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By changing habitat conditions, ecosystem engineers increase niche diversity and have profound effects on the distribution and abundances of other organisms. Although many ecosystems contain several engineering species, it is still unclear how the coexistence of multiple engineers affects the physical habitat and the structure of the community on a landscape scale. Here, we investigated through a large-scale field manipulation how three coexisting engineers on intertidal flats (cockles Cerastoderma edule; lugworms Arenicola marina; blue mussels Mytilus edulis) influence the functional composition of the local macrobenthic community and what the consequences are at the landscape level. By using biological trait analysis (BTA), we show that on the local scale biogenic changes in sediment accumulation and organic matter content translated into specific shifts in the distribution of functional traits within the community. At a landscape scale, the co-occurrence of multiple ecosystem engineers resulted in the spatial separation of different functional groups, i.e. different functional groups dominated unique complementary habitats. Our results emphasize the role of co-occurring multiple engineers in shaping natural communities, thus contributing to a better knowledge of community assembly rules. This understanding can profitably be used to improve ecosystem-based management and conservation actions.

Facilitation by foundation species can play a critical role in structuring ecological communities. As environmental stress increases, generally more organisms become dependent on the stress buffering provided by foundation species. As such, foundation species may even facilitate their own predators, an interaction that can influence the functioning and structure of the foundation species population and thereby the facilitated organisms. This work presents a case study on the blue mussels and sea stars, where we tested to what extent a foundation species (i.e. blue mussels) may facilitate its own predator (sea star) when exposed to a gradient of environmental stress (hydrodynamic forces). Amelioration of hydrodynamic stress by mussels facilitated sea stars, allowing them to persist on a soft bottom in highly dynamic environment, which would not be possible in the absence of mussels. Moreover, sea stars continue preying on mussels when environmental stress increases. The results suggest that a foundation species may interact with its own predator beyond the role of food source, by ameliorating environmental stress, creating an additional dependence link between the foundation species and the predator, which potentially has major implications for ecosystem structure and stability.

Recently, Lévy walks have been put forward as a new paradigm for animal search and many cases have been made for its presence in nature. However, it remains debated whether Lévy walks are an inherent behavioural strategy or emerge from the animal reacting to its habitat. Here, we demonstrate signatures of Lévy behaviour in the search movement of mud snails (Hydrobia ulvae) based on a novel, direct assessment of movement properties in an experimental set-up using different food distributions. Our experimental data uncovered clusters of small movement steps alternating with long moves independent of food encounter and landscape complexity. Moreover, size distributions of these clusters followed truncated power laws. These two findings are characteristic signatures of mechanisms underlying inherent Lévy-like movement. Thus, our study provides clear experimental evidence that such multi-scale movement is an inherent behaviour rather than resulting from the animal interacting with its environment

Grazers can have a large impact on ecosystem processes and are known to changevegetation composition. However, knowledge of how the long-term presence of grazers affectssoil carbon sequestration is limited. In this study, we estimated total accumulated organic carbonin soils of a back-barrier salt marsh and determined how this is affected by long-term grazing byboth small and large grazers in relation to age of the ecosystem. In young marshes, where smallgrazers predominate, hare and geese have a limited effect on total accumulated organic carbon.In older, mature marshes, where large grazers predominate, cattle substantially enhanced carboncontent in the marsh soil. We ascribe this to a shift in biomass distribution in the local vegetationtowards the roots in combination with trampling effects on the soil chemistry. These large grazersthus act as ecosystem engineers: their known effect on soil compaction (based on a previous study)enhances anoxic conditions in the marsh soil, thereby reducing the oxygen available for organiccarbon decomposition by the local microbial community. This study showed that the indirecteffects of grazing can significantly enhance soil carbon storage through changing soil abiotic conditions.This process should be taken into account when estimating the role of ecosystems inreducing carbon dioxide concentration in the atmosphere. Ultimately, we propose a testableconceptual framework that includes 3 pathways by which grazers can alter carbon storage:(1) through above-ground biomass removal, (2) through alteration of biomass distribution towardsthe roots and/or (3) by changing soil abiotic conditions that affect decomposition.

Mounting evidence indicates that spatial interactions are important in structuring coastal ecosystems. Until recently, however, most of this work has been focused on seemingly exceptional systems that are characterized by regular, self-organized patterns. In this review, we document that interactions that operate at long distances, beyond the direct neighborhood of individual organisms, are more common and have much more far-reaching implications for coastal ecosystems than was previously realized. We review studies from a variety of ecosystem types—including cobble beaches, mussel beds, coral reefs, seagrass meadows, and mangrove forests—that reveal a startling interplay of positive and negative interactions between habitats across distances of up to a kilometer. In addition to classical feeding relations, alterations of physical conditions constitute an important part of these long-distance interactions. This entanglement of habitats has crucial implications for how humans manage coastal ecosystems, and evaluations of anthropogenic impact should explicitly address long-distance and system-wide effects before we deem these human activities to be causing little harm.

