Habitat persistence for sedentary organisms in managed rivers: the case for the federally endangered dwarf wedgemussel (Alasmidonta heterodon) in the Delaware River

1. To manage the environmental flow requirements of sedentary taxa, such as mussels and aquatic insects with fixed retreats, we need a measure of habitat availability over a variety of flows (i.e. a measure of persistent habitat). Habitat suitability measures in current environmental flow assessments are measured on a ‘flow by flow’ basis and thus are not appropriate for these taxa. Here, we present a novel measure of persistent habitat suitability for the dwarf wedgemussel (Alasmidonta heterodon), listed as federally endangered in the U.S.A., in three reaches of the Delaware River. 2. We used a two‐dimensional hydrodynamic model to quantify suitable habitat over a range of flows based on modelled depth, velocity, Froude number, shear velocity and shear stress at three scales (individual mussel, mussel bed and reach). Baseline potentially persistent habitat was quantified as the sum of pixels that met all thresholds identified for these variables for flows ≥40 m³ s^?1, and we calculated the loss of persistently suitable habitat by sequentially summing suitable habitat estimates at lower flows. We estimated the proportion of mussel beds exposed at each flow and the amount of change in the size of the mussel bed for one reach. 3. For two reaches, mussel beds occupied areas with lower velocity, shear velocity, shear stress and Froude number than the reach average at all flows. In the third reach, this was true only at higher flows. Together, these results indicate that beds were possible refuge areas from the effects of these hydrological parameters. Two reaches showed an increase in the amount of exposed mussel beds with decreasing flow. 4. Baseline potentially persistent habitat was less than half the areal extent of potentially suitable habitat, and it decreased with decreasing flow. Actually identified beds and modelled persistent habitat showed good spatial overlap, but identified beds occupied only a portion of the total modelled persistent habitat, indicating either that additional suitable habitat is available or the need to improve habitat criteria. At one site, persistent beds (beds where mussels were routinely collected) were located at sites with stable substratum, whereas marginal beds (beds where mussels were infrequently collected or that were lost following a large flood event) were located in scoured areas. 5. Taken together, these model results support a multifaceted approach, which incorporates the effects of low and high flow stressors, to quantify habitat suitability for mussels and other sedentary taxa. Models of persistent habitat can provide a more holistic environmental flow assessment of rivers.     

Impacts of sea star predation on mussel bed restoration

Predation by sea stars has the potential to cause elevated levels of mortality in reestablished populations of bivalves relative to levels of recruitment. Recent efforts to restore beds of the nearly extirpated green‐lipped mussel within the Hauraki Gulf, New Zealand, resulted in high abundances of sea stars occupying those beds and it is unknown whether predation poses a potential limitation to the success of restoration in this bivalve species. The contribution of predation by sea stars to the mortality of mussels across four experimental mussel beds over a 2‐year period was estimated using data from regular assessments of sea star abundance and an experimentally determined consumption rate of sea stars upon mussels. In addition, the potential size selection of mussels by sea stars was assessed to determine if large sea stars selected for recent settlers. Sea stars' abundance within the mussel beds grew to an average of 0.57 sea stars/m² within 9 months, remaining at similar levels throughout the study. These predominantly large sea stars were estimated to have consumed 30.1% of the mussels over a 25‐month period, representing a contribution of 40.4% of the mussel mortality. Sea stars predominantly selected for larger mussels, and their predation likely contributes little to any lack in mussel recruitment. Sea star predation is clearly a limiting factor on the survival of transplanted adult mussels and the present study highlights the need to continually assess predation risk to determine if remediation is necessary for the persistence of the restored beds.     

Modeling the influence of a young mussel bed on fine sediment dynamics on an intertidal flat in the Wadden Sea

Mussel beds are known to affect fine sediment dynamics and morphology on mudflat scale, a clear example of ecosystem engineering. Current research into possible ecological engineering applications of mussel beds makes quantitative modeling desirable. In this study a process-based model of the interaction between a young mussel bed and fine sediment was set up for use in the hydrodynamic and morphological model Delft3D-FLOW. The model encompasses the hydraulic roughness of the mussel bed, active capture of suspended sediment by filter feeding and changed bed properties due to biodeposited matter. The mussel bed implementation in Delft3D-FLOW was applied in a test case: a Wadden Sea intertidal mudflat model. It was concluded that a combination of active deposition via filtration and slow down of the flow due to increased roughness leads to high net deposition in the mussel bed. In addition, the ability of young mussels to quickly climb on top of deposited material results in rapid trapping of large amounts of fine sediment. In the wake of the mussel bed, deposition is also high because of reduced flow velocities. The effects of different existing mussel bed patterns were also evaluated. Patchiness and specifically striped patterns cause mussel beds to experience less sedimentation than uniformly covered beds of the same size and may therefore be favorable to mussels.     

