Christine Angelini

A trophic cascade triggers collapse of a salt‐marsh ecosystem with intensive recreational fishing

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.

A keystone mutualism underpins resilience of a coastal ecosystem to drought.

Droughts are increasing in severity and frequency, yet the mechanisms that strengthen ecosystem resilience to this stress remain poorly understood. Here, we test whether positive interactions in the form of a mutualism between mussels and dominant cordgrass in salt marshes enhance ecosystem resistance to and recovery from drought. Surveys spanning 250 km of southeastern US coastline reveal spatially dispersed mussel mounds increased cordgrass survival during severe drought by 5- to 25-times. Surveys and mussel addition experiments indicate this positive effect of mussels on cordgrass was due to mounds enhancing water storage and reducing soil salinity stress. Observations and models then demonstrate that surviving cordgrass patches associated with mussels function as nuclei for vegetative re-growth and, despite covering only 0.1–12% of die-offs, markedly shorten marsh recovery periods. These results indicate that mutualisms, in supporting stress-resistant patches, can play a disproportionately large, keystone role in enhancing ecosystem resilience to climatic extremes.

How habitat-modifying organisms structure the food web of two coastal ecosystems

The diversity and structure of ecosystems has been found to depend both on trophic interactions in food webs and on other species interactions such as habitat modification and mutualism that form non-trophic interaction networks. However, quantification of the dependencies between these two main interaction networks has remained elusive. In this study, we assessed how habitat-modifying organisms affect basic food web properties by conducting in-depth empirical investigations of two ecosystems: North American temperate fringing marshes and West African tropical seagrass meadows. Results reveal that habitat-modifying species, through non-trophic facilitation rather than their trophic role, enhance species richness across multiple trophic levels, increase the number of interactions per species (link density), but decrease the realized fraction of all possible links within the food web (connectance). Compared to the trophic role of the most highly connected species, we found this non-trophic effects to be more important for species richness and of more or similar importance for link density and connectance. Our findings demonstrate that food webs can be fundamentally shaped by interactions outside the trophic network, yet intrinsic to the species participating in it. Better integration of non-trophic interactions in food web analyses may therefore strongly contribute to their explanatory and predictive capacity.

Livestock as a potential biological control agent for an invasive wetland plant

Invasive species threaten biodiversity and incur costs exceeding billions of US

Mutualistic interactions amplify saltmarsh restoration success

. Eradication efforts, however, are nearly always unsuccessful. Throughout much of North America, land managers have used expensive, and ultimately ineffective, techniques to combat invasive Phragmites australis in marshes. Here, we reveal that Phragmites may potentially be controlled by employing an affordable measure from its native European range: livestock grazing. Experimental field tests demonstrate that rotational goat grazing (where goats have no choice but to graze Phragmites) can reduce Phragmites cover from 100 to 20% and that cows and horses also readily consume this plant. These results, combined with the fact that Europeans have suppressed Phragmites through seasonal livestock grazing for 6,000 years, suggest Phragmites management can shift to include more economical and effective top-down control strategies. More generally, these findings support an emerging paradigm shift in conservation from high-cost eradication to economically sustainable control of dominant invasive species.

Methods to test the interactive effects of drought and plant invasion on ecosystem structure and function using complementary common garden and field experiments

Summary Mounting evidence shows that the functioning and stability of coastal ecosystems often depends critically on habitat-forming foundation species such as seagrasses, mangroves and salt marsh grasses that engage in facultative mutualistic interactions. However, although restoration science is now gradually expanding its long-standing paradigm of minimizing competition to including intraspecific, or within species, facilitation in its designs, the potential of harnessing mutualistic interactions between species for restoration purposes remains uninvestigated. Here, we experimentally tested whether a previously documented mutualism between marsh-forming Spartina alterniflora (cordgrass) and Geukensia demissa (mussels) can increase restoration success in degraded US salt marshes. We found that co-transplanted mussels locally increased nutrients and reduced sulfide stress, thereby increasing cordgrass growth and clonal expansion by 50%. We then removed aboveground vegetation and mussels to simulate a disturbance event and discovered that cordgrass co-transplanted with mussels experienced 3-times greater survival than control transplants. Synthesis and applications. Our findings indicate that mussels amplify cordgrass re-colonization and resilience over spatial and temporal scales that exceed those of their actual mutualistic interaction. By experimentally demonstrating that mutualistic partners can enable foundation species to overcome stress barriers to establish and persist, we highlight that coastal restoration needs to evolve beyond the sole inclusion of intraspecific positive interactions. In particular, we suggest that integrating mutualisms in restoration designs may powerfully enhance long-term restoration success and ecosystem resilience in the many coastal ecosystems where mutualisms involving foundation species are important ecosystem-structuring interactions. This article is protected by copyright. All rights reserved.

Multiple stressors and the potential for synergistic loss of New England salt marshes

Abstract 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.

Spillover of Secondary Foundation Species Transforms Community Structure and Accelerates Decomposition in Oak Savannas

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.

Secondary foundation species enhance biodiversity

Secondary foundation species facilitate biodiversity by providing complex habitat within the protective structures of many primary foundation species; whether they retain this role when they are dispersed in novel environments is unclear. Here, we explore how secondary foundation species’ density within hosts controls the rate at which individuals accrue and test if dislodged secondary foundation species then bolster the abundance and diversity of associated species and decomposition in a widespread system: southern live oaks, Quercus virginiana, hosting festoons of the bromeliad, Tillandsia usneoides, in southeastern US savannas. One year of monitoring showed that as Tillandsia density within oaks increases, Tillandsia deposition beneath oaks increases linearly in kind. Once on the ground, two manipulative experiments revealed that: on small scales, individual festoons retain moisture, reduce temperature, and facilitate invertebrates and fungi which combine to accelerate litter decomposition and, on larger scales, continual deposition of festoons dramatically reduces grass cover and increases litter layer depth, invertebrate and fungi richness and density, and decomposition. Surveys further suggest that Tillandsia populations within oaks, in moderating festoon accumulation, are shaping understory communities throughout this geographic region. Thus, disturbance-induced spillover enables secondary foundation species to define biodiversity and ecosystem functioning beyond the boundaries of their hosts.

A natural history model of New England salt marsh die-off

It has long been recognized that primary foundation species (FS), such as trees and seagrasses, enhance biodiversity. Among the species facilitated are secondary FS, including mistletoes and epiphytes. Case studies have demonstrated that secondary FS can further modify habitat-associated organisms (‘inhabitants’), but their net effects remain unknown. Here we assess how inhabitants, globally, are affected by secondary FS. We extracted and calculated 2,187 abundance and 397 richness Hedges’ g effect sizes from 91 and 50 publications, respectively. A weighted meta-analysis revealed that secondary FS significantly enhanced the abundance and richness of inhabitants compared to the primary FS alone. This indirect facilitation arising through sequential habitat formation was consistent across environmental and experimental conditions. Complementary unweighted analyses on log response ratios revealed that the magnitude of these effects was similar to the global average strength of direct facilitation from primary foundation species and greater than the average strength of trophic cascades, a widely recognized type of indirect facilitation arising through sequential consumption. The finding that secondary FS enhance the abundance and richness of inhabitants has important implications for understanding the mechanisms that regulate biodiversity. Integrating secondary FS into conservation practice will improve our ability to protect biodiversity and ecosystem function.Secondary foundation species, such as epiphytes, form structurally complex habitats on primary foundation species. A meta-analysis shows that they significantly enhance the abundance and richness of inhabitants compared to primary foundation species alone.