Human exploitation of bivalve populations has changed intertidal landscapes worldwide. Many bivalves are ecosystem engineers that modify the physical environment, affecting the conditions for their survival. Here we argue that lack of recovery of overexploited intertidal bivalve beds may be partly caused by the loss of important biological feedbacks from depleted populations. In a large-scale experiment we investigated engineering effects of cockles (Cerastoderma edule L.) and lugworms (Arenicola marina L.) on juvenile cockles by adding high densities of either species to 5 × 5 m plots in areas with different hydrodynamic and sediment conditions in the intertidal flats of the Wadden Sea. We hypothesized that cockles would facilitate the new generations by increasing sediment stability, while lugworms would have negative effects on juvenile cockles through sediment disturbance. We found that in sandy areas with high wave and current energy cockles enhanced sediment accumulation and promoted local densities of young cockles, while lugworms did not have any effect on juvenile cockles. In muddy sites sheltered from the tidal currents by mussel reefs (Mytilus edulis L.), juvenile cockle densities were generally high, demonstrating the general importance of biological engineering for recruitment processes in the intertidal. We suggest that the acknowledgement of positive feedbacks between bivalves and sediment stability is essential to achieve long-term restoration goals in coastal ecosystems.

Invasive species threaten biodiversity and incur costs exceeding billions of US$. 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.

All living organisms modify their biotic and abiotic environment. Niche construction theory posits that organism-mediated modifications to the environment can change selection pressures and influence the evolutionary trajectories of natural populations. While there is broad support for this proposition in general, there is considerable uncertainty about how niche construction is related to other similar concepts in ecology and evolution. Comparative studies dealing with certain aspects of niche construction are increasingly common, but there is a troubling lack of experimental tests of the core concepts of niche construction theory. Here, we propose an operational framework to evaluate comparative and experimental evidence of the evolutionary consequences of niche construction, and suggest how such research can improve our understanding of ecological and evolutionary dynamics in ecosystems. We advocate for a shift toward explicit experimental tests of how organism-mediated environmental change can influence the selection pressures underlying evolutionary responses, as well as targeted field-based comparative research to identify the mode of evolution by niche construction and assess its importance in natural populations.

Theoretical models highlight that spatially self-organized patterns can have important emergent effects on the functioning of ecosystems, for instance by increasing productivity and affecting the vulnerability to catastrophic shifts. However, most theoretical studies presume idealized homogeneous conditions, which are rarely met in real ecosystems. Using self-organized mussel beds as a case study, we reveal that spatial heterogeneity, resulting from the large-scale effects of mussel beds on their environment, significantly alters the emergent properties predicted by idealized self-organization models that assume homogeneous conditions. The proposed model explicitly considers that the suspended algae, the prime food for the mussels, are supplied by water flow from the seaward boundary of the bed, which causes in combination with consumption a gradual depletion of algae over the simulated domain. Predictions of the model are consistent with properties of natural mussel patterns observed in the field, featuring a decline in mussel biomass and a change in patterning. Model analyses reveal a fundamental change in ecosystem functioning when this self-induced algal depletion gradient is included in the model. First, no enhancement of secondary productivity of the mussels comparing with non-patterns states is predicted, irrespective of parameter setting; the equilibrium amount of mussels is entirely set by the input of algae. Second, alternate stable states, potentially present in the original (no algal gradient) model, are absent when gradual depletion of algae in the overflowing water layer is allowed. Our findings stress the importance of including sufficiently realistic environmental conditions when assessing the emergent properties of self-organized ecosystems.

Marine protected areas (MPAs) are key tools for combatting the global over-exploitation of endangered species. The prevailing paradigm is that MPAs are beneficial in helping to restore ecosystems to more 'natural' conditions. However, MPAs may have unintended negative effects when increasing densities of protected species exert destructive effects on their habitat. Here, we report on severe seagrass degradation in a decade-old MPA where hyper-abundant green turtles adopted a previously undescribed below-ground foraging strategy. By digging for and consuming rhizomes and roots, turtles create abundant bare gaps, thereby enhancing erosion and reducing seagrass regrowth. A fully parametrized model reveals that the ecosystem is approaching a tipping point, where consumption overwhelms regrowth, which could potentially lead to complete collapse of the seagrass habitat. Seagrass recovery will not ensue unless turtle density is reduced to nearly zero, eliminating the MPA's value as a turtle reserve. Our results reveal an unrecognized, yet imminent threat to MPAs, as sea turtle densities are increasing at major nesting sites and the decline of seagrass habitat forces turtles to concentrate on the remaining meadows inside reserves. This emphasizes the need for policy and management approaches that consider the interactions of protected species with their habitat.

Bivalves are key components of coastal ecosystems because they link pelagic and benthic food webs, and shape the landscape through habitat modification. Nevertheless, many bivalve stocks have dramatically declined, and recruitment failure due to (anthropogenically-) increased predation by mesopredators and loss of facilitation mechanisms have been separately hypothesized as underlying causes. Here, we tested the interactive effects of predation and habitat modification on bivalve recruitment in a large-scale experiment in the Wadden Sea, one of the world's largest intertidal soft-sediment ecosystems. We applied anti-erosion mats to simulate biotic attachment and substrate stabilization by commonly found tubeworm beds, crossed this with addition of adult mussels, and manipulated shrimp and crab predation using exclosures within these treatments. Epibenthic mussel recruits were only found in treatments with manipulated substrates, attached to either the anti-erosion mat or adult mussels. Three out of four endobenthic species were facilitated by the mat, but were inhibited by adult mussels. In contrast, invasive surf-dwelling American razor clams were inhibited by both substrate manipulations, indicating a preference for unstable sediments. These facilitation and inhibition effects, however, only clearly emerged when predators were excluded, demonstrating strong synergistic effects between predation and habitat modification. Our findings suggest that disturbance of trophic interactions and loss of habitat modifying species interactively affect bivalve recruitment dynamics in coastal ecosystems. We conclude that conservation and restoration of bivalves should focus on protecting and restoring internal facilitation mechanisms, and should simultaneously reduce excessive mesopredator predation by restoring natural food web dynamics, including the role of top-predators.

