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.     

Inshore–offshore comparison of parasite infestation in Mytilus edulis: implications for open ocean aquaculture

Offshore production offers a new perspective for mussel aquaculture in the German Bight (North Sea) as no expansion is possible in the intertidal and subtidal zone of the Wadden Sea because of restrictions on the number of licenses. The development of offshore wind farms offers a unique opportunity because of the associated infrastructure. Service platforms, as well as the pylons themselves, offer perfect structures for mooring longlines and other culture units. One of the advantages of offshore culture may be a lower parasite load in offshore mussels compared with mussels produced under traditional inshore bottom culture. By sampling mussel spat from offshore suspended buoys or collectors, we simulated an offshore culture situation and compared parasite infestation rates with those in mussel spat obtained from suspended inshore buoys or collectors, in mussels from inshore benthic subtidal beds and from inshore benthic intertidal mussel beds. Mussels from offshore sites were free of trematodes and shell‐boring polychaetes. Parasitic copepods only occurred at a single offshore site, on a 20‐year‐old research platform, but not on buoys or collectors exposed for shorter time periods. All three monitored parasite taxa were present at all other sites. The highest prevalence was found for trematodes in inshore benthic intertidal mussels (78.7 ± 6.4%) and locally reached 100%. Through a variety of detrimental effects, trematodes, parasitic copepods and shell‐boring polychaetes are known to affect growth performance and product quality. We therefore propose that offshore mussel production could be a promising culture procedure because it seems to result in lower parasite burden than at traditional culture sites. Whether offshore production also results in better survival and growth, compared with inshore mussel culture on a commercial scale, needs to be investigated further.     

The surf zone: a semi-permeable barrier to onshore recruitment of invertebrate larvae?

The supply of larvae to the shore is important for population replenishment and intertidal community dynamics but its variability at most scales is not well understood. We tested the relationship between nearshore mussel larval abundance and intertidal settlement rates over several years at multiple spatiotemporal scales in Oregon and New Zealand. Abundance of competent larvae nearshore and intertidal recruitment rates were simultaneously quantified using collectors mounted at different depths on moorings 50-1100 m from shore, and at adjacent rocky intertidal sites. Total mussel larval abundance and oceanographic conditions were also measured in some locations. At all scales, abundance of nearshore mussel larvae was unrelated to intertidal recruitment rates. In the intertidal, patterns of mussel recruitment were persistent in space, with sites of consistently high or low recruitment. In contrast, nearshore competent larval abundance showed generally similar abundances among sites except for a high, and spatially-inconsistent, variability in Oregon during 1998 only. On moorings, recruitment tended to be greater on midwater collectors than shallower or deeper. Finally, on moorings larval abundance in traps and recruitment on collectors was unrelated. These results suggest that (1) among sites, the size of the nearshore larval pool is relatively uniform while onshore recruitment varies and is unrelated to larval abundance, (2) temporal variability in nearshore larval availability is not strongly expressed onshore, (3) vertical stratification of competent larvae nearshore is strong and may influence transport and recruitment, and (4) within-coast variability in onshore recruitment is strongly driven by processes occurring locally within the surf zone that need to be studied to understand coastal recruitment dynamics.     

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.     

Mussel disturbance dynamics: signatures of oceanographic forcing from local interactions

Local interactions, biotic and abiotic, can have a strong influence on the large-scale properties of ecosystems. However, ecological models often explore the influence of local biotic interactions where physical disturbance is included as a large-scale and imposed source of variability but is not allowed to interact with biotic processes at the local scale. In marine intertidal communities dominated by mussels, wave disturbances create gaps in the mussel bed that recover through a successional sequence. We present a lattice model of mussel disturbance dynamics that allows local interactions between wave disturbance and mussel recolonization, in which each cell of the lattice can be empty, occupied by a mussel bed element, or disturbed (which corresponds to a newly disturbed cell that has unstable edges). As in natural ecosystems, wave disturbance can also spread from disturbed to adjacent occupied cells, and recolonization can also spread from occupied to adjacent empty cells. We first validate the local rules from artificial gap experiments and from natural gap monitoring along the Oregon coast. We analyze the properties of the model system as a function of different oceanographic forcings of productivity and disturbance. We show that the mussel bed can go through phase transitions characterized by a large sensitivity of mussel cover and patterns to oceanographic forcings but also that criticality (scale invariance) is observed over wide ranges of parameters, which suggests self-organization. We also show that spatial patterns in the intertidal can provide a robust signature of local processes and can inform about oceanographic regimes. We do so by comparing the large-scale patterns of the simulation (scaling exponents) with field data, which suggest that some experimental sites are close to criticality. Our results suggest that regional patterns in disturbed populations can be explained by local biotic and abiotic processes submitted to oceanographic forcing.     

