Larval recruitment of the blue mussel Mytilus edulis: The effect of flow and algae

The mussel Mytilus edulis settlement and distribution was studied on plastic panels with manipulated flow regime (faired, bluff, split and angled) with or without water soluble metabolites of the green alga Cladophora rupestris. The panels were exposed vertically on a device (hydrovane) that ensures their constant orientation in the current during the peak of larval settlement at 1 m depth. In order to investigate larval distribution on the panels, half of them were coated with a silicone vacuum grease that prevents larvae from de-attachment. This grease was not toxic and did not attract or repel larvae. Low densities of larvae on the un-greased plates compared to the greased ones suggested that some of larvae left the substratum. The blue mussel larvae initially settled in regions of reduced shear velocity and then redistribute to the regions of high shear velocity. The presence of the alga increased the density of blue mussel larvae and changed their distribution on the panels. Overall, our results demonstrated that larval recruitment of M. edulis is an active process affected both by boundary-layer hydrodynamics and algal waterborne compounds.     

Seasonal and interannual dynamics of larval abundance of <Emphasis Type="Italic">Mytilus coruscus</Emphasis> Gould, 1861 (Bivalvia: Mytilidae) in Amursky Bay (Peter the Great Bay,Sea of Japan)

Based on an analysis of the gonads, as well as the size composition and abundance of larvae in plankton, we determined the seasonal and long-term dynamics of the reproduction of the mussel Mytilus coruscus in Amursky Bay (Peter the Great Bay, Sea of Japan). Mussel gonads were in a mature condition in late July-early August; the spawning season continued from mid-August to mid-September. The larvae of this species occurred in plankton from June to September, under surface water temperatures of 14.5–23.5°C. The density of larvae in different years varied from 350 to 4500 ind/m³. Two peaks of larval abundance were observed annually, in June and September. The first peak was apparently provided by allochthonous larvae, the second, by larvae of the local population. The high density of larvae of M. coruscus in the plankton of Peter the Great Bay enabled us to consider this species as a potential object for mariculture.     

Pharmacological Induction of Larval Settlement and Metamorphosis in the Blue Mussel Mytilus edulis L.

The blue mussel Mytilus edulis L. is an important aquaculture and fouling species in northern seas. Although the general role of chemical cues for settlement of larvae of the blue mussel has been proposed, few studies have focused on induction of settlement and metamorphosis by pharmacological agents. In this study, the induction of larval settlement of the blue mussel by pharmacological compounds was investigated through a series of laboratory experiments with an aim of identifying artificial cues for laboratory bioassay systems in fouling and antifouling research. Gamma-aminobutiric acid (GABA), dihydroxyphenyl L-alanine (DOPA), isobutyl methylxanthine (IBMX) and acetylcholine chloride (ACH) at 10^{m 7}-10^{m 2} M as well as KCl at 10-40 mM K⁺ in excess of the level in normal seawater were tested for their inductive effect on larval settlement. In filtered seawater (FSW) <9% of the larvae settled after 48 h. Elevated K⁺ and GABA levels had no effect on larval settlement and metamorphosis. DOPA at 10^{m 5} M and IBMX at 10^{m 6}-10^{m 4} M induced 41-83% larval settlement and ACH at 10^{m 7}-10^{m 5} M induced < 40% larval settlement. While the highest settlement rates were observed after 48 h exposure to the chemicals, most of the larvae settled within 24 h. Compounds at concentrations of 10^{m 3}-10^{m 2} M were either toxic to larvae or retarded the growth of the post-larvae shell. Juveniles resulting from induction by lower concentrations of chemicals had a very high survival rate, completed metamorphosis and grew as well as the juveniles that metamorphosed spontaneously. IBMX at 10^{m 6}-10^{m 4} M and L-DOPA at 10^{m 5} M are effective agents for induction of settlement and metamorphosis for future studies using juvenile M. edulis.     

Density Dynamics and Size Composition of Bivalve Mollusk Larvae of the Genus Mya in Peter the Great Bay (Sea of Japan)

The seasonal population size of planktonic larvae of mollusks of the soft-shell clam genus Mya were studied in Vostok Bay. The larvae occurred in plankton from the first third of June to early September. The highest density of larvae (279.7 ± 98.7 specimens/m³) was found in late June. In early July, larval numbers decreased sharply, and at the end of the first third of July, only single Mya larvae were found in the plankton. Another increase in larval population was observed in early and late August. The larvae varied in length from 175 to 325 m in June–July and from 150 to 250 m in August. It is likely that the larvae of two species of Mya, M. arenaria and M. uzenensis, are present in the plankton of Vostok Bay.     