Ecological theory uses Brownian motion as a default template for describing ecological movement, despite limited mechanistic underpinning. The generality of Brownian motion has recently been challenged by empirical studies that highlight alternative movement patterns of animals, especially when foraging in resource-poor environments. Yet, empirical studies reveal animals moving in a Brownian fashion when resources are abundant. We demonstrate that Einstein's original theory of collision-induced Brownian motion in physics provides a parsimonious, mechanistic explanation for these observations. Here, Brownian motion results from frequent encounters between organisms in dense environments. In density-controlled experiments, movement patterns of mussels shifted from Lévy towards Brownian motion with increasing density. When the analysis was restricted to moves not truncated by encounters, this shift did not occur. Using a theoretical argument, we explain that any movement pattern approximates Brownian motion at high-resource densities, provided that movement is interrupted upon encounters. Hence, the observed shift to Brownian motion does not indicate a density-dependent change in movement strategy but rather results from frequent collisions. Our results emphasize the need for a more mechanistic use of Brownian motion in ecology, highlighting that especially in rich environments, Brownian motion emerges from ecological interactions, rather than being a default movement pattern., Ecological theory uses Brownian motion as a default template for describing ecological movement, despite limited mechanistic underpinning. The generality of Brownian motion has recently been challenged by empirical studies that highlight alternative movement patterns of animals, especially when foraging in resource-poor environments. Yet, empirical studies reveal animals moving in a Brownian fashion when resources are abundant. We demonstrate that Einstein's original theory of collision-induced Brownian motion in physics provides a parsimonious, mechanistic explanation for these observations. Here, Brownian motion results from frequent encounters between organisms in dense environments. In density-controlled experiments, movement patterns of mussels shifted from Lévy towards Brownian motion with increasing density. When the analysis was restricted to moves not truncated by encounters, this shift did not occur. Using a theoretical argument, we explain that any movement pattern approximates Brownian motion at high-resource densities, provided that movement is interrupted upon encounters. Hence, the observed shift to Brownian motion does not indicate a density-dependent change in movement strategy but rather results from frequent collisions. Our results emphasize the need for a more mechanistic use of Brownian motion in ecology, highlighting that especially in rich environments, Brownian motion emerges from ecological interactions, rather than being a default movement pattern.

The food webs of terrestrial soils and of freshwater and marine sediments depend on adjacent aboveground or pelagic ecosystems for organic matter input that provides nutrients and energy. There are important similarities in the flow of organic matter through these food webs and how this flow feeds back to primary production. In both soils and sediments, trophic interactions occur in a cycle in which consumers stimulate nutrient cycling such that mineralized resources are made available to the primary producers. However, aquatic sediments and terrestrial soils differ greatly in the connectivity between the production and the consumption of organic matter. Terrestrial soils and shallow aquatic sediments can receive organic matter within hours of photosynthesis when roots leak carbon, whereas deep oceanic sediments receive organic matter possibly months after carbon assimilation by phytoplankton. This comparison has implications for the capacity of soils and sediments to affect the global carbon balance., The food webs of terrestrial soils and of freshwater and marine sediments depend on adjacent aboveground or pelagic ecosystems for organic matter input that provides nutrients and energy. There are important similarities in the flow of organic matter through these food webs and how this flow feeds back to primary production. In both soils and sediments, trophic interactions occur in a cycle in which consumers stimulate nutrient cycling such that mineralized resources are made available to the primary producers. However, aquatic sediments and terrestrial soils differ greatly in the connectivity between the production and the consumption of organic matter. Terrestrial soils and shallow aquatic sediments can receive organic matter within hours of photosynthesis when roots leak carbon, whereas deep oceanic sediments receive organic matter possibly months after carbon assimilation by phytoplankton. This comparison has implications for the capacity of soils and sediments to affect the global carbon balance.

The importance of positive effects of ecosystem engineers on associated communities is predicted to increase with environmental stress. However, incorporating such non-trophic interactions into ecological theory is not trivial because facilitation of associated species is conditional on both the type of engineer and the type of abiotic stress. We tested the influence of two allogenic ecosystem engineers (lugworms, Arenicola marina L. and cockles, Cerastoderma edule L.) on the main primary producers (microphytobenthos) of the tidal flats, under different abiotic stresses controlled by reefs of blue mussels (Mytilus edulis L.). We added 25,000 cockles or 2,000 lugworms to 5 × 5 m plots, both in a muddy site with high sedimentation rates located coastward of a mussel bed, and in a sandy site without mussels and characterized by high hydrodynamic stress. After a year, cockles increased algal biomass in the sandy area, but not in the mussel bed site, where high values were measured in all plots. However, lugworms did not affect algal biomass in any of the sites. Field measurements suggest that cockles outweighed negative effects of water currents in the site without mussels by locally increasing sediment stability, whereas mussels overruled the effects of cockles in the wake of the reefs through hydrodynamic stress alleviation and/or biodeposition. Our results suggest that non-trophic interactions by ecosystem engineering bivalves control primary production of intertidal areas, and that the sediment-stabilizing effect of cockles plays a crucial role where the overruling effects of mussel beds are not present.

