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.     

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.     

Impact assessment of an invasive flatworm, Convoluta convoluta, in the Southern Gulf of Maine

Convoluta convoluta (Abildgaard 1806) is a small (2-3 mm long) acoellous turbellarian flatworm from Europe that has invaded the Gulf of Maine within the last 5 years. Although it has been reported in densities of up to 19 individuals/cm², its ecological impact remains unknown. In its native habitat, it consumes harpacticoid copepods and primary settling mussels <0.5-mm shell length. This study estimated the impact of C. convoluta on juvenile blue mussel populations (Mytilus edulis Linnaeus 1758) around the Isles of Shoals in the southern Gulf of Maine, USA as well as looking at their distribution in their new habitat. We surveyed worm densities at sites of differing wave exposure over three substrates (hard substrates, bladed algae, filamentous algae) to quantify patterns of worm abundance. We found worms on all substrate types with their highest abundances occurring in areas of maximal sunlight exposure and minimal physical disturbance. We showed a definite pattern of consumption of mussels in the lab and found C. convoluta to consume up to 35% of primary settling mussels in the field, but only under certain conditions. Per capita impact on juvenile mussels was found to vary greatly in correlation with mussel recruitment rate and water temperature, but not with the consumption of harpacticoids. Our results also suggested that per capita interaction strength was reduced by intraspecific density-dependent competition and water temperature. The overall impact of C. convoluta on mussel populations in the southern Gulf of Maine is therefore estimated to be minimal.     

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.     

Mussels as ecosystem engineers: Their contribution to species richness in a rocky littoral community

Mussels are important ecosystem engineers in marine benthic systems because they aggregate into beds, thus modifying the nature and complexity of the substrate. In this study, we evaluated the contribution of mussels (Brachidontes rodriguezii, Mytilus edulis platensis, and Perna perna) to the benthic species richness of intertidal and shallow subtidal communities at Cerro Verde (Uruguay). We compared the richness of macro-benthic species between mussel-engineered patches and patches without mussels but dominated by algae or barnacles at a landscape scale (all samples), between tidal levels, and between sites distributed along a wave exposition gradient. Overall, we found a net increase in species richness in samples with mussels (35 species), in contrast to samples where mussels were naturally absent or scarce (27 species). The positive trend of the effect did not depend upon tidal level or wave exposition, but its magnitude varied between sites. Within sites, a significant positive effect was detected only at the protected site. Within the mussel-engineered patches, the richness of all macro-faunal groups (total, sessile and mobile) was positively correlated with mussel abundance. This evidence indicates that the mussel beds studied here were important in maintaining species richness at the landscape-level, and highlights that beds of shelled bivalves should not be neglected as conservation targets in marine benthic environments.     

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.     

Using patterns of variability to test for multiple community states on rocky intertidal shores

Predictions based on theory of multiple stable states suggest that larger perturbations should lead to more unpredictable patterns of succession. This prediction was tested in the Gulf of Maine using data from 60 intertidal plots of varying size that were experimentally cleared of the rockweed Ascophyllum nodosum and from 14 benchmark sites from throughout the Gulf. Rockweed was removed from the experimental clearings ranging from 1 to 8 m in diameter in 1996 and data collected in 2004 were used to test effects of clearing size and location on divergence and variability in species composition. Benchmark data were collected in 2005, and the 14 sites were from a dataset on 53 sites throughout the Gulf of Maine. The selected sites were randomly chosen from all sites with > 80% canopy cover by A. nodosum and were expected to be similar to uncleared control plots from the experiment. Experimental removal of A. nodosum resulted in clearings at 12 sites within 4 bays. Abundances of gastropods, barnacles, mussels, and fucoid algae and the percentage cover of barnacles, mussels, fucoid algae, bare space, and other species were sampled. CAP and PERMDISP analyses revealed significant differences in multivariate dispersion and variability with both clearing size and location. Variability generally increased with clearing size and location effects were related to the north-south positioning of the sites. Benchmark sites were similar to the experimental control plots but as variable as the largest clearings. Results suggest that succession in larger clearings has been more unpredictable than in small clearings. The pattern of variability in the experimental clearings is consistent with the predictions of multiple stable states. However, the large amount of variation among the benchmark sites was due to mussels and was unexpected. This unexpected variability underscores the importance of sampling benchmark sites as part of experiments.     

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.     

Spatial patterns of growth in 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 variability in the region around PC affects the growth of the competitively dominant species in the rocky intertidal assemblage: the mussel Mytilus californianus. 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 oceanographic gradients in temperature, upwelling, wave exposure and productivity around PC would exert strong bottom–up influences on growth rates of mussels, potentially underlying large-scale differences in community structure around the PC region. We evaluated these predictions by measuring mussel growth rates across the PC region both in the intertidal and offshore on moorings. Intertidal mussels grew at much higher rates at sites south relative to north of PC and growth rates decreased in a gradient from south to north. The gradient in intertidal mussel growth around PC was uncorrelated with inshore concentrations of chlorophyll-a, and was most strongly correlated with the alongshore gradient in wave exposure and intertidal temperature. Mussels on moorings offshore from the intertidal sites grew at much higher rates than those in the corresponding intertidal areas, and mussel growth rates did not differ significantly among moored locations around PC. The gradient of increasing temperature from north to south among mooring sites was correlated with a decreasing gradient in productivity in the same direction, potentially contributing to equal and opposite effects on mussel growth at offshore moorings. This study suggests that environmental factors such as cold temperatures and high wave exposure contribute to the spatial pattern of decreasing mussel growth rates from south to north around PC, underlying large-scale patterns of community structure in this region.     

