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Coastal ecosystems, such as sand dunes, salt marshes, and mangroves, stabilize shorelines and protect coastal populations. In New England, salt marshes have experienced widespread cordgrass (Spartina alterniflora) die-off and habitat loss, and it is unknown how this has affected their ability to provide coastal protection. We quantified wave attenuation and shoreline stability on healthy, die-off and recovered marsh creek banks. We found that coastal protection has been severely compromised by salt marsh die-off, and that to date, S. alterniflora recovery, while superficially impressive, has not returned this ecosystem service to the levels of intact marshes. Climate driven sea-level rise and predicted increases in the frequency and severity of storms over the next century will likely further increase the vulnerability of coastal populations. Therefore, recovery of coastal protection is essential for maintaining the ecological and economic wellbeing of coastal communities. Our results suggest that quantification of the recovery of ecosystem services should be employed in order to successfully measure recovery in degraded ecosystems.

Understanding ecosystem resilience to human impacts is critical for conserva- tion and restoration. The large-scale die-off of New England salt marshes was triggered by overfishing and resulted from decades of runaway crab grazing. In 2009, however, cordgrass began to recover, decreasing die-off ;40% by 2010. We used surveys and experiments to test whether plant-substrate feedbacks underlie marsh resilience. Initially, grazer-generated die-off swept through the cordgrass, creating exposed, stressful peat banks that inhibited plant growth. This desertification cycle broke when banks eroded and peat transitioned into mud with fewer herbivores, less grazing, and lower physical stress. Cordgrass reestablished in these areas through a feedback where it engineered a recovery zone by further ameliorating physical stresses and facilitating additional revegetation. Our results reveal that feedbacks can play a critical role in rapid, reversible ecosystem shifts associated with human impacts, and that the interplay of facilitative and consumer interactions should be incorporated into resilience theory.

Coastal areas are among the world's most productive and highly affected ecosystems. Centuries of human activity on coastlines have led to overexploitation of marine predators, which in turn has led to cascading ecosystem-level effects. Human effects and approaches to mediating them, however, differ regionally due to gradients in biotic and abiotic factors. Salt marsh die-off on Cape Cod, Massachusetts (U.S.A.), triggered by a recreational-fishing-induced trophic cascade that has released herbivorous crabs from predator control, has been ongoing since 1976. Similar salt marsh die-offs have been reported in Long Island Sound and Narragansett Bay (U.S.A.), but the driving mechanism of these die-offs has not been examined. We used field experiments to assess trophic interactions and historical reconstructions of 24 New England marshes to test the hypotheses that recreational fishing and predator depletion are a regional trigger of salt marsh die-off in New England and that die-offs in Long Island Sound and Narragansett Bay are more recent than those on Cape Cod. Predator depletion was the general trigger of marsh die-off and explained differences in herbivorous crab abundance and the severity of die-off across regions. Die-offs in Long Island Sound and Narragansett Bay are following a trajectory similar to die-off on Cape Cod, but are approximately 20 years behind those on Cape Cod. As a result, die-off currently affects 31.2% (SE 2.2) of low-marsh areas in Long Island Sound and Narragansett Bay, less than half the severity of die-off on Cape Cod. Our results contribute to the growing evidence that recreational fishing is an increasing threat to coastal ecosystems and that studying the effects of human activity at regional scales can provide insight into local effects and aid in early detection and potential remediation. Ontogenia Regional de un Incremento en la Mortandad en una Marisma Salada de Nueva Inglaterra Resumen Las zonas costeras se encuentran entre los ecosistemas mas productivos y mas afectados del mundo. Los siglos de actividad humana sobre las lineas costeras han ocasionado la sobreexplotacion de depredadores marinos, lo que ha llevado a efectos en cascada a nivel de ecosistema. Sin embargo, los efectos humanos y las aproximaciones para mediarlos varian regionalmente debido a los gradientes en los factores bioticos y abioticos. El incremento en la mortandad en una marisma salada en Cape Cod, Massachusetts (E.U.A.), iniciada por una cascada trofica inducida por pesca recreativa que ha liberado a los cangrejos herbivoros del control por depredadores, ha estado sucediendo desde 1976. Se han reportado incrementos similares en las marismas saladas de Long Island Sound y Narragansett Bay (E.U.A), pero el mecanismo que las causa no ha sido examinado. Usamos experimentos de campo para estudiar las interacciones troficas y reconstrucciones historicas de 24 marismas de Nueva Inglaterra para probar la hipotesis de que la pesca recreativa y la disminucion de depredadores son un detonante regional del incremento en la mortandad de las marismas saladas en Nueva Inglaterra y que los incrementos en Long Island Sound y Narragansett Bay son mas recientes que los de Cape Cod. La disminucion de depredadores fue el detonante general del incremento en mortandad de las marismas y explico las diferencias entre la abundancia de cangrejos herbivoros y la severidad del incremento a lo largo de las regiones. Los incrementos en Long Island Sound y Narragansett Bay estan siguiendo una trayectoria similar al de Cape Cod, pero estan aproximadamente 20 anos atras del de esa localidad. Como resultado, el incremento en la mortandad actualmente afecta 31.2% (SE 2.2) de las areas de marismas bajas en Long Island Sound y Narragansett Bay, menos de la mitad de la severidad del incremento en Cape Cod. Nuestros resultados contribuyen a la creciente evidencia de que la pesca recreativa es una amenaza que va en aumento para los ecosistemas costeros y que el estudio de los efectos de la actividad humana en escalas regionales puede proporcionar una introspectiva a los efectos locales y auxiliar en la deteccion temprana y la remediacion potencial.