Topographically generated fronts, very nearshore oceanography and the distribution and settlement of mussel larvae and barnacle cyprids

Flow patterns adjacent to shore may prevent or aid shorewardmigration of benthic invertebrate larvae. We hypothesized thata front at the mouth of Sunset Bay, Oregon, prevents shorewarddispersal of larvae, significantly altering settlement of mussellarvae and barnacle cyprids. Settlement was measured at threesets of moorings (three moorings per site) distributed acrossthe front at Sunset Bay. From 6 July to 4 September 2000, sampleswere collected roughly every other day. Concurrently, we madevertical zooplankton tows adjacent to each mooring site andcollected physical oceanographic data. During upwelling-favorablewinds, the front was always present at the bay mouth, separatingsignificantly cooler, saltier and denser offshore water fromthat within the bay. During downwelling winds, the front brokedown and we found no significant difference in the surface physicaloceanographic parameters across the bay mouth. During upwelling,the concentration of mussel larvae was higher seaward of thefront than landward, but there was no significant differencein concentration during downwelling, suggesting that the frontmay act as a barrier to the shoreward dispersal of mussels.Mussel settlement was too low and sporadic to allow statisticalanalysis. There was no difference in cyprid concentrations acrossthe bay mouth whether the front was present or not. Cyprid settlementwas, however, nearly an order of magnitude lower at mooringsseaward of the front than at those landward. A significant cross-correlationwas found between settlement at the offshore mooring and tidalrange (r = 0.464, lag = 0 days) and between settlement at themid and inner moorings and downwelling winds (r = 0.532 midbay, r = 0.532 inner bay, lag = 0 days). Seaward of the front,settlement varied with tidal range, while landward of the front,most settlement occurred as brief pulses during downwellingwinds, periods when the front was not present. We found largedifferences in the distribution of cyprids, and mussel larvaeand cyprid settlement relative to the front; larval distributionsand settlement varied with upwelling versus downwelling windsand was due to differences in the very nearshore (i.e. within100–1000 m of shore) coastal oceanography.     

A GRADIENT IN BENTHIC INTERTIDAL ALGAL ASSEMBLAGES ALONG THE SOUTHERN CALIFORNIA COAST1

Based on an analysis floristic data, a gradient appears to exist in the composition of intertidal algae along the 450 km of southern California coastline immediately south of Point Conception. Reciprocal averaging ordination of the algal flora at 51 sites in this area suggests that the gradient is not strictly latitudinal. Variation from a latitudinal gradient occurs ca. 60-80 km south of Point Conception. An ordination of frequently occurring species indicated that compositional changes are substantial when sites at the extreme north and south are compared, but that the changes are gradual and continuous. Possible explanations for the gradient are discussed, and these include: sea temperature, upwelling, sand movement, human disturbance (i.e., pollution, trampling), and wave action.     

The combination of selection and dispersal helps explain genetic structure in intertidal mussels

Understanding patterns of gene flow, selection and genetic diversity within and among populations is a critical element of predicting how long-term changes in environmental conditions are likely to affect species distribution. The intertidal mussel Perna perna consists of two distinct genetic lineages in South Africa, but the mechanisms maintaining these lineages remains obscure. We used regional oceanography and lineage-specific responses to environmental conditions as proxies for gene flow and local selection, respectively, to test how these mechanisms could shape population genetic structure. Laboratory experiments supported the field findings that mussels on the east coast (eastern lineage) are physiologically more tolerant of sand inundation and high temperatures than those on the south coast (western lineage). Temperature loggers mimicking mussel body temperatures revealed that mussels experience higher body temperatures during aerial exposure on the subtropical east coast than on the temperate south coast. Translocations showed that, on the east coast, the western lineage suffered higher mortality rates than local individuals, while on the south coast, mortality rates did not differ significantly between the lineages. Nearshore drogues showed remarkably little overlap between the trajectories of drifters released off the south coast and those released off the east coast. Physiological tolerances can thus explain the exclusion of western individuals from the east coast, but they cannot explain the exclusion of the eastern lineage from the south coast. In contrast, however, ocean dynamics may limit larval dispersal between the two lineages, helping to explain the absence of eastern individuals from the south coast. We emphasise the importance of a multidisciplinary approach in a macro-ecological context to understand fully the mechanisms promoting evolutionary divergence between genetic entities. Our results suggest that phylogeographic patterns of Perna perna may be maintained by a combination of local conditions and the isolating effect of the Agulhas Current that reduces gene exchange.     

