Sediment destabilizing and stabilizing bio-engineers on tidal flats: cascading effects of experimental exclusion

Bioturbating lugworms (Arenicola marina) were excluded from 400 m2 plots of intertidal sand which initiated sequences of direct and indirect changes in the structure of the benthic community. The sessile, tube-building species Polydora cornuta and Lanice conchilega took advantage of the absence of lugworms and settled preferentially on lugworm exclusion plots. The protruding tubes provided attachment for an ephemeral development of algal tufts (Berkeleya colonies and Enteromorpha thalli) which in turn enhanced settlement of the juvenile drifting clams Mya arenaria and Macoma balthica. This causal chain of enhanced bivalve settlement in the presence of above-ground structures, like animal tubes and algae, on lugworm exclusion plots occurred in 2 years at different tidal zones with different tube builders, algae and juvenile clams. A significant response of L. conchilega in a year with relatively low lugworm abundances at the entire site suggests that not only the actual absence of large bioturbators was responsible for the establishment of tube-dwelling species, but also a cumulative change of the sediment in exclusion plots since the onset of the experiment. While the sediment on lugworm plots remained permeable, fine particles and organic matter accumulated at exclusion plots. It is suggested that these differences in sediment characteristics were the product of divergent benthic engineering by sediment destabilizing lugworms on control plots and sediment stabilizing species on exclusion plots. Cumulative changes of the sedimentary habitat and cascading effects in the benthic community may explain the persistence of patches that are dominated by either sediment stabilizing or destabilizing species in the assemblage mosaic of intertidal sediments.     

Non-trophic Interactions Control Benthic Producers on Intertidal Flats

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.     

Habitat-mediated facilitation and counteracting ecosystem engineering interactively influence ecosystem responses to disturbance

Recovery of an ecosystem following disturbance can be severely hampered or even shift altogether when a point disturbance exceeds a certain spatial threshold. Such scale-dependent dynamics may be caused by preemptive competition, but may also result from diminished self-facilitation due to weakened ecosystem engineering. Moreover, disturbance can facilitate colonization by engineering species that alter abiotic conditions in ways that exacerbate stress on the original species. Consequently, establishment of such counteracting engineers might reduce the spatial threshold for the disturbance, by effectively slowing recovery and increasing the risk for ecosystem shifts to alternative states. We tested these predictions in an intertidal mudflat characterized by a two-state mosaic of hummocks (humps exposed during low tide) dominated by the sediment-stabilizing seagrass Zostera noltii) and hollows (low-tide waterlogged depressions dominated by the bioturbating lugworm Arenicola marina). In contrast to expectations, seagrass recolonized both natural and experimental clearings via lateral expansion and seemed unaffected by both clearing size and lugworm addition. Near the end of the growth season, however, an additional disturbance (most likely waterfowl grazing and/or strong hydrodynamics) selectively impacted recolonizing seagrass in the largest (1 m(2)) clearings (regardless of lugworm addition), and in those medium (0.25 m(2)) clearings where lugworms had been added nearly five months earlier. Further analyses showed that the risk for the disturbance increased with hollow size, with a threshold of 0.24 m(2). Hollows of that size were caused by seagrass removal alone in the largest clearings, and by a weaker seagrass removal effect exacerbated by lugworm bioturbation in the medium clearings. Consequently, a sufficiently large disturbance increased the vulnerability of recolonizing seagrass to additional disturbance by weakening seagrass engineering effects (sediment stabilization). Meanwhile, the counteracting ecosystem engineering (lugworm bioturbation) reduced that threshold size. Therefore, scale-dependent interactions between habitat-mediated facilitation, competition and disturbance seem to maintain the spatial two-state mosaic in this ecosystem.     

Multi‐scale habitat modification by coexisting ecosystem engineers drives spatial separation of macrobenthic functional groups

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.     

