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. Expected final online publication date for the Annual Review of Marine Science Volume 7 is January 03, 2015. Please see http://www.annualreviews.org/catalog/pubdates.aspx for revised estimates.
Despite being a highly valuable key-stone ecosystem, seagrass meadows are threatened and declining worldwide, creating urgent need for indicators of their health status. We compared two indicators for seagrass health: standing leaf area index versus relative recovery from local disturbance. Disturbance was created by removing aboveground biomass and recording the rate of regrowth for Zostera marina meadows exposed to contrasting wave regimes and nutrient stress levels. Within the experimental period, relative regrowth in gaps was around 50% in most plots, except for the ambient nutrient treatment at the sheltered site, where it exceeded 100%. The two indicators showed an opposite response to disturbance: the higher the standing leaf area index, the lower the relative recovery from disturbance. This conflicting response raises the question on the proper interpretation of such indicators to estimate seagrass health and resilience, and how to ideally monitor seagrass ecosystems in order to predict collapse.
As a result of anthropogenic disturbances and natural stressors, seagrass beds are often patchy and heterogeneous. The effects of high loads of nutrients and organic matter in patch development and expansion in heterogeneous seagrass beds have, however, poorly been studied. We experimentally assessed the in situ effects of sediment quality on seagrass (Zostera noltii) patch dynamics by studying patch (0.35 m diameter) development and expansion for 4 sediment treatments: control, nutrient addition (NPK), organic matter addition (OM) and a combination (NPK+OM). OM addition strongly increased porewater sulfide concentrations whereas NPK increased porewater ammonium, nitrate and phosphate concentrations. As high nitrate concentrations suppressed sulfide production in NPK+OM, this treatment was biogeochemically comparable to NPK. Sulfide and ammonium concentrations differed within treatments, but over a 77 days period, seagrass patch survival and expansion were impaired by all additions compared to the control treatment. Expansion decreased at porewater ammonium concentrations >2,000 ?mol L(-1). Mother patch biomass was not affected by high porewater ammonium concentrations as a result of its detoxification by higher seagrass densities. Sulfide concentrations >1,000 ?mol L(-1) were toxic to both patch expansion and mother patch. We conclude that patch survival and expansion are constrained at high loads of nutrients or organic matter as a result of porewater ammonium or sulfide toxicity.
Seagrass beds are globally declining due to human activities in coastal areas. We here aimed to identify threats from eutrophication to the valuable seagrass beds of Curaçao and Bonaire in the Caribbean, which function as nursery habitats for commercial fish species. We documented surface- and porewater nutrient concentrations, and seagrass nutrient concentrations in 6 bays varying in nutrient loads. Water measurements only provided a momentary snapshot, due to timing, tidal stage, etc., but Thalassia testudinum nutrient concentrations indicated long-term nutrient loads. Nutrient levels in most bays did not raise any concern, but high leaf % P values of Thalassia in Piscadera Bay (?0.31%) and Spanish Water Bay (?0.21%) showed that seagrasses may be threatened by eutrophication, due to emergency overflow of waste water and coastal housing. We thus showed that seagrasses may be threatened and measures should be taken to prevent loss of these important nursery areas due to eutrophication.
Seagrass beds are highly productive coastal ecosystems providing a large array of ecosystem services including fisheries and carbon sequestration. As seagrasses are known to be highly sensitive to anthropogenic forcing, we evaluated the use of trace metal concentrations in seagrasses as bioindicators for trace metal pollution of coastal regions at both global and local scale. We carried out a meta-analysis based on literature data to provide a global benchmark list for trace metal accumulation in seagrasses, which was lacking in literature. We subsequently carried out a case study at the Caribbean islands of Curaçao and Bonaire to test for local-scale differences in trace metal concentrations in seagrasses, and internal metal allocation. The benchmark and local study show that trace metal concentrations in seagrass leaves, regardless of the species, can vary over a 100-1000-fold range, and are related to the level of anthropogenic pressure, making seagrasses highly valuable indicators.