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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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.     

Age-dependent zonation of the periwinkle Littorina littorea (L.) in the Wadden Sea

On sedimentary tidal flats near the island of Sylt (German Bight, North Sea) abundance and size distribution of periwinkles, Littorina littorea L., were studied in low intertidal and in shallow and deep subtidal mussel beds (Mytilus edulis L.). In low intertidal mussel beds, surveys revealed that high densities (1,369±571 m^–2) of juvenile snails (≤13 mm) were positively correlated with strong barnacle epigrowth (Semibalanus balanoides L. and Balanus crenatus Bruguière) on mussels. A subsequent field experiment showed that recruitment of L. littorea was restricted to the intertidal zone. Abundances of periwinkles (213±114 m^–2) and barnacles abruptly decreased in the adjacent shallow subtidal zone, which served as a habitat for older snails (>13 mm). L. littorea was completely absent from disjunct deep (5 m) subtidal mussel beds. Snail abundance varied seasonally with maxima of >4,000 m^–2 in low intertidal mussel beds in October and minima in July, just before the onset of new recruitment. I suggest that the presence of cracks and crevices among the dense barnacle overgrowth in intertidal mussel beds favoured recruitment and survival of juvenile snails. Larger (older) specimens are assumed to actively migrate to the less favourable adjacent subtidal. Therefore, intertidal mussel beds are considered as nurseries for the population of L. littorea in the Wadden Sea. Received in revised form: 25 September 2000 Electronic Publication     

Reproduction and early life history of an invasive ascidian Styela clava Herdman in Prince Edward Island, Canada

In the past decade, many aquatic invasive species have been introduced into Prince Edward Island, Canada, coastal waters, and they have had adverse effects on the aquaculture and fisheries industry. For instance, the mussel aquaculture industry has been overwhelmed by extremely high abundances of the invasive ascidian Styela clava in some areas. To understand and minimize the risk of spread of S. clava to other shellfish aquaculture sites, basic information on the reproductive biology of the ascidian is needed. This study investigated Styela gonad development, larval abundance and recruitment. Gonad development, using histological methods, indicated ovaries were ripe from late June through late October, 2004. Weekly water samples indicated larvae were present from 24 June to 29 October, with a distinct abundance peak in mid-August. Recruitment occurred from 24 June to 21 October. Timing of larval sampling was judged to be a crucial part of evaluating recruitment potential for monitoring purposes and a field experiment was carried out to document spatio-temporal larval concentrations. Data were gathered over 30 h sampling periods on 4 August and 14 September 2004. Water samples were taken simultaneously on an hourly basis at three depths: surface, 2 m and 4 m below the surface. Results indicated peaks in larval abundance were, in most cases, limited to a 3 h time interval in the early afternoon. Larval concentrations increased with water depth.     

Abundance and distribution of invertebrate larvae in the Bellingshausen Sea (West Antarctica)

The central part of the Bellingshausen Sea has been poorly studied, partly because of the presence of ice during most of the year. The main aim of this study was to analyse the abundance and distribution of meroplankton, and the influence of oceanographic properties were investigated in the Bellingshausen Sea (West Antarctica) during the BENTART-06 cruise carried out in January–February 2006. Zooplankton samples were collected with a 80-μm mesh plankton net hauled vertically from a depth of 200?m to the surface at fifteen stations across the Bellingshausen Sea. Fifteen types of larval benthic invertebrates were found, with echinospira and nudibranch veligers being the most abundant. Hierarchical analysis and non-metric multidimensional scaling revealed a high degree of spatial variability in both larval abundance and larval types across the Bellingshausen Sea. The variability was significantly correlated with total chlorophyll-a and the contribution of large (>5?μm) phytoplankton to total chlorophyll, indicating the availability of food as an important factor determining the larval distribution observed. Nudibranch veligers, nemertean pilidia, echinoderm and planula larvae were more abundant at stations in the central Bellingshausen Sea, which was characterized by low phytoplankton biomass and production. Higher abundances of echinospira veligers and polychaete larvae were found at the more productive stations close to Peter I Island and the Antarctic Peninsula. The abundance and diversity of larval types found in the Bellingshausen Sea during the BENTART-06 cruise support the hypothesis that indirect development through larval swimming stages plays a key role in benthic recruitment in polar areas.     