The food webs of terrestrial soils and of freshwater and marine sediments depend on adjacent aboveground or pelagic ecosystems for organic matter input that provides nutrients and energy. There are important similarities in the flow of organic matter through these food webs and how this flow feeds back to primary production. In both soils and sediments, trophic interactions occur in a cycle in which consumers stimulate nutrient cycling such that mineralized resources are made available to the primary producers. However, aquatic sediments and terrestrial soils differ greatly in the connectivity between the production and the consumption of organic matter. Terrestrial soils and shallow aquatic sediments can receive organic matter within hours of photosynthesis when roots leak carbon, whereas deep oceanic sediments receive organic matter possibly months after carbon assimilation by phytoplankton. This comparison has implications for the capacity of soils and sediments to affect the global carbon balance.

Increasing evidence shows that spatial interactions between sedentary organisms can structure communities and promote landscape complexity in many ecosystems. Here we tested the hypothesis that reef-forming mussels (Mytilus edulis L.), a dominant intertidal ecosystem engineer in the Wadden Sea, promote abundances of the burrowing bivalve Cerastoderma edule L. (cockle) in neighboring habitats at relatively long distances coastward from mussel beds. Field surveys within and around three mussel beds showed a peak in cockle densities at 50-100 m toward the coast from the mussel bed, while cockle abundances elsewhere in the study area were very low. Field transplantation of cockles showed higher survival of young cockles (2-3 years old) and increased spat fall coastward of the mussel bed compared to within the bed and to areas without mussels, whereas growth decreased within and coastward of the mussel bed. Our measurements suggest that the observed spatial patterns in cockle numbers resulted from (1) inhibition effects by the mussels close to the beds due to preemptive algal depletion and deteriorated sediment conditions and (2) facilitation effects by the mussels farther away from the beds due to reduction of wave energy. Our results imply that these spatial, scale-dependent interactions between reef-forming ecosystem engineers and surrounding communities of sedentary benthic organisms can be an important determinant of the large-scale community structure in intertidal ecosystems. Understanding this interplay between neighboring communities of sedentary species is therefore essential for effective conservation and restoration of soft-bottom intertidal communities.

The origin of regular spatial patterns in ecological systems has long fascinated researchers. Turing's activator-inhibitor principle is considered the central paradigm to explain such patterns. According to this principle, local activation combined with long-range inhibition of growth and survival is an essential prerequisite for pattern formation. Here, we show that the physical principle of phase separation, solely based on density-dependent movement by organisms, represents an alternative class of self-organized pattern formation in ecology. Using experiments with self-organizing mussel beds, we derive an empirical relation between the speed of animal movement and local animal density. By incorporating this relation in a partial differential equation, we demonstrate that this model corresponds mathematically to the wellknown Cahn-Hilliard equation for phase separation in physics. Finally, we show that the predicted patterns match those found both in field observations and in our experiments. Our results reveal a principle for ecological self-organization, where phase separation rather than activation and inhibition processes drives spatial pattern formation.

There is a growing recognition of the important role of scale-dependent feedback for biogeomorphological landscape formation, where organisms locally improve survival and growth but at the same time negatively affect organisms at larger distance. However, little is known on how scale-dependent bio-geomorphic feedback is influenced by organism traits in combination with abiotic forcing. This was studied by measuring in a flume, the flow patterns around patches of three contrasting marsh species (Spartina anglica, Puccinellia maritima and Salicornia procumbens), using the flow acceleration around vegetation patches and deceleration within vegetation patches as quantitative proxy for the negative and positive feedback to the vegetation performance. The importance of external forcing was assessed by comparing three realistic current velocities: 0.1, 02 and 0.3 m s(-1). Our results showed that the dense clonal growth of stiff Spartina anglica shoots caused strongest flow deviations, irrespective of the applied current velocity. In contrast, the more sparsely growing, shorter stiff shoots of Salicornia procumbens induced much less flow deviation, allowing more water to pass through and over the vegetation canopy. The dense but highly flexible shoots of Puccinellia maritima caused strong flow deviations at low velocities, which diminished at higher velocities due to bending of the vegetation. Overall, these hydrodynamic results demonstrate that plant species traits interact with environmental conditions in creating scale-dependent feedbacks explaining why the effects of vegetation on landscape formation in saltmarshes are species specific.

Salt marshes are delicate landforms at the boundary between the sea and land. These ecosystems support a diverse biota that modifies the erosive characteristics of the substrate and mediates sediment transport processes. Here we present a broad overview of recent numerical models that quantify the formation and evolution of salt marshes under different physical and ecological drivers. In particular, we focus on the coupling between geomorphological and ecological processes and on how these feedbacks are included in predictive models of landform evolution. We describe in detail models that simulate fluxes of water, organic matter, and sediments in salt marshes. The interplay between biological and morphological processes often produces a distinct scarp between salt marshes and tidal flats. Numerical models can capture the dynamics of this boundary and the progradation or regression of the marsh in time. Tidal channels are also key features of the marsh landscape, flooding and draining the marsh platform and providing a source of sediments and nutrients to the marsh ecosystem. In recent years, several numerical models have been developed to describe the morphogenesis and long-term dynamics of salt marsh channels. Finally, salt marshes are highly sensitive to the effects of long-term climatic change. We therefore discuss in detail how numerical models have been used to determine salt marsh survival under different scenarios of sea level rise.

