Mussels matter: postlarval dispersal dynamics altered by a spatially complex ecosystem engineer

This study investigated postlarval dispersal of soft-bottom macrofauna at a spatially complex intertidal mudflat comprising patches of bare sediment and an ecosystem engineer, the mussel Mytilus edulis. At each of four sites in Guard Point Cove, Maine, USA, we took core samples and deployed bedload traps in bare sediment and mussel bed habitats to estimate ambient densities, rates of sediment flux, and several measures of postlarval dispersal. Univariate and multivariate nonmetric multidimensional scaling (nMDS) results showed few significant site effects and no habitat×site interactions. In contrast, there were numerous significant habitat effects. Compared to the bare sediment, the mussel bed habitat had: fewer species; higher ambient density and proportional abundance of the oligochaete Tubificoides benedeni (the dominant species in both habitats); lower ambient densities and proportional abundances of major taxa and the nonoligochaetes as a group; and higher sediment flux and relative (i.e., per capita) dispersal of nonoligochaetes. Macrofauna species dispersed in relative proportions that were different from those in the ambient assemblage. Per capita T. benedeni transport rates were low in mussel beds compared to those for nonoligochaetes, consistent with the view that beds represent favorable habitat for oligochaetes. The number of total macrofauna individuals trap⁻¹ day⁻¹ was negatively correlated with ambient density and positively correlated with sediment flux in both habitats, but these relationships were significant only in the mussel bed. The results indicate that altered transport rates of sediment and postlarvae are important mechanisms by which mussels act as ecosystem engineers to modify soft-bottom habitats. Differential transport rates caused by aggregations of mussels and other foundation species must be considered in explanations of spatial pattern in soft-bottom communities.     

High survival and growth rates of introduced Pacific oysters may cause restrictions on habitat use by native mussels in the Wadden Sea

Pacific oysters Crassostrea gigas (Thunberg, 1793) were introduced to the northern Wadden Sea (North Sea, Germany) by aquaculture in 1986 and finally became established. Even though at first recruitment success was rare, three consecutive warm summers led to a massive increase in oyster abundances and to the overgrowth of native mussel beds (Mytilus edulis L.). These mussels constitute biogenic reefs on the sand and mud flats in this area. Survival and growth of the invading C. gigas were investigated and compared with the native mussels in order to predict the further development of the oyster population and the scope for coexistence of both species. Field experiments revealed high survival of juvenile C. gigas (approximately 70%) during the first three months after settlement. Survival during the first winter varied between > 90% during a mild and 25% during a cold winter and was independent of substrate (i.e., mussels or oysters) and tide level. Within their first year C. gigas reached a mean length of 35-53 mm, and within two years they grew to 68-82 mm, which is about twice the size native mussels would attain during that time. Growth of juvenile oysters was not affected by substrate (i.e., sand, mussels, and other oysters), barnacle epibionts and tide level, but was facilitated by fucoid algae. By contrast, growth of juvenile mussels was significantly higher on sand flats than on mussel or oyster beds and higher in the subtidal compared to intertidal locations. Cover with fucoid algae increased mussel growth but decreased their condition expressed as dry flesh weight versus shell weight. High survival and growth rates may compensate for years with low recruitment, and may therefore allow a fast population increase. This may lead to restrictions on habitat use by native mussels in the Wadden Sea.     

Vulnerability of newly settled plaice (Pleuronectes platessa L.) to predation: effects of habitat structure and predator functional response

Plaice (Pleuronectes platessa) nursery grounds on the Swedish west coast have been subject to increasing cover of annual green macroalgae during recent years, with growth of algae starting at the time of plaice settlement in April to May. A laboratory experiment was performed to investigate how the vulnerability to predation of metamorphosing plaice was affected by the presence of filamentous algae. Predation by shrimps (Crangon crangon) on settling plaice larvae was higher on sand than among algae, whereas predation by crabs (Carcinus maenas) was unaffected by habitat type, suggesting a lower overall mortality of plaice in the vegetated habitat. When predators and prey were presented with a combination of the two habitats, predation by shrimps was as high as that in the sand treatment alone, whereas predation by crabs was lower than that in the two treatments with one habitat. Based on these results, an additional experiment was performed, investigating the functional response of shrimps to six densities of juvenile plaice in a sand habitat with alternative prey present. The proportional mortality of juvenile plaice (12-16 mm total length (TL)) was density-dependent and was best described by a type III (sigmoid) functional response of the predatory shrimps. The results suggested that the combined predation pressure from shrimps and crabs was lower among algae than on sand, but settling plaice and predatory shrimps chose the sand habitat. Plaice densities in the sigmoid part of the obtained functional response curve represented normal to high field densities of plaice on the Swedish west coast, suggesting that shrimp predation could have a stabilising effect on plaice recruitment. The formation of macroalgae mats could therefore lead to a concentration of plaice juveniles in the remaining sand habitat and increased mortality through density-dependent predation by shrimps.     