When two ecosystem engineers share the same natural environment, the outcome of their interaction will be unclear if they have contrasting habitat-modifying effects (e.g., sediment stabilization vs. sediment destabilization). The outcome of the interaction may depend on local environmental conditions such as season or sediment type, which may affect the extent and type of habitat modification by the ecosystem engineers involved. We mechanistically studied the interaction between the sediment-stabilizing seagrass Zostera noltii and the bioturbating and sediment-destabilizing lugworm Arenicola marina, which sometimes co-occur for prolonged periods. We investigated (1) if the negative sediment destabilization effect of A. marina on Z. noltii might be counteracted by positive biogeochemical effects of bioirrigation (burrow flushing) by A. marina in sulfide-rich sediments, and (2) if previously observed nutrient release by A. marina bioirrigation could affect seagrasses. We tested the individual and combined effects of A. marina presence and high porewater sulfide concentrations (induced by organic matter addition) on seagrass biomass in a full factorial lab experiment. Contrary to our expectations, we did not find an effect of A. marina on porewater sulfide concentrations. A. marina activities affected the seagrass physically as well as by pumping nutrients, mainly ammonium and phosphate, from the porewater to the surface water, which promoted epiphyte growth on seagrass leaves in our experimental set-up. We conclude that A. marina bioirrigation did not alleviate sulfide stress to seagrasses. Instead, we found synergistic negative effects of the presence of A. marina and high sediment sulfide levels on seagrass biomass.
Due to increasing development Southeast Asia's coastlines are undergoing massive changes, but the associated impacts on marine habitats are poorly known. Singapore, a densely populated island city-state, is a quintessential example of coastal modification that has resulted in the (hitherto undocumented) loss of seagrass. We reconstructed the historic extent and diversity of local seagrass meadows through herbarium records and backwards extrapolation from contemporary seagrass locations. We also determined the current status of seagrass meadows using long-term monitoring data and identified the main threats to their presence in Singapore. Results show that, even though ?45% of seagrass has been lost during the last five decades, species diversity remains stable. The main cause of seagrass loss was, and continues to be, land reclamation. We conclude that strict controls on terrestrial runoff and pollution have made it possible for seagrass to persist adjacent to this highly urbanised city-state.
Ecosystems in the tropical coastal zone exchange particulate organic matter (POM) with adjacent systems, but differences in this function among ecosystems remain poorly quantified. Seagrass beds are often a relatively small section of this coastal zone, but have a potentially much larger ecological influence than suggested by their surface area. Using stable isotopes as tracers of oceanic, terrestrial, mangrove and seagrass sources, we investigated the origin of particulate organic matter in nine mangrove bays around the island of Phuket (Thailand). We used a linear mixing model based on bulk organic carbon, total nitrogen and ?13C and ?15N and found that oceanic sources dominated suspended particulate organic matter samples along the mangrove-seagrass-ocean gradient. Sediment trap samples showed contributions from four sources oceanic, mangrove forest/terrestrial and seagrass beds where oceanic had the strongest contribution and seagrass beds the smallest. Based on ecosystem area, however, the contribution of suspended particulate organic matter derived from seagrass beds was disproportionally high, relative to the entire area occupied by mangrove forests, the catchment area (terrestrial) and seagrass beds. The contribution from mangrove forests was approximately equal to their surface area, whereas terrestrial contributions to suspended organic matter under contributed compared to their relative catchment area. Interestingly, mangrove forest contribution at 0 m on the transects showed a positive relationship with the exposed frontal width of the mangrove, indicating that mangrove forest exposure to hydrodynamic energy may be a controlling factor in mangrove outwelling. However we found no relationship between seagrass bed contribution and any physical factors, which we measured. Our results indicate that although seagrass beds occupy a relatively small area of the coastal zone, their role in the export of organic matter is disproportional and should be considered in coastal management especially with respect to their importance as a nutrient source for other ecosystems and organisms.