Theoretical models predict that spatial self-organization can have important, unexpected implications by affecting the functioning of ecosystems in terms of resilience and productivity. Whether and how these emergent effects depend on specific formulations of the underlying mechanisms are questions that are often ignored. Here, we compare two alternative models of regular spatial pattern formation in mussel beds that have different mechanistic descriptions of the facilitative interactions between mussels. The first mechanism involves a reduced mussel loss rate at high density owing to mutual protection between the mussels, which is the basis of prior studies on the pattern formation in mussels. The second mechanism assumes, based on novel experimental evidence, that mussels feed more efficiently on top of mussel-generated hummocks. Model simulations point out that the second mechanism produces very similar types of spatial patterns in mussel beds. Yet the mechanisms predict a strikingly contrasting effect of these spatial patterns on ecosystem functioning, in terms of productivity and resilience. In the first model, where high mussel densities reduce mussel loss rates, patterns are predicted to strongly increase productivity and decrease the recovery time of the bed following a disturbance. When pattern formation is generated by increased feeding efficiency on hummocks, only minor emergent effects of pattern formation on ecosystem functioning are predicted. Our results provide a warning against predictions of the implications and emergent properties of spatial self-organization, when the mechanisms that underlie self-organization are incompletely understood and not based on the experimental study.

More than 2 y have passed since the BP-Deepwater Horizon oil spill in the Gulf of Mexico, yet we still have little understanding of its ecological impacts. Examining effects of this oil spill will generate much-needed insight into how shoreline habitats and the valuable ecological services they provide (e. g., shoreline protection) are affected by and recover from large-scale disturbance. Here we report on not only rapid salt-marsh recovery (high resilience) but also permanent marsh area loss after the BP-Deepwater Horizon oil spill. Field observations, experimental manipulations, and wave-propagation modeling reveal that (i) oil coverage was primarily concentrated on the seaward edge of marshes; (ii) there were thresholds of oil coverage that were associated with severity of salt-marsh damage, with heavy oiling leading to plant mortality; (iii) oil-driven plant death on the edges of these marshes more than doubled rates of shoreline erosion, further driving marsh platform loss that is likely to be permanent; and (iv) after 18 mo, marsh grasses have largely recovered into previously oiled, noneroded areas, and the elevated shoreline retreat rates observed at oiled sites have decreased to levels at reference marsh sites. This paper highlights that heavy oil coverage on the shorelines of Louisiana marshes, already experiencing elevated retreat because of intense human activities, induced a geomorphic feedback that amplified this erosion and thereby set limits to the recovery of otherwise resilient vegetation. It thus warns of the enhanced vulnerability of already degraded marshes to heavy oil coverage and provides a clear example of how multiple human-induced stressors can interact to hasten ecosystem decline.

Seagrasses evolved from terrestrial plants into marine foundation species around 100 million years ago. Their ecological success, however, remains a mystery because natural organic matter accumulation within the beds should result in toxic sediment sulfide levels. Using a meta-analysis, a field study, and a laboratory experiment, we reveal how an ancient three-stage symbiosis between seagrass, lucinid bivalves, and their sulfide-oxidizing gill bacteria reduces sulfide stress for seagrasses. We found that the bivalve-sulfide-oxidizer symbiosis reduced sulfide levels and enhanced seagrass production as measured in biomass. In turn, the bivalves and their endosymbionts profit from organic matter accumulation and radial oxygen release from the seagrass roots. These findings elucidate the long-term success of seagrasses in warm waters and offer new prospects for seagrass ecosystem conservation.

Avian influenza virus reveals persistent and recurrent outbreaks in North American wild waterfowl, and exhibits major outbreaks at 2-8 years intervals in duck populations. The standard susceptible-infected-recovered (SIR) framework, which includes seasonal migration and reproduction, but lacks environmental transmission, is unable to reproduce the multi-periodic patterns of avian influenza epidemics. In this paper, we argue that a fully stochastic theory based on environmental transmission provides a simple, plausible explanation for the phenomenon of multi-year periodic outbreaks of avian flu. Our theory predicts complex fluctuations with a dominant period of 2 to 8 years which essentially depends on the intensity of environmental transmission. A wavelet analysis of the observed data supports this prediction. Furthermore, using master equations and van Kampen system-size expansion techniques, we provide an analytical expression for the spectrum of stochastic fluctuations, revealing how the outbreak period varies with the environmental transmission.

Complexity theory proposes that spatial self-organization, the process whereby small-scale, localized interactions among the components of a system generate complex spatial structures at large spatial scales, explains the formation of autogenic spatial patterns in ecosystems. We question this premise by reviewing three estuarine ecosystems - mussel beds, mudflats and salt marshes - where self-organization has been put forward to explain spatial patterns. Our review highlights that these self-organized estuarine systems are shaped by the combination of small-scale interactions between ecological and physical processes on the one hand, and large-scale physical forcing on the other. More specifically, local interactions generate patchiness at small spatial scales, whereas landscape forcing determines the shape and orientation of these patches in the landscape. We present a framework that illustrates how self-organized ecosystems are shaped by interactions between organisms and physical processes occurring at multiple spatial scales. Moreover, the present review of estuarine systems underlines that scale-dependent feedbacks are capable of explaining spatial patterns that are much more complex than the regular patterns to which they have been applied so far.