Habitat modification inhibits conspecific seedling recruitment in populations of an invasive ecosystem engineer

Invasive species that strongly modify their physical habitat are a particular management concern. Theoretical models predict that habitat modification could speed spread rates or allow invasion of sites that would otherwise resist invasion. There are few empirical tests of this hypothesis, however. We tested whether habitat modification by invading Spartina alterniflora populations facilitates conspecific seedling recruitment and spatial spread in Willapa Bay, WA, USA. Established S. alterniflora individuals strongly modified their local physical environment. Hydrologic flow, porewater salinity, and light availability were decreased while sediment NH₄ ⁺ increased with increasing S. alterniflora stem density. The S. alterniflora seed bank was greater and spring seedlings were denser within meadows of S. alterniflora than on unvegetated tideflats. However, almost all seedling recruitment after 1 year occurred on tideflats or on meadow edge plots where the above ground S. alterniflora biomass had been removed. Instead of facilitating invasive spread, ecosystem engineering in this system appears to create conditions that inhibit local seedling recruitment. These results suggest that the influence of ecosystem engineering on invasive spread is highly contingent on the relative spatial scales of habitat modification, environmental heterogeneity, and propagule availability. Control activities could change these spatial relationships, however, inadvertently promoting invasive recruitment.     

Interactive effects of porewater nutrient enrichment, bioturbation and sediment characteristics on benthic assemblages in sandy sediments

Effective management of eutrophic ecosystems requires an understanding of how nutrient input affects the structure and function of benthic communities. The effects of nutrients in soft sediment habitats can be influenced by a variety of factors including sediment characteristics, hydrodynamic exposure, and the presence of bioturbating macroinvertebrates. We used a large scale exclusion experiment (400 m² areas, n = 6) to test if bioturbating lugworms, Arenicola marina mediate the effects of nutrient enrichment. We incorporated small plots (30 × 30 cm) dosed with household garden fertilizer within the lugworm exclusion and corresponding control areas and predicted that the effects of nutrient enrichment would be greater in the absence of lugworms. We found that the increases in nutrient concentrations were higher in the absence of lugworms, but only in the less permeable sediment in the low intertidal zone compared to the more permeable sediment in the high intertidal. Contrary to expectations, the accumulation of nutrients in the plots did not affect the organic matter and chlorophyll levels in the sediment. Interestingly, there were overall negative effects of nutrient additions on some of the most abundant molluscs, Hydrobia ulvae, Retusa obtusa and juvenile Cerastoderma edule. Possible explanations for these adverse effects such as the changes in the sediment chemistry or the physical presence of the fertilizer in the sediment caused by the nutrient additions are discussed. We conclude that the effects of nutrient enrichment in soft sediment habitats on benthic assemblages are determined by the interplay between the presence of bioturbating macroinvertebrates, tidal height and sediment characteristics.     

Seagrasses are negatively affected by organic matter loading and Arenicola marina activity in a laboratory experiment

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.     

Ecosystem engineering by seagrasses interacts with grazing to shape an intertidal landscape

Self-facilitation through ecosystem engineering (i.e., organism modification of the abiotic environment) and consumer-resource interactions are both major determinants of spatial patchiness in ecosystems. However, interactive effects of these two mechanisms on spatial complexity have not been extensively studied. We investigated the mechanisms underlying a spatial mosaic of low-tide exposed hummocks and waterlogged hollows on an intertidal mudflat in the Wadden Sea dominated by the seagrass Zostera noltii. A combination of field measurements, an experiment and a spatially explicit model indicated that the mosaic resulted from localized sediment accretion by seagrass followed by selective waterfowl grazing. Hollows were bare in winter, but were rapidly colonized by seagrass during the growth season. Colonized hollows were heavily grazed by brent geese and widgeon in autumn, converting these patches to a bare state again and disrupting sediment accretion by seagrass. In contrast, hummocks were covered by seagrass throughout the year and were rarely grazed, most likely because the waterfowl were not able to employ their preferred but water requiring feeding strategy ('dabbling') here. Our study exemplifies that interactions between ecosystem engineering by a foundation species (seagrass) and consumption (waterfowl grazing) can increase spatial complexity at the landscape level.     

