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Climate change causes both chronic and pulsed environmental changes to ecosystems. In estuaries, tidal freshwater marshes experience both extended and episodic periods of elevated salinities due to sea level rise, reduced river discharge during drought and storm surge, but most research has focused on extended (press) perturbations.Over a 4‐year period, we added diluted seawater to replicated plots in a tidal freshwater marsh in Georgia, USA to raise porewater salinities from freshwater to oligohaline. We assessed the ecosystem effects of press (continuous) and pulse (2 months per year) changes in salinity by measuring the responses of dominant angiosperms, benthic microalgae and dominant macro‐invertebrates (fiddler crabs). We transplanted angiosperms typical of oligohaline and mesohaline conditions into plots as bioindicators (phytometers) to assess potential for vegetation changes over longer time periods.In the press treatment, the cover of all common plant species decreased. Ludwigia repens almost disappeared within the first month; Polygonum hydropiperoides nearly disappeared within the first growing season; Pontederia cordata and Zizaniopsis miliacea declined over the 4 years but did not completely disappear. The decline in the four most dominant plant species decreased total above‐ground plant biomass, leading to an increase in light penetration and increased densities of benthic diatoms and cyanobacteria. The density of fiddler crab burrows was not significantly affected by the treatments, but was positively related to above‐ground plant biomass across all replicates. Transplant experiments indicated that plants typical of higher salinity conditions (e.g. Schoenoplectus sp.) performed well under conditions prevalent in the press plots.In the pulse treatment, only L. repens declined, and there was no effect on community‐level above‐ground biomass or other community variables.Synthesis. Our results indicate that tidal freshwater marsh plant and animal communities are vulnerable to extended periods of salinization but resilient to short saline pulses. Although saline pulses did not impair most ecosystem functions, the decline in a single species (L. repens) in the pulse treatment was associated with reduced marsh accretion and no elevation gain—factors which must be positive for wetland survival in an era of rising seas. Thus, periodic salinization may threaten the long‐term persistence of freshwater wetlands even before dramatic changes in community structure occurs. [ABSTRACT FROM AUTHOR]

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Disturbances are increasing in size and frequency with climate change, facilitating species that opportunistically exploit areas where habitat‐forming foundation species have been removed. Although it is well‐recognized that consumers, disease and weedy space‐holders can affect foundation species' resistance to and recovery from disturbance, how predators influence their resilience is less clear.In salt marshareas de‐vegetated by drought and intensive snail Littoraria irrorata grazing (hereafter, 'die‐offs'), we monitored bird use and experimentally manipulated bird and nekton access to the vegetated borders of die‐off mudflats across periods of both vegetation die‐off and regrowth to explore how these predators mediate the resilience of cordgrass Spartina alterniflora, the foundation species that structures US Atlantic coast salt marshes.Surveys revealed that birds, especially probers that agitate soils, forage year‐round for invertebrates in die‐off mudflats in our study area but not in adjacent vegetated areas.During periods of die‐off, cordgrass borders accessible to bird and nekton predators retreated >3‐times slower and snail densities were halved, relative to predator exclusion cages. In predator‐accessible plots, slower border retreat corresponded to greater snail infection by a bird host‐dependent trematode parasite. During recovery, cordgrass borders revegetated more quickly, and snail densities declined faster over time in unmanipulated controls relative to predator exclusions.Synthesis. These findings suggest that birds, through their transmission of parasites to snails, appear to act synergistically with snail‐consuming nekton to slow cordgrass loss after drought‐snail disturbances. Predator access also corresponds to faster cordgrass recovery as environmental conditions improve, although the mechanisms behind this need further investigation. Thus, predators that opportunistically forage within disturbances have the potential to suppress consumer impacts through multiple mechanisms, including consumption and disease transmission, thereby bolstering foundation species' resilience and modulating whole ecosystem responses to climate change. [ABSTRACT FROM AUTHOR]

Ecosystems worldwide depend on habitat‐forming foundation species that often facilitate themselves with increasing density and patch size, while also engaging in facultative mutualisms. Anthropogenic global change (e.g., climate change, eutrophication, overharvest, land‐use change), however, is causing rapid declines of foundation species‐structured ecosystems, often typified by sudden collapse. Although disruption of obligate mutualisms involving foundation species is known to precipitate collapse (e.g., coral bleaching), how facultative mutualisms (i.e., context‐dependent, nonbinding reciprocal interactions) affect ecosystem resilience is uncertain. Here, we synthesize recent advancements and combine these with model analyses supported by real‐world examples, to propose that facultative mutualisms may pose a double‐edged sword for foundation species. We suggest that by amplifying self‐facilitative feedbacks by foundation species, facultative mutualisms can increase foundation species' resistance to stress from anthropogenic impact. Simultaneously, however, mutualism dependency can generate or exacerbate bistability, implying a potential for sudden collapse when the mutualism's buffering capacity is exceeded, while recovery requires conditions to improve beyond the initial collapse point (hysteresis). Thus, our work emphasizes the importance of acknowledging facultative mutualisms for conservation and restoration of foundation species‐structured ecosystems, but highlights the potential risk of relying on mutualisms in the face of global change. We argue that significant caveats remain regarding the determination of these feedbacks, and suggest empirical manipulation across stress gradients as a way forward to identify related nonlinear responses. [ABSTRACT FROM AUTHOR]