Environmental correlates of phenotypic variation: do variable tidal regimes influence morphology in intertidal seaweeds?

Seaweed morphology is often shaped by the hydrodynamic environment. However, exposure to air at low tide represents an additional factor potentially affecting the morphology of intertidal species. Here, we examined the relationships between the morphology of Hormosira banksii, an important intertidal habitat‐forming seaweed in southern Australia, and environmental factors across multiple spatial scales around the island of Tasmania, Australia. Tasmania is surrounded by a diverse coastline with differences in wave exposure, tidal parameters, and temperature. We sampled Hormosira from four regions (100s km apart), three sites (10s km apart) within each region, and two zones (meters apart; eulittoral and sublittoral) at each site, and measured multiple morphological variables to test for differences in morphology at those different spatial scales. Thirteen environmental variables reflecting wave exposure, tidal conditions, and temperature for each site were generated to assess the relationship between Hormosira morphology and environmental variation. Morphology varied at all spatial scales examined. Most notably, north coast individuals had a distinct morphology, generally having smaller vesicles and shorter fronds, compared to other regions. Tidal conditions were the main environmental factors separating north coast sites from other sites and tidal regime was identified as the best predictor of morphological differences between regions. In contrast to other studies, we found little evidence that wave exposure was associated with morphological variation. Overall, our study emphasizes the role of tidal conditions, associated with emersion stress during low tide, in affecting the morphology of intertidal seaweeds.     

Gradients of intertidal primary productivity around the coast of South Africa and their relationships with consumer biomass

The structure of rocky intertidal communities may be influenced by large-scale patterns of productivity. In this study we examine the in situ rates of production by intertidal epilithic microalgae (chlorophyll a production per unit area per month), intertidal nutrient concentrations (nitrates, nitrites, phosphates and silicates), and standing stocks of different functional-form groups of macroalgae around the South African coast, and their relationships to consumer biomass. Clear gradients of in situ intertidal primary production and nutrient concentrations were recorded around the South African coast, values being highest on the west coast, intermediate on the south and lowest on the east coast. Primary production by intertidal epilithic microalgae was correlated with nutrient availability and could also be related to nearshore phytoplankton production. The dominance patterns of different functional forms of macroalgae changed around the coast, with foliose algae prevalent on the west coast and coralline algae on the east coast. However, overall macroalgal standing stocks did not reflect the productivity gradient, being equally high on the east and west coasts, and low in the south. Positive relationships existed between the average biomass of intertidal intertebrate consumers (grazers and filter-feeders) and intertidal productivity, although only the grazers were directly connected to in situ production by epilithic intertidal microalgae. The maximum body size of a widely distributed limpet, Patella granularis, was also positively correlated with level of in situ primary production. The maximal values of biomass attained by intertidal filter-feeders were not related to intertidal primary production, and were relatively constant around the coast. At a local scale, filter-feeder biomass is known to be strongly influenced by wave action. This implies that the local-scale water movements over-ride any effects that large-scale gradients of primary production may have on filter-feeders. The large-scale gradient in intertidal productivity around the coast is thus strongly linked with grazer biomass and individual body size, but any effect it has on filter-feeder biomass seems subsidiary to the local effects of wave action.     

Shell morphometrics and growth rate of the invasive bivalve mollusc Mytilus galloprovincialis in the Knysna estuarine embayment,South Africa

Shell morphometrics of the invasive mussel Mytilus galloprovincialis were compared at five sites and growth rate at four sites (in four seasons) in the Knysna estuarine embayment. Mussels from two sites (The Heads, Leisure Isle) where wave action was present had shells significantly lower for any length when compared with other more sheltered sites. There was no significant difference in shell width of mussels for any given length among sites. Mussels from The Heads had thicker shells than other sites, and those from Leisure Isle thicker shells than three other embayment sites where shells did not differ in thickness. Growth rate of mussels at two embayment sites (Thesen’s Wharf and Thesen Islands Marina) was greatest in autumn and summer whereas at The Heads and Leisure Isle there was little seasonal difference in growth rate. Growth rate of mussels at Thesen’s Wharf and Thesen Islands Marina was mainly greater in all seasons when compared with mussels at The Heads and Leisure Isle. The more rapid growth rate of mussels at the sheltered embayment sites might in part explain why M. galloprovincialis now dominates the mid- to lower intertidal on hard substrata in this region of the Knysna estuary.     