Isolated and combined impacts of blue mussels (Mytilus edulis) and barnacles (Balanus improvisus) on structure and diversity of a fouling community

Effects of two presumably dominant competitors, the blue mussel Mytilus edulis and the barnacle Balanus improvisus on recruitment, population dynamics and community structure on hard substrata were experimentally investigated in the subtidal Kiel Fjord, Western Baltic. The hypothesis that blue mussels and/or barnacles are local dominants and strongly influence succession and community structure was tested by monitoring succession in the presence and absence of simulated predation on either or both species. Manipulations included blue mussel removal, barnacle removal, combined blue mussel and barnacle removal, as well as a control treatment for natural (non-manipulated) succession. In the second part of the experiment, recovery from the treatments was monitored over 1 year.During the manipulative phase of the experiment, blue mussels had a negative effect on recruitment of species, whereas barnacles had no significant effect. Even so, a negative synergistic effect of blue mussels and barnacles was detected. Calculation of species richness and diversity H′ (Shannon Index) showed a negative synergistic effect of blue mussels and barnacles on community structure. Additionally, diversity H′ was negatively affected by the dominant competitor M. edulis. These effects were also detectable in the ANOSIM-Analysis. The non-manipulative phase of the experiment brought about a drastic loss of diversity and species richness. Blue mussels dominated all four communities. Barnacles were the only other species still being able to coexist with mussels. Effects of simulated predation disappeared fast.Thus, in the absence of predation on blue mussels, M. edulis within a few months dominates available space, and diversity of the benthic community is low. In contrast, when mussel dominance is controlled by specific predators, more species may persist and diversity remains high.     

Mussel and dogwhelk distribution along the north‐west Atlantic coast: testing predictions derived from the abundant‐centre model

Aim We performed the first test of predictions from the abundant‐centre model using north‐west Atlantic coastal organisms. We tested the hypotheses that the density of intertidal mussels (Mytilus edulis and M. trossulus) and dogwhelks (Nucella lapillus) and mussel age and size would peak at an intermediate location along their distribution range. We also assessed the latitudinal variation in critical aerial exposure time. Location North‐west Atlantic coast between Newfoundland (Canada) and New York (USA), covering 1800 km of shoreline. Methods Using a nested design, we measured mussel density, age and size and dogwhelk density in 60 wave‐exposed rocky intertidal sites spread evenly in six regions. Critical aerial exposure times were determined using online data. Results Mytilus edulis peaked in abundance in Maine and was much less abundant in the other regions. Mytilus trossulus peaked in abundance in southern Nova Scotia and Maine, was less abundant in the other regions to the north, and was absent in the southernmost region (New York). Both mussel species were least abundant in a northern region (Cape Breton), although not in the northernmost region (Newfoundland). Critical aerial exposure times were negatively correlated with overall mussel density. Mussel age and size were similar among regions. Dogwhelks peaked in abundance in Maine and were much less abundant in the other regions, being positively correlated with overall mussel density across regions. Main conclusions Density data for M. edulis and N. lapillus provide limited support for an abundant‐centre pattern, while M. trossulus shows a clear ramped‐south distribution. Critical aerial exposure times suggest that physiological stress during summer and winter low tides may be lowest in Maine and southern Nova Scotia, which might partially explain mussel predominance in those regions. Winter ice scour in Cape Breton may explain the abundance trough observed there. Mussel size and age may be more limited by wave exposure at our sites (as they all face open waters) than by regional differences in environmental stress. Dogwhelks, which prey on mussels, seem to respond positively to prey density at the regional scale. Our study supports the notion that, while the abundant‐centre model is a useful starting point for research, it often represents an oversimplification of reality.     

Blue mussels (Mytilus edulis) in the diet of roach (Rutilus rutilus) in outer archipelago areas of the western Gulf of Finland,Baltic Sea

The blue mussel (Mytilus edulis) is one of the key species in the Baltic Sea ecosystem and it is living at the edge of its range in the western Gulf of Finland. Roach (Rutilus rutilus) is a freshwater fish species that has benefited from recent coastal eutrophication and is at present highly abundant in the outer archipelago of the Gulf of Finland. In 2000 and 2001, a total of 516 roach were sampled for diet analysis in three study areas. Shelled molluscs formed over 95% of the diet of roach, blue mussels being the dominant single species. The proportion of this species in the food of roach in the three study areas ranged between 38–61% for smaller roach (<225 mm) and 39–85% for larger (>225 mm) roach, indicating that blue mussel is a highly important food source for roach in outer archipelago areas of the western Gulf of Finland, in contrast to reports from other parts of the northern Baltic Sea. The scarcity of large blue mussels in mussel beds in the easternmost study area was reflected in the lower proportion of blue mussels in the diet of larger roach. However, the growth of roach was not affected by the availability of blue mussels. The twofold differences observed in the annual growth of roach between warm and cold years demonstrated that temperature is an important factor controlling the growth of roach in the western Gulf of Finland.     

