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Spatiotemporal variation in herbivory is a major driver of intraspecific variation in plant defense. Comparatively little is known, however, about how changes in herbivory regime affect the balance of constitutive and induced resistance, which are often considered alternative defensive strategies. Here, we investigated how nearly a decade of insect herbivore suppression affected constitutive and induced resistance in horsenettle (Solanum carolinense), a widespread herbaceous perennial. We allowed replicated horsenettle populations to respond to the presence or absence of herbivores by applying insecticide to all plants in half of 16 field plots. Horsenettle density rapidly increased in response to insecticide treatment, and this effect persisted for at least 4 years after the cessation of herbivore suppression. We subsequently grew half-sibling families from seeds collected during and shortly after insecticide treatment in a common garden and found strong effects of insect suppression on induced resistance. Feeding trials in field mesocosms with false Colorado potato beetles (Leptinotarsa juncta), a common specialist herbivore, revealed that multiyear herbivore suppression drove rapid attenuation of induced resistance: offspring of plants from insect-suppression plots exhibited a near-complete loss of induced resistance to beetles, whereas those from control plots incurred ~70% less damage after experimental induction. Plants from insect-suppression plots also had ~40% greater constitutive resistance compared with those from control plots, although this difference was not statistically significant. We nonetheless detected a strong trade-off between constitutive and induced resistance across families. In contrast, the constitutive expression of trypsin inhibitors (TI), an important chemical defense trait in horsenettle, was reduced by 20% in the offspring of plants from insect-suppression plots relative to those from control plots. However, TIs were induced to an equal extent whether or not insect herbivores had been historically suppressed. Although several defense and performance traits (prickle density, TI concentration, resistance against false Colorado potato beetles and flea beetles, biomass, and seed mass) varied markedly across families, no traits exhibited significant pairwise correlations. Overall, our results indicate that, whereas the divergent responses of multiple defense traits to insect suppression led to comparatively small changes in overall constitutive resistance, they significantly reduced induced resistance against false Colorado potato beetle.

Diverse communities of large mammalian herbivores (LMH), once widespread, are now rare. LMH exert strong direct and indirect effects on community structure and ecosystem functions, and measuring these effects is important for testing ecological theory and for understanding past, current, and future environmental change. This in turn requires long-term experimental manipulations, owing to the slow and often nonlinear responses of populations and assemblages to LMH removal. Moreover, the effects of particular species or body-size classes within diverse LMH guilds are difficult to pinpoint, and the magnitude and even direction of these effects often depends on environmental context. Since 2008, we have maintained the Ungulate Herbivory Under Rainfall Uncertainty (UHURU) experiment, a series of size-selective LMH exclosures replicated across a rainfall/productivity gradient in a semiarid Kenyan savanna. The goals of the UHURU experiment are to measure the effects of removing successively smaller size classes of LMH (mimicking the process of size-biased extirpation) and to establish how these effects are shaped by spatial and temporal variation in rainfall. The UHURU experiment comprises three LMH-exclusion treatments and an unfenced control, applied to nine randomized blocks of contiguous 1-ha plots (n = 36). The fenced treatments are MEGA (exclusion of megaherbivores, elephant and giraffe), MESO (exclusion of herbivores ≥40 kg), and TOTAL (exclusion of herbivores ≥5 kg). Each block is replicated three times at three sites across the 20-km rainfall gradient, which has fluctuated over the course of the experiment. The first 5 years of data were published previously (Ecological Archives E095-064) and have been used in numerous studies. Since that publication, we have (1) continued to collect data following the original protocols, (2) improved the taxonomic resolution and accuracy of plant and small-mammal identifications, and (3) begun collecting several new data sets. Here, we present updated and extended raw data from the first 12 years of the UHURU experiment (2008-2019). Data include daily rainfall data throughout the experiment; annual surveys of understory plant communities; annual censuses of woody-plant communities; annual measurements of individually tagged woody plants; monthly monitoring of flowering and fruiting phenology; every-other-month small-mammal mark-recapture data; and quarterly large-mammal dung surveys. There are no copyright restrictions; notification of when and how data are used is appreciated and users of UHURU data should cite this data paper when using the data.