Trophodynamics of the plankton community at Dogger Bank: predatory impact by larval fish

The trophodynamics of a coastal plankton community were studied,focusing on fish larvae and their copepod prey. The major objectiveswere to describe distributional overlap and evaluate the predatoryimpact by larval fish. The study was carried out across DoggerBank in the North Sea, August-September 1991. Sampling transectscrossed tidal fronts off the Bank and plankton at all trophiclevels showed peak abundance within frontal zones. Also Verticallythere was a significant overlap in distributional patterns ofthe plankton. Seven species of fish larvae were abundant, ofthese sprat (Sprattus sprattus) dominated. The abundance ofone group of fish larvae peaked in the shallow water close tothe Bank, whereas other species, including sprat, were foundin deeper water. Prey preference and predation pressure of fishlarvae were assessed using information on prey sizes and growthrates of larvae and the copepod prey. We estimated larval removalof preferred prey sizes to 3–4% day^–1, counterbalancedby a 3–7% day^–1' replenishment from copepod productionand growth. Additional predation pressure on copepods by aninvertebrate predator was estimated to 1–3%day^–1.In conclusion, the dynamics of fish larvae and other zooplankterswere closely linked. At peak abundances of fish larvae (>35mg dry weight m^–2), the accumulated predation on specificsize ranges of copepods, exerted by larvae and other predators,could exceed the ability of copepod replenishment and intra-/interspecificcompetition among predators might take place.     

Mechanisms used by the inhabitants of a sand flat in the White Sea to colonize aggregates of Mytilus edulis Linnaeus, 1758 (Bivalvia: Mytilidae)

The colonization of artificially created aggregates of the mussel Mytilus edulis by organisms that inhabit an intertidal sand and mud flat was studied in a field experiment. The sediment of 40 experimental plots was cleared of macrofauna. Thirty of these plots were covered with fishing nets, on which either live mussels (ten plots, type M) or mussel dummies (ten plots, type D) were placed; on ten plots, the nets were left empty (type Z). The remaining ten plots without nets but with a cleared surface (type E) were the control. For comparison, samples were taken from the ambient intact community. After 16 days of exposure, the community on the plots differed from the ambient one. In the E-Z-D-M series, only 7 out of 29 taxa that were found in the samples differed significantly in abundance. Chironomid larvae, adults, and spat of M. edulis and Mya arenaria were most abundant on the plots with a hard substrate (Z, D, M). Adult Jaera sp., Oligochaeta, Littorina saxatilis, and Nemertea were more abundant on the type M plots. The abundance of spat of Macoma balthica and Hydrobia ulvae did not differ significantly. The results indicate that the biological activity of mussels M. edulis in dense assemblages facilitated immigration of adult animals of other species. The M. edulis aggregates either had no influence on the spat of mollusks or they attracted it as a hard substrate for settlement.     

Distribution and drift of the crab Carcinus maenas (L.) (Decapoda, Portunidae) larvae over the continental shelf off northern Portugal in April 1991

The coastal distribution and abundance of Carcinus maenas larvalstages were studied from plankton samples collected in a gridof 79 sampling station organized into six cross-shore transectsextending from the coast to –170 km offshore, betweenthe Mondego and Ave rivers, on the northwest coast of Portugal.The samples were collected in April 1991 with a modified Bémultinet plankton sampler, which was towed from a maximum depthof 200 m and provided a vertical resolution of up to five standardstrata at each station. Current and wind data were availablefor a period that extended beyond the period covered by theobservations All the species' larval stages were found in thesamples, but their distribution was confined to the inner andmiddle shelf stations Vertically, 88% of the larvae occurredin the top 30 m and another 11% in the 30–60 m stratum.The zonal stages I and II were concentrated (–90%) inthe surface layer, but a gradual ontogenic displacement to deeperwaters was observed from then on, the megalopa being equallydistributed between the 0–30 and the 30–60 m strata.Horizontally, there was a dear association of the first zoeawith the estuarine inlet while the older meal stages were dispersedprogressively offshore. Evidence was found that the megalopaexperienced an onshore transport that did not affect the previousstages.This transport is wnsistent with the observation of anonshore flow component at 40 m. It was not possible, however,to examine the hypothesis that this flow conveys the last larvalstage to the coast, but not the previous ones. The dependenceof the along-shore flow component on wind stress lends supportto the hypothesis that the larvae are advected from the northas the upwelling season progresses.     