Understanding the interactive effects of species invasions and climate change is essential for predicting future shifts in biodiversity. Because multiple stressors can interact in synergistic or antagonistic ways, it is notoriously difficult to anticipate their combined effects on species assemblages. However, some hypotheses predict that plant invasions will become increasingly problematic as climate change improves conditions for invaders or lowers the biotic resistance of native communities. In a 4‐yr field experiment, we quantified the individual and interactive effects of invasion by a globally problematic C4 grass, Imperata cylindrica, and chronic simulated drought imposed by rainout shelters on the whole plant communities of regenerating longleaf pine forest. Invasion both inhibited plant colonization and enhanced plot‐level extinctions, resulting in a severe (60%) loss of plant diversity across all functional groups, including perennial grasses and forbs, annual forbs, and woody species and dramatic shifts in community composition. Experimental drought reduced diversity by 20%, and caused a shift in the dominant functional groups, but had no significant effect on cover of the invader. The invader partially ameliorated water stress in the drought treatment such that invaded plots had higher soil moisture than uninvaded plots. Consequently, the combined effects of invasion and drought were lower than expected from an additive model of multiple stressors. These findings, which may have broader implications for how other C4 grass invaders will interact with drought to shift native community dynamics, challenge the perception that climate change will exacerbate invasions. In revealing that invasive species pose a major threat to the diversity and structure of native communities despite their moderating effects on abiotic stress, this work also highlights that management of aggressive invaders may be critical to preserving biodiversity regardless of future climate. [ABSTRACT FROM AUTHOR]

Climate change and consumer outbreaks are driving ecosystem collapse worldwide. Although much research has demonstrated that these factors can interact, how heterogeneity in top-down control intensity and physical forcing modulates ecosystem resilience to climate stress remains poorly understood. Here, we explore whether the nocturnal herbivorous crab Sesarma reticulatum can control spatially dominant cordgrass (Spartina alterniflora) growth and how its top-down effects vary with crab density, drought stress, and large-scale disturbance in southeastern US salt marshes. In multiple field experiments and surveys, we show that Sesarma depresses cordgrass growth and that its effects increase in a saturating manner with increasing crab density, such that the highest naturally occurring densities of this consumer can trigger local cordgrass die-off. This top-down effect of Sesarma is similar in magnitude to what is thought to be the dominant grazer in the system, the marsh periwinkle snail Littoraria irrorata. In a drought stress by Sesarma density experiment, we further show that salinity stress and intensive crab herbivory additively suppress cordgrass drought resistance. After drought subsides, surveys and experiments reveal that Sesarma also stifles cordgrass re-growth into existing die-off areas. Together, these results show that multiple grazers powerfully regulate the productivity and drought resilience of these intertidal grasslands and that heterogeneity in physical stress and consumer density can dictate when and where top-down forcing is important. More generally, this work provides a rare, experimental demonstration of the critical role top-down control can play across the initiation and recovery stages of ecosystem die-off. [ABSTRACT FROM AUTHOR]

Climate change-driven range expansion of black mangroves ( Avicennia germinans) is predicted along the northern Gulf of Mexico, where sea level rise is also driving conversion of freshwater forest islands to salt marsh. While climate-driven A. germinans range expansion has garnered considerable scientific attention, the role of top-down controls on colonization is largely overlooked. We investigated the effects of abiotic (flooding frequency, soil depth, soil salinity) and biotic (predation, herbivory) controls on A. germinans establishment at its northern range limit along Florida's Gulf Coast by comparing fates of caged and non-caged propagules across four landscape positions (from creek edge to forest island interior) and at three sites along a tidal flooding frequency gradient. Within 12 days, grapsid crab, Sesarma reticulatum, consumed 99% of non-caged propagules. Among caged propagules, establishment increased with increasing flooding frequency; however, cages did not entirely prevent predation, which remained a primary cause of mortality, except in the rarely flooded island. Propagules that survived to seedlings experienced mild to fatal herbivory across landscape positions and sites. This study revealed that while relict forest islands and surrounding marshes can support A. germinans, predation and herbivory strongly suppress colonization, suggesting that mangrove expansion models should incorporate biotic controls. [ABSTRACT FROM AUTHOR]