Can we predict the effects of alien species? A case-history of the invasion of South Africa by Mytilus galloprovincialis (Lamarck)

The Mediterranean mussel Mytilus galloprovincialis invaded the shores of South Africa in about the mid-1970s. It was first detected at a harbour in Saldanha on the west coast and apparently arrived accidentally. From there, it spread at a rate of about 115 km·year⁻¹ and now occupies the whole of the west coast of South Africa and at least the southern half of Namibia. It was deliberately introduced from the west coast to the south coast for mariculture. In this case study, we record its effects on intertidal rocky shores, cast in terms of predictions based on (a) the history of its invasions elsewhere, (b) its mode of dispersal, (c) its physiological performance relative to indigenous mussels, (d) the role of wave action as a moderator of competition, (e) the influence of relative body sizes, (f) the projected effects of the mussel on infauna, (g) consumption by higher trophic levels, and (h) rates of parasitism.

Several properties of M. galloprovincialis itself, and of the recipient community, conspired to favour the spread and establishment of this alien mussel, including high productivity on the west coast of South Africa, prevalently strong wave action, a sparsity of predators, an absence of parasites, the mussel's fast growth and high reproductive output, and its possession of a planktotrophic larva. It competitively displaces several species because of its physiological performance. Some of the species gain a substitute substratum on the mussels themselves, but only if they are small enough to live and reproduce on the mussels. M. galloprovincialis has had little effect on infauna, but has provided an additional source of food for higher predators, including the rare and endangered African Black Oystercatcher (Haematopus moquini).

Nearly all these effects and conditions were forecasted (or, at the least, explainable with hindsight), but despite these successes in predicting the impacts of M. galloprovincialis, its spread was not only unavoidable but was encouraged by its transfer to the south coast for mariculture. Moreover, there was one completely unpredictable effect of M. galloprovincialis—which led to mass mortalities of a swimming crab. Given the failure of even quite detailed and accurate predictions to allow control of M. galloprovincialis once it arrived, prevention rather than cure must be the prime means of avoiding future unwanted introductions of invasive species.     

128 Capturing the Light Fantastic: Oceanographic and Tidal Influences on Intertidal Algae

Seaweeds experience many challenges to their persistence in intertidal zone habitats. Their growth rates must exceed losses associated with a range of ecological and physiological factors including desiccation, herbivory and wave forces. Growth rates depend on an alga's ability to capture and process light to build carbon-based molecules. We examined local (tidal height) and large (oceanographic) scale influences on algal photosynthetic efficiency and light climate, respectively. At the local scale, we combined periodic measurements of physiological state using PAM fluorometry with traditional demographic monitoring of a Postelsia palmaeformis , population over a tidal height gradient. Parameter estimates derived from rapid fluorescence-irradiance curves were correlated with longer-term ecological performance measures including growth rate, morphology, survivorship and reproductive output. At larger scales, we made continuous in situ measurements of chlorophyll fluorescence and light attenuation in the intertidal zone at six sites during 2001 and 2002. Light attenuation to the benthos was sharply reduced at sites when chl-a fluorescence was high. Long-term, large-scale monitoring of intertidal zone chl-a and macroalgal abundances documents that striking differences among sites are persistent and associated with oceanographic factors. The light saturation parameter and maximum photosynthetic rate calculated for several common intertidal macrophytes, along with published values of the irradiance needed to saturate their growth rates, suggest that underwater light levels may limit intertidal algal growth where phytoplankton blooms are common and persistent. We conclude that physiological stress associated with tidal and oceanographic factors contribute to macroalgal distributions.     

Biogeographical patterns of rocky intertidal communities along the Pacific coast of North America