Distribution, abundance, size and recruitment of the mussel, Mytilus californianus, across a major oceanographic and biogeographic boundary at Point Conception, California, USA

The Point Conception, California, USA region (hereafter PC) is one of the most important biogeographic and oceanographic discontinuities on the US west coast. Here we address how mesoscale oceanographic and environmental variability in the region around PC, CA may influence the distribution, abundance and size of the mussel Mytilus californianus, a competitively dominant species in rocky intertidal assemblages along the northeast Pacific. Strong upwelling and high wave exposure dominate the California coast north of PC, and weak, seasonal upwelling and warmer water temperatures are characteristic of the region south/east of PC. We hypothesized that the gradient in temperature, upwelling, and wave exposure around PC would greatly influence patterns of recruitment and abundance of mussels, potentially underlying large-scale differences in community structure. We evaluated these predictions by surveying intertidal community structure, mussel distribution, size, abundance and recruitment at a range of intertidal sites around PC. We found that intertidal communities north of PC were dominated mainly by macrophytes, while mussels and barnacles were relatively scarce. Intertidal communities south of PC were dominated by mussels and barnacles, with a low abundance of macrophytes. Mussels were larger and mussel beds were more expansive and extended lower in elevation at sites ranging from north to south around PC. At northern sites, high abundances of sea star predators and elevated wave exposure effectively displaced the entire mussel zone upwards. We found no differences in the numbers of mussel recruits to sites around PC, suggesting that spatial patterns of mussel abundance were not driven by differential recruitment. These results suggest that unlike other well-studied systems, supply of benthic larvae does not underly the large-scale gradient in community structure around PC. We suggest that environmental conditions favorable to macroalgal growth north of PC, and conditions favorable to filter-feeder growth south of PC may underly mesoscale patterns of intertidal community structure in this region.     

Effects of environmental stress on intertidal mussels and their sea star predators

Consumer stress models of ecological theory predict that predators are more susceptible to stress than their prey. Intertidal mussels, Mytilus californianus, span a vertical stress gradient from the low zone (lower stress) to the high zone (higher thermal and desiccation stress), while their sea star predators, Pisaster ochraceus, range from the low zone only into the lower edge of the mussel zone. In summer 2003, we tested the responses of sea stars and mussels to environmental stress in an experiment conducted on the Oregon coast. Mussels were transplanted from the middle of the mussel bed to cages in the low and high edges of the mussel bed. Sea star predators were added to half of the mussel cages. Mussels and sea stars were sampled between June and August for indicators of sublethal stress. Mussel growth was measured, and tissues were collected for heat shock protein (Hsp70) analyses and histological analyses of reproduction. Sea stars were weighed, and tissues were sampled for Hsp70 analyses. Mussels in high-edge cages had higher levels of total Hsp70 and exhibited spawning activity earlier in the summer than mussels in the low-edge cages. Sea stars suffered high mortality in the high edge, and low-edge sea stars lost weight but showed no differences in Hsp70 production. These results suggest that stress in the intertidal zone affected the mobile predator more than its sessile prey, which is consistent with predictions of consumer stress models. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Impact scenario for the invasive red king crab <Emphasis Type="Italic">Paralithodes camtschaticus</Emphasis> (Tilesius, 1815) (Reptantia,Lithodidae) on Norwegian,native, epibenthic prey