Climate change and other anthropogenic stressors are converging on coastal ecosystems worldwide. Understanding how these stressors interact to affect ecosystem structure and function has immediate implications for coastal planning, however few studies quantify stressor interactions. We examined past and potential future interactions between two leading stressors on New England salt marshes: sea-level rise and marsh crab (Sesarma reticulatum) grazing driven low marsh die-off. Geospatial analyses reveal that crab-driven die-off has led to an order of magnitude more marsh loss than sea-level rise between 2005 and 2013. However, field transplant experimental results suggest that sea-level rise will facilitate crab expansion into higher elevation marsh platforms by inundating and gradually softening now-tough high marsh peat, exposing large areas to crab-driven die-off. Taking interactive effects of marsh softening and concomitant overgrazing into account, we estimate that even modest levels of sea-level rise will lead to levels of salt marsh habitat loss that are 3x greater than the additive effects of sea-level rise and crab-driven die-off would predict. These findings highlight the importance of multiple stressor studies in enhancing mechanistic understanding of ecosystem vulnerabilities to future stress scenarios and encourage managers to focus on ameliorating local stressors to break detrimental synergisms, reduce future ecosystem loss, and enhance ecosystem resilience to global change. [ABSTRACT FROM AUTHOR]

Abiotic global change drivers affect ecosystem structure and function, but how they interact with biotic factors such as invasive plants is understudied. Such interactions may be additive, synergistic, or offsetting, and difficult to predict. We present methods to test the individual and interactive effects of drought and plant invasion on native ecosystems. We coupled a factorial common garden experiment containing resident communities exposed to drought (imposed with rainout shelters) and invasion with a field experiment where the invader was removed from sites spanning a natural soil moisture gradient. We detail treatments and their effects on abiotic conditions, including soil moisture, light, temperature, and humidity, which shape community and ecosystem responses. Ambient precipitation during the garden experiment exceeded historic norms despite severe drought in prior years. Soil moisture was 48% lower in drought than ambient plots, but the invader largely offset drought effects. Additionally, temperature and light were lower and humidity higher in invaded plots. Field sites spanned up to a 10-fold range in soil moisture and up to a 2.5-fold range in light availability. Invaded and resident vegetation did not differentially mediate soil moisture, unlike in the garden experiment. Herbicide effectively removed invaded and resident vegetation, with removal having site-specific effects on soil moisture and light availability. However, light was generally higher in invader-removal than control plots, whereas resident removal had less effect on light, similar to the garden experiment. Invasion mitigated a constellation of abiotic conditions associated with drought stress in the garden experiment. In the field, where other factors co-varied, these patterns did not emerge. Still, neither experiment suggested that drought and invasion will have synergistic negative effects on ecosystems, although invasion can limit light availability. Coupling factorial garden experiments with field experiments across environmental gradients will be effective for predicting how multiple stressors interact in natural systems. [ABSTRACT FROM AUTHOR]

Consumer fronts occur when grazers or predators aggregate in bands along the edges of a resource. Our review reveals that consumer fronts are a common phenomenon in nature, occur in many different ecosystems, and are triggered by universal mechanisms: External forces locally increase top-down control beyond prey carrying and/or renewal capacity, and resource-dependent movement leads to consumer aggregation along the edge of the remaining prey population. Once formed, consumer fronts move through systems as spatially propagating waves, self-reinforced via intense overexploitation and amplified by density-dependent feedbacks. When consumer fronts are spatially restricted, they generate patchiness. In contrast, when consumer fronts are expansive, they can lead to runaway responses that cause large-scale ecosystem degradation and regime shifts. We conceptualize a synergistic stress hypothesis and model that highlight how coupled intensification of physical stress and enhanced consumer pressure can trigger increased occurrence of consumer fronts and decreased system stability and resilience. With escalating climate change and food-web modification, the physical and biological conditions favoring consumer-front formation will likely become a common feature of many ecosystems. [ABSTRACT FROM AUTHOR]

Massive anthropogenic and climate-related disturbances are now common in ecosystems worldwide, generating widespread die-off and subsequent community recovery dominated by remnant-patch dynamics rather than open-gap dynamics. Whether communities can recover and, if so, which factors mediate recolonization rate and extent remain unresolved. Here we evaluate recolonization dynamics of southern U.S. salt marshes that experienced extensive, drought-induced die-off of the foundation species Spartina altern(flora over the previous decade. Surveys of Georgia (USA) salt marshes showed little seedling recruitment in die-off areas but persistence of Spartina particularly in large, rather than small, remnant patches. Given this natural variation in remnant patch size, we conducted field experiments to test whether key plant-controlling biotic (grazing, plant neighbor presence) and abiotic (water availability) factors differentially impact Spartina recolonization at small and large-patch scales. In the small-patch (20 m2) borders advanced significantly over the same time period regardless of grazer or neighbor removal. We continued the large-patch experiments in 2010, a year that experienced drought, and also added freshwater or salt to borders to modify ambient drought stress; overall, borders advanced less than the previous year but significantly more where neighbors were removed or freshwater added. Thus, water availability appears to mediate Spartina recovery by fueling large-patch expansion during wet summers and intensifying interspecific competition during drought. Combined, these findings suggest ecosystems can recover from massive disturbance if remnant foundation species' patches are large enough to overcome biotic inhibition and successfully expand during periods of relaxed abiotic stress. [ABSTRACT FROM AUTHOR]