Aim Our aim in this paper is to present the first broad‐scale quantification of species abundance for rocky intertidal communities along the Pacific coast of North America. Here we examine the community‐level marine biogeographical patterns in the context of formerly described biogeographical regions, and we evaluate the combined effects of geographical distance and environmental conditions on patterns of species similarity across this region. Location Pacific coast of North America. Methods Data on the percentage cover of benthic marine organisms were collected at 67 rocky intertidal sites from south‐eastern Alaska, USA, to central Baja California Sur, Mexico. Cluster analysis and non‐metric multidimensional scaling were used to evaluate the spatial patterns of species similarity among sites relative to those of previously defined biogeographical regions. Matrices of similarity in species composition among all sites were computed and analysed with respect to geographical distance and long‐term mean sea surface temperature (SST) as a measure of environmental conditions. Results We found a high degree of spatial structure in the similarity of intertidal communities along the coast. Cluster analysis identified 13 major community structure ‘groups’. Although breaks between clusters of sites generally occurred at major biogeographical boundaries, some of the larger biogeographical regions contained several clusters of sites that did not group according to spatial position or identifiable coastal features. Additionally, there were several outliers – sites that grouped alone or with sites outside their region – for which localized features may play an important role in driving community structure. Patterns of species similarity at the large scale were highly correlated with geographical distance among sites and with SST. Importantly, we found community similarity to be highly correlated with long‐term mean SST while controlling for the effects of geographical distance. Main conclusions These findings reveal a high degree of spatial structure in the similarity of rocky intertidal communities of the north‐east Pacific, and are generally consistent with those of previously described biogeographical regions, with some notable differences. Breaks in similarity among clusters are generally coincident with known biogeographical and oceanographic discontinuities. The strong correlations between species similarity and both geographical position and SST suggest that both geography and oceanographic conditions have a large influence on patterns of intertidal community structure along the Pacific coast of North America.     

Does rock type account for variation in mussel attachment strength? A test with <Emphasis Type="Italic">Brachidontes rodriguezii</Emphasis> in the southwestern Atlantic

Mussel attachment strength varies in space and time, frequently in association with variations in wave exposure. Yet, it remains uninvestigated whether different rock types can contribute to variation in mussel attachment. Here we compared the attachment strength of the mussel Brachidontes rodriguezii between soft and hard intertidal rock substrates that are typical of coastal Buenos Aires Province, Argentina: Pampean loess cemented by calcium carbonate and orthoquartzite, respectively. Overall comparisons of mussel attachment across natural platforms of either rock type (10 loess sites and 4 orthoquartzite sites) indicated stronger mussel attachment to orthoquartzite. However, mussel attachment strength did not differ when compared across natural loess platforms and introduced orthoquartzite blocks (i.e., groins and revetments) occurring within the same site. Mussels attaching to loess showed more byssal threads than those attaching to orthoquartzite at the same site. These findings suggest, first, that rock type does not influence mussel attachment strength in our study system, secondly, that overall differences in mussel attachment strength with rock type across natural platforms in our study range are due to confounding influences of co-varying factors (e.g., wave exposure) and, finally, that mussels can increase byssus production to counteract potential substrate failure when attaching to soft, friable rock. The latter likely explains the ability of mussels to maintain relatively stable cover across rocks of contrasting hardness.     

Climate‐driven trends and ecological implications of event‐scale upwelling in the California Current System

Eastern boundary current systems are among the most productive and lucrative ecosystems on Earth because they benefit from upwelling currents. Upwelling currents subsidize the base of the coastal food web by bringing deep, cold and nutrient‐rich water to the surface. As upwelling is driven by large‐scale atmospheric patterns, global climate change has the potential to affect a wide range of significant ecological processes through changes in water chemistry, water temperature, and the transport processes that influence species dispersal and recruitment. We examined long‐term trends in the frequency, duration, and strength of continuous upwelling events for the Oregon and California regions of the California Current System in the eastern Pacific Ocean. We then associated event‐scale upwelling with up to 21 years of barnacle and mussel recruitment, and water temperature data measured at rocky intertidal field sites along the Oregon coast. Our analyses suggest that upwelling events are changing in ways that are consistent with climate change predictions: upwelling events are becoming less frequent, stronger, and longer in duration. In addition, upwelling events have a quasi‐instantaneous and cumulative effect on rocky intertidal water temperatures, with longer events leading to colder temperatures. Longer, more persistent upwelling events were negatively associated with barnacle recruitment but positively associated with mussel recruitment. However, since barnacles facilitate mussel recruitment by providing attachment sites, increased upwelling persistence could have indirect negative impacts on mussel populations. Overall, our results indicate that changes in coastal upwelling that are consistent with climate change predictions are altering the tempo and the mode of environmental forcing in near‐shore ecosystems, with potentially severe and discontinuous ramifications for ecosystem structure and functioning.     