Self-organized spatial patterns have been proposed as possible indicators for regime shifts in ecosystems. Until now this hypothesis has only been tested in drylands. Here, we focus on intertidal mudflats where regular spatial patterns develop in early spring from the interaction between diatom growth and sedimentation but disappear when benthic herbivore abundance increases in early summer, accompanied by a dramatic shift to a bare mudflat. We followed the patch-size distributions of diatom biofilms during this degradation process. As time progressed, we found a temporal change in the spatial configuration occurring simultaneously with the loss of the diatom-sediment feedback. This indicates a gradual failure in time of the self-organization process that underlies regular patterning in this ecosystem. The path to degradation co-occurred with the loss of the larger patches in the ecosystem, which resulted in a decrease of the truncation in the patch-size distribution. Hence, our study in mudflat ecosystems confirms the general hypothesis that spatial patterns can provide important clues about the level of degradation. Nevertheless, our study highlights the need for thorough study about the type of spatial patterns and the nature of the underlying feedbacks before a reliable assessment of ecosystem status can be made, as changes in patch-size distribution differed markedly with those observed in other ecosystems.

We agree with Jansen et al. that a composite movement model provides a better statistical description of mussel movement than any simple movement strategy. This does not undermine the take-home message of our paper, which addresses the feedback between individual movement patterns and spatial complexity. Simple movement strategies provide more insight in the eco-evolutionary analysis and are therefore our model of choice., We agree with Jansen et al. that a composite movement model provides a better statistical description of mussel movement than any simple movement strategy. This does not undermine the take-home message of our paper, which addresses the feedback between individual movement patterns and spatial complexity. Simple movement strategies provide more insight in the eco-evolutionary analysis and are therefore our model of choice.

Ecological theory predicts that animal movement is shaped by its efficiency of resource acquisition. Focusing solely on efficiency, however, ignores the fact that animal activity can affect resource availability and distribution. Here, we show that feedback between individual behavior and environmental complexity can explain movement strategies in mussels. Specifically, experiments show that mussels use a Lévy walk during the formation of spatially patterned beds, and models reveal that this Lévy movement accelerates pattern formation. The emergent patterning in mussel beds, in turn, improves individual fitness. These results suggest that Lévy walks evolved as a result of the selective advantage conferred by autonomously generated, emergent spatial patterns in mussel beds. Our results emphasize that an interaction between individual selection and habitat complexity shapes animal movement in natural systems., Ecological theory predicts that animal movement is shaped by its efficiency of resource acquisition. Focusing solely on efficiency, however, ignores the fact that animal activity can affect resource availability and distribution. Here, we show that feedback between individual behavior and environmental complexity can explain movement strategies in mussels. Specifically, experiments show that mussels use a Lévy walk during the formation of spatially patterned beds, and models reveal that this Lévy movement accelerates pattern formation. The emergent patterning in mussel beds, in turn, improves individual fitness. These results suggest that Lévy walks evolved as a result of the selective advantage conferred by autonomously generated, emergent spatial patterns in mussel beds. Our results emphasize that an interaction between individual selection and habitat complexity shapes animal movement in natural systems.

How the properties of ecosystems relate to spatial scale is a prominent topic in current ecosystem research. Despite this, spatially explicit models typically include only a limited range of spatial scales, mostly because of computing limitations. Here, we describe the use of graphics processors to efficiently solve spatially explicit ecological models at large spatial scale using the CUDA language extension. We explain this technique by implementing three classical models of spatial self-organization in ecology: a spiral-wave forming predator-prey model, a model of pattern formation in arid vegetation, and a model of disturbance in mussel beds on rocky shores. Using these models, we show that the solutions of models on large spatial grids can be obtained on graphics processors with up to two orders of magnitude reduction in simulation time relative to normal pc processors. This allows for efficient simulation of very large spatial grids, which is crucial for, for instance, the study of the effect of spatial heterogeneity on the formation of self-organized spatial patterns, thereby facilitating the comparison between theoretical results and empirical data. Finally, we show that large-scale spatial simulations are preferable over repetitions at smaller spatial scales in identifying the presence of scaling relations in spatially self-organized ecosystems. Hence, the study of scaling laws in ecology may benefit significantly from implementation of ecological models on graphics processors. (C) 2011 Elsevier B.V. All rights reserved., How the properties of ecosystems relate to spatial scale is a prominent topic in current ecosystem research. Despite this, spatially explicit models typically include only a limited range of spatial scales, mostly because of computing limitations. Here, we describe the use of graphics processors to efficiently solve spatially explicit ecological models at large spatial scale using the CUDA language extension. We explain this technique by implementing three classical models of spatial self-organization in ecology: a spiral-wave forming predator-prey model, a model of pattern formation in arid vegetation, and a model of disturbance in mussel beds on rocky shores. Using these models, we show that the solutions of models on large spatial grids can be obtained on graphics processors with up to two orders of magnitude reduction in simulation time relative to normal pc processors. This allows for efficient simulation of very large spatial grids, which is crucial for, for instance, the study of the effect of spatial heterogeneity on the formation of self-organized spatial patterns, thereby facilitating the comparison between theoretical results and empirical data. Finally, we show that large-scale spatial simulations are preferable over repetitions at smaller spatial scales in identifying the presence of scaling relations in spatially self-organized ecosystems. Hence, the study of scaling laws in ecology may benefit significantly from implementation of ecological models on graphics processors. (C) 2011 Elsevier B.V. All rights reserved.