Understanding human impacts on ecosystems is critical for conservation, but can be complicated by interactions between multiple impacts occurring at different times. Using historical patterns of ditch construction on Cape Cod, Massachusetts, we tested the hypothesis that mosquito ditches have exacerbated salt marsh die-offs. Ditching activities occurred in the 1930s and were followed by post-World War II shoreline development, which created > 90% of current shoreline infrastructure on Cape Cod. Recently, predator depletion caused by recreational fishing has allowed populations of a native herbivorous crab (Sesarma reticulatum) to increase dramatically, triggering herbivore-driven cordgrass (Spartina alterniflora) die-off at developed sites. Depression-era mosquito ditching had little effect for decades, but accelerated subsequent die-offs by expanding cordgrass habitat. Despite occurring decades apart, ditching interacted synergistically with shoreline development and recreational fishing to devastate ~ 5...

Overexploitation of predators has been linked to the collapse of a growing number of shallow-water marine ecosystems. However, salt-marsh ecosystems are often viewed and managed as systems controlled by physical processes, despite recent evidence for herbivore-driven die-off of marsh vegetation. Here we use field observations, experiments, and historical records at 14 sites to examine whether the recently reported die-off of northwestern Atlantic salt marshes is associated with the cascading effects of predator dynamics and intensive recreational fishing activity. We found that the localized depletion of top predators at sites accessible to recreational anglers has triggered the proliferation of herbivorous crabs, which in turn results in runaway consumption of marsh vegetation. This suggests that overfishing may be a general mechanism underlying the consumer-driven die-off of salt marshes spreading throughout the western Atlantic. Our findings support the emerging realization that consumers play a dominant role in regulating marine plant communities and can lead to ecosystem collapse when their impacts are amplified by human activities, including recreational fishing.

Salt marsh habitat loss to vegetation die-offs has accelerated throughout the western Atlantic in the last four decades. Recent studies have suggested that eutrophication, pollution and/or disease may contribute to the loss of marsh habitat. In light of recent evidence that predators are important determinants of marsh health in New England, we performed a total predator exclusion experiment. Here, we provide the first experimental evidence that predator depletion can cause salt marsh die-off by releasing the herbivorous crab Sesarma reticulatum from predator control. Excluding predators from a marsh ecosystem for a single growing season resulted in a >100% increase in herbivory and a >150% increase in unvegetated bare space compared to plots with predators. Our results confirm that marshes in this region face multiple, potentially synergistic threats.

