Growth and mortality rates of the fan mussel <Emphasis Type="Italic">Pinna nobilis</Emphasis> in Lake Vouliagmeni (Korinthiakos Gulf,Greece): a generalized additive modelling approach

The temporal patterns and the effect of shell size and depth on growth and mortality rates of the endangered fan mussel Pinna nobilis were investigated in the marine Lake Vouliagmeni (Korinthiakos Gulf, Greece). A total of 160 individuals were tagged and monitored monthly for a period of 17 months. At each visit, the size of the tagged individuals (shell width, w) was measured in situ and recorded. Any mortality event was also recorded and attributed to natural causes or (illegal) fishing. Growth and mortality rates were modeled with generalized additive models, which are non-parametric flexible models that free the researcher from the limiting concept of a strict parametric shape. The use of GAMs allowed the exploration of shapes of growth and mortality response curves in relation to predictor variables and allowed the fitting of statistical models that better agree with ecological theory. Growth rates had a seasonal pattern, with an extended period of very slow growth between late autumn and early spring, i.e., during the cold season, another short period of slow growth during August (when water temperatures reached their maximum values exceeding 29°C), and a peak in growth rates during late spring–early summer, probably related to an optimum combination of temperature and food availability. Growth rates varied with shell size, with a peak at w ∼4.5 cm, followed by a sharp decline to an approximately constant level, with sizes ranging from 9 to 15 cm, and a further decline with larger sizes down to almost zero for w > 20 cm. Growth rates did not vary substantially with depth. Although P. nobilis is a protected species in the EU and its fishing is strictly prohibited, fishing mortality was very high in Lake Vouliagmeni (much greater than natural mortality), especially during the hot season when the lake was crowded by summer visitors. The fan mussels were poached exclusively by free-diving and due to the high turbidity of the lake’s water, fishing mortality was higher in shallow areas (and mostly for large individuals) and was practically zero at depths >9 m. Due to fishing mortality, a size segregation of P. nobilis was observed in the lake: large individuals were restricted to deeper areas, while young and small individuals were more abundant in shallow areas where there was preferential recruitment. Natural mortality was strikingly size dependent and P. nobilis suffered high natural mortality during the first year of life; the probability of death by natural causes quickly diminished as the fan mussels grew in size. No depth-related differences in natural mortality were found.     

High intertidal community organization on a rocky headland in Maine,USA

A mosaic patchwork of the barnacle Balanus balanoides L., the mussel Mytilus edulis L., and the alga Fucus vesiculosus L. was found in the transitional region between the mid and high intertidal zones on a rocky headland on Mount Desert Island, Maine, USA. The development of the mosaic was observed by following recruitment and survival of B. balanoides in denuded patches located at the same tidal level along a 60 m stretch of shore. Barnacle recruitment was least under canopies of F. vesiculosus and greatest in open areas kept moist at low tide by surf. Barnacle survival after settlement was least under the F. vesiculosus canopy due to the whiplash effect of the algal fronds in the surf and greatest in open areas free from competition from mussels. In open areas, early mortality was correlated with settlement density. In areas of dense settlement (60 spat cm^-2) up to 90% mortality resulted within 5 months from crowding associated with growth. In older individuals crowding produced hummocks of elongated, weakly attached barnacles which were more prone to removal by surf than uncrowded barnacles. Mussels exerted competitive dominance over barnacles for space and the presence of mussel beds prevented further barnacle recruitment. Mussels suffered extensive mortality during winter storms when surf removed dense mats of weakly attached mussels. The patchy distribution of mussels and barnacles results from irregular rock substrata producing numerous environmental patches with respect to wave exposure and drainage at low tide, and from densitydependent mortality of both mussels and barnacles which creates patches of new colonizable space within each environmental patch.     