Large invasive predators like the king crab, Paralithodes camtschaticus, deserve particular attention due to their potential for catastrophic ecological impact on recipient communities. Conspicuous, epibenthic prey species, such as the slow growing commercial scallop Chlamys islandica, are particularly exposed to the risk of local extinction. A research program integrating experiments and field monitoring is attempting to predict and track the impact of invasive king crab on scallop beds and associated fauna along the north Norwegian coast. The claw gape of the crab shows no limitations in handling the flat-bodied scallop. However, the potential impact of the crab on scallop may depend on the availability of other calcified prey associated with scallop beds, such as the sea star, sea urchin, and blue mussel, all species recorded in the diet of P. camtschaticus. To address this issue, a laboratory experiment on foraging behaviour of P. camtschaticus was conducted. The experimental results show that all size classes of red king crab prefer scallops, but small juveniles and medium sized crabs demonstrate active selection for starfish (Asterias rubens) that equals or surpasses the electivity of the large crab. The selection of sea urchin (Strongylocentrotus droebachiensis) and blue mussel (Mytilus edulis) is slightly positive or neutral for the three crab size classes. These results suggest that scallop beds with a rich associated fauna are less vulnerable to red king crabs predation and possibly more resilient than beds with few associated species. Also, crab size distribution is likely relevant for invasion impact, with increasing abundance of small and medium sized crabs being detrimental for alternative calcified prey associated with scallop beds. Successive stages of crab invasion will see an acceleration of scallop mortality rates associated with (i) decreasing availability of alternative prey, due to protracted predation pressure intensified by recruitment of juvenile crabs, and (ii) increased number of large crabs. Estimates of crab density and intake rates suggest that the accelerated loss rates will eventually endanger scallop beds persistence.     

An analysis of mussel bed habitats in the Dutch Wadden Sea

A habitat suitability analysis for littoral mussel beds in the Dutch Wadden Sea was carried out. The analysis was based on the presence of mussel beds in the years 1960–1970, and a number of environmental characteristics: wave action, flow velocity, median grain size, emersion times and distance to a gully border. The habitat model describes mussel bed appearance quantitatively. It predicts the distribution of mussel beds quite well, as well as the distribution of spatfall in the years 1994 and 1996. From the analysis we found that wave action (maximum orbital velocity) was the main structuring factor. A low orbital velocity was preferred. Neither very low, nor maximum flow velocities were favourable for mussel beds. Very coarse sands or silty environments were not preferred. Sites close to the low water line showed lower mussel bed appearance; when emersion time was above 50% , hardly any mussel beds could be found. The habitat suitability analysis and the construction of a habitat suitability map was performed in the framework of the discussions on a further or reduced exploitation of the tidal flats in the Dutch Wadden Sea by cockle and mussel fishery activities. Electronic Publication     

Investigation of the effects of current velocity on mussel feeding and mussel bed stability using an annular flume

An annular flume was used to measure the effect of increasing current velocity on mussel (Mytilus edulis) feeding rate and the stability of mussel beds sampled from the mouth of the Exe estuary (SW England). It was found that, in contrast to earlier flume studies, the feeding rates of mussels from open coast sites were unaffected by current velocities up to 0.8 m s^–1. Algal cell depletion in the water column above mussels was a function of current velocity, increasing with declining currents below 0.05 m s^–1. The erodability/stability of the mussel bed, measured in terms of critical erosion velocity, sediment mass eroded and mean erosion rate, was found to be a function of the nature of the substrate and the density of the mussels. Erosion of mussel beds on sandy substrate showed a non-linear relationship with mussel bed density. In comparison with the sand (0% mussel cover), sediment resuspension was about five and four times higher for 25% and 50% cover, respectively. This was due to the increased turbulence and scouring around the clumps of mussels in low-density parts of the bed, and this resulted in some mussels detaching from the bed. At ~100% mussel cover, the sandy bed was more protected by the dense surface layer of mussels, and none became detached during erosion due to the high number of byssal attachments between individuals. The sediment resuspension from the 100% mussel cover was about three times lower than the 0% cover. Erosion of the bed with 50% cover resulted in burial of a large proportion of the mussels, with a 6 cm increase in sediment level. However, the mussels returned to the surface and recovered in 1–2 days, due to a combination of migration upwards and substrate settlement. Channels on the edge of the main Exmouth mussel bed were characterised by a more stable substrate comprising pebbles and sand with varying mussel densities. At these sites, where mussels experience high current velocities on spring tides (up to 0.9 m s^–1), there was no difference between the erodability of pebble/sand substrate with 0% and 100% mussel cover. The sediment erosion was also lower than the 100% mussel cover on the sandy substrate, particularly at currents >0.4 m s^–1. Sampling of different parts of the mussel bed at Exmouth showed mussels at low densities were made up of smaller clumps with a lower mass ratio of mussels to attached substrate (pebbles/sand), thus providing a greater degree of anchorage. Electronic Publication     