Direct and indirect human impacts on coastal ecosystems have increased over the last several centuries, leading to unprecedented degradation of coastal habitats and loss of ecological services. Here we document a two-century temporal disparity between salt marsh accretion and subsequent loss to indirect human impacts. Field surveys, manipulative experiments and GIS analyses reveal that crab burrowing weakens the marsh peat base and facilitates further burrowing, leading to bank calving, disruption of marsh accretion, and a loss of over two centuries of sequestered carbon from the marsh edge in only three decades. Analogous temporal disparities exist in other systems and are a largely unrecognized obstacle in attaining sustainable ecosystem services in an increasingly human impacted world. In light of the growing threat of indirect impacts worldwide and despite uncertainties in the fate of lost carbon, we suggest that estimates of carbon emissions based only on direct human impacts may significantly underestimate total anthropogenic carbon emissions.

With global increases in human impacts, invasive species have become a major threat to ecosystems worldwide. While they have been traditionally viewed as harmful, invasive species may facilitate the restoration of degraded ecosystems outside their native ranges. In New England (USA) overfishing has depleted salt marsh predators, allowing the herbivorous crab Sesarma reticulatum to denude hundreds of hectares of low marsh. Here, using multiple site surveys and field caging experiments, we show that the subsequent invasion of green crabs, Carcinus maenas, into heavily burrowed marshes partially reverses decades of cordgrass die-off. By consuming Sesarma, eliciting a nonlethal escape response, and evicting Sesarma from burrows, Carcinus reduces Sesarma herbivory and promotes cordgrass recovery. These results suggest that invasive species can contribute to restoring degraded ecosystems and underscores the potential for invasive species to return ecological functions lost to human impacts.

Predator depletion on Cape Cod (USA) has released the herbivorous crab Sesarma reticulatum from predator control leading to the loss of cordgrass from salt marsh creek banks. After more than three decades of die-off, cordgrass is recovering at heavily damaged sites coincident with the invasion of green crabs ( Carcinusmaenas ) into intertidal Sesarma burrows. We hypothesized that Carcinus is dependent on Sesarma burrows for refuge from physical and biotic stress in the salt marsh intertidal and reduces Sesarma functional density and herbivory through consumptive and non-consumptive effects, mediated by both visual and olfactory cues. Our results reveal that in the intertidal zone of New England salt marshes, Carcinus are burrow dependent, Carcinus reduce Sesarma functional density and herbivory in die-off areas and Sesarma exhibit a generic avoidance response to large, predatory crustaceans. These results support recent suggestions that invasive Carcinus are playing a role in the recovery of New England salt marshes and assertions that invasive species can play positive roles outside of their native ranges.

Belowground herbivory is commonly overlooked as a mechanism of top-down control in vegetated habitats, particularly in aquatic ecosystems. Recent research has revealed that increased densities of the herbivorous crab Sesarma reticulatum have led to runaway herbivory and widespread salt marsh die-off on Cape Cod, Massachusetts, USA. Aboveground herbivory is a major driver of this cordgrass habitat loss, but the role of belowground grazing is poorly understood. Sesarma live in communal burrows typically consisting of 1-2 openings and containing 2-3 crabs. However, at die-off sites, burrow complexes can cover > 90% of the low marsh zone, with crab densities as high as 50 crabs/m2 and burrow opening densities of 170 openings/m2. The magnitude of belowground Sesarma activity in association with salt marsh die-off provides an excellent opportunity to extend our knowledge of belowground herbivory impacts in coastal wetlands. Since Sesarma burrows allow access to cordgrass roots and rhizomes, and Sesarma are frequently restricted to burrows by thermal stress and predation, we hypothesized that belowground herbivory would be widespread in die-off areas. We experimentally demonstrate that Sesarma readily eat belowground roots and rhizomes in addition to aboveground cordgrass leaves. We then partitioned above- and belowground herbivory with field manipulations and found that belowground grazing is not only common, but can cause total plant mortality. Additional experiments revealed that plants remain vulnerable to belowground herbivory even after reaching a size refuge from aboveground grazing. This suggests that belowground herbivory contributes to salt marsh die-offs and adds to growing evidence that belowground herbivory is a widespread structuring force in plant communities that can limit habitat persistence.