Human infrastructures can modify ecosystems, thereby affecting the occurrence and spatial distribution of organisms, as well as ecosystem functionality. Sustainable development requires the ability to predict responses of species to anthropogenic pressures. We investigated the large scale, long term effect of important human alterations of benthic habitats with an integrated approach combining engineering and ecological modelling. We focused our analysis on the Oosterschelde basin (The Netherlands), which was partially embanked by a storm surge barrier (Oosterscheldekering, 1986). We made use of 1) a prognostic (numerical) environmental (hydrodynamic) model and 2) a novel application of quantile regression to Species Distribution Modeling (SDM) to simulate both the realized and potential (habitat suitability) abundance of four macrozoobenthic species: Scoloplos armiger, Peringia ulvae, Cerastoderma edule and Lanice conchilega. The analysis shows that part of the fluctuations in macrozoobenthic biomass stocks during the last decades is related to the effect of the coastal defense infrastructures on the basin morphology and hydrodynamics. The methodological framework we propose is particularly suitable for the analysis of large abundance datasets combined with high-resolution environmental data. Our analysis provides useful information on future changes in ecosystem functionality induced by human activities.
Physical habitat complexity regulates the structure and function of biological communities, although the mechanisms underlying this relationship remain unclear. Urbanisation, pollution, unsustainable resource exploitation and climate change have resulted in the widespread simplification (and loss) of habitats worldwide. One way to restore physical complexity to anthropogenically simplified habitats is through the use of artificial substrates, which also offer excellent opportunities to explore the effects of different components (variables) of complexity on biodiversity and community structure that would be difficult to separate in natural systems. Here, we describe a software program (CASU) that enables users to visualise static, physical complexity. CASU also provides output files that can be used to create artificial substrates for experimental and/or restoration studies. It has two different operational modes: simple and advanced. In simple mode, users can adjust the five main variables of informational complexity (i.e. the number of object types, relative abundance of object types, density of objects, variability and range in the objects' dimensions, and their spatial arrangement) and visualise the changes as they do so. The advanced mode allows users to design artificial substrates by fine-tuning the complexity variables as well as alter object-specific parameters. We illustrate how CASU can be used to create tiles of different designs for application in a marine environment. Such an ability to systematically influence physical complexity could greatly facilitate ecological restoration by allowing conservationists to rebuild complexity in degraded and simplified habitats.
Marine protected areas (MPAs) are key tools for combatting the global overexploitation 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.
Seagrasses have substantial capacity to survive long periods of light reduction, but how acclimation to chronic low light environments may influence their ability to cope with additional stress is poorly understood. This study examines the effect of temporal light reduction by adding two levels of shading to Halophila ovalis plants in two meadows with different light histories, one characterized by a low light (turbid) environment and the other by a relatively high light (clear) environment. Additional shading resulted in complete mortality for both shading treatments at the turbid site while the clear site showed a pattern of decreased shoot density and increased photochemical efficiency (Fv/Fm) with increased shading. These contrasting results for the same species in two different locations indicate that acclimation to chronic low light regimes can affect seagrass resilience and highlights the importance of light history in determining the outcome of exposure to further (short-term) stress.
The risk of flood disasters is increasing for many coastal societies owing to global and regional changes in climate conditions, sea-level rise, land subsidence and sediment supply. At the same time, in many locations, conventional coastal engineering solutions such as sea walls are increasingly challenged by these changes and their maintenance may become unsustainable. We argue that flood protection by ecosystem creation and restoration can provide a more sustainable, cost-effective and ecologically sound alternative to conventional coastal engineering and that, in suitable locations, it should be implemented globally and on a large scale.
1. Vegetated biogeomorphic systems (e.g. mangroves, salt marshes, dunes, riparian vegetation) have been intensively studied for the impact of the biota on sediment transport processes and the resulting self-organization of such landscapes. However, there is a lack of understanding of physical disturbance mechanisms that limit primary colonization in active sedimentary environments. 2. This study elucidates the effect of sediment disturbance during the seedling stage of pioneer vegetation, using mangroves as a model system. We performed mesocosm experiments that mimicked sediment disturbance as (i) accretion/burial of plants and (ii) erosion/excavation of plants of different magnitudes and temporal distribution in combination with water movement and inundation stress. 3. Cumulative sediment disturbance reduced seedling survival, with the faster-growing Avicennia alba showing less mortality than the slower-growing Sonneratia alba. The presence of the additional stressors (inundation and water movement) predominantly reduced the survival of S. alba. 4. Non-lethal accretion treatments increased shoot biomass of the seedlings, whereas non-lethal erosion treatments increased root biomass allocation. This morphological plasticity in combination with the abiotic disturbance history determined how much maximum erosion the seedlings were able to withstand. 5. Synthesis and applications. Seedling survival in dynamic sedimentary environments is determined by the frequency and magnitude of sediment accretion or erosion events, with non-lethal events causing feedbacks to seedling stability. Managers attempting restoration of mangroves, salt marshes, dunes and riparian vegetation should recognize sediment dynamics as a main bottleneck to primary colonization. The temporal distribution of erosion and accretion events has to be evaluated against the ability of the seedlings to outgrow or adjust to disturbances. Our results suggest that selecting fast-growing pioneer species and measures to enhance seedling growth or temporary reduction in sediment dynamics at the restoration site can aid restoration success for vegetated biogeomorphic ecosystems.
