Vegetation controls on channel network complexity in coastal wetlands.

Channel networks are key to coastal wetland functioning and resilience under climate change. Vegetation affects sediment and hydrodynamics in many different ways, which calls for a coherent framework to explain how vegetation shapes channel network geometry and functioning. Here, we introduce an idealized model that shows how coastal wetland vegetation creates more complexly branching networks by increasing the ratio of channel incision versus topographic diffusion rates, thereby amplifying the channelization feedback that recursively incises finer-scale side-channels. This complexification trend qualitatively agrees with and provides an explanation for field data presented here as well as in earlier studies. Moreover, our model demonstrates that a stronger biogeomorphic feedback leads to higher and more densely vegetated marsh platforms and more extensive drainage networks. These findings may inspire future field research by raising the hypothesis that vegetation-induced self-organization enhances the storm surge buffering capacity of coastal wetlands and their resilience under sea-level rise.

A probabilistic framework for windows of opportunity: the role of temporal variability in critical transitions.

The establishment of young organisms in harsh environments often requires a window of opportunity (WoO). That is, a short time window in which environmental conditions drop long enough below the hostile average level, giving the organism time to develop tolerance and transition into stable existence. It has been suggested that this kind of establishment dynamics is a noise-induced transition between two alternate states. Understanding how temporal variability (i.e. noise) in environmental conditions affects establishment of organisms is therefore key, yet not well understood or included explicitly in the WoO framework. In this paper, we develop a coherent theoretical framework for understanding when the WoO open or close based on simple dichotomous environmental variation. We reveal that understanding of the intrinsic timescales of both the developing organism and the environment is fundamental to predict if organisms can or cannot establish. These insights have allowed us to develop statistical laws for predicting establishment probabilities based on the period and variance of the fluctuations in naturally variable environments. Based on this framework, we now get a clear understanding of how changes in the timing and magnitude of climate variability or management can mediate establishment chances.

Sediment availability provokes a shift from Brownian to Lévy-like clonal expansion in a dune building grass.

In biogeomorphic landscapes, plant traits can steer landscape development through plant-mediated feedback interactions. Interspecific differences in clonal expansion strategy can therefore lead to the emergence of different landscape organisations. Yet, whether landscape-forming plants adopt different clonal expansion strategies depending on their physical environment remains to be tested. Here, we use a field survey and a complementary mesocosm approach to investigate whether sediment deposition affects the clonal expansion strategy employed by dune-building marram grass individuals. Our results reveal a consistent shift in expansion pattern from more clumped, Brownian-like, movement in sediment-poor conditions, to patchier, Lévy-like, movement under high sediment supply rates. Additional model simulations illustrate that the sediment-dependent shift in movement strategies induces a shift in optimisation of the cost-benefit relation between landscape engineering (i.e. dune formation) and expansion. Plasticity in expansion strategy may therefore allow landscape-forming plants to optimise their engineering ability depending on their physical landscape.

A Lévy expansion strategy optimizes early dune building by beach grasses.

Lifeforms ranging from bacteria to humans employ specialized random movement patterns. Although effective as optimization strategies in many scientific fields, random walk application in biology has remained focused on search optimization by mobile organisms. Here, we report on the discovery that heavy-tailed random walks underlie the ability of clonally expanding plants to self-organize and dictate the formation of biogeomorphic landscapes. Using cross-Atlantic surveys, we show that congeneric beach grasses adopt distinct heavy-tailed clonal expansion strategies. Next, we demonstrate with a spatially explicit model and a field experiment that the Lévy-type strategy of the species building the highest dunes worldwide generates a clonal network with a patchy shoot organization that optimizes sand trapping efficiency. Our findings demonstrate Lévy-like movement in plants, and emphasize the role of species-specific expansion strategies in landscape formation. This mechanistic understanding paves the way for tailor-made planting designs to successfully construct and restore biogeomorphic landscapes and their services.