Ecological restoration of mussel beds onto soft‐sediment using transplanted adults

The success of restoration initiatives to restore bivalve beds relies on sufficient recruitment of larvae to offset mortality of re‐established populations. Individuals of the nearly extirpated green‐lipped mussel are capable of surviving within the current environment of the Hauraki Gulf, New Zealand; however, it is uncertain what potential factors might inhibit the establishment and persistence of restored mussel beds. Four experimental mussel beds were established within a shallow soft‐sediment embayment and assessments of population dynamics were conducted approximately every 6 months over a 2‐year period. Deployed mussels quickly congregated into contiguous mussel beds that persisted throughout the study; however, only 26.2% of mussels that were initially established survived until the end of the study. The cause of this overall loss of mussels can be attributed to a near lack of observed recruitment, with only three individual recruiting mussels observed throughout the entire study. Despite similar mortality rates within the restored mussel beds to that of natural populations, these populations will be unsustainable long term given the lack of recruitment. Potential causes of the observed mortality and lack of recruitment are discussed, including environmental factors affecting non‐natal mussel stock and sea star predation. This research provides a foundation for the development of best‐practice methods in the restoration of green‐lipped mussels. However, further investigation into recruitment pathways and sources of mortality for adult mussels will be necessary to overcome the observed limitations if future restoration is to be successful.     

Mussel beds — amensalism or amelioration for intertidal fauna?

The faunal assemblages of a mussel bed (Mytilus edulis L.) and ambient sandflat were compared to study how a bioherm of suspension feeding organisms affects benthic communities in a tidal flat. During a survey of mussel beds in the Wadden Sea at the island of Sylt (North Sea), a total of 52 macrofaunal species and 44 meiobenthic plathelminth species were detected. They occupied different microhabitats in the mussel bed. 56% of the macrofauna species were dwelling in the sediment beneath the mussels and 42% were epibenthic or epiphytic. The latter were restricted in their occurrence to the mussel bed. Along a transect from the sandflat to the mussel bed the mean species densities of macrofauna did not differ significantly, while abundances were significantly lower in the mussel bed than in the sandflat. The composition of the assemblages shifted from a dominance of Polychaeta in the sandflat to Oligochaeta in the mussel bed. Surface filter-feeding polychaetes of the sandflat (Tharyx marioni) were displaced by deposit feeding polychaetes under the mussel cover (Capitella capitata, Heteromastus filiformis). The total meiobenthic density was lower and single taxa (Ostracoda, Plathelminthes, Nematoda) were significantly less abundant in the mud of the mussel bed. The plathelminth assemblage was dominated by grazing species (Archaphanostoma agile), and differed in community structure from a sandflat aseemblage. An amensalistic relationship was found between the suspension-feeding mussels and suspension-feeding infauna, while deposit-feeders were enhanced. The presence of epibenthic microhabitats results in a variety of trophic groups co-occurring in a mussel bed. This is hypothesized as trophic group amelioration and described as an attribute of heterotrophic reefs.     

<Emphasis Type="Italic">Eisenia bicyclis</Emphasis> bed coverage off Oshika Peninsula,Japan, in relation to sporophyte survival and <Emphasis Type="Italic">Strongylocentrotus nudus</Emphasis> abundance

It has recently been reported that the Eisenia bicyclis beds on the coast of Tomarihama in the Oshika Peninsula in Miyagi Prefecture (northern part of Japan) have been progressively receding from the offshore side of the beds every year. In this study, to determine the factors causing the change in the distribution of E. bicyclis beds as well as the extent of the change, an ecological survey of E. bicyclis and grazing animals was conducted. A 150 m-long fixed line was set up in the rocky coastal area on the Oshika Peninsula in Miyagi Prefecture, and the number of E. bicyclis individuals inhabiting a 1-m width was then measured. In addition, the numbers of individuals of sea urchins (Strongylocentrotus nudus) and abalone (Haliotis discus hannai) were measured. Although the E. bicyclis bed extended to the 64-m point on the fixed line at the time of the survey in August 1999, by July 2001, its lower limit was at the 53-m point; thus, a retreat of the outer edge of the bed of 11 m was observed in 2 years. The density of adult E. bicyclis sporophytes in the bed was reduced from 1.08 to 1.80 individuals (ind.) m⁻², respectively. Juvenile sporophytes were also observed, but most of those that had settled near the lower limit of the bed failed to survive to become adult sporophytes. There were few sea urchins, the main grazing animal on the site, inside the E. bicyclis bed (3.5 ind. m⁻² in October 2001) while there were many near the bed lower limit and on the deep-site side (12.9 ind. m⁻² in October 2001). These results indicate that the high grazing pressure of sea urchins reduced the survival of juvenile sporophytes near the bed lower limit, thus impeding the recruitment of juvenile sporophytes to replace adult sporophytes that died due to senescence, storm action and/or grazing.     