Trophic rewilding seeks to rehabilitate degraded ecosystems by repopulating them with large animals, thereby re-establishing strong top-down interactions. Yet there are very few tests of whether such initiatives can restore ecosystem structure and functions, and on what timescales. Here we show that war-induced collapse of large-mammal populations in Mozambique's Gorongosa National Park exacerbated woody encroachment by the invasive shrub Mimosa pigra-considered one of the world's 100 worst invasive species-and that one decade of concerted trophic rewilding restored this invasion to pre-war baseline levels. Mimosa occurrence increased between 1972 and 2015, a period encompassing the near extirpation of large herbivores during the Mozambican Civil War. From 2015 to 2019, mimosa abundance declined as ungulate biomass recovered. DNA metabarcoding revealed that ruminant herbivores fed heavily on mimosa, and experimental exclosures confirmed the causal role of mammalian herbivory in containing shrub encroachment. Our results provide mechanistic evidence that trophic rewilding has rapidly revived a key ecosystem function (biotic resistance to a notorious woody invader), underscoring the potential for restoring ecological health in degraded protected areas.

Biological invasions are both a pressing environmental challenge and an opportunity to investigate fundamental ecological processes, such as the role of top predators in regulating biodiversity and food-web structure. In whole-ecosystem manipulations of small Caribbean islands on which brown anole lizards (Anolis sagrei) were the native top predator, we experimentally staged invasions by competitors (green anoles, Anolis smaragdinus) and/or new top predators (curly-tailed lizards, Leiocephalus carinatus). We show that curly-tailed lizards destabilized the coexistence of competing prey species, contrary to the classic idea of keystone predation. Fear-driven avoidance of predators collapsed the spatial and dietary niche structure that otherwise stabilized coexistence, which intensified interspecific competition within predator-free refuges and contributed to the extinction of green-anole populations on two islands. Moreover, whereas adding either green anoles or curly-tailed lizards lengthened food chains on the islands, adding both species reversed this effect—in part because the apex predators were trophic omnivores. Our results underscore the importance of top-down control in ecological communities, but show that its outcomes depend on prey behaviour, spatial structure, and omnivory. Diversity-enhancing effects of top predators cannot be assumed, and non-consumptive effects of predation risk may be a widespread constraint on species coexistence. Whole-ecosystem manipulations of Caribbean islands occupied by brown anoles, involving the addition of competitors (green anoles) and/or top predators (curly-tailed lizards), demonstrate that predator introductions can alter the ecological niches and destabilize the coexistence of competing prey species.

African savannas are the last stronghold of diverse large-mammal communities, and a major focus of savanna ecology is to understand how these animals affect the relative abundance of trees and grasses. However, savannas support diverse plant life-forms, and human-induced changes in large-herbivore assemblages-declining wildlife populations and their displacement by livestock-may cause unexpected shifts in plant community composition. We investigated how herbivory affects the prevalence of lianas (woody vines) and their impact on trees in an East African savanna. Although scarce (