Regular, self-organized spatial patterns in primary producers have been described in a wide range of ecosystems and are predicted to affect community production and resilience. Although consumers are abundant in most systems, the effect of trophic interactions on pattern formation in primary producers remains unstudied. We studied the effects of top-down control by herbivores on a self-organized landscape of regularly spaced, diatom-covered hummocks alternating with water-filled hollows on an intertidal mudflat in The Netherlands. Spatial patterns developed during spring but were followed by a rapid collapse in summer, leading to a flat landscape with low diatom densities and little variation in sediment bed level. This dramatic decline co-occurred with a gradual increase of benthic herbivores. A manipulative field experiment, where benthic herbivores were removed from the sediment, revealed that both diatom growth and hummock formation were inhibited by the activity of benthic herbivores. Our study provides clear evidence of top-down control of spatial selforganized patterns by benthic herbivores within a biological–geomorphic landscape., Regular, self-organized spatial patterns in primary producers have been described in a wide range of ecosystems and are predicted to affect community production and resilience. Although consumers are abundant in most systems, the effect of trophic interactions on pattern formation in primary producers remains unstudied. We studied the effects of top-down control by herbivores on a self-organized landscape of regularly spaced, diatom-covered hummocks alternating with water-filled hollows on an intertidal mudflat in The Netherlands. Spatial patterns developed during spring but were followed by a rapid collapse in summer, leading to a flat landscape with low diatom densities and little variation in sediment bed level. This dramatic decline co-occurred with a gradual increase of benthic herbivores. A manipulative field experiment, where benthic herbivores were removed from the sediment, revealed that both diatom growth and hummock formation were inhibited by the activity of benthic herbivores. Our study provides clear evidence of top-down control of spatial selforganized patterns by benthic herbivores within a biological–geomorphic landscape.

Natural ecosystems can show regular spatial vegetation patterns, which develop from small-scale ecological interactions. Some studies suggest that grazers can play a major role in controlling vegetation distribution in ecosystems with regular vegetation patterns, but the distribution of grazers and the effects of grazing on vegetation in spatially patterned ecosystems remains poorly understood. Here, we study how macrofaunal grazers are distributed on a spatially patterned intertidal flat and how they interact with benthic microalgae. The study was carried out on an intertidal flat where each year a regular spatial pattern of diatom-covered hummocks and water-filled hollows develops. In 2 consecutive years, benthic algal biomass was 5-fold lower in the hollows compared with the hummocks, whereas benthic macrofauna was equally abundant on both hummocks and hollows. In the laboratory we evaluated the interactive effect of landscape morphology (hummocks and hollows) and food abundance. Experiments with 2 abundant grazers, Corophium volutator and Hydrobia ulvae showed that food availability was the main driver for the spatial distribution of benthic herbivores rather than landscape morphology. Laboratory experiments, where we analyzed the grazing effects of H. ulvae and C. volutator on benthic algal biomass, revealed that both species increased biomass-specific primary production. This could indicate that benthic grazers can stimulate algal growth, which may explain the high grazer abundances in nearly bare hollows. Increased production of benthic algae may be a key factor uncoupling herbivore density from benthic algal biomass on a spatially patterned intertidal flat., Natural ecosystems can show regular spatial vegetation patterns, which develop from small-scale ecological interactions. Some studies suggest that grazers can play a major role in controlling vegetation distribution in ecosystems with regular vegetation patterns, but the distribution of grazers and the effects of grazing on vegetation in spatially patterned ecosystems remains poorly understood. Here, we study how macrofaunal grazers are distributed on a spatially patterned intertidal flat and how they interact with benthic microalgae. The study was carried out on an intertidal flat where each year a regular spatial pattern of diatom-covered hummocks and water-filled hollows develops. In 2 consecutive years, benthic algal biomass was 5-fold lower in the hollows compared with the hummocks, whereas benthic macrofauna was equally abundant on both hummocks and hollows. In the laboratory we evaluated the interactive effect of landscape morphology (hummocks and hollows) and food abundance. Experiments with 2 abundant grazers, Corophium volutator and Hydrobia ulvae showed that food availability was the main driver for the spatial distribution of benthic herbivores rather than landscape morphology. Laboratory experiments, where we analyzed the grazing effects of H. ulvae and C. volutator on benthic algal biomass, revealed that both species increased biomass-specific primary production. This could indicate that benthic grazers can stimulate algal growth, which may explain the high grazer abundances in nearly bare hollows. Increased production of benthic algae may be a key factor uncoupling herbivore density from benthic algal biomass on a spatially patterned intertidal flat.

Shallow soft-sediment systems are mostly dominated by species that, by strongly affecting sediment dynamics, modify their local environment. Such ecosystem engineering species can have either sediment-stabilizing or sediment-destabilizing effects on tidal flats. They interplay with abiotic forcing conditions (wind, tide, nutrient inputs) in driving the community structure and generating spatial heterogeneity, determining the composition of different communities of associated species, and thereby affecting the channelling of energy through different compartments in the food web. This suggests that, depending on local species composition, tidal flats may have conspicuously different geomorphology and biological functions under similar external conditions. Here we use a historical reconstruction of benthic production in the Wadden Sea to construct a framework for the relationships between human impacts, ecosystem engineering and sediment dynamics. We propose that increased sediment disturbances by human exploitation interfere with biological controls of sediment dynamics, and thereby have shifted the dominant compartments of both primary and secondary production in the Wadden Sea, transforming the intertidal from an internally regulated and spatially heterogeneous, to an externally regulated and spatially homogenous system. This framework contributes to the general understanding of the interaction between biological and environmental control of ecosystem functioning, and suggests a general framework for predicting effects of human impacts on soft-bottom ecosystems., Shallow soft-sediment systems are mostly dominated by species that, by strongly affecting sediment dynamics, modify their local environment. Such ecosystem engineering species can have either sediment-stabilizing or sediment-destabilizing effects on tidal flats. They interplay with abiotic forcing conditions (wind, tide, nutrient inputs) in driving the community structure and generating spatial heterogeneity, determining the composition of different communities of associated species, and thereby affecting the channelling of energy through different compartments in the food web. This suggests that, depending on local species composition, tidal flats may have conspicuously different geomorphology and biological functions under similar external conditions. Here we use a historical reconstruction of benthic production in the Wadden Sea to construct a framework for the relationships between human impacts, ecosystem engineering and sediment dynamics. We propose that increased sediment disturbances by human exploitation interfere with biological controls of sediment dynamics, and thereby have shifted the dominant compartments of both primary and secondary production in the Wadden Sea, transforming the intertidal from an internally regulated and spatially heterogeneous, to an externally regulated and spatially homogenous system. This framework contributes to the general understanding of the interaction between biological and environmental control of ecosystem functioning, and suggests a general framework for predicting effects of human impacts on soft-bottom ecosystems.