Population dynamics of the green-lipped mussel, Perna canaliculus, at various spatial and temporal scales in northern New Zealand

Ecological processes that differentiate and maintain intertidal populations of mussels, Perna canaliculus, were studied within three sites at Ninety Mile Beach, northern New Zealand. At these three sites (Scott Point, The Bluff and Tonatona Beach), the dynamics of larval availability, primary and secondary settlement, recruitment and mortality rates were investigated at various spatial and temporal scales. (1) Mussel concentrations in seawater were variable with respect to study site and time of year, with highest abundances at the northernmost population (Scott Point) and lowest concentrations at the middle population (The Bluff). In seawater at all three sites, small mussels (< 0.25 mm in shell length) were more abundant in August 2000, while larger mussels (> 0.5 mm in shell length) were more abundant in March 2001. (2) Primary and secondary settlement patterns were investigated during short-term (daily) and long-term (monthly) settlement experiments, within quadrats that were cleared of all mussels in both the mussel bed and in adjacent algal band habitats. At all sites, primary settlement (< 0.5 mm in shell length) was high within the algal band habitat in August 1999, 2000 and 2001. Conversely, secondary settlement (> 2.0 mm in shell length) was high within cleared areas in the mussel beds in November-March 1999-2000 and 2000-2001. Abundance of mussels settling on artificial substrates placed in the intertidal did not differ greatly from comparable areas of natural substrates (bare rock or algae within cleared quadrats). (3) Recruitment and mortality rates were recorded during monthly surveys of the adult populations. Within three mussel size classes (< 24, 25-74 and > 75 mm in shell length), peak recruitment coincided with high mortality in August of the 2 years studied. However, the most dramatic turnover of the population was observed at Scott Point in both years, following a spawning event. In adjacent waters at Scott Point, large accumulations of drift algae covered (up to 100% cover) with juvenile mussels may deplete food supplies usually delivered to intertidal adult mussels, causing their demise. Mats of adult mussels were observed “peeling-off” from the rocky shore at this time of the year, making space available to the new recruits. Where nearshore algal accumulations were moderate to low, only moderate to low mussel turnovers were observed (e.g. Tonatona Beach and The Bluff).     

Elemental Fingerprinting of Mussel Shells to Predict Population Sources and Redistribution Potential in the Gulf of Maine

As the climate warms, species that cannot tolerate changing conditions will only persist if they undergo range shifts. Redistribution ability may be particularly variable for benthic marine species that disperse as pelagic larvae in ocean currents. The blue mussel, Mytilus edulis, has recently experienced a warming-related range contraction in the southeastern USA and may face limitations to northward range shifts within the Gulf of Maine where dominant coastal currents flow southward. Thus, blue mussels might be especially vulnerable to warming, and understanding dispersal patterns is crucial given the species'' relatively long planktonic larval period (>1 month). To determine whether trace elemental “fingerprints” incorporated in mussel shells could be used to identify population sources (i.e. collection locations), we assessed the geographic variation in shell chemistry of blue mussels collected from seven populations between Cape Cod, Massachusetts and northern Maine. Across this ∼500 km of coastline, we were able to successfully predict population sources for over two-thirds of juvenile individuals, with almost 80% of juveniles classified within one site of their collection location and 97% correctly classified to region. These results indicate that significant differences in elemental signatures of mussel shells exist between open-coast sites separated by ∼50 km throughout the Gulf of Maine. Our findings suggest that elemental “fingerprinting” is a promising approach for predicting redistribution potential of the blue mussel, an ecologically and economically important species in the region.     