A facultative mutualistic feedback enhances the stability of tropical intertidal seagrass beds.

Marine foundation species such as corals, seagrasses, salt marsh plants, and mangrove trees are increasingly found to engage in mutualistic interactions. Because mutualisms by their very nature generate a positive feedback between the species, subtle environmental impacts on one of the species involved may trigger mutualism breakdown, potentially leading to ecosystem regime shifts. Using an empirically parameterized model, we investigate a facultative mutualism between seagrass and lucinid bivalves with endosymbiotic sulfide-oxidizing gill bacteria in a tropical intertidal ecosystem. Model predictions for our system show that, by alleviating the build-up of toxic sulfide, this mutualism maintains an otherwise intrinsically unstable seagrass ecosystem. However, an increase in seagrass mortality above natural levels, due to e.g. desiccation stress, triggers mutualism breakdown. This pushes the system in collapse-and-recovery dynamics ('slow-fast cycles') characterized by long-term persistent states of bare and seagrass-dominated, with rapid transitions in between. Model results were consistent with remote sensing analyses that suggest feedback-mediated state shifts induced by desiccation. Overall, our combined theoretical and empirical results illustrate the potential of mutualistic feedbacks to stabilize ecosystems, but also reveal an important drawback as small environmental changes may trigger shifts. We therefore suggest that mutualisms should be considered for marine conservation and restoration of seagrass beds.

Vegetation recovery in tidal marshes reveals critical slowing down under increased inundation.

A declining rate of recovery following disturbance has been proposed as an important early warning for impending tipping points in complex systems. Despite extensive theoretical and laboratory studies, this 'critical slowing down' remains largely untested in the complex settings of real-world ecosystems. Here, we provide both observational and experimental support of critical slowing down along natural stress gradients in tidal marsh ecosystems. Time series of aerial images of European marsh development reveal a consistent lengthening of recovery time as inundation stress increases. We corroborate this finding with transplantation experiments in European and North American tidal marshes. In particular, our results emphasize the power of direct observational or experimental measures of recovery over indirect statistical signatures, such as spatial variance or autocorrelation. Our results indicate that the phenomenon of critical slowing down can provide a powerful tool to probe the resilience of natural ecosystems.

Sprouting as a gardening strategy to obtain superior supplementary food: evidence from a seed-caching marine worm.

Only a handful of non-human animals are known to grow their own food by cultivating high-yield fungal or algal crops as staple food. Here we report an alternative strategy utilized by an omnivorous marine worm, Hediste diversicolor, to supplement its diet: gardening by sprouting seeds. In addition to having many other known feeding modes, we showed using video recordings and manipulative mesocosm experiments that this species can also behave like gardeners by deliberately burying cordgrass seeds in their burrows, which has been previously shown to reduce the loss of seeds to water. These seeds, however, are protected by the seed husk, and we used feeding experiments to show that they were not edible for H. diversicolor until they had sprouted or the seed husk had been artificially removed. Additionally, sprouts were shown to be highly nutritious, permitting higher growth rates in H. diversicolor than the low-quality basal food, detritus. We propose both a proximate cause (seed husk as a physical barrier) and ultimate cause (nutritional demand) for this peculiar feeding behavior. Our findings suggest that sprouting may be a common strategy used by seed-collecting animals to exploit nutrients from well-protected seeds.

Drought, Mutualism Breakdown, and Landscape-Scale Degradation of Seagrass Beds.