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.
Intertidal wetlands such as saltmarshes and mangroves provide numerous important ecological functions, though they are in rapid and global decline. To better conserve and restore these wetland ecosystems, we need an understanding of the fundamental natural bottlenecks and thresholds to their establishment and long-term ecological maintenance. Despite inhabiting similar intertidal positions, the biological traits of these systems differ markedly in structure, phenology, life history, phylogeny and dispersal, suggesting large differences in biophysical interactions. By providing the first systematic comparison between saltmarshes and mangroves, we unravel how the interplay between species-specific life-history traits, biophysical interactions and biogeomorphological feedback processes determine where, when and what wetland can establish, the thresholds to long-term ecosystem stability, and constraints to genetic connectivity between intertidal wetland populations at the landscape level. To understand these process interactions, research into the constraints to wetland development, and biological adaptations to overcome these critical bottlenecks and thresholds requires a truly interdisciplinary approach.
External mechanical forces resulting from the pressure exerted by wind or water movement are a major stress factor for plants and may cause regular disturbances in many ecosystems. A plants ability to resist these forces relies either on minimizing the forces encountered by the plant (avoidance strategy), or on maximizing its resistance to breakage (tolerance strategy). We investigated plant resistance strategies using aquatic vegetation as a model, and examined whether avoidance and tolerance are negatively correlated. We tested the avoidance-tolerance correlation across 28 species using a phylogenetically corrected analysis, after construction of a molecular phylogeny for the species considered. Different species demonstrated contrasting avoidance and tolerance and we demonstrated a significant negative relationship between the two strategies, which suggests an avoidance-tolerance trade-off. Negative relationships may result from costs that each strategy incurs or from constraints imposed by physical laws on plant tissues. The existence of such a trade-off has important ecological and evolutionary consequences. It would lead to constraints on the evolution and variation of both strategies, possibly limiting their evolution and may constrain many morphological, anatomical and architectural traits that underlie avoidance and tolerance.
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.
In a seagrass restoration project, we explored the potential for enhancing the restoration process by excluding antagonistic engineering interactions (i.e., biomechanical warfare) between two ecosystem engineers: the bioturbating lugworm Arenicola marina and the sediment-stabilizing seagrass Zostera noltii Hornem. Applying a shell layer underneath half of our seagrass transplants successfully reduced adult lugworm density by over 80% and reduced lugworm-induced microtopography (a proxy for lugworm disturbance) at the wave-sheltered site. At the wave-exposed site adult lugworm densities and microtopography were already lower than at the sheltered site but were further reduced in the shell-treated units. Excluding lugworms and their bioengineering effects corresponded well with a strongly enhanced seagrass growth at the wave-sheltered site, which was absent at the exposed site. Enhanced seagrass growth in the present study was fully assigned to the removal of lugworms negative engineering effects and not to any (indirect) evolving effects such as an altered biogeochemistry or sediment-stabilizing effects by the shell layer. The context-dependency implies that seagrass establishment at the exposed site is not constrained by negative ecosystem-engineering interactions only, but also by overriding physical stresses causing poor growth conditions. Present findings underline that, in addition to recent emphasis on considering positive (facilitating) interactions in ecological theory and practice, it is equally important to consider negative engineering interactions between ecosystem-engineering species. Removal of such negative interactions between ecosystem-engineering species can give a head start to the target species at the initial establishment phase, when positive engineering feedbacks by the target species on itself are still lacking. Though our study was carried out in a marine environment with variable levels of wave disturbance, similar principles may be expected to apply to other ecosystems that are inhabited by ecosystem engineers.
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.
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