PREMISE Mismatches between light conditions and light-capture strategy can reduce plant performance and prevent colonization of novel habitats. Although light-capture strategies tend to be highly conserved among closely related species, evolutionary transitions from shaded to unshaded habitats (and vice versa) occur in numerous plant lineages. METHODS We combined phylogenetic approaches with field and greenhouse experiments to investigate evolutionary constraints on light-capture strategy in North American milkweeds (genus Asclepias) and to determine whether colonization of shaded habitats in this heliophilic clade is associated with reduced plasticity and attenuation of the shade avoidance response. RESULTS Colonization of shaded habitats has occurred at least 10 times in this genus, including at least once in each major North American clade. Evolutionary transitions between habitats exhibit strong directional bias, with shifts from full-sun to shaded habitats occurring at least three times as often as the opposite transition. In field and greenhouse experiments, sun species responded to shade by increasing internode length, height, and specific leaf area, consistent with the shade avoidance response; paired shade species exhibited reduced plasticity overall, and only one trait (specific leaf area) responded to experimental shade. CONCLUSIONS Our results suggest that milkweeds colonized shaded environments multiple times using a light-capture strategy distinct from the ancestral (putatively shade avoidant) strategy, including a general attenuation of plasticity in response to variable light conditions. This pattern bolsters the notion that shade avoidance and tolerance represent divergent evolutionary strategies for maximizing performance under qualitatively different types of shade.

Trophic rewilding seeks to rehabilitate degraded ecosystems by repopulating them with large animals, thereby re-establishing strong top-down interactions. Yet there are very few tests of whether such initiatives can restore ecosystem structure and functions, and on what timescales. Here we show that war-induced collapse of large-mammal populations in Mozambique's Gorongosa National Park exacerbated woody encroachment by the invasive shrub Mimosa pigra-considered one of the world's 100 worst invasive species-and that one decade of concerted trophic rewilding restored this invasion to pre-war baseline levels. Mimosa occurrence increased between 1972 and 2015, a period encompassing the near extirpation of large herbivores during the Mozambican Civil War. From 2015 to 2019, mimosa abundance declined as ungulate biomass recovered. DNA metabarcoding revealed that ruminant herbivores fed heavily on mimosa, and experimental exclosures confirmed the causal role of mammalian herbivory in containing shrub encroachment. Our results provide mechanistic evidence that trophic rewilding has rapidly revived a key ecosystem function (biotic resistance to a notorious woody invader), underscoring the potential for restoring ecological health in degraded protected areas.

Positive indirect effects of consumers on their resources can stabilize food webs by preventing overexploitation, but the coupling of trophic and non-trophic interactions remains poorly integrated into our understanding of community dynamics. Elephants engineer African savanna ecosystems by toppling trees and breaking branches, and although their negative effects on trees are well documented, their effects on small-statured plants remain poorly understood. Using data on 117 understory plant taxa collected over 7 yr within 36 1-ha experimental plots in a semi-arid Kenyan savanna, we measured the strength and direction of elephant impacts on understory vegetation. We found that elephants had neutral effects on most (83-89%) species, with a similar frequency of positive and negative responses among the remainder. Overall, estimated understory biomass was 5-14% greater in the presence of elephants across a range of rainfall levels. Whereas direct consumption likely accounts for the negative effects, positive effects are presumably indirect. We hypothesized that elephants create associational refuges for understory plants by damaging tree canopies in ways that physically inhibit feeding by other large herbivores. As predicted, understory biomass and species richness beneath elephant-damaged trees were 55% and 21% greater, respectively, than under undamaged trees. Experimentally simulated elephant damage increased understory biomass by 37% and species richness by 49% after 1 yr. Conversely, experimentally removing elephant damaged branches decreased understory biomass by 39% and richness by 30% relative to sham-manipulated trees. Camera-trap surveys revealed that elephant damage reduced the frequency of herbivory by 71%, whereas we detected no significant effect of damage on temperature, light, or soil moisture. We conclude that elephants locally facilitate understory plants by creating refuges from herbivory, which countervails the direct negative effects of consumption and enhances larger-scale biomass and diversity by promoting the persistence of rare and palatable species. Our results offer a counterpoint to concerns about the deleterious impacts of elephant "overpopulation" that should be considered in debates over wildlife management in African protected areas: understory species comprise the bulk of savanna plant biodiversity, and their responses to elephants are buffered by the interplay of opposing consumptive and non-consumptive interactions.