An ecosystem engineer,the beaver,increases species richness at the landscape scale

Ecosystem engineering - the physical modification of habitats by organisms - has been proposed as an important mechanism for maintaining high species richness at the landscape scale by increasing habitat heterogeneity. Dams built by beaver (Castor canadensis) dramatically alter riparian landscapes throughout much of North America. In the central Adirondacks, New York, USA, ecosystem engineering by beaver leads to the formation of extensive wetland habitat capable of supporting herbaceous plant species not found elsewhere in the riparian zone. We show that by increasing habitat heterogeneity, beaver increase the number of species of herbaceous plants in the riparian zone by over 33% at a scale that encompasses both beaver-modified patches and patches with no history of beaver occupation. We suggest that ecosystem engineers will increase species richness at the landscape scale whenever there are species present in a landscape that are restricted to engineered habitats during at least some stages of their life cycle.     

Facilitation and Competition among Invasive Plants: A Field Experiment with Alligatorweed and Water Hyacinth

Ecosystems that are heavily invaded by an exotic species often contain abundant populations of other invasive species. This may reflect shared responses to a common factor, but may also reflect positive interactions among these exotic species. Armand Bayou (Pasadena, TX) is one such ecosystem where multiple species of invasive aquatic plants are common. We used this system to investigate whether presence of one exotic species made subsequent invasions by other exotic species more likely, less likely, or if it had no effect. We performed an experiment in which we selectively removed exotic rooted and/or floating aquatic plant species and tracked subsequent colonization and growth of native and invasive species. This allowed us to quantify how presence or absence of one plant functional group influenced the likelihood of successful invasion by members of the other functional group. We found that presence of alligatorweed (rooted plant) decreased establishment of new water hyacinth (free-floating plant) patches but increased growth of hyacinth in established patches, with an overall net positive effect on success of water hyacinth. Water hyacinth presence had no effect on establishment of alligatorweed but decreased growth of existing alligatorweed patches, with an overall net negative effect on success of alligatorweed. Moreover, observational data showed positive correlations between hyacinth and alligatorweed with hyacinth, on average, more abundant. The negative effect of hyacinth on alligatorweed growth implies competition, not strong mutual facilitation (invasional meltdown), is occurring in this system. Removal of hyacinth may increase alligatorweed invasion through release from competition. However, removal of alligatorweed may have more complex effects on hyacinth patch dynamics because there were strong opposing effects on establishment versus growth. The mix of positive and negative interactions between floating and rooted aquatic plants may influence local population dynamics of each group and thus overall invasion pressure in this watershed.     

When does ecosystem engineering cause invasion and species replacement?

Introduced exotic species can dominate communities and replace native species that should be better adapted to their local environment, a paradox that is usually explained by the absence of natural enemies and by habitat alteration resulting from anthropogenic disturbance. Additionally, introduced species can enhance their invasion success and impact on native species by modifying selection pressures in their new environment through ecosystem engineering. We analyse a simple dynamic model of indirect competition for habitat between a non-engineering resident species and an engineering exotic species. The conditions for invasion and competitive exclusion of the resident by the exotic species and the range of dynamic outcomes suggested by the model are determined by the form of density dependence. We give simple criteria for the success of the invading species on dimensionless quantities involving rates of ecosystem engineering and of habitat degradation. The model's predictions offer an additional explanation for a range of invasion dynamics reported in the literature, including lag times between introduction and establishment. One intriguing result is that a series of failed invasions may successively reduce environmental resistance to subsequent invasion, through a cumulative effect of habitat transformation. More work is needed to determine the frequency and conditions in which engineering is required for successful establishment, and whether highly-successful (or high-impact) invaders are more likely to possess ecosystem engineering traits.     

Species effects on ecosystem processes are modified by faunal responses to habitat composition