Recruitment of the amphidromous goby <Emphasis Type="Italic">Sicyopterus japonicus</Emphasis> to the estuary of the Ota River,Wakayama, Japan

Ecological aspects of recruitment in the amphidromous goby, Sicyopterus japonicus, were studied from larval collections made with a set net in the estuary of the Ota River, Wakayama, Japan. The abundance patterns of the 12,766 larvae collected from 18 April to 26 August 2006 showed several peaks during the recruitment season. Their body sizes at recruitment ranged from 23.5 to 30.0 mm standard length (mean ± SD, 26.3 ± 1.1 mm), 0.11 to 0.49 g body weight (0.22 ± 0.05 g), and 8 to 20 condition factor (11 ± 2). The standard length of the goby larvae tended to decrease with the season, while their body weight slightly increased and resulted in an increase in condition factor. The recruitment of larvae occurred mainly during the daytime. Otolith growth increment analysis of 30 larvae collected by a square lift net on 30 April 2005 revealed that the oceanic larval duration after downstream migration ranged from 173 to 253 days (208 ± 22) after hatching. A limited time of recruitment in early summer and a considerably long duration of oceanic life (about a half year) appeared to be unique characteristics of this Sicyopterus species that lives in a temperate region in comparison to other tropical species of the genus Sicyopterus that all have year-round recruitment.     

Different zooplankton structures in the German Bight

In August 1982, a net of 48 stations with altogether 208 samples was investigated in the eastern German Bight with respect to temperature, salinity, as well as the amount and species composition of the mesozooplankton (>80 μm). The data were arranged into different structures by means of a cluster analysis. Four different clusters were found: (a) a “Wadden sea water” with few holoplankton organisms but a higher amount of spionid larvae; (b) a “German Bight water” with a maximum occurrence of turbellaria (Alaurina composita) and medium concentrations of copepods; (c) a mixing area between these two water masses with highest amounts ofOikopleura dioica, Temora longicornis, Acartia sp., mussel larvae and larvae of the spionid worms; (d) a “North Sea water” mass with highest concentrations ofPseudocalanus elongatus, Paracalanus parvus undOithona similis. The differences in the concentrations of the species mentioned between the four clusters were significant on the 0.1%-level.     

Phytoplankton composition of the Kandalaksha Gulf, Russian White Sea: Dinophysis and lipophilic toxins in the blue mussel (Mytilus edulis)

Dinophysis acuminata and D. norvegica were observed in plankton net samples during the summer of 2002 from the Kandalaksha Gulf in the White Sea (North European Russia). Prorocentrum lima was found as an epiphyte on subtidal macroalgae in August, but not observed in plankton net samples. Protein phosphatase 2A (PP2A) inhibition measured 127.8 ng OA-equivalent/g of mussel (Mytilus edulis) hepatopancreas from samples collected a few days after when Dinophysis was recorded at a density of 1550 cells L⁻¹. Liquid chromatography–mass spectrometry confirmed presence of several classes of lipophilic shellfish toxins associated with Dinophysis spp. in the mussels including okadaic acid, dinophysistoxin-1, pectenotoxins and yessotoxins. No azaspiracid was detected. This represents the first identification of phycotoxicity in the White Sea.     

Proteomic approach to probe for larval proteins of the mussel Mytilus galloprovincialis

A proteomic approach was used to search for larval proteins specific to the mussel Mytilus galloprovincialis from Galicia in northwest Spain. The study included both a comparative analysis, through two-dimensional electrophoresis, of protein expression maps of the larvae of the mussel and of 5 abundant and commercially important bivalve species from the region (Ostrea edulis, Cerastoderma edule, Pecten maximus, Tapes decussatus, and Venarupis pullastra) and subsequent mass spectrometric analysis of some of the protein spots. A total of 18 spots were selected and isolated from gels of M. galloprovincialis larvae. From their relative position on the electrophoresis gels, 6 of these were clearly exclusive to the mussel species. However, it was not clear whether the other spots were shared by other species. To overcome this ambiguity, first an analysis using matrix assisted laser desorption ionization with time-of-flight (MALDI-TOF) was conducted on the 6 spots of Mytilus that could possibly be shared with only one species. The peptide mass fingerprinting was completely different for the proteins compared. This result confirmed that the 6 proteins were exclusively mussel proteins, but demonstrated the utility of this approach when working with species that are poorly represented at the protein level in databases.     