The contribution of marine snow to the particle food supply of the benthic suspension feeder, Mytilus edulis

In order to examine the importance of the settling of large particles to the food supply and feeding behavior of a benthic culture of the blue mussel, Mytilus edulis, we investigated the tidal dynamics of large (>0.5 mm diameter) marine aggregates, commonly known as marine snow, during three tidal cycles in July 1998 at a shallow, subtidal, low current flow regime site along the coast of Maine (Shorey Cove, Roque Island, Englishman's Bay, Maine).In situ, optically measured marine snow showed a distinct tidal signal displaying an increase in size and abundance through high tide with a peak on the early ebb tide as it settled to the bottom. Marine snow volume ranged an order of magnitude through the tidal cycle, from under 8 to over 80 mm³ l⁻¹. An increase in the in situ marine snow volume corresponded with an increase in benthic mussel feeding activity (from 20% to 60% of maximum exhalant siphon area, which is an estimate of pumping rate) and maximum rates of pseudofeces production by the mussels during periods of low tidal current speeds. In contrast, mussels from the same population feeding on surface waters in shipboard chambers produced no pseudofeces and had high pumping rates (80-100% maximum exhalant siphon area) over the whole tidal period. A second peak in benthic mussel pumping rates also occurred during flood tide.Food quality was lower in the bottom waters due to significantly higher particulate inorganic matter (PIM, >2 mg l⁻¹) when compared with the surface waters. PIM accounted for 95% of the total settled mass flux of 3.4 g m⁻² day⁻¹ measured in sediment traps deployed 1 m off the bottom, with organic carbon representing only 2.5% of the mass flux during the mid-summer conditions. At low-current sites such as Shorey Cove, Roque Island, Maine, the settling of marine snow provides an important additional source of food, albeit of low quality, to benthic populations of blue mussels.     

Effects of epibiotic algae on the survival, biomass and recruitment of mussels, Mytilus L. (Bivalvia: Mollusca)

Habitats composed of living ‘ecosystem engineers’, such as mussels, are subject to direct and indirect interactions with organisms that live among them. These interactions may affect the presence and structure of habitat, and hence, the associated taxa. We examined the direct effects of epibiotic algae on the survival, biomass and recruitment of mussels (Mytilus L.) on the west coast of Ireland. A field experiment showed that the presence of epibiotic fucoid algae reduced the likelihood of survival of mussels during storms. We also found that the strength of attachment of mussels did not increase in the presence of epibionts. Another in situ experiment revealed that the presence of ephemeral epibiotic algal mats had no effect on the biomass of host mussels, suggesting no effect on mussel growth or production. The abundance of small mussels (< 5 mm) covaried with cover of epibiotic algae at one location, but experimental results suggest no direct effect of the algae on recruitment unless through the action of chemical cues which linger after the removal of algae. We have identified a negative direct interaction between epibiotic algae and mussels on exposed rocky shores, which may often be characterised by bottom-up regulation. It is thought that positive interactions may be more important on more sheltered or sedimentary shores where top-down processes are more likely to dominate.     

Increase in density of genetically diverse invasive Asian shore crab (<Emphasis Type="Italic">Hemigrapsus sanguineus</Emphasis>) populations in the Gulf of Maine

Hemigrapsus sanguineus, the Asian shore crab, has rapidly replaced Carcinus maenas, the green crab, as the most abundant crab on rocky shores in the northwest Atlantic since its introduction to the United States (USA) in 1988. The northern edge of this progressing invasion is the Gulf of Maine, where Asian shore crabs are only abundant in the south. We compared H. sanguineus population densities to those from published 2005 surveys and quantified genetic variation using the cytochrome c oxidase subunit I gene. We found that the range of H. sanguineus had extended northward since 2005, that population density had increased substantially (at least 10-fold at all sites), and that Asian shore crabs had become the dominant intertidal crab species in New Hampshire and southern Maine. Despite the significant increase in population density of H. sanguineus, populations only increased by a factor of 14 in Maine compared to 70 in southern New England, possibly due to cooler temperatures in the Gulf of Maine. Genetically, populations were predominantly composed of a single haplotype of Japanese, Korean, or Taiwanese origin, although an additional seven haplotypes were found. Six of these haplotypes were of Asian origin, while two are newly described. Large increases in population sizes of genetically diverse individuals in Maine will likely have a large ecological impact, causing a reduction in populations of mussels, barnacles, snails, and other crabs, similar to what has occurred at southern sites with large populations of this invasive crab species.     