Biological invasions are both a pressing environmental challenge and an opportunity to investigate fundamental ecological processes, such as the role of top predators in regulating biodiversity and food-web structure. In whole-ecosystem manipulations of small Caribbean islands on which brown anole lizards (Anolis sagrei) were the native top predator, we experimentally staged invasions by competitors (green anoles, Anolis smaragdinus) and/or new top predators (curly-tailed lizards, Leiocephalus carinatus). We show that curly-tailed lizards destabilized the coexistence of competing prey species, contrary to the classic idea of keystone predation. Fear-driven avoidance of predators collapsed the spatial and dietary niche structure that otherwise stabilized coexistence, which intensified interspecific competition within predator-free refuges and contributed to the extinction of green-anole populations on two islands. Moreover, whereas adding either green anoles or curly-tailed lizards lengthened food chains on the islands, adding both species reversed this effect-in part because the apex predators were trophic omnivores. Our results underscore the importance of top-down control in ecological communities, but show that its outcomes depend on prey behaviour, spatial structure, and omnivory. Diversity-enhancing effects of top predators cannot be assumed, and non-consumptive effects of predation risk may be a widespread constraint on species coexistence.

BACKGROUND AND AIMS: Herbivory by large mammals imposes a critical recruitment bottleneck on plants in many systems. Spines defend plants against large herbivores, and how early they emerge in saplings may be one of the strongest predictors of sapling survival in herbivore-rich environments. Yet little effort has been directed at understanding the variability in spine emergence across saplings. METHODS: We present a multispecies study examining whether and how sapling size, spine type and species' environmental niche (light and precipitation environment) influence early emergence and biomass investment in spines. A phylogenetically diverse pool of 45 species possessing different spine types (spines, prickles and thorns; that are derived from distinct plant organs: leaf, epidermis or cortex, and branch, respectively), were grown under common-garden conditions, and patterns of spine emergence and biomass allocation to spines at 5 and 15 weeks after transplanting were characterized. KEY RESULTS: Spine type and species' resource niche were the main factors driving early emergence and investment patterns. Spines emerged earliest in leaf spine-bearing species, and latest in thorn-bearing species. The probability of early spine emergence increased with decreasing precipitation, and was greater in species from open than from closed habitats. Sapling investment in spines changed with plant mass but was contingent on spine type and habitat type. CONCLUSIONS: Different spine types have strikingly different timing of expression, suggesting that developmental origins of spines play a critical role in sapling defences. Furthermore, species from different precipitation and light environments (open vs. closed habitats) showed contrasting patterns of early spine expression, suggesting that resource limitation in their native range may have driven divergent evolution of early defence expression.

Niche partitioning facilitates species coexistence in a world of limited resources, thereby enriching biodiversity. For decades, biologists have sought to understand how diverse assemblages of large mammalian herbivores (LMH) partition food resources. Several complementary mechanisms have been identified, including differential consumption of grasses versus nongrasses and spatiotemporal stratification in use of different parts of the same plant. However, the extent to which LMH partition food-plant species is largely unknown because comprehensive species-level identification is prohibitively difficult with traditional methods. We used DNA metabarcoding to quantify diet breadth, composition, and overlap for seven abundant LMH species (six wild, one domestic) in semiarid African savanna. These species ranged from almost-exclusive grazers to almost-exclusive browsers: Grass consumption inferred from mean sequence relative read abundance (RRA) ranged from >99% (plains zebra) to

Self-organized spatial vegetation patterning is widespread and has been described using models of scale-dependent feedback between plants and water on homogeneous substrates. As rainfall decreases, these models yield a characteristic sequence of patterns with increasingly sparse vegetation, followed by sudden collapse to desert. Thus, the final, spot-like pattern may provide early warning for such catastrophic shifts. In many arid ecosystems, however, termite nests impart substrate heterogeneity by altering soil properties, thereby enhancing plant growth. We show that termite-induced heterogeneity interacts with scale-dependent feedbacks to produce vegetation patterns at different spatial grains. Although the coarse-grained patterning resembles that created by scale-dependent feedback alone, it does not indicate imminent desertification. Rather, mound-field landscapes are more robust to aridity, suggesting that termites may help stabilize ecosystems under global change.