Heterogeneity is a well-recognized feature of natural environments, and the spatial distribution and movement of individual species is primarily driven by resource requirements. In laboratory experiments designed to explore how different species drive ecosystem processes, such as nutrient release, habitat heterogeneity is often seen as something which must be rigorously controlled for. Most small experimental systems are therefore spatially homogeneous, and the link between environmental heterogeneity and its effects on the redistribution of individuals and species, and on ecosystem processes, has not been fully explored. In this paper, we used a mesocosm system to investigate the relationship between habitat composition, species movement and sediment nutrient release for each of four functionally contrasting species of marine benthic invertebrate macrofauna. For each species, various habitat configurations were generated by selectively enriching patches of sediment with macroalgae, a natural source of spatial variability in intertidal mudflats. We found that the direction and extent of faunal movement between patches differs with species identity, density and habitat composition. Combinations of these factors lead to concomitant changes in nutrient release, such that habitat composition effects are modified by species identity (in the case of NH4-N) and by species density (in the case of PO4-P). It is clear that failure to accommodate natural patterns of spatial heterogeneity in such studies may result in an incomplete understanding of system behaviour. This will be particularly important for future experiments designed to explore the effects of species richness on ecosystem processes, where the complex interactions reported here for single species may be compounded when species are brought together in multi-species combinations.     

Complementary influences of co-occurring physical ecosystem engineers on species richness: insights from a Patagonian rocky shore

Structural modification of the environment by physical ecosystem engineers often allows for the occurrence of species that are not able to establish in unengineered habitats, thus leading to increased species richness at the landscape-level (i.e., areas encompassing engineered and unengineered habitats). Unlike previous studies that focused on the contribution of a single engineering species to landscape-level species richness, this study evaluates whether co-occurring engineers—i.e., intertidal mussels (primarily Perumytilus purpuratus) and rock boring bivalves (Lithophaga patagonica)—contribute to landscape-level species richness in a similar or complementary way. Our results show that both mussel and L. patagonica patches harbor a substantial number of invertebrate species in addition to those occurring in the unenegineered rock substrate. However, the distinctive habitat patches created by each engineer add exclusive subsets of species to the study area, which implies that mussel and L. patagonica patches contribute complementarily to overall species richness in our intertidal landscape. Here we postulate that complementary engineering effects on landscape-level species richness will occur when the engineered patches structurally differ from each other and, thus, vary in their relative ability to modulate two or more abiotic conditions and/or resources that prevent species establishment in the unengineered state. In spite of its inherently small spatial scale (500 m), our study highlights the potential for complementary engineering impacts at the larger scales that are usually implied in biodiversity conservation and management (tens to hundreds of kilometers) and outlines a simple conceptual basis and approach to address them.     

Remedial habitat creation: does Nereis diversicolor play a confounding role in the colonisation and establishment of the pioneering saltmarsh plant, Spartina anglica?

Increasing concerns over global warming and expected sea level rises have led to the adoption of new coastal management strategies around the south-east coast of England. This paper explores the role played by the estuarine invertebrate Nereis diversicolor in limiting the colonisation and establishment of the invasive pioneering salt marsh plant, Spartina anglica. The biology of N. diversicolor is briefly reviewed and data from field experiments are presented demonstrating significant negative effects of worm abundance on transplanted S. anglica biomass. Laboratory-based experiments demonstrated significant negative effects of N. diversicolor abundance on the survival of S. anglica seeds transplanted to sediment cores. The importance of estuarine invertebrates in engineering the mudflat habitat may confound the foreseen ecosystem services and function provided by saltmarsh management schemes. Received: 15 February 1999 / Received in revised form: 4 June 1999 / Accepted: 11 June 1999     

Ecosystem engineering and biodiversity in coastal sediments: posing hypotheses

Coastal sediments in sheltered temperate locations are strongly modified by ecosystem engineering species such as marsh plants, seagrass, and algae as well as by epibenthic and endobenthic invertebrates. These ecosystem engineers are shaping the coastal sea and landscape, control particulate and dissolved material fluxes between the land and sea, and between the benthos and the passing water or air. Above all, habitat engineering exerts facilitating and inhibiting effects on biodiversity. Despite a strongly growing interest in the functional role of ecosystem engineering over the recent years, compared to food web analyses, the conceptual understanding of engineering-mediated species interactions is still in its infancy. In the present paper, we provide a concise overview on current insights and propose two hypotheses on the general mechanisms by which ecosystem engineering may affect biodiversity in coastal sediments. We hypothesise that autogenic and allogenic ecosystem engineers have inverse effects on epibenthic and endobenthic biodiversity in coastal sediments. The primarily autogenic structures of the epibenthos achieve high diversity at the expense of endobenthos, whilst allogenic sediment reworking by infauna may facilitate other infauna and inhibits epibenthos. On a larger scale, these antagonistic processes generate patchiness and habitat diversity. Due to such interaction, anthropogenic influences can strongly modify the engineering community by removing autogenic ecosystem engineers through coastal engineering or bottom trawling. Another source of anthropogenic influences comes from introducing invasive engineers, from which the impact is often hard to predict. We hypothesise that the local biodiversity effects of invasive ecosystem engineers will depend on the engineering strength of the invasive species, with engineering strength defined as the number of habitats it can invade and the extent of modification. At a larger scale of an entire shore, biodiversity need not be decreased by invasive engineers and may even increase. On a global scale, invasive engineers may cause shore biota to converge, especially visually due to the presence of epibenthic structures.     

