of widespread degradation in marine and coastal ecosystems. Although intense drought is thought to be the primary cause of this die-off, we found snail grazing to be a major contributing factor. Survey of marsh die-off areas in three states revealed high-density fronts of snails on die-off edges at 11 of 12 sites. Exclusion experiments demonstrated that snails actively converted marshes to exposed mudflats. Salt addition and comparative field studies suggest that drought-induced stress and grazers acted synergistically and to varying degrees to cause initial plant death. After these disturbances, snail fronts formed on die-off edges and subsequently propagated through healthy marsh, leading to cascading vegetation loss. These results, combined with model analyses, reveal strong interactions between increasing climatic stress and grazer pressure, both potentially related to human environmental impacts, which amplify the likelihood and intensity of runaway collapse in these coastal systems.

PHYSICAL AND BIOTIC DRIVERS OF PLANT DISTRIBUTION ACROSS ESTUARINE SALINITY GRADIENTS

Although it has long been recognized that marsh plant community composition shifts across estuarine salinity gradients, the mechanisms responsible for this species zonation have never been experimentally examined. In southern New England marshes of the United States, we investigated the relative importance of physical and biotic factors in generating estuarine species distribution patterns. Greenhouse studies revealed that all of the common plants in this system grow better in fresh water than in full-strength salt water. To test the hypothesis that the spatial segregation of these plants is driven by differential tolerance to salt stress and plant competition, we performed transplant experiments with 10 common plants in the system. When freshwater marsh plants were transplanted to salt marshes, they did poorly and generally died with or without neighbors present. In contrast, when saltmarsh plants were transplanted to freshwater marshes, they thrived in the absence of neighbors, growing better than they did in salt marshes, but when neighbors were present, they were strongly suppressed. These results suggest that the spatial segregation of plants across estuarine salinity gradients is driven by competitively superior freshwater marsh plants displacing salt-tolerant plants to physically harsh saltmarsh habitats, whereas freshwater marsh plants are limited from living in salt marshes by physical factors (e.g., high salinities). These results contribute to our understanding of the organization and assembly of tidal marsh plant communities and have important implications for understanding how marsh plant communities will respond to human modification of estuarine hydrology and climate change.

TOP‐DOWN CONTROL OF SPARTINA ALTERNIFLORA PRODUCTION BY PERIWINKLE GRAZING IN A VIRGINIA SALT MARSH

Top-down forces, such as grazing and predation, have long been thought to be unimportant in controlling plant growth in salt marshes. Instead, bottom-up forces, such as porewater ammonium and oxygen availability, are thought to be the primary regulating factors. In the field, we observed the periwinkle, Littoraria irrorata, grazing on live saltmarsh cordgrass, Spartina alterniflora. To examine the relative importance of periwinkle grazing and nutrient availability in controlling marsh grass growth, we manipulated snail and nitrogen levels in a season-long field experiment in a Virginia salt marsh. Snails differentially affected plant growth at varying nitrogen levels. In unfertilized plots, snail removal increased S. alterniflora aboveground production by 38%, whereas in nitrogen addition plots, snail removal increased cordgrass growth by 78%. Snail addition decreased aboveground production by 51% in unfertilized stands, while in fertilized stands, snail addition led to even greater reductions (66%). By comparison, nitrogen addition increased S. alterniflora production by 443% in snail removal treatments, while in controls and snail addition treatments, fertilization effects were dampened significantly, as nitrogen enrichment increased marsh grass growth by 322% and 189%, respectively. Feeding assays examining the rate at which snails ingested live S. alterniflora revealed that snail consumption alone could not be responsible for these large biomass reductions. Experimental results suggest that grazer-induced defoliation and grazer control of plant demand for nitrogen fertilizer were the likely mechanisms involved. A survey of salt marshes from Maryland to Georgia showed that periwinkle grazing on live S. alterniflora is widespread. Our results show that L. irrorata can exert strong top-down control of S. alterniflora production, and that this effect increases with increasing nitrogen avaliability. Together, these findings question the widely accepted notion that grazers play a relatively unimportant role in the salt marsh community.

Degradation and resilience in Louisiana salt marshes after the BP–Deepwater Horizon oil spill

More than 2 y have passed since the BP–Deepwater Horizon oil spill in the Gulf of Mexico, yet we still have little understanding of its ecological impacts. Examining effects of this oil spill will generate much-needed insight into how shoreline habitats and the valuable ecological services they provide (e.g., shoreline protection) are affected by and recover from large-scale disturbance. Here we report on not only rapid salt-marsh recovery (high resilience) but also permanent marsh area loss after the BP–Deepwater Horizon oil spill. Field observations, experimental manipulations, and wave-propagation modeling reveal that (i) oil coverage was primarily concentrated on the seaward edge of marshes; (ii) there were thresholds of oil coverage that were associated with severity of salt-marsh damage, with heavy oiling leading to plant mortality; (iii) oil-driven plant death on the edges of these marshes more than doubled rates of shoreline erosion, further driving marsh platform loss that is likely to be permanent; and (iv) after 18 mo, marsh grasses have largely recovered into previously oiled, noneroded areas, and the elevated shoreline retreat rates observed at oiled sites have decreased to levels at reference marsh sites. This paper highlights that heavy oil coverage on the shorelines of Louisiana marshes, already experiencing elevated retreat because of intense human activities, induced a geomorphic feedback that amplified this erosion and thereby set limits to the recovery of otherwise resilient vegetation. It thus warns of the enhanced vulnerability of already degraded marshes to heavy oil coverage and provides a clear example of how multiple human-induced stressors can interact to hasten ecosystem decline.