Wave exposure effects on population structure and recruitment in the mussel <Emphasis Type="Italic">Perna perna</Emphasis> suggest regulation primarily through availability of recruits and food,not space

Recruitment and population structure of Perna perna in low shore mussel beds were investigated over 15 months at six sites along the south coast of South Africa. Initial, subjective classification of sites as wave exposed or wave sheltered (three of each) was confirmed using the dissolution of cement blocks to measure average water flux and dynamometers for maximum wave force. Recruitment occurred throughout the year, but recruit (1–5 mm) densities were significantly higher from January to April 1996 on both shore types. Recruit densities were positively correlated with adult (>15 mm) densities for both shore types (P < 0.05) but the correlations were extremely weak (r ² < 0.06 in each case). In areas with 100% cover, adult size (mean and maximum lengths) was greater on exposed sites, but density showed the reverse and was negatively correlated with maximum wave strength (r = −0.84). Despite differences in adult densities and sizes, biomass, which is a product of the two, showed no significant difference between the two shore types (ANOVA P > 0.05). Thus wave exposure dramatically affects density, recruitment and mussel size, but not recruitment timing or biomass where there is 100% cover, and mediates a three-way interaction among food supply, larval supply and intraspecific competition for space. In contrast to shores with saturation recruitment, mussel biomass here appears to be limited by recruit supply and constraints of food, especially on sheltered shores, while density is regulated through intraspecific competition for space primarily on exposed shores and at small spatial scales.     

Age,growth and population structure of <Emphasis Type="Italic">Modiolus barbatus</Emphasis> from the Adriatic

Age, growth and population structure of Modiolus barbatus from Mali Ston Bay, Croatia were determined using modal size (age) classes in length frequency distributions, annual pallial line scars on the inner shell surface, internal annual growth lines in shell sections of the middle nacreous layer and Calcein marked and transplanted mussels. The length frequency distributions indicated that M. barbatus attain a length of ∼40 mm in 5–6 years indicating that a large proportion of the population in Mali Ston Bay is <5 years old. Some mussels of ∼60 mm were predicted to be 14 years old using the Von Bertalanffy growth (VBG) equation. Up to the first 6 pallial line scars were visible in young (<6 years) mussels but in older shells the first scars became obscured by nacre deposition as the mussel increased in length and age. The age of the older shells (>6 years) was determined from the middle nacreous lines in shell section, which formed annually in winter between February and March; the wider dark increments forming during summer (June to September). The oldest mussel, determined from the middle nacreous lines, was >12 years, with the majority of mussels aged between 3 and 6 years of age. The ages of mussels ascertained using the growth lines were not dissimilar to the ages predicted from the length frequency distributions. Age at length curves produced using modal size class data were not different from the data obtained using the pallial scar rings and internal growth lines. Taken together these data suggest that M. barbatus attains a length of 40 and 50 mm within 5 and 8 years, respectively. Eighty one percent of individual M. barbatus injected with a Calcein seawater solution (300 mg Calcein l⁻¹), into their mantle cavity successfully deposited a fluorescent line, which was visible in suitably prepared shell sections under ultra violet light. Incorporation of Calcein into the mussel shells was seasonally variable with the lowest frequency of incorporation in mussels marked in February and recovered in May. Seasonal shell growth was observed with significantly higher growth rates in mussels marked in May and removed in August (ANCOVA, F _3,149 = 23.11, P < 0.001). Mussels (∼18 to 22 mm) marked in May and recovered in August displayed maximal growth rates of >2.5 mm month⁻¹ compared with a mean mussel growth rate of 1.2 ± 0.6 mm month⁻¹. At other times of the year mussel shell growth ranged from immeasurable to 1.48 mm month⁻¹.     

Prevalence, patterns, and effects of shell damage on Geukensia demissa in South Carolina estuarine habitats

Shelled molluscs frequently exhibit a record of damage on exterior surfaces that can evidence past predation attempts and may affect survival and growth. In South Carolina populations of the ribbed marsh mussel, Geukensia demissa, >90% of the individuals and up to 60% of the total shell area are damaged. A trend toward greater amounts of damage occurred on mid-marsh compared to oyster reef mussels from the barrier beach side of inlets. Shell damage effects on survivorship and shell and tissue growth were assessed seasonally during multi- and single-season field experiments. Mussels from a common mid-marsh site were divided into size classes (~50 or 70 mm), treated to create two damage levels (undamaged and damaged), and replaced within mid-marsh exclusion cages to minimize additional shell damage. In both multi- and single-season experiments increased shell damage resulted in significantly greater mortality. Linear shell growth was unaffected by increased damage, but 50 mm mussels grew twice as fast. Shell mass increased 16–50% in the multi-season and single-season winter period, but decreased 7–12% during the single-season summer period. Tissue mass significantly decreased 31–43% in 50 mm damaged mussels, but increased by 33% for 70 mm mussels in both multi-season and the single-season winter period experiments. Shell damage did reduce tissue mass 43% in 70 mm single-season summer mussels. Experimental results indicate shell damage from a simulated increase in predation can affect negatively both survival and growth of marsh mussels. Seasonal timing of shell damage and initial mussel size also influenced the effects of sublethal predation on shell and tissue growth. The previously unrecognized importance of sublethal predation and the resultant significant negative effects of shell damage on survival and growth will affect the distribution and population dynamics of G. demissa in coastal marshes and will influence the overall contribution of ribbed mussels to estuarine ecosystems.     