Mesoscale advection of Upogebia pusilla larvae through an upwelling filament in the Canaries Coastal Transition Zone (CTZ)

Larval transport has a strong influence on marine populations and ecosystem function. Traditional hypotheses establish that larvae of coastal species are swept offshore during strong upwelling periods producing low recruitment rates; however, recent studies have demonstrated that this hypothesis is not supported by larval distribution data. The present study examines the influence of upwelling filaments on larval advection and wastage. During August 1993, the transport of Upogebia pusilla was analysed in relation to an offshore filament off Northwest Africa. This mesoscale structure was generated by a trapped, quasi-permanent cyclonic eddy located between the Canary Islands and the African shelf and extended 150?km westward into the oligotrophic open ocean waters. The horizontal distribution depicted by U. pusilla larvae was strongly influenced by the oceanographic features of the Canaries Coastal Transition Zone (CTZ). Specifically, the larvae were closely associated with upwelling front and filament position, showing that these neritic larvae can be advected 10–100s of kilometres offshore through the filament. However, the low larval densities observed in the samples suggest that many larvae might remain close to the coast, thus avoiding seaward transport. This implies that filaments probably do not significantly affect the recruitment success of intertidal invertebrates such us U. pusilla in upwelling systems.     

Effect of air exposure on lysosomal tissues of Mytilus edulis L. from natural intertidal wild beds and submerged culture ropes

Blue mussels collected from suspended culture ropes and from three natural intertidal wild beds from different areas of the German Bight were tested for their ability to cope with hypoxic conditions. During the experiment mussels were exposed to air from 0 to 72 h. Mussels from all sampling sites displayed high tolerance to aerial exposure with moderate levels of mortality after 12 to 48 h of exposure. Lysosomal membrane stability (LMS), a biomarker of general stress, changed notably between minimum values after 12 h and maximum values after 24 h of aerial exposure in intertidal mussels. In contrast, labilization times of mussels from the hanging culture increased continuously up to 48 h of exposure. Intertidal mussels from the island of Heligoland exhibited significantly decreased membrane stability after 72 h of air exposure, correlating to higher mortality rates. Intertidal mussels, although adapted to daily aerial exposure in their natural environment, showed a similar pattern of mortality and lower LMS values during the experiment than mussels from the suspended culture site. The increase of LMS values of mussels under hypoxic conditions at the beginning of the experiment at all sites was tested for the influence of macro-autophagic processes using immune labelling techniques. With this approach it could be demonstrated that high LMS values significantly correlate with low autophagic activity. However, hypoxic conditions do not enhance autophagic processes during the early periods of aerial exposure. Only at the end of the experiment, high values for autophagy were measured in mussels from an intertidal site accompanied with high mortalities. The results indicate that autophagic processes are not involved in the early adaptive processes that enable the mussel to cope with periods of aerial exposure.     

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.     

Evidence of cannibalism and bentho-pelagic coupling within the life cycle of the mussel, Perna canaliculus

The feeding ecology of the green-lipped mussel, Perna canaliculus, was investigated within three intertidal mussel beds along Ninety Mile Beach, northern New Zealand, between August 2000 and March 2001. Adult mussels of different sizes (45-105 mm in shell length) were collected from the intertidal sites about 30 min after being submerged by the incoming tide for gut content analyses. Results of these analyses indicate that mussels consume a variety of phytoplankton, micro- and mesozooplankton, including mussel larvae and post-larvae. Cannibalism of juveniles of up to 620 μm was recorded for intertidal mussels, and conspecifics of up to 2.4 mm were found within the stomachs of additional mussels collected in August 2000 from a nearby subtidal site. For all three intertidal populations, mussel larvae and juveniles contribute about 70% of the food particle consumption during the spawning peak in August, while phytoplankton and other zooplankton constitute the majority of the food source (about 99%) in March, during gametogenesis. Larger intertidal mussels tended to have more food particles in their stomachs than smaller mussels within all three populations. Distinctive differences in food consumption among intertidal populations directly coincide with variations in total particulate matter (TPM), particulate organic matter (POM) and percent organic matter (OM) in the adjacent seawater.Separate experiments designed to test the feeding behavior of mussels feeding at different times during the incoming tide were conducted at one of the intertidal sites during August 2000 and March 2001. Results from these experiments indicate a marked shift in food consumption from bivalves to other mesozooplankton in August, and from phytoplankton to mesozooplankton in March. The observed combination of mussel predatory and grazing behavior over the incoming tide and through the year provides evidence for a strong food-web link between the benthic and pelagic life stages of this species. Furthermore, the high rate of cannibalism during some months of the year suggests that this source of food may significantly contribute to the energy budget of wild populations, with potential implications for evolutionary adaptive success.