Benefits and limitations of isolated floral patches in a pollinator restoration project in Arizona

This study examined invertebrate floral visitor responses to floral richness, floral abundance, and distance between floral patches within a newly planted pollinator restoration habitat in an arid ecosystem in central Arizona, United States. We created a pollinator habitat experiment consisting of a large central garden (11‐m diameter) surrounded by concentric rings of smaller habitat patches (1‐m diameter), separated from one another by 1, 8, 13, and 21 m, respectively, and including four flowering species. We observed plant and visitor interactions via structured 10‐minute flower visitation observations over a 3‐month period. Key findings included: (1) each plant species interacted with a variety of flower visitors, but flower visitor groups differed only marginally among the plant species; (2) floral patches outside the central garden exhibited reduced quantities of floral structures; and (3) number of floral structures per patch, but not isolation of floral patches within the habitat, affected the number of visitors and visitor taxa richness. For practitioners and land managers looking to restore pollination systems in arid ecosystems with low establishment via seeding, the results of this study suggest that installing species‐rich and florally abundant patches of flowering plant species within a habitat could efficiently support plant‐pollinator interactions.     

Complex Effects of Ecosystem Engineer Loss on Benthic Ecosystem Response to Detrital Macroalgae

Ecosystem engineers change abiotic conditions, community assembly and ecosystem functioning. Consequently, their loss may modify thresholds of ecosystem response to disturbance and undermine ecosystem stability. This study investigates how loss of the bioturbating lugworm Arenicola marina modifies the response to macroalgal detrital enrichment of sediment biogeochemical properties, microphytobenthos and macrofauna assemblages. A field manipulative experiment was done on an intertidal sandflat (Oosterschelde estuary, The Netherlands). Lugworms were deliberately excluded from 1× m sediment plots and different amounts of detrital Ulva (0, 200 or 600 g Wet Weight) were added twice. Sediment biogeochemistry changes were evaluated through benthic respiration, sediment organic carbon content and porewater inorganic carbon as well as detrital macroalgae remaining in the sediment one month after enrichment. Microalgal biomass and macrofauna composition were measured at the same time. Macroalgal carbon mineralization and transfer to the benthic consumers were also investigated during decomposition at low enrichment level (200 g WW). The interaction between lugworm exclusion and detrital enrichment did not modify sediment organic carbon or benthic respiration. Weak but significant changes were instead found for porewater inorganic carbon and microalgal biomass. Lugworm exclusion caused an increase of porewater carbon and a decrease of microalgal biomass, while detrital enrichment drove these values back to values typical of lugworm-dominated sediments. Lugworm exclusion also decreased the amount of macroalgae remaining into the sediment and accelerated detrital carbon mineralization and CO₂ release to the water column. Eventually, the interaction between lugworm exclusion and detrital enrichment affected macrofauna abundance and diversity, which collapsed at high level of enrichment only when the lugworms were present. This study reveals that in nature the role of this ecosystem engineer may be variable and sometimes have no or even negative effects on stability, conversely to what it should be expected based on current research knowledge.     