Effects of native predators and eelgrass habitat structure on the introduced Asian mussel Musculista senhousia (Benson in Cantor) in southern California

The ability of predators to control the abundance of non-native species has been little explored in marine systems. Native predators may be used to control non-native species or may confer invasion resistance to communities if predation rates on invaders are density-dependent. We studied the response of southern California native predators to the density of Musculista senhousia (Benson in Cantor, 1842), a small, fast growing mussel that has been introduced from Japan to several coastlines worldwide. We performed field experiments to determine if M. senhousia proportional mortality is density-dependent and if eelgrass Zostera marina L. habitat structure influenced mussel density-dependent mortality. We also evaluated the effect of seagrass habitat structure on the aggregative and functional responses of the predatory gastropod Pteropurpura festiva (Hinds, 1844) to Asian mussel density. In the summer of 2002, P. festiva aggregated in plots with high mussel density and was responsible for nearly all predation on M. senhousia. However, M. senhousia proportional mortality was inversely density-dependent at all levels of eelgrass above-ground and below-ground habitat structure. Asian mussel proportional mortality also was inversely density-dependent and was not influenced by eelgrass habitat structure in the spring of 2004 when wading birds were the chief predator of mussels. In contrast to results for mussel proportional mortality, the aggregative and functional responses of P. festiva varied with seagrass habitat structure. P. festiva density increased with Asian mussel density in plots with low simulated habitat structure, but the relationship between P. festiva density and Asian mussel density was parabolic at zero, intermediate and high levels of habitat structure. In field enclosures, P. festiva exhibited a Type I (linear) functional response to Asian mussel density at low levels of eelgrass structure, and a Type II (hyperbolic) functional response to mussel density at high levels of eelgrass structure. Our results and those of others suggest that the degree to which local benthic communities in southern California are resistant to Asian mussel invasion depends on habitat structure, mussel settlement rates, and the density and diversity of predators.     

Recruitment of shore crabsCarcinus maenas on tidal flats: Mussel clumps as an important refuge for juveniles

During the late summer and early fall, juvenile shore crabs (Carcinus maenas L.) occurred in high abundances in mussel clumps scattered on tidal flats of the Wadden Sea. Abundances were much lower on bare tidal flats without mussel clumps and decreased substantially from July to November, whereas numbers in mussel clumps remained high. Large crabs left the tidal flats in early fall, whereas juveniles undertook tidal migrations only in the late fall. In March very few shore crabs were found in the intertidal area. The size of juvenile shore crabs living between mussels did not increase significantly during fall. On the bare tidal flats surrounding the mussels, a size increase was observed. Mussel beds and mussel clumps serve as a spatial refuge for the early benthic phases of juvenile shore crabs. Between mussels they can hide effectively from their epibenthic predators. Juvenile shore crabs do not leave the intertidal area and the mussel habitats before their major predators have left the area. Mussel clumps scattered over the tidal flats may be a critical refuge for juvenile shore crabs settling on tidal flats. Intensified efforts in mussel culturing in the European Wadden Sea during recent decades may have caused an increased abundance of mussel clumps on tidal flats, thus enhancing habitat availability for some mussel-clump inhabitants.     

Effects of depth, sediment and grazers on the degradation of drifting filamentous algae (Cladophora glomerata and Pilayella littoralis)

Eutrophication in the northern Baltic Sea promotes growth of annual filamentous algae. The algae detach, gather at the bottom and give rise to algal mats of varying size, density, composition and condition. Dense mats of filamentous algae induce anoxia, which in turn leads to faunal mortality. By a set of field experiments, we have studied the fate of the abundant Cladophora glomerata after detaching from the rocky substrate, and the effect of water depth and sediment on its decay. Further, we have studied the importance of common mesograzers (Gammarus and Idotea) on the rate of degradation of C. glomerata and Pilayella littoralis.Our results show that loose algae at shallow sites (8 m) decompose faster than algae in deeper (18 m) areas. Drifting C. glomerata on the sediment is more rapidly broken down and dissolved than algae floating in the water column, which depends on higher microbiological activity. Dominant amphipods (Gammarus spp) colonise near-shore drift algae quickly, and juvenile bivalves (Cerastoderma glaucum) utilise algae in the water column for settling. Moderate natural densities of grazers (Gammarus spp and Idotea baltica) in the drifting algae did not increase the degradation rates of C. glomerata and P. littoralis. C. glomerata was completely decomposed in 4 months.Our experiments demonstrate the effects of position (depth, water/sediment) and grazing on the degradation of drifting filamentous algae. Mass developments of opportunistic algae occur annually in the study area, and information on the destiny of detached drift algae may help us to predict their longevity and the damage they cause, and hence, to decide on long-term measures needed to improve environmental conditions.     