Detrimental effects of the isopod, <Emphasis Type="Italic">Edotia doellojuradoi,</Emphasis> on gill morphology and host condition of the mussel, <Emphasis Type="Italic">Mytilus edulis platensis</Emphasis>

There are many reported associations between mussels and other invertebrates, such as pea crabs, polychaetes, turbellarians and copepods, which live in their mantle cavities. The boundary between commensalism and parasitism is often indistinguishable because of insufficient knowledge or because the interaction is variable. Preliminary evidence led to a closer examination of the relationship between the mussel, Mytilus edulis platensis, and an isopod, Edotia doellojuradoi, previously described as commensalism. Monthly intertidal samples of mussels were taken from September 2004 to August 2005 at Caleta Cordova Norte (45°43′S, 67°22′W) in southern Argentina and assessed for the prevalence and abundance levels of isopods. Mussels with and without isopods were measured, examined for evidence of gill damage and their condition (soft tissue dry weight) was determined. The overall isopod prevalence in mussels was 57.9% and infestation varied with mussel length, with maximum occurence at 30.2 mm (medium-sized mussels). Experimental evidence indicated that the position of the isopod inside the mussel depended on the feeding activity of the mussel. Female isopods were observed grasping the ventral food groove of the gill demibranchs and feeding on the mucous food strand produced by the mussel. Juveniles and males were observed clustered together on the dorsal side of the single female in each occupied mussel, suggesting extended maternal care. Gill damage was observed in 58.2% of mussels at the Argentine site and was significantly associated with isopod occurrence. Categorical regression analysis showed that the most important factor associated with the degree of gill lesions was the number of male and juvenile isopods per mussel, followed by the length of female isopods and the developmental stage of juveniles. Conversely, the degree of gill damage decreased with increasing mussel length. Overall, E. doellojuradoi had a significant effect on mussel condition throughout the year, with low flesh weight in mussels with isopods, except during the austral summer and early autumn. In contrast to previous studies, which concluded that the isopod was a commensal, the present study clearly demonstrates that E. doellojuradoi is a parasite of M. edulis platensis. Other symbiotic interactions formerly classified as commensal might not be innocuous on further investigation, especially if samples are taken at multiple sites and at different times of the year.     

Growth and survival differences among native,introduced and hybrid blue mussels (<Emphasis Type="Italic">Mytilus</Emphasis> spp.): genotype,environment and interaction effects

The Mytilus species complex consists of three closely related mussel species: Mytilus trossulus, Mytilus edulis, and Mytilus galloprovincialis, which are found globally in temperate intertidal waters. Introduction of one or more of these species have occurred world-wide via shipping and aquaculture. Stable hybrid zones have developed in areas where these species have come into contact, making the invasion process complex. On the east coast of Vancouver Island (VI), British Columbia (BC), Canada, the native (M. trossulus) and introduced species (M. edulis and M. galloprovincialis), as well as their hybrid offspring, occur sympatrically. This study used a common environment experiment to quantify growth and survival differences among native, introduced, and introgressed mussels on VI. Mussels were collected from an area of known hybridization and reared in cages from May to August 2006. The cages were deployed at a local site as well as a remote site (approximately 150 km apart), and the mussels were genotyped at two species-specific loci. Growth and survival, as fitness measures, were monitored: native, introduced, and introgressed individuals were compared between and within sites to determine whether growth and survival were independent of site and genotype. Overall, mussels reared at Quadra Island performed better than locally-reared mussels at Ladysmith. Specifically, introgressed mussels reared at Quadra Island performed better than all genotypes reared at Ladysmith, as well as better than native mussels reared at Quadra Island. Differences in survival and growth among the native, introduced and introgressed mussels may serve to explain the complex hybridization patterns and dynamics characteristic of the VI introgression zone.     

