The reed Phragmites australis Cav. is aggressively invading salt marshes along the Atlantic Coast of North America. We examined the interactive role of habitat alteration (i.e., shoreline ...development) in driving this invasion and its consequences for plant richness in New England salt marshes. We surveyed 22 salt marshes in Narragansett Bay Rhode Island, and quantified shoreline development, Phragmites cover, soil salinity, and nitrogen availability. Shoreline development, operationally defined as removal of the woody vegetation bordering marshes, explained >90% of intermarsh variation in Phragmites cover. Shoreline development was also significantly correlated with reduced soil salinities and increased nitrogen availability, suggesting that removing woody vegetation bordering marshes increases nitrogen availability and decreases soil salinities, thus facilitating Phragmites invasion. Soil salinity (64%) and nitrogen availability (56%) alone explained a large proportion of variation in Phragmites cover, but together they explained 80% of the variation in Phragmites invasion success. Both univariate and aggregate (multidimensional scaling) analyses of plant community composition revealed that Phragmites dominance in developed salt marshes resulted in an almost three-fold decrease in plant species richness. Our findings illustrate the importance of maintaining integrity of habitat borders in conserving natural communities and provide an example of the critical role that local conservation can play in preserving these systems. In addition, our findings provide ecologists and natural resource managers with a mechanistic understanding of how human habitat alteration in one vegetation community can interact with species introductions in adjacent communities (i.e., flow-on or adjacency effects) to hasten ecosystem degradation.
Bacterial symbionts are integral to the health and homeostasis of invertebrate hosts. Notably, members of the Rickettsiales genus Wolbachia influence several aspects of the fitness and evolution of ...their terrestrial hosts, but few analogous partnerships have been found in marine systems. We report here the genome, phylogenetics, and biogeography of a ubiquitous and novel Rickettsiales species that primarily associates with marine organisms. We previously showed that this bacterium was found in scleractinian corals, responds to nutrient exposure, and is associated with reduced host growth and increased mortality. This bacterium, like other Rickettsiales, has a reduced genome indicative of a parasitic lifestyle. Phylogenetic analysis places this Rickettsiales within a new genus we define as "Candidatus Aquarickettsia." Using data from the Earth Microbiome Project and SRA databases, we also demonstrate that members of "Ca. Aquarickettsia" are found globally in dozens of invertebrate lineages. The coral-associated "Candidatus A. rohweri" is the first finished genome in this new clade. "Ca. A. rohweri" lacks genes to synthesize most sugars and amino acids but possesses several genes linked to pathogenicity including Tlc, an antiporter that exchanges host ATP for ADP, and a complete Type IV secretion system. Despite its inability to metabolize nitrogen, "Ca. A. rohweri" possesses the NtrY-NtrX two-component system involved in sensing and responding to extracellular nitrogen. Given these data, along with visualization of the parasite in host tissues, we hypothesize that "Ca. A. rohweri" reduces coral health by consuming host nutrients and energy, thus weakening and eventually killing host cells. Last, we hypothesize that nutrient enrichment, which is increasingly common on coral reefs, encourages unrestricted growth of "Ca. A. rohweri" in its host by providing abundant N-rich metabolites to be scavenged.
Parasites are more diverse and numerous than their hosts and commonly control population dynamics. Whether parasites also regulate key ecosystem processes, such as resistance to climate stress, is ...unclear. In southern U.S. salt marshes, drought interacts synergistically with keystone grazing to generate extensive ecosystem die-off. Field manipulations of parasite prevalence and salt stress in sediments in healthy marshes demonstrated that trematode parasites, by suppressing feeding activity of grazers that overgraze on drought-stressed plants, have the potential to slow the rate of ecosystem loss. Surveys along 1,000 km of coastline during an intense drought event revealed parasitism is common in grazers on die-off borders and that increasing infection prevalence along marsh die-off borders is negatively correlated with per capita grazing. Combined, results from this field experiment and survey suggested, but did not show, that parasites could affect rates of drought-driven salt marsh dieoff. To test whether parasites can indeed protect marshes under real drought conditions, we experimentally manipulated parasite prevalence in grazers over a month-long period on active die-off borders in three North Carolina marshes. Experimentally reducing parasite prevalence markedly increased the rate of plant ecosystem decline, an effect that scaled positively with prevalence. Thus parasites, by generating a trophic cascade, indirectly enhanced ecosystem resistance to overgrazing under intense drought in these North Carolina marshes. The generality of these results across the entire range of this keystone grazer in the southeastern United States needs to be tested, employing both experiments and extensive surveys that examine how the rate of ecosystem decline is mediated by parasitism. Given the ubiquity of parasites in ecosystems, our results suggest that more research effort should be invested in examining the possible roles for parasitism in regulating ecosystem function and stability.