In this study, we investigated the emergence of spatial self-organized patterns on intertidal flats, resulting from the interaction between biological and geomorphological processes. Autocorrelation analysis of aerial photographs revealed that diatoms occur in regularly spaced patterns consisting of elevated hummocks alternating with water-filled hollows. Hummocks were characterized by high diatom content and a high sediment erosion threshold, while both were low in hollows. These results highlight the interaction between diatom growth and sedimentary processes as a potential mechanism for spatial patterning. Several alternative mechanisms could be excluded as important mechanisms in the formation of spatial patterns. We developed a spatially explicit mathematical model that revealed that scale-dependent interactions between sedimentation, diatom growth, and water redistribution explain the observed patterns. The model predicts that areas exhibiting spatially self-organized patterns have increased sediment accretion and diatom biomass compared with areas lacking spatial patterns, a prediction confirmed by empirical evidence. Our study on intertidal mudflats provides a simple but clear-cut example of how the interaction between biological and sedimentary processes, through the process of self-organization, induces spatial patterns at a landscape level., In this study, we investigated the emergence of spatial self-organized patterns on intertidal flats, resulting from the interaction between biological and geomorphological processes. Autocorrelation analysis of aerial photographs revealed that diatoms occur in regularly spaced patterns consisting of elevated hummocks alternating with water-filled hollows. Hummocks were characterized by high diatom content and a high sediment erosion threshold, while both were low in hollows. These results highlight the interaction between diatom growth and sedimentary processes as a potential mechanism for spatial patterning. Several alternative mechanisms could be excluded as important mechanisms in the formation of spatial patterns. We developed a spatially explicit mathematical model that revealed that scale-dependent interactions between sedimentation, diatom growth, and water redistribution explain the observed patterns. The model predicts that areas exhibiting spatially self-organized patterns have increased sediment accretion and diatom biomass compared with areas lacking spatial patterns, a prediction confirmed by empirical evidence. Our study on intertidal mudflats provides a simple but clear-cut example of how the interaction between biological and sedimentary processes, through the process of self-organization, induces spatial patterns at a landscape level.

The spatial structure of seagrass landscapes is typically ascribed to the direct influence of physical factors such as hydrodynamics, light, and sediment transport. We studied regularly interspaced banded patterns, formed by elongated patches of seagrass, in a small-scale intertidal ecosystem. We investigated (1) whether the observed spatial patterns may arise from feedback interactions between seagrass and its abiotic environment and (2) whether changes in abiotic conditions may lead to predictable changes in these spatial patterns. Field measurements, experiments, and a spatially explicit computer model identified a “scale-dependent feedback” (a mechanism for spatial self-organization) as a possible cause for the banded patterns. Increased protection from uprooting by improved anchoring with increasing seagrass density caused a local positive feedback. Sediment erosion around seagrass shoots increased with distance through the seagrass bands, hence causing a long-range negative feedback. Measurements across the depth gradient of the intertidal, together with model simulations, demonstrated that seagrass cover and mean patch size were predictably influenced by additional external stress caused by light limitation and desiccation. Thus, our study provides direct empirical evidence for a consistent response of spatial self-organized patterns to changing abiotic conditions, suggesting a potential use for self-organized spatial patterns as stress indicators in ecosystems., The spatial structure of seagrass landscapes is typically ascribed to the direct influence of physical factors such as hydrodynamics, light, and sediment transport. We studied regularly interspaced banded patterns, formed by elongated patches of seagrass, in a small-scale intertidal ecosystem. We investigated (1) whether the observed spatial patterns may arise from feedback interactions between seagrass and its abiotic environment and (2) whether changes in abiotic conditions may lead to predictable changes in these spatial patterns. Field measurements, experiments, and a spatially explicit computer model identified a “scale-dependent feedback” (a mechanism for spatial self-organization) as a possible cause for the banded patterns. Increased protection from uprooting by improved anchoring with increasing seagrass density caused a local positive feedback. Sediment erosion around seagrass shoots increased with distance through the seagrass bands, hence causing a long-range negative feedback. Measurements across the depth gradient of the intertidal, together with model simulations, demonstrated that seagrass cover and mean patch size were predictably influenced by additional external stress caused by light limitation and desiccation. Thus, our study provides direct empirical evidence for a consistent response of spatial self-organized patterns to changing abiotic conditions, suggesting a potential use for self-organized spatial patterns as stress indicators in ecosystems.