Quantitative studies on gonad cycle,fecundity, reproductive output and recruitment in a baltic Mytilus edulis population

Since Mytilus edulis L. is a biomass dominant in the Baltic much interest is focused on the ecology of the species. In this paper an attempt is made to quantitatively cover the reproductive cycle of a Baltic M. edulis population in order to provide data for energy flow models and to discuss aspects of recruitment in this species. Histological preparations of gonads showed that gametogenesis started with declining temperatures in autumn and proceeded very slowly through winter. At the beginning of March when food was supplied during the spring phytoplankton bloom, rapid maturation took place. This was also revealed by an increase in meat weight of the mussels. Only one spawning period was recorded, from the middle of May until the beginning of June, due to food being strongly limited to the population during the rest of the year. The length of the larval period was estimated as being 5 to 6 wk and settlement was registered from the end of June through July. In general the large annual variations found in the Baltic with regard to temperature and food abundance give rise to a more marked annual pattern in the reproductive cycle than is encountered in other seas. Fecundity was assessed for two populations from 4-and 15-m depths from studies covering two annual cycles of the changes in the relation of shell length — meat weight. The size-related fecundity was found to be equal in both populations and related to food abundance and not to growth or age. Fecundity, expressed as weight loss at spawning, ranged from 0% in 2-mm mussels and increased from 38 to 52% in 10-to 30-mm mussels. The fecundity as percentage of biomass in full-grown Baltic M. edulis is of similar magnitude as in full-grown mussels from other areas despite the smaller size of Baltic mussels. The reproductive output for the total 160 km² research area was calculated as being 1 200 tons dry weight or 80% of the standing stock, which, due to the particular features of the Baltic M. edulis population probably represents the larger part of the total mussel production. This reproductive output, calculated as 8·10⁷ eggs·m^-2 and corresponding to 50% of the total annual zooplankton production, may thus consitute an important food source for herring larvae and carnivorous zooplankton. Recruitment was divided into two phases: (1) Recruitment of juveniles (=settlement of larvae), and (2) recruitment to breeding stock. Monitoring studies of settlement on ropes and the year round presence of high abundances of mussels <2 mm indicate that settlement is in excess of the demands for maintaining population size and that most settled mussels form a pool of competitively suppressed non-growing individuals. Not until death of an already established mussel will these become recruited to the breeding population. Thus recruitment is possible throughout the year which stabilizes the population and maintains it near the carrying capacity of the area with regard to food and space availability.     

Local extinction of a foundation species in a hypoxic estuary: integrating individuals to ecosystem

We integrated across individual, population, community, and ecosystem levels to understand the impact of environmental stress by tracking the foundation species Mytilus edulis in the hypoxic estuary Narragansett Bay, Rhode Island, USA. Our initial surveys revealed that the mussels occurred in nine extensive (2-28 ha) dense (814-9943 individuals/m2) subtidal reefs that attracted a diverse suite of predators (sea stars, crabs, gastropods). Hypoxia occurred in the summer of 2001, and a mussel transplant experiment revealed overall reduced growth rates of individuals, and higher mortality rates among larger mussels. At the population level, large decreases in densities and cover of mussels were correlated with dissolved oxygen concentrations, leading to extinction at one site and reductions of over an order of magnitude at others. Within one year, seven of the eight remaining populations were edged to extinction, and the previously extinct population was recolonized. At the community level, a predator exclusion experiment indicated that predation was an unimportant source of mussel mortality during the hypoxic period, in part due to the emigration of sea stars, as predicted by the Consumer Stress Model. However, mussels were too intolerant to hypoxia to have a net benefit from the predation refuge. The seasonal (summer) occurrence of hypoxia allowed sea stars to return following a lag, as predicted by a stress return time model, and the resumption of predation contributed to the subsequent extinction of mussel populations. At the ecosystem level, the initial filtration rate of the mussel reefs was estimated at 134.6 x 10(6) m3/d, equivalent to filtering the volume of the bay 1.3 times during the 26-d average residence time. That function was reduced by >75% following hypoxia. The effect of hypoxia on each level of organization had consequences at others. For example, size-specific mortality and decreased growth of individuals, and reduced filtration capacity of reefs, indicated a loss of the ability of mussels to entrain planktonic productivity and potential to control future eutrophication and hypoxia. Our study quantified patterns of loss and identified pathways within an integrative framework of feedbacks, summarized in a conceptual model that is applicable to similar foundation species subjected to environmental stress.     