Hierarchical spatial structure in soft-bottom mussel beds

Mussels (Mytilus edulis L.) are unusual because they thrive in both rocky shore and soft-bottom habitats. Despite their ecological and economic importance, little is known about their spatial structure. Mussels do not generally recruit to bare soft substrate because larvae and postlarvae cannot attach to a bottom of small sediment particles. They attach to hard objects on the sediment surface (especially other mussels), so soft-bottom mussel beds may be spatially organized in ways that are fundamentally different from those on rocky shores. The purpose of our study was to characterize the scales of spatial variability for several mussel abundance parameters in soft-bottom, intertidal M. edulis beds in coastal Maine. We used a random factor nested-ANOVA design of 200 cm² Cores within 1 m² Quadrats within 6 m Transects within Positions within bed Sites along 70 km (euclidean distance) of the Maine coast. Based on the literature and our field observations, we hypothesized that Sites and Positions account for most of the spatial variance in soft-botttom mussel beds. We rejected this hypothesis. Sites and Positions were not important in explaining variation in total mussel density, density of new recruits, or density of larger mussels. Although most of the variance in surface silt-clay fraction did occur at these levels, most mussel variation occurred at smaller spatial scales, specifically at the Quadrat scale for new recruits and total mussels and at the Transect scale for larger mussels. Variance in mussel parameters was not closely linked to the silt-clay fraction of surface sediment or to Site rankings of wind exposure and tidal flow. Variance in total mussel density was due primarily to variance in recruitment. No single scale explained more than about half the mussel variance, and no single scale was best at explaining all the mussel parameters. Greater knowledge about mussel bed spatial variability would be useful because it can help direct scale-dependent sampling regimes, field experiments, and coastal management practices.     

Divergent succession and implications for alternative states on rocky intertidal shores

It has been hypothesized that rockweed stands and mussel beds in sheltered bays in the Gulf of Maine, USA, are alternative community states. As a test of this hypothesis, experimental clearings of different sizes were established in stands of the rockweed Ascophyllum nodosum (L.) Le Jolis to determine if successional changes in large clearings developed species assemblages distinctly different from the surrounding A. nodosum stands. Clearings ranging from 1 to 8 m in diameter were created at 12 sites in 4 bays on Swan's Island, Maine, in 1996 to mimic the effects of ice scour, and abundances of gastropods, barnacles, mussels and fucoid algae were monitored until 2002. ANOVAs and MDS showed strong effects of clearing size and divergent successional changes in large clearings. Large clearings were quickly filled in and remained dominated by the alga Fucus vesiculosus L. and the barnacle Semibalanus balanoides (L). There was no evidence for site-specific effects, and Mantel tests showed clearing size was a better predictor of species composition than geographic distances among sites. Results suggest that large pulse disturbances using clearings of 8 m in diameter can initiate divergent successional pathways and have a protracted effect on species composition. Results are also consistent with the hypothesis that mussel beds and rockweed stands in sheltered bays may be alternative community states.     

Predator size interacts with habitat structure to determine the allometric scaling of the functional response

While both predator body size and prey refuge provided by habitat structure have been established as major factors influencing the functional response (per capita consumption rate as a function of prey density), potential interactions between these factors have rarely been explored. Using a crab predator (Panopeus herbstii) – mussel prey (Brachidontes exustus) system, we examined the allometric scaling of the functional response in oyster (Crassostrea virginica) reef habitat, where crevices within oyster clusters provide mussels refuge from predation. A field survey of mussel distribution showed that mussels attach closer to the cluster periphery at high mussel density, indicating the potential for saturation of the refuge. In functional response experiments, the consumption rate of large crabs was depressed at low prey density relative to small crabs, while at high prey density the reverse was true. Specifically, the attack rate coefficient and handling time both decreased non‐linearly with crab size. An additional manipulation revealed that at low prey densities, the ability of large crabs to maneuver their claws and bodies to extract mussels from crevices was inhibited relative to small crabs by the structured habitat, reducing their attack rate. At high prey densities, crevices were saturated, forcing mussels to the edge of clusters where crabs were only limited by handling time. Our study illuminates a potentially general mechanism where the quality of the prey refuge provided by habitat structure is dependent on the relative size of the predator. Thus anthropogenic influences that alter the natural crab size distribution or degrade reef habitat structure could threaten the long‐term stability of the crab –mussel interaction in reefs.     