In many marine ecosystems, biodiversity critically depends on foundation species such as corals and seagrasses that engage in mutualistic interactions [1-3]. Concerns grow that environmental disruption of marine mutualisms exacerbates ecosystem degradation, with breakdown of the obligate coral mutualism ("coral bleaching") being an iconic example [2, 4, 5]. However, as these mutualisms are mostly facultative rather than obligate, it remains unclear whether mutualism breakdown is a common risk in marine ecosystems, and thus a potential accelerator of ecosystem degradation. Here, we provide evidence that drought triggered landscape-scale seagrass degradation and show the consequent failure of a facultative mutualistic feedback between seagrass and sulfide-consuming lucinid bivalves that in turn appeared to exacerbate the observed collapse. Local climate and remote sensing analyses revealed seagrass collapse after a summer with intense low-tide drought stress. Potential analysis-a novel approach to detect feedback-mediated state shifts-revealed two attractors (healthy and degraded states) during the collapse, suggesting that the drought disrupted internal feedbacks to cause abrupt, patch-wise degradation. Field measurements comparing degraded patches that were healthy before the collapse with patches that remained healthy demonstrated that bivalves declined dramatically in degrading patches with associated high sediment sulfide concentrations, confirming the breakdown of the mutualistic seagrass-lucinid feedback. Our findings indicate that drought triggered mutualism breakdown, resulting in toxic sulfide concentrations that aggravated seagrass degradation. We conclude that external disturbances can cause sudden breakdown of facultative marine mutualistic feedbacks. As this may amplify ecosystem degradation, we suggest including mutualisms in marine conservation and restoration approaches.

Management of local stressors can improve the resilience of marine canopy algae to global stressors.

Coastal systems are increasingly threatened by multiple local anthropogenic and global climatic stressors. With the difficulties in remediating global stressors, management requires alternative approaches that focus on local scales. We used manipulative experiments to test whether reducing local stressors (sediment load and nutrient concentrations) can improve the resilience of foundation species (canopy algae along temperate rocky coastlines) to future projected global climate stressors (high wave exposure, increasing sea surface temperature), which are less amenable to management actions. We focused on Fucoids (Cystoseira barbata) along the north-western Adriatic coast in the Mediterranean Sea because of their ecological relevance, sensitivity to a variety of human impacts, and declared conservation priority. At current levels of sediment and nutrients, C. barbata showed negative responses to the simulated future scenarios of high wave exposure and increased sea surface temperature. However, reducing the sediment load increased the survival of C. barbata recruits by 90.24% at high wave exposure while reducing nutrient concentrations resulted in a 20.14% increase in the survival and enhanced the growth of recruited juveniles at high temperature. We conclude that improving water quality by reducing nutrient concentrations, and particularly the sediment load, would significantly increase the resilience of C. barbata populations to projected increases in climate stressors. Developing and applying appropriate targets for specific local anthropogenic stressors could be an effective management action to halt the severe and ongoing loss of key marine habitats.

Low-canopy seagrass beds still provide important coastal protection services.

One of the most frequently quoted ecosystem services of seagrass meadows is their value for coastal protection. Many studies emphasize the role of above-ground shoots in attenuating waves, enhancing sedimentation and preventing erosion. This raises the question if short-leaved, low density (grazed) seagrass meadows with most of their biomass in belowground tissues can also stabilize sediments. We examined this by combining manipulative field experiments and wave measurements along a typical tropical reef flat where green turtles intensively graze upon the seagrass canopy. We experimentally manipulated wave energy and grazing intensity along a transect perpendicular to the beach, and compared sediment bed level change between vegetated and experimentally created bare plots at three distances from the beach. Our experiments showed that i) even the short-leaved, low-biomass and heavily-grazed seagrass vegetation reduced wave-induced sediment erosion up to threefold, and ii) that erosion was a function of location along the vegetated reef flat. Where other studies stress the importance of the seagrass canopy for shoreline protection, our study on open, low-biomass and heavily grazed seagrass beds strongly suggests that belowground biomass also has a major effect on the immobilization of sediment. These results imply that, compared to shallow unvegetated nearshore reef flats, the presence of a short, low-biomass seagrass meadow maintains a higher bed level, attenuating waves before reaching the beach and hence lowering beach erosion rates. We propose that the sole use of aboveground biomass as a proxy for valuing coastal protection services should be reconsidered.