The effect of selection treatments on <Emphasis Type="Italic">Mytilus edulis</Emphasis>, modifications of genetic and physiological characteristics

This study examined the effects of two selection treatments (elevated water temperature and air exposure) on the genetic and physiological characteristics of the juvenile marine mussel, Mytilus edulis (<10 mm). Genetic effects were measured on five allozymes and fitness assessed using physiological tests to estimate energy balance (scope for growth) as well as size, growth and survival. The in vitro treatments resulted in 48% mortality from an air exposure of 11 h at 27°C and 76% mortality from a 6-h exposure to 33°C water. Survivors (n = 1,152) of each treatment along with controls (n = 2,304) were measured and randomly placed in compartmentalized cages. Mussels were deployed to three bays in Prince Edward Island, Canada and monitored over a 10-month period. Initially, both of the treatments had an effect on mussel size and increased the heterozygosity of the surviving mussels. Physiological analyses after 3 months in the field showed that the two treated mussels showed lower metabolic rate that the control group. After 10 months in the field, the treated mussels were larger and had lower mortality than the untreated control mussels. Unexplained environmental interaction in each of the bays had an effect on allelic frequencies and heterozygosity. Overall, the results demonstrate that simple husbandry techniques can be used to increase the productivity of mussel seed and heterozygosity measures can be used to assess fitness. However, more field data is needed to determine the consistency of the increased productivity and if the increased productivity justifies the costs of a selective treatment. Furthermore, because the level of heterozygosity in juvenile mussel populations can vary considerably, both spatially and temporally, it may be effective as a warning of future natural mass mortality when overall heterozygosity levels are found to be low.     

Demographic variation in the kelp<Emphasis Type="Italic"> Laminaria hyperborea</Emphasis> along a latitudinal gradient

We compared populations of Laminaria hyperborea from different regions along the Norwegian coast (58–71°N). The aim of the study was to examine possible regional differences in population demography and to relate these to latitudinal gradients in ambient temperature and light. A number of population parameters were examined in understory and canopy kelp individuals. We found latitudinal differences in stipe growth rate and growth pattern, mortality, longevity, recruitment and density. Stipe growth rate was higher for young individuals in mid-Norway than in south and north Norway. Mean stipe length was related to temperature and light, and correlated significantly with indices combining temperature during the growth season and daylength in summer. Mortality decreased and longevity increased with increasing latitude, probably because of a temperature decrease with increasing latitude and a general reduction in metabolic rate. Greater longevity may also cause the observed decrease in recruitment rate with increasing latitude, since there are fewer opportunities for understory individuals to replace dead canopy individuals. The development of particularly large kelp in mid-Norway appears to be explained by high growth rates and not a particularly long life span, as earlier assumed. This has consequences for decisions of the optimal harvesting regime for the species in the region, with regard to yield and maintenance of associated biodiversity.Communicated by M. Kühl, Helsingør     

Ingestion of bivalve larvae by<Emphasis Type="Italic"> Mytilus edulis</Emphasis>: experimental and field demonstrations of larviphagy in farmed blue mussels

Ingestion of bivalve larvae by Mytilus edulis was investigated. Laboratory experiments revealed that ~ 90% of bivalve larvae offered to mussels was ingested and apparently fully digested. The shell of the bivalve larvae offered no protection against digestive processes, resulting in high larval mortality once inside the stomach. Stomach content analysis (September 2001–January 2003) showed that bivalve larvae were ingested by farmed mussels year-round, with the exception of March 2002. Numbers of ingested larvae were highest in October 2001 and May 2002, which coincides with known spawning times of farmed mussels in Ireland. Mussels ingested a large size-range of bivalve larvae, suggesting that all stages of the bivalve life cycle are vulnerable to predation. It is suggested that adult bivalves routinely filter larvae from the surrounding water and that, given the high biomass of mussels present in mussel farms, filtration by adult bivalves significantly reduces numbers of bivalve larvae in nearby waters.Communicated by J.P. Thorpe, Port Erin     