Using polyclonal antibodies to detect predation by dogwhelks Nucella lapillus (L.)

Interactions between predators and their multiple prey species can vary greatly among locations where they coexist. As a method to assess spatial variation in predation by intertidal dogwhelks on their dominant prey, immunoassays of dogwhelk gut contents from experimental populations and field collected individuals were evaluated using polyclonal antibodies raised separately to soluble proteins from Mytilus edulis L. mussels and Semibalanus balanoides (L.) barnacles. Both antisera produced strong reactions against their homologous antigens but no cross reactions between prey species. Experimental trials tested the critical hypothesis that prey species had equal detection intervals in dogwhelk guts. Two groups of 225 dogwhelks were starved for 14 days, provided with either mussels or barnacles for five days, and then sampled over 22 days. Independent immunoassays of dogwhelk gut contents against each antibody revealed a consistent, weak cross reaction between the anti-mussel antibody and dogwhelk gut tissues. After accounting for this cross reaction, the strength of immunoassays against both prey species declined exponentially and at similar rates. The proportions of dogwhelks that tested positive for their provided prey species declined linearly through time and were not significantly influenced by prey type. Prey were detectable throughout the sampled post-feeding period and were projected to have detection limits of 24.4 days (barnacles) and 26.5 days (mussels), demonstrating that immunoassay results are not biased by dissimilar prey detection intervals. Reactions against the antibody from the non-provided prey were time invariant and occurred at relatively low frequencies. Immunoassays of dogwhelks collected from five intertidal sites on Swans Island, Maine, USA revealed patterns similar to field observations, though immunoassays classified far fewer individuals as non-feeders and more as barnacle feeders than indicated by direct field observations. Unlike single observations, immunoassays also revealed the presence of both prey in dogwhelks from four sites, though most individuals tested positive for only a single prey type. Immunoassays facilitate concurrent collections of predation data from many individuals and will enable further local- to regional-scale assessments of dogwhelk predation at additional sites around the Gulf of Maine.     

Community dynamics of intertidal soft-bottom mussel beds over two decades

Macrozoobenthos communities in the North Sea showed pronounced changes over the past decade in relation to an increasing number of invasive species and climate change. We analysed data sets spanning 22 years on abundance, biomass and species composition of intertidal soft bottom mussel beds near the island of Sylt (German Bight) in the Northern Wadden Sea, based on surveys from 1983/1984, 1990, 1993 and from 1999 to 2005. Mussel bed area and blue mussel biomass decreased, and a change in the dominance structure in the associated community comparing 1984 to mid-1990s with the period from 1999 to 2005 was observed. Coverage of the mussel beds with the algae Fucus vesiculosus decreased since the end of the 1990s. Within the study period biomass and densities of the associated community increased significantly. Dominance structure changed mainly because of increasing abundances of associated epibenthic taxa. Apart from the Pacific oyster Crassostrea gigas all other alien species were already present in the area during the study period. Community changes already started before Pacific oysters became abundant. An attempt is made to evaluate effects on the observed changes of decreasing mussel biomass, ageing of mussel beds, decreasing fucoid coverage and increasing abundances of invader. All four factors are assumed to contribute to changing community structure of intertidal mussel beds.     

Top-down and bottom-up factors in tidepool communities

Recent studies suggest that nutrient variation influences rocky intertidal community structure, however empirical evidence is rare. In the Gulf of Maine, tidepools that occur on seagull feeding roosts are potentially subjected to regular nutrient loading from seagull guano. The results of a survey conducted on Swan's Island, ME show that roost tidepools have very low macroinvertebrate and macroalgal diversity as well as very high phytoplankton biomass compared to non-roost tidepools. An experiment presented here tested basic food chain hypotheses in tidepool communities. These basic food chain models predict that in a tidepool with one trophic level (phytoplankton only), phytoplankton biomass will increase when nutrients are enriched. In contrast, these models predict that in two trophic level tidepools (phytoplankton and mussels) herbivory will prevent an increase in phytoplankton biomass when nutrients are enriched. A short term 2×2 factorially designed field experiment was used to test this basic conceptual model using herbivory by mussels and enrichment with nitrogen as the main effects. The results of this investigation are consistent with the predictions of basic food chain models, and indicate that over the short time interval of a few days, herbivory by mussels is sufficient to maintain low phytoplankton levels following enrichment with nitrogen. Experimental enrichment with phosphorus in this study had no effect on phytoplankton biomass. The results of this study suggest that periodic pulses of nitrogen into tidepools will have little effect on phytoplankton biomass when mussels are present and that longer-term chronic nitrogen influxes may be driving the patterns of community structure in tidepools occurring on roosts.     

The influence of pre‐settlement and early post‐settlement processes on the adult distribution and relative dominance of two invasive mussel species

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Experimental confirmation of multiple community states in a marine ecosystem

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