Intraspecific variation in plant defense phenotype is common and has wide-ranging ecological consequences. Yet prevailing theories of plant defense allocation, which primarily account for interspecific differences in defense phenotype, often fail to predict intraspecific patterns. Furthermore, although individual variation in defense phenotype is often attributed to ecological interactions, few general mechanisms have been proposed to explain the ubiquity of variable defense phenotype within species. Here, we show experimentally that associational refuges and induced resistance interact to create predictable intraspecific variation in defense phenotype in African savanna plants. Physically defended species from four families (Acanthaceae, Asparagaceae, Cactaceae, and Solanaceae) growing in close association with spinescent Acacia trees had 39-78% fewer spines and thorns than did isolated conspecifics. For a subset of these species, we used a series of manipulative experiments to show that this variability is maintained primarily by a reduction in induced responses among individuals that seldom experience mammalian herbivory, whether due to association with Acacia trees or to experimental herbivore exclusion. Unassociated plants incurred 4- to 16-fold more browsing damage than did associated individuals and increased spine density by 16-38% within one month following simulated browsing. In contrast, experimental clipping induced no net change in spine density among plants growing beneath Acacia canopies or inside long-term herbivore exclosures. Associated and unassociated individuals produced similar numbers of flowers and seeds, but seedling recruitment and survival were vastly greater in refuge habitats, suggesting a net fitness benefit of association. We conclude that plant-plant associations consistently decrease defense investment in this system by reducing both the frequency of herbivory and the intensity of induced responses, and that inducible responses enable plants to capitalize on such associations in heterogeneous environments. Given the prevalence of associational and induced defenses in plant communities worldwide, our results suggest a potentially general mechanism by which biotic interactions might predictably shape intraspecific variation in plant defense phenotype.

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.

Self-organized spatial vegetation patterning is widespread and has been described using models of scale-dependent feedback between plants and water on homogeneous substrates. As rainfall decreases, these models yield a characteristic sequence of patterns with increasingly sparse vegetation, followed by sudden collapse to desert. Thus, the final, spot-like pattern may provide early warning for such catastrophic shifts. In many arid ecosystems, however, termite nests impart substrate heterogeneity by altering soil properties, thereby enhancing plant growth. We show that termite-induced heterogeneity interacts with scale-dependent feedbacks to produce vegetation patterns at different spatial grains. Although the coarse-grained patterning resembles that created by scale-dependent feedback alone, it does not indicate imminent desertification. Rather, mound-field landscapes are more robust to aridity, suggesting that termites may help stabilize ecosystems under global change.

Despite their value, coastal ecosystems are globally threatened by anthropogenic impacts, yet how these impacts are driven by economic development is not well understood. We compiled a multifaceted dataset to quantify coastal trends and examine the role of economic growth in China’s coastal degradation since the 1950s. Although China’s coastal population growth did not change following the 1978 economic reforms, its coastal economy increased by orders of magnitude. All 15 coastal human impacts examined increased over time, especially after the reforms. Econometric analysis revealed positive relationships between most impacts and GDP across temporal and spatial scales, often lacking dropping thresholds. These relationships generally held when influences of population growth were addressed by analyzing per capita impacts, and when population density was included as explanatory variables. Historical trends in physical and biotic indicators showed that China’s coastal ecosystems changed little or slowly between the 1950s and 1978, but have degraded at accelerated rates since 1978. Thus economic growth has been the cause of accelerating human damage to China’s coastal ecosystems. China’s GDP per capita remains very low. Without strict conservation efforts, continuing economic growth will further degrade China’s coastal ecosystems.

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.

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.