Local and meso-scale patterns of recruitment and abundance of two intertidal crab species that compete for refuges

Interference competition for limited habitat or refuges is known to produce density-dependent mortality and generate patterns of micro-habitat distribution. While in mobile species the outcome of interference at a local scale can usually be determined from differences in body size and behavior, the population-level consequences of such interactions vary depending on rates of settlement and recruitment at a site, which are not directly correlated to local reproductive success. Previous experimental studies in central Chile demonstrated that interference competition for refuges is the primary factor driving microhabitat segregation between the predatory crabs Acanthocyclus gayi and Acanthocyclus hassleri, with the latter species monopolizing galleries inside mussel beds and excluding A. gayi to rock crevices. Between April 2001 and March 2006 we quantified monthly recruitment rates in artificial collectors at 17 sites over 900 km of the central coast of Chile. Results show that recruitment rates of A. hassleri are almost two orders of magnitude lower than those of A. gayi, and that they are tightly and positively correlated among sites across the region, suggesting that at scales of kilometers larval stages of these species are affected by similar oceanographic processes. Total crab densities per site were also positively correlated between species and strongly associated to mussel cover, with overall low crab densities at all sites where mussel cover was lower than about 60%. At all sites with mussel cover >60%, the ratio of A. gayi to A. hassleri density progressively decreased from recruits (2.6) to juveniles (0.5) to adults (0.04), overcoming initial differences in recruitment rates. The relative success of the inferior competitor at sites with low mussel cover does not appear to provide a potential mechanism favoring regional coexistence through dispersal to other sites (“mass effects”), because their densities were lower than at sites of high mussel cover. Yet, at many sites of low mussel cover the dominant competitor is virtually absent, allowing A. gayi to attain larger population sizes at the scale of the region. Thus, the factors limiting the dominant competitor from successfully utilizing other microhabitats seem to be the most critical factor in promoting both local and regional coexistence between these species.     

Adaptive host choice and avoidance of superparasitism in the spawning decisions of bitterling (Rhodeus sericeus)

Choice of a site for oviposition can have fitness consequences. We investigated the consequences of female oviposition decisions for offspring survival using the bitterling, Rhodeus sericeus, a freshwater fish that spawns inside living unionid mussels. A field survey of nine bitterling populations in the Czech Republic revealed a significantly lower rate of release of juvenile bitterling from Anodonta cygnea compared to three other mussel species. A field experiment demonstrated that female bitterling show highly significant preferences for spawning in A. anatina, Unio pictorum, and U. tumidus. Within a species, female bitterling avoided mussels containing high numbers of bitterling embryos. Mortality rates of bitterling embryos in mussels were strongly density dependent and the strength of density dependence varied significantly among mussel species. Female preferences for mussels matched survival rates of embryos within mussels and females distributed their eggs among mussels such that embryo mortalities conformed to the predictions of an ideal free distribution model. Thus, female oviposition choice is adaptive and minimizes individual embryo mortality. Received: 6 October 1999 / Received in revised form: 7 January 2000 / Accepted: 13 March 2000     

Positive effects of the introduced green alga, Codium fragile ssp. tomentosoides, on recruitment and survival of mussels

The green macroalga, Codium fragile ssp. tomentosoides, is an important component of sheltered low-shore assemblages on breakwaters along sandy shores in the northern Adriatic Sea. Macroscopic thalli of C. fragile are not perennial, but develop from propagules and/or undifferentiated forms in early spring, when the dominant native space-occupier, the mussel Mytilus galloprovincialis, recruits. By mid-summer, rapid growth of C. fragile leads to the formation of a dense canopy. We investigated the effects of juvenile and adult thalli of C. fragile on recruitment, survival and growth of mussels. Two experiments tested the hypotheses: (1) that recruitment of mussels is greater within patches of juvenile thalli (primordia) of C. fragile than on adjacent bare surfaces; (2) that the presence of a canopy of C. fragile affects the survival and growth of mussel recruits. The number of recruits of mussels was significantly larger within clumps of primordia of C. fragile than on bare surfaces. The removal of the canopy of C. fragile affected negatively the density of mussels after 2 months from the start of the experiment, but there were no effects on the mean size of individuals, nor on the size–frequency distribution. The same trend persisted after 4 months from the start of the experiment. These results show that re-colonisation of space by mussels is enhanced by C. fragile. Given that mussels, in turn, have the potential to reduce recruitment rates of C. fragile, quick recovery of mussel beds after disturbances could be crucial for controlling the abundance of this alga on breakwaters. Results also suggest that the effects of introduced species on native assemblages can be explained only through studies encompassing different life-stages of interacting organisms.     