Short-term impact of blue mussel dredging (Mytilus edulis L.) on a benthic community

The short-term effect of mussel dredging in a brackish Danish sound was studied. A commercial dredging track was identified and an analysis of the species composition inside the track and at an adjacent control area showed that dredging changed the community structure by reducing the density of polychaetes. In order to investigate the extent and the duration of the dredging impact experimental dredging was conducted. The experimental dredging removed 50% of the mussels in two dredged areas. Immediately after dredging, a significantly lower number of species was measured inside the mussel beds in dredged areas compared to control and boundary areas. This effect lasted for at least 40 days. The analysis of the species composition showed that the dredged area had a significantly lower density, particularly of polychaetes compared to the boundary area. An increased number of species was recorded outside the mussel beds just after dredging, but this effect lasted for less than 7 days. After dredging, brown shrimps, C. crangon invaded the dredged areas. This species is an important predator of smaller invertebrates, and it is suspected that it was feeding on small vulnerable polychaetes exposed at the sediment surface after dredging. The dredging process was observed to form 2–5-cm deep furrows in the seabed, but the sediment texture and the organic content of the sediment was not affected. The biomass accumulation of individual blue mussels was significantly lower in the dredged area compared to the boundary area. This indicates that the disturbance of the mussel bed structure reduced growth and that the lowering of intraspecific food competition caused by a reduced density of mussels did not increase the accumulation of biomass in the mussels which remained in the dredged area.     

Cheating the Locals: Invasive Mussels Steal and Benefit from the Cooling Effect of Indigenous Mussels

The indigenous South African mussel Perna perna gapes during periods of aerial exposure to maintain aerobic respiration. This behaviour has no effect on the body temperatures of isolated individuals, but when surrounded by conspecifics, beneficial cooling effects of gaping emerge. It is uncertain, however, whether the presence of the invasive mussel Mytilus galloprovincialis limits the ability of P. perna for collective thermoregulation. We investigated whether varying densities of P. perna and M. galloprovincialis influences the thermal properties of both natural and artificial mussel beds during periods of emersion. Using infrared thermography, body temperatures of P. perna within mixed artificial beds were shown to increase faster and reach higher temperatures than individuals in conspecific beds, indicating that the presence of M. galloprovincialis limits the group cooling effects of gaping. In contrast, body temperatures of M. galloprovincialis within mixed artificial mussel beds increased slower and exhibited lower temperatures than for individuals in beds comprised entirely of M. galloprovincialis. Interestingly, differences in bed temperatures and heating rates were largely dependent on the size of mussels, with beds comprised of larger individuals experiencing less thermal stress irrespective of species composition. The small-scale patterns of thermal stress detected within manipulated beds were not observed within naturally occurring mixed mussel beds. We propose that small-scale differences in topography, size-structure, mussel bed size and the presence of organisms encrusting the mussel shells mask the effects of gaping behaviour within natural mussel beds. Nevertheless, the results from our manipulative experiment indicate that the invasive species M. galloprovincialis steals thermal properties as well as resources from the indigenous mussel P. perna. This may have significant implications for predicting how the co-existence of these two species may change as global temperatures continue to rise.     