Annual spawning of the hydrothermal vent mussel, Bathymodiolus azoricus, under controlled aquarium, conditions at atmospheric pressure

Dense macrofaunal communities of modioliform mussels are a major component of many hydrothermal vent and cold seep ecosystems. The hydrothermal vent mussel Bathymodiolus azoricus, that dominates hydrothermal vent communities near the Azores Triple Junction, can be maintained in aquaria at atmospheric pressure. Cages containing these mussels were placed over diffuse vent outlets and recovered at different times. Cages recovered in January 2003 contained mussels with ripe gonads while those recovered in July, August and November 2001 and in April 2003 did not. Mussels collected post-spawning in April 2003 spawned in the aquaria in January 2004. Young mussels recruited to the cages in April 2003. The data support a main single period of spawning and of juvenile recruitment for B. azoricus.     

Body size and food thresholds for zero growth in Dreissena polymorpha: a mechanism underlying intraspecific competition

1. Dreissena polymorpha is an extraordinarily successful invasive species that shows high recruitment of small juvenile mussels on established mussel banks. Such juvenile settlement on, and overgrowth of, large adult mussels; however, leads to competition with adults, and often at high densities and low‐food concentrations. 2. The concept of food thresholds for zero growth has been a powerful approach to explaining size‐related exploitative competition in different zooplankton species. We applied it to investigate whether food threshold concentrations for zero growth (C₀) differ between juvenile and adult zebra mussels. 3. By determining body mass growth at various concentrations of a diet mixture (Nannochloropsis limnetica and Isochrysis aff. galbana) we demonstrate that the threshold food concentration for growth of juvenile mussels (C₀ = 0.08 mg C L⁻¹) is substantially lower than that for adults (C₀ = 0.36 mg C L⁻¹). 4. This indicates that, at low food availability, juvenile zebra mussels are competitively superior to their larger conspecifics. Within zebra mussel banks plankton food is substantially depleted and so the observed mechanism might ensure juvenile success and therefore the regeneration of mussel banks in nature.     

Larval settlement of the green sea urchin, Strongylocentrotus droebachiensis, in the southern Gulf of Maine

Abstract. Settlement timing and differential settlement for the larval stage of the green sea urchin, Strongylocentrotus droebachiensis , in the southern Gulf of Maine was studied during the summer of 1996. Settlement densities on astroturf panels were highest in June and early July (13 to 37 m ⁻² d⁻¹), and peaked in mid-June (199 m⁻² d⁻¹). Settlement was low to nonexistent from mid-July through August (0 to 2 m⁻² d⁻¹). During the peak in settlement, no selection for substrate type was observed. In the remainder of the settlement period, differential settlement occurred, with a preference for substrate covered with live coralline algae. Test diameter of newly settled urchins varied among the substrates, with urchins settling on live coralline algae having the largest test diameter (0.43 ± 0.01 mm). There were no differences in test diameter among the different weeks in which sampling was done. Sustained onshore winds occurred only during peak settlement, suggesting that wind drift currents may concentrate larvae and influence patterns of larval settlement.     

Effects of Adult Dreissena polymorpha on Settling Juveniles and Associated Macroinvertebrates

The impact of Dreissena polymorpha settlement on recruitment of juvenile mussels and density of other macroinvertebrates was studied in field experiments using blank concrete blocks and tiles (control), blocks and tiles with attached empty zebra mussel shells, and blocks and tiles with attached living mussels. On blocks, dominant invertebrate taxa showed colonization patterns coinciding with increased habitat complexity owing to zebra mussel settlement or the biodeposition of faeces and pseudofaeces. Adult and especially juvenile zebra mussels preferred blocks with empty shells to blank blocks and blocks with living mussels; this might possibly be caused by a chemical cue that induces gregarious settlement. Lower recruitment on blocks with attached living mussels compared to blocks with only shells could be the consequence of ingestion of larvae by adult mussels and of competition for food. On tiles, the sediments deposited and the organic content of the sediment were investigated. Sedimentation was significantly higher on shell‐only and live‐mussel tiles compared to blank tiles. Organic matter differed significantly between blank and live‐mussel tiles.     