Mechanisms of habitat segregation between an invasive and an indigenous mussel: settlement,post-settlement mortality and recruitment

The mussel Mytilus galloprovincialis is highly invasive worldwide, but displays varying degrees of local and regional coexistence with indigenous mussels through spatial habitat segregation. We investigated the roles of settlement, post-settlement mortality, juvenile growth and recruitment in partial habitat segregation between the invasive M. galloprovincialis and the indigenous mussel Perna perna on the south coast of South Africa. We used two study locations, Plettenberg Bay and Tsitsikamma, 70 km apart, with two sites (separated by 300–400 m) per location, each divided into three vertical zones. There were no significant effects in Tsitsikamma, where daily settlement and monthly recruitment were significantly lower than in Plettenberg Bay. In Plettenberg Bay, settlement (primary and secondary) and recruitment of both species decreased upshore. Post-settlement mortality was measured over two consecutive 6-day periods during a spring tide and a neap tide. For both species mortality was low on the low-shore. High-shore mortality was consistently low for M. galloprovincialis, but increased dramatically for P. perna during spring tide. No data were obtained for growth of P. perna, but juvenile M. galloprovincialis grew more slowly farther upshore. P. perna recruited mainly in spring and summer, with a peak in summer far greater than for M. galloprovincialis. Recruitment of M. galloprovincialis was more protracted, continuing through autumn and winter. Thus local coexistence is due to a combination of pre- and post-recruitment factors differing in importance for each species. P. perna is excluded from the high-shore by recruitment failure (low settlement, high mortality). High survival and slow growth in juveniles may allow large densities of M. galloprovincialis to accumulate there, despite low settlement rates. With no differences between species in settlement or mortality on the low-shore, exclusion of M. galloprovincialis from that zone is likely to be by post-recruitment processes, possibly strengthened by periodic heavy recruitments of P. perna. At larger scales, larval retention and protracted recruitment contribute to the success of M. galloprovincialis at Plettenberg Bay, while recruitment limitation may explain why M. galloprovincialis is less successful at other sites.     

Growth and population dynamic model for the non-zooxanthellate temperate solitary coral Leptopsammia pruvoti (Scleractinia, Dendrophylliidae)

In corals where complex life history processes decoupling age from size (e.g., fragmentation, fusion, partial colony mortality) are rare or clearly detectable, individual age may be determined from size, and age-based growth and population dynamic models may be applied. One example is the solitary Mediterranean coral Leptopsammia pruvoti Lacaze-Duthiers 1897, whose population size and structure, and growth rates were determined at Calafuria (43°28′N and 10°20′E), Ligurian Sea, from December 2007 to June 2009. Growth rate decreased with increasing size. The growth curve derived from field measurements confirmed the one obtained by growth band analysis. The frequency of individuals decreased exponentially with age, indicating a steady state population. Turnover time was 2.3 years. Maximum life span was 13 years. Most reproductive output was from intermediate age classes (6 years), while older individuals (>7 years), although having higher fecundity, were rare and accounted for a minority of population reproductive output. In comparison with other solitary dendrophylliids, L. pruvoti life strategy was characterized by a reproduction with r-strategy correlates (high fecundity, short embryo incubation, small planula size, fast achievement of sexual maturity), and a rate of demographic renewal occurring halfway along the r–K continuum (intermediate turnover time and maximum longevity).     