Context dependence of marine ecosystem engineer invasion impacts on benthic ecosystem functioning

Introduced ecosystem engineers can severely modify the functioning on invaded systems. Species-level effects on ecosystem functioning (EF) are context dependent, but the effects of introduced ecosystem engineers are frequently assessed through single-location studies. The present work aimed to identify sources of context-dependence that can regulate the impacts of invasive ecosystem engineers on ecosystem functioning. As model systems, four locations where the bivalve Ruditapes philippinarum (Adams and Reeve) has been introduced were investigated, providing variability in habitat characteristics and community composition. As a measure of ecosystem engineering, the relative contribution of this species to community bioturbation potential was quantified at each site. The relevance of bioturbation to the local establishment of the mixing depth of marine sediments (used as a proxy for EF) was quantified in order to determine the potential for impact of the introduced species at each site. We found that R. philippinarum is one of the most important bioturbators within analysed communities, but the relative importance of this contribution at the community level depended on local species composition. The net contribution of bioturbation to the establishment of sediment mixing depths varied across sites depending on the presence of structuring vegetation, sediment granulometry and compaction. The effects of vegetation on sediment mixing were previously unreported. These findings indicate that the species composition of invaded communities, and the habitat characteristics of invaded systems, are important modulators of the impacts of introduced species on ecosystem functioning. A framework that encompasses these aspects for the prediction of the functional impacts of invasive ecosystem engineers is suggested, supporting a multi-site approach to invasive ecology studies concerned with ecosystem functioning.     

Sewage,green algal mats anchored by lugworms,and the effects on Turbellaria and small Polychaeta

On sandy tidal flats at the Island of Sylt (North Sea) ephemeral mats of green algae covered wide areas in the vicinity of sewage outflows. Algae became anchored in the feeding funnels of lugworms (Arenicola marina) and thus were able to resist displacement by tidal currents. Below the algal mats anoxic conditions extend to the sediment surface. After about one month a rough sea removed all algae. Polychaetes endured this short-term environmental deterioration, while the more sensitive Turbellaria decreased in abundance and species richness. Diatom-feeders were affected most, predators to a medium extent, and bacteria-feeders the least affected. Rare and very abundant species were more affected than moderately abundant ones. None of the turbellarian species increased in abundance and none colonized the algal mats above the sediment. In a semicontrolled experiment with daily hand-removal of drift algae from a 100-m² plot within an extensive field of algal mats, this cleaned "island" served as a refuge to Turbellaria escaping from their algal covered habitat. Here abundance doubled relative to initial conditions and was 5-times higher than below algal mats.     

Influence of an ecosystem engineer,the emergent macrophyte Sparganium erectum,on seed trapping in lowland rivers and consequences for landform colonisation

1. Plant physical ecosystem engineers can influence vegetation population and community dynamics by modifying, maintaining or creating habitats. They may also have the potential to act upon biotic processes, such as seed dispersal. 2. Examples exist of reduction in seed dispersal distances in vegetated compared to unvegetated terrestrial environments, and concentration of seed deposits associated with plant patches. Such effects in aquatic environments have been little studied, but the engineering effect of plant patches on patterns of flow velocity and sediment deposition in streams suggests that they may play a similar role. 3. In this study, we assess the potential of an emergent aquatic species, Sparganium erectum, to play a role in physically modifying river habitats and trapping seeds by examining patterns of seed deposition and substrate type in 47 river reaches across England and southern Scotland, U.K. 4. Areas of the river channel within or adjacent to S. erectum patches harboured more plant seeds and more species than unvegetated areas and had finer, sandier substrates with higher organic matter, total nitrogen and total phosphorus content. Most seed species were competitive, indicating that they were well suited to colonise the competitive environment of an S. erectum patch, and could potentially further stabilise accumulated sediments and contribute to landform development. 5. We demonstrate that S. erectum patches influence both the physical environment and the retention of seeds, in consistent patterns across the channel bed, for a range of lowland rivers that vary in stream power and geology and which can be expected to vary in levels of supply of fine sediment and seeds. 6. Our findings support the hypothesis that the fundamental influence of a riverine ecosystem‐engineering species on slowing fluid flow links the habitat creation process of sediment sorting and retention to seed trapping. We suggest the process is applicable to a wide range of aquatic and riparian vegetation. We also suggest that the mono‐specific and competitive growth, which is typical of these engineering species, will strongly influence the recruitment of trapped seeds.