Survivorship of juvenile barnacles and mussels: spatial dependence and the origin of alternative communities

Mussels, barnacles, and rockweeds often form a distinct mosaic of patches on rocky intertidal shores, and it has been suggested that these communities may represent alternative community states. One way that alternative community states can arise is if early successional events are scale-dependent, but it is not known if juvenile survivorships of mussels and barnacles are, in fact, scale-dependent. Scale-dependence of barnacles (Semibalanus balanoides (L.)) and mussels (Mytilus edulis L.) was tested in the Gulf of Maine, USA. In winter 1997, clearings of 1, 2, 4 and 8 m in diameter and uncleared controls were made in stands of the rockweed Ascophyllum nodosum (L.) Le Jolis at 12 sites spread evenly across four bays on Swan's Island, ME. Summer and fall-winter survivorship of barnacles, which recruited in spring 1997, were estimated by tracking the 1997 cohort until late winter 1998. Survivorship of mussels was estimated from following the fate of transplanted juveniles over 4 days in late August 1997. Both barnacles and mussels showed better survival in 4 and 8 m clearings than in small clearings and controls. There was also significant variation in survivorship among sites. Densities of gastropods in the clearings did not reflect survivorship patterns of barnacles and mussels. Barnacle survivorship increased in fall and winter, and in large clearings was comparable to survivorship in barnacle-dominated habitats. Mussel survivorship was low in all clearing sizes suggesting that mussel beds develop slowly.     

Assessing the potential benthic-pelagic coupling in episodic blue mussel (Mytilus edulis) settlement events within eelgrass (Zostera marina) communities

Coastal marine seagrass ecosystems are important nursery grounds for commercially and recreationally important species, and they serve as key settlement and recruitment sites for other species. We investigated several years (2001-2003) where episodic settlement events of blue mussels (Mytilus edulis) occurred in Barnegat Bay, NJ, USA. Population assessment indicated that blue mussels settled in eelgrass beds (Zostera marina) in late spring with peak densities exceeding 170,000 m⁻². Based on calculated filtration rates of M. edulis, we determined that for at least 53 days in 2001, the density and size distribution of M. edulis were sufficient to filter the water column volume in excess of twice a day, with maximum calculated filtration rates exceeding 8 m³ water m⁻² day⁻¹. While the settlement event in 2001 was very localized, in 2003, the settlement event was considerably more widespread throughout the bay, with maximum settling densities exceeding 175,000 individuals m⁻². Associated with these high densities, maximum calculated filtration rates exceeded 15 m³ water m⁻² day⁻¹. This filtration potential may have impeded the localized development of a brown-tide (Aureococcus anophagefferens) bloom in 2001, which occurred in other regions of the bay, but the widespread settlement event seen in 2003 may have impeded the development of any brown-tide blooms in Barnegat Bay during that summer. The decline in mussel densities throughout the summer may be a result of elevated water temperatures in this back bay, but at one site, the high settlement of M. edulis was followed by a substantial migration (>40 individuals m⁻²) of small sea stars (Asterias forbesii). In 2001, A. forbesii was a significant factor in reducing M. edulis density by the end of the summer at the Barnegat Inlet site and a community level assessment showed significant positive correlations between mussel aggregations and sea star densities (r=0.68-0.73, P<0.001). At this same site in 2003, the sea stars were again present in high densities (26 m⁻²) and were a potential mechanism for mussel decline. In other regions of the bay, sea star densities are very low, but numerous other predatory species exist, including blue crabs (Callinectes sapidus), green crabs (Carcinus maenus), spider crabs (Libinia spp.), and several Xanthid crabs. Given the high mussel densities seen in this study and the considerable predation by sea stars and other benthic predators, the benthic-pelagic coupling which these mussels provide in this system contributes to the high secondary production in these grass beds.