Roach (<Emphasis Type="Italic">Rutilus rutilus</Emphasis>) as an important predator on blue mussel (<Emphasis Type="Italic">Mytilus edulis</Emphasis>) populations in a brackish water environment,the northern Baltic Sea

Suspension-feeding bivalves are organisms of major functional importance in several aquatic environments around the world. They are also important food items for many fish and benthivorous seabirds. It has commonly been thought that predation pressure on blue mussel (Mytilus edulis) populations is negligible in the Baltic Sea, owing to the scarcity of major invertebrate predators such as starfish and crabs. It has recently been shown, however, that the blue mussel is the main food item for roach (Rutilus rutilus) in the archipelago areas of the western Gulf of Finland, where this freshwater fish species has become increasingly abundant, mainly due to increased eutrophication. To quantify the influence of roach predation on blue mussel populations we measured the standing biomass and size structure of the local blue mussel population and used a bioenergetic model to estimate mussel consumption by individual roach during two consecutive summers, 1997 and 1998. The results of the model were combined with existing data on roach abundance, giving annual consumption estimates of 75–105 kg blue mussel dry weight ha^–1 in the study area, approximately two-thirds of these consumed mussels being >10 mm. This corresponds to approximately one-third of the standing population of mussels >10 mm in the area. Our results suggest that the predation effects of vertebrates on Baltic blue mussel populations are not insignificant, as previously believed. Predation by roach and other predators may have an important structuring effect on unstable blue mussel communities within the Gulf of Finland, where the species lives at the edge of its range.Communicated by M. Kühl, Helsingør     

Toxic metal health risk by mussel consumption

Seafood is a major dietary food worldwide. However, seafood consumption by humans can induce health risk because seafood may be contaminated by various pollutants. The mussel Mytilus galloprovincialis is widely distributed in the coastal waters of Montenegro, SouthEast Adriatic Sea. Here, Zn, Fe, Cu, Ni, Cd, Pb, As, and Hg contents in M. galloprovincialis from ten sites were analyzed to investigate health risks associated with the consumption of wild and cultivated mussels. Since there is a lack of data on the mussel consumption rate in Montenegro, the amount of mussels that can be ingested weekly over a lifetime with no risk of negative health effects was calculated using provisional tolerable weekly intakes (PTWI). We found that Cd concentrations were the limiting factor for mussels as a food. The weekly consumptions of 0.64–1.2 kg of fresh wild and 0.84–1.2 kg of fresh cultivated mussel would be sufficient to reach the PTWI_Cd, which may result in a risky weekly intake of Cd for long-term exposure. Moreover, weekly intake of 125 g mussels was used to calculate the dietary intake of trace elements by mussel consumption and compared with the prescribed PTWIs. Here, we found that there is no risk for human health for all investigated elements. In this case, the highest Cd level obtained in wild and in cultivated mussels represents 19.8 and 14.9% of the PTWI_Cd, respectively. This is the first study in Montenegro giving an assessment of the health risk from trace elements via the consumption of wild and cultivated M. galloprovincialis.     

The relationship between allele frequency and tidal height in a mussel hybrid zone: a test of the differential settlement hypothesis

The blue mussels Mytilus edulis L. and M. galloprovincialis Lmk. hybridize in western Europe. Within hybrid populations nuclear alleles specific to M. galloprovincialis increase in frequency with age and size. This relationship changes with tidal height; alleles from M. galloprovincialis occur more frequently high in the intertidal zone, while M. edulis alleles predominate in the low intertidal zone. We tested the hypotheses that larvae with M. galloprovincialis alleles tend to settle higher in the intertidal zone, or that mussels redistribute themselves with respect to tidal height after initial larval settlement. We sampled recently metamorphosed mussels every 2 weeks in a hybrid mussel population at Whitsand Bay in southwest England throughout the summer of 1996. We observed four cohorts of newly settled mussels. There was no evidence of differential settlement of mussels with different genotypes in connection with tidal height, or into shaded versus unshaded microsites. Therefore, we rejected the preferential settlement hypothesis. There was substantial movement of juvenile mussels in the first 4 weeks following initial settlement, but this “secondary settlement” did not result in genetic differentiation with respect to tidal height. Further, significant differences in allele frequencies were found between primary and secondary spat. This allele frequency change was in the opposite direction of that seen in the adult population, suggesting newly settled larvae may be experiencing different selective pressures than adults. We propose that the genetic structure of hybrid mussel populations with respect to tidal height is the consequence of differences in selection intensity. Received: 30 April 1999 / Accepted: 5 May 2000