Seagrasses are a valuable environmental resource. They provide a multitude of ecosystem services including nursery habitat, improved water quality, coastal protection, and carbon sequestration. ...However, seagrasses are in crisis as global coverage is declining at an accelerating rate. Recently, restoration has increased in popularity as a primary conservation action that may help re-establish degraded seagrass beds. With the elevation of restoration efforts as a conservation strategy, new methods that enhance restoration yields need to be explored. Terrestrially, incorporating positive species interactions and feedbacks into planting designs have proven beneficial to enhance restoration success at little additional cost. Decades of work in coastal plant ecosystems, including seagrasses, has shown that positive species relationships and feedbacks are critical for ecosystem stability, expansion, and recovery from disturbance. We reviewed the restoration literature on seagrasses and found, despite the critical role positive interactions play in marine plant systems, less than 8% of studies have tested for the beneficial effects of including positive interactions in seagrass restoration designs. Here we review the full suite of positive species interactions that have been documented in seagrass ecosystems, where they occur, and how they might be integrated into seagrass restoration. The few studies in marine plant communities that have explicitly incorporated positive species interactions and feedbacks have found an increase in plant growth with little additional resource investment. As oceans continue to change and stressors become more prevalent, pioneering restoration methods, such as harnessing positive feedbacks between species, will be key in rehabilitating populations of seagrasses.
Coastal ecosystems have drastically declined in coverage and condition across the globe. To combat these losses, marine conservation has recently employed habitat restoration as a strategy to enhance ...depleted coastal ecosystems. For restoration to be a successful enterprise, however, it is necessary to identify and address potential knowledge gaps and review whether the field has tracked scientific advances regarding best practices. This enables managers, researchers, and practitioners alike to more readily establish restoration priorities and goals. We synthesized the peer-reviewed, published literature on habitat restoration research in salt marshes, oyster reefs, and seagrasses to address three questions related to restoration efforts: (i) How frequent is cross-sector authorship in coastal restoration research? (ii) What is the geographic distribution of coastal restoration research? and (iii) Are abiotic and biotic factors equally emphasized in the literature, and how does this vary with time? Our vote-count survey indicated that one-third of the journal-published studies listed authors from at least two sectors, and 6% listed authors from all three sectors. Across all habitat types, there was a dearth of studies from Africa, Asia, and South America. Finally, despite many experimental studies demonstrating that species interactions can greatly affect the recovery and persistence of coastal foundation species, only one-fourth of the studies we examined discussed their effects on restoration. Combined, our results reveal gaps and discrepancies in restoration research that should be addressed in order to further propel coastal restoration science.
Ecosystem restoration has traditionally focused on re-establishing vegetation and other foundation species at basal trophic levels, with mixed outcomes. Here, we show that threatened shorebirds could ...be important to restoring coastal wetland multifunctionality. We carried out surveys and manipulative field experiments in a region along the Yellow Sea affected by the invasive cordgrass Spartina alterniflora. We found that planting native plants alone failed to restore wetland multifunctionality in a field restoration experiment. Shorebird exclusion weakened wetland multifunctionality, whereas mimicking higher predation before shorebird population declines by excluding their key prey - crab grazers - enhanced wetland multifunctionality. The mechanism underlying these effects is a simple trophic cascade, whereby shorebirds control crab grazers that otherwise suppress native vegetation recovery and destabilize sediments (via bioturbation). Our findings suggest that harnessing the top-down effects of shorebirds - through habitat conservation, rewilding, or temporary simulation of consumptive or non-consumptive effects - should be explored as a nature-based solution to restoring the multifunctionality of degraded coastal wetlands.