Phytoplankton blooms and fish recruitment rate: Effects of spatial distribution

We consider the spatio-temporal dynamics of a spatially-structured generalization of the phytoplankton-zooplankton-fish larvae model system proposed earlier (Biktashev et al., 2003, J. Plankton Res. 5, 21–33; James et al., 2003, Ecol. Model. 160, 77–90). In contrast to Pitchford and Brindley (2001, Bull. Math. Biol. 63, 527–546), who were concerned with small scale patchiness (i.e., 1–10m), on which the (stochastic) raptorial behaviour of individual larvae is important, we address here the much larger scale ‘patchy’ problems (i.e., 10–100 km), on which both larvae and plankton may be regarded as passive tracers of the fluid motion, dispersed and mixed by the turbulent diffusion processes. In particular, we study the dependence of the fish recruitment on carrying capacities of the plankton subsystem and on spatio-temporal evolution of that subsystem with respect to the larvae hatching site(s). It is shown that the main features found both in the nonstructured and age-structured spatially uniform models are observed in the spatially structured case, but that spatial effects can significantly modify the overall quantitative outcome. Spatial patterns in the metamorphosed fish distribution are a consequence of quasi-local interaction of larvae with plankton, in which the dispersion of larvae by large scale turbulent eddies plays little part due to the relatively short timescale of the larvae development. As a result, in a strong phyto/zooplankton subsystem, with fast reproduction rate and large carrying capacity of phytoplankton and high conversion ratio of zooplankton, recruitment success depends only on the localization and timing of the hatching with respect to the plankton patches. In a weak phyto/zooplankton system, with slow reproduction rate and small carrying capacity of phytoplankton and low conversion ratio of zooplankton, the larvae may significantly influence the evolution of the plankton patches, which may lead to nontrivial cooperative effects between different patches of larvae. However, in this case, recruitment is very low.     

Can bivalve veligers escape feeding currents of adult bivalves?

While the stock of introduced Pacific oysters (Crassostrea gigas) increased in the Oosterschelde estuary (SW Netherlands), so did the filtration pressure of all bivalve species together. In the same period, stocks of native bivalves declined slightly. The expansion of Pacific oysters in Dutch estuaries might be partially due to better abilities of their larvae to avoid or escape filtration, compared to larvae of native bivalves. In this context, escape and swimming abilities of Pacific oyster larvae and the larvae of the native blue mussel (Mytilus edulis) were compared.Swimming behaviour of C. gigas larvae and larvae of M. edulis was recorded in still water and in a suction current mimicking a bivalve feeding current, in a horizontal and in a vertical plane. Larval swimming behaviour in a suction flow field was reconstructed by subtracting local water movement vectors from the total movement of larvae, yielding movement paths due to larval swimming alone.Swimming speeds and the rate of displacement in vertical direction of C. gigas and M. edulis larvae were related to larval shell length, and to the pitch of up- or downward swimming.Larvae of both species did not show escape reactions in a suction flow field. With increasing shell length, larval swimming speeds of both species increased significantly. Swimming speeds of C. gigas larvae were significantly higher than swimming speeds of M. edulis larvae, resulting in a faster vertical displacement. The ability to migrate to more favourable water layers faster may offer C. gigas an advantage over native bivalves with slower swimming larvae.     

Distribution and abundance of fish larvae in the northern Ionian Sea (Eastern Mediterranean)

The purpose of this paper was to study the spatial distribution, abundance and composition of fish larvae in the northern Ionian Sea. Samples were collected to the 600 m depth with an electronic multinet BIONESS during the “INTERREG Italia-Grecia” oceanographic cruise carried out in March 2000 off the Apulian Italian coast. A total of 46 species of teleost early stages were collected, belonging to 38 genera and 22 families. Over 52% of the larvae identified were mesopelagic species, almost 27% were demersal and about 21% pelagic. A total of 307 myctophids, 69 clupeids and 61 gadid post-larvae dominated the community. Benthosema glaciale (mean 6.1 mm SL) was the most abundant species (21.6%), the most frequent in the samples (28.8%), and dominant in the whole study area (mean 1.4 ind/100 m³). Particular attention was given to the horizontal and vertical distribution and abundance of the three dominant post-larval species: Benthosema glaciale, Sprattus sprattus sprattus and Notoscopelus elongatus. The Pearson coefficient (R = 0.734) showed a high correlation between total zooplankton and fish larval assemblages in terms of spatial distribution abundance values. Regarding the vertical distribution of fish larvae, Sorensen’s index (S = 0.69) showed that fish larvae and total zooplankton abundance peaks co-occurred along the water column.     

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

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