Documentation of sites of intertidal blue mussel (Mytilus edulis L.) beds of the Lower Saxonian Wadden Sea, southern North Sea (as of 2003) and the role of their structure for spatfall settlement

Field surveys (dating back to 1950) and aerial photograph series (dating back to 1966) were evaluated to determine sites of intertidal blue mussel (Mytilus edulis) beds at the Wadden Sea coast of Lower Saxony. Maps were prepared indicating sites of blue mussel beds during the last decades. A table gives additional information on the presence (or absence) of blue mussel beds at each site at the time of large-scale surveys. Altogether 187 sites of M. edulis beds were recorded in the investigation area. In spring 1996, there were still only 19 sites where mussel beds still occurred, although at 51 sites residual mussel-bed structures were present, e.g. shell bases of former beds or protruding patches (which had been occupied by M. edulis before the beds vanished) and open spaces. At that time, the majority of the sites contained neither mussel beds nor mussel-bed structures. The analysis of recent data confirmed that mussel larvae have preferred to settle in sites of present mussel beds and sites with bases of former mussel beds. There was no preferential selection of one of these categories (settled beds vs. shell bases). On the other hand, the presence of mussel beds or mussel bed structures is not obligatory for settlement, because sites without those structures were also re-settled by the spatfall in 1996, even though on a smaller scale.     

Large-Scale Spatial Dynamics of Intertidal Mussel (Mytilus edulis L.) Bed Coverage in the German and Dutch Wadden Sea

Intertidal blue mussel beds are important for the functioning and community composition of coastal ecosystems. Modeling spatial dynamics of intertidal mussel beds is complicated because suitable habitat is spatially heterogeneously distributed and recruitment and loss are hard to predict. To get insight into the main determinants of dispersion, growth and loss of intertidal mussel beds, we analyzed spatial distributions and growth patterns in the German and Dutch Wadden Sea. We considered yearly distributions of adult intertidal mussel beds from 36 connected tidal basins between 1999 and 2010 and for the period 1968–1976. We found that in both periods the highest coverage of tidal flats by mussel beds occurs in the sheltered basins in the southern Wadden Sea. We used a stochastic growth model to investigate the effects of density dependence, winter temperature and storminess on changes in mussel bed coverage between 1999 and 2010. In contrast to expectation, we found no evidence that cold winters consistently induced events of synchronous population growth, nor did we find strong evidence for increased removal of adult mussel beds after stormy winter seasons. However, we did find synchronic growth within groups of proximate tidal basins and that synchrony between distant groups is mainly low or negative. Because the boundaries between synchronic groups are located near river mouths and in areas lacking suitable mussel bed habitat, we suggest that the metapopulation is under the control of larval dispersal conditions. Our study demonstrates the importance of moving from simple habitat suitability models to models that incorporate metapopulation processes to understand spatial dynamics of mussel beds. The spatio-dynamic structure revealed in this paper will be instrumental for that purpose.     

LIFE CYCLE AND HABITAT SHIFT OF THE TROCHID SNAIL DILOMA SUAVIS (PHILIPPI) WITHIN INTERTIDAL MUSSEL ZONES

The life cycle of the trochid snail Diloma suavis (Philippi, 1849),was studied on an intertidal rocky shore at Shirahama, Wakayama Prefecture,where two mytilid bivalves, Septifer virgatus (Wiegmann) andHormomya mutabilis (Gould), formed vertically contiguous musselbeds in the upper-middle and lower zones, respectively. At lowtide in April, the snail density increased with decreasing shoreheight and was greatest at the middle level of the H. mutabilisbed. Then, the density decreased towards the lower littoralfringe. Newly settled juveniles smaller than 2 mm in shell heightappeared abundantly in late summer and autumn within algal turfon the lower shore. As snails grew larger than 2 mm, they appearedwithin the gaps of the H. mutabilis bed and the S. virgatusbed. They increased in size monotonically towards the next summer,but rate of growth in shell height tended to be great in autumnand small in winter. Seasonal change in the density of snailsfound within the gaps of the mussel beds was remarkable during athree year period, increasing from autumn to winter and then decreasingtowards next summer. Reproduction occurred in summer, and adultsnails disappeared by September. It is thus suggested that this specieshas a one year lifespan and shows a habitat shift from algal turfto the gaps of the mussel beds with growth. (Received 12 October 1998; accepted 2 March 1999)     

The role of the blue mussel Mytilus edulis in the cycling of nutrients in the Oosterschelde estuary (The Netherlands)