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 alterniflora
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 (<1 m
2
) experiment in 2009, removing grazers and plant neighbors prompted dramatically higher expansion and growth of
Spartina
relative to controls, while adding freshwater to reduce water limitation had little effect. In contrast, large-patch (>20 m
2
) 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.
Coastal wetlands are among the most productive ecosystems in the world. They generate critical services for humans including shoreline protection, carbon storage, pollution mitigation, and fisheries ...production. Restoration of coastal wetlands has historically been viewed as a secondary conservation strategy, but recently—given the continued loss of wetlands worldwide—many non-governmental and governmental organizations have elevated habitat restoration to be a primary method for wetland conservation. The long-held paradigm in coastal wetland restoration has been to restore target habitats by reducing physical stressors and avoiding competition among outplants, such as mangrove saplings or Spartina plugs. Recent ecological research, however, reveals that positive species interactions, such as facilitation, are critical to wetland recovery after disturbance. Here, we review the scientific evidence for the importance of positive species interactions in the recovery of salt-marsh and mangrove ecosystems and assess the extent to which they have been integrated into restoration studies. We found that only a small proportion of studies of marsh and mangrove restoration examined the effects of positive species interactions, despite the important role they play in the regrowth of coastal wetlands. We outline how positive species interactions can be systematically incorporated into future restoration work and discuss how this incorporation can help the reestablishment of coastal wetland biota through: (1) trophic facilitation, (2) stress reduction, and (3) associational defenses. The absence of positive interactions in restoration designs may partially explain the significant disparities between the functioning of natural and restored coastal plant ecosystems.
Invasive consumers can cause extensive ecological damage to native communities but effects on ecosystem resilience are less understood. Here, we use drone surveys, manipulative experiments, and ...mathematical models to show how feral hogs reduce resilience in southeastern US salt marshes by dismantling an essential marsh cordgrass-ribbed mussel mutualism. Mussels usually double plant growth and enhance marsh resilience to extreme drought but, when hogs invade, switch from being essential for plant survival to a liability; hogs selectively forage in mussel-rich areas leading to a 50% reduction in plant biomass and slower post-drought recovery rate. Hogs increase habitat fragmentation across landscapes by maintaining large, disturbed areas through trampling of cordgrass during targeted mussel consumption. Experiments and climate-disturbance recovery models show trampling alone slows marsh recovery by 3x while focused mussel predation creates marshes that may never recover from large-scale disturbances without hog eradication. Our work highlights that an invasive consumer can reshape ecosystems not just via competition and predation, but by disrupting key, positive species interactions that underlie resilience to climatic disturbances.
Impacts of marine invaders on local biodiversity have not been analyzed across invasive species and invaded habitats. We conducted a meta-analysis of 56 field experiments published in 29 papers that ...examined the effects of marine invaders on local species richness, diversity, and/or evenness. We show that invaders, across studies, typically have negative effects on biodiversity within a trophic level but positive effects on biodiversity of higher trophic levels. For example, both plants and sessile filter-feeders had positive effects on richness and diversity of mobile consumers. The contrasting negative and positive effects on similar versus higher trophic levels are potentially manifested through community-wide antagonism (competition and consumption) versus facilitation (habitat and food provisioning) interactions, respectively. These relation ships extended to functional interactions, as sessile invaders had negative effects on the biodiversity of sessile communities (intra-functional interactions) but positive effects on the biodiversity of mobile communities (inter-functional interactions). Our analyses highlight the importance of pairing attributes of the invader and the impacted organisms to obtain simple predictions of how the diversity of entire communities may respond to species invasions on local scales. We also note that our analysis did not require information on co-evolutionary history but that such data, coupled with long-term large-scale mensurative data, are needed to gain a comprehensive predictive insight into invasion impact.
PDF