The fluxes of particulate and dissolved material between bivalve beds and the water column in the Oosterschelde estuary have been measured in situ with a Benthic Ecosystem Tunnel. On mussel beds uptake of POC, PON and POP was observed. POC and PON fluxes showed a significant positive correlation, and the average C:N ratio of the fluxes was 9.4. There was a high release of phosphate, nitrate, ammonium and silicate from the mussel bed into the water column. The effluxes of dissolved inorganic nitrogen and phosphate showed a significant correlation, with an average N:P ratio of 16.5. A comparison of the in situ measurements with individual nutrient excretion rates showed that excretion by the mussels contributed 31–85% to the total phosphate flux from the mussel bed. Ammonium excretion by the mussels accounted for 17–94% of the ammonium flux from the mussel bed. The mussels did not excrete silicate or nitrate. Mineralization of biodeposition on the mussel bed was probably the main source of the regenerated nutrients.From the in situ observations net budgets of N, P and Si for the mussel bed were calculated. A comparison between the uptake of particulate organic N and the release of dissolved inorganic N (ammonium + nitrate) showed that little N is retained by the mussel bed, and suggested that denitrification is a minor process in the mussel bed sediment. On average, only 2/3 of the particulate organic P, taken up by the mussel bed, was recycled as phosphate. A net Si uptake was observed during phytoplankton blooms, and a net release dominated during autumn. It is concluded that mussel beds increase the mineralization rate of phytoplankton and affect nutrient ratios in the water column. A comparison of N regeneration by mussels in the central part of the Oosterschelde estuary with model estimates of total N remineralization showed that mussels play a major role in the recycling of nitrogen.     

Reduction and recovery of keystone predation pressure after disease‐related mass mortality

Disturbances such as disease can reshape communities through interruption of ecological interactions. Changes to population demographics alter how effectively a species performs its ecological role. While a population may recover in density, this may not translate to recovery of ecological function. In 2013, a sea star wasting syndrome outbreak caused mass mortality of the keystone predator Pisaster ochraceus on the North American Pacific coast. We analyzed sea star counts, biomass, size distributions, and recruitment from long‐term intertidal monitoring sites from San Diego to Alaska to assess regional trends in sea star recovery following the outbreak. Recruitment, an indicator of population recovery, has been spatially patchy and varied within and among regions of the coast. Despite sea star counts approaching predisease numbers, sea star biomass, a measure of predation potential on the mussel Mytilus californianus, has remained low. This indicates that post‐outbreak populations have not regained their full predation pressure. The regional variability in percent of recovering sites suggested differences in factors promoting sea star recovery between regions but did not show consistent patterns in postoutbreak recruitment on a coast‐wide scale. These results shape predictions of where changes in community composition are likely to occur in years following the disease outbreak and provide insight into how populations of keystone species resume their ecological roles following mortality‐inducing disturbances.     

Building Resilience Against Climate-Driven Shifts in a Temperate Reef System: Staying Away from Context-Dependent Ecological Thresholds

As climate-driven environmental changes and anthropogenic perturbations increasingly affect ecological systems, the number of abrupt phase shifts in ecosystem dynamics is rising, with far-reaching ecological, economic and social effects. These shifts are notoriously difficult to study, anticipate and manage. Although indicators of impending phase shifts in ecosystems have been described theoretically, they have only been observed empirically either after the fact or under controlled experiments. Here we demonstrate the usefulness of case-specific simulation models to estimate tipping points in the dynamics of real ecological systems, characterise how these thresholds may vary depending on local conditions and derive safe management targets associated with low risk of undesirable phase shifts. Under the combined effects of ocean changes and fishing, inshore rocky reefs in eastern Tasmania can transition from dense seaweed beds to sea urchin ‘barrens’ habitat, realising severe local loss of habitat, productivity and valuable fisheries. Using Monte-Carlo simulations with a validated model that realistically captures reef dynamics, we characterise the hysteresis in community dynamics and the variability in ecological thresholds along the gradient of environmental conditions. Simulation suggests that prevention of ongoing sea urchin destructive grazing of macroalgal beds is achievable but the yet-to-be-observed restoration of seaweed beds from extensive sea urchin barrens is highly unlikely. To guide management against undesirable phase shifts, we define target points associated with low risk of widespread barrens formation and show that, along with sea urchin culling, recognising the role of lobsters in mitigating sea urchin destructive grazing through predation is key to maintain reef productivity.