Desiccation,higher temperatures and upper intertidal limits of three species of sea mussels (Mollusca: Bivalvia) in New Zealand

Distribution, abundance, and resistance adaptations to higher temperature and desiccation of three species of intertidal mussels (Mytilus edulis aoteanus, Perna canaliculus and Aulacomya maoriana) were studied in New Zealand. M. edulis aoteanus generally was more abundant upshore, with P. canaliculus dominating downshore. M. edulis aoteanus was more common than P. canaliculus on the outside of mixed-species clumps. Abundance of A. maoriana was variable, with individuals favouring damp habitats such as inside mussel clumps. In moving air at 75% relative humidity and at 20°C or 30°C, median lethal levels of water loss were similar for P. canaliculus and M. edulis aoteanus but lower for A. maoriana. Rates of desiccation varied inversely with size and were higher for P. canaliculus, due mainly to valve gaping with resultant loss of water from the mantle cavity. M. edulis aoteanus was more tolerant of higher water temperatures than were the other species. Success in colonizing upshore or more aerially exposed habitats seems to be related to ability of small mussels to tolerate desiccation, especially during hot, windy weather.     

Biochemical genetic variation at a leucine aminopeptidase (LAP) locus in blue (Mytilus galloprovincialis) and greenshell (Perna canaliculus) mussel populations along a salinity gradient

In several marine bivalve species, biochemical genetic variation at a leucine aminopeptidase (LAP) locus is associated with environmental variability, primarily salinity fluctuation. Population genetic variation at a LAP locus was investigated here in two sympatric mussel species (Mytilus galloprovincialis and Perna canaliculus) from three locations along a salinity gradient in Wellington Harbour, New Zealand. The data for M. galloprovincialis and P. canaliculus do not support the hypothesis that the LAP polymorphism in either species is associated with salinity variation among adult mussels. Due to the absence of small mussels among the samples it is not possible to discount the hypothesis that selection acts primarily against juveniles, as it does for M. edulis in Long Island Sound, USA. Wellington Harbour populations of M. galloprovincialis exhibited large and often highly significant heterozygote excesses at the LAP locus, whereas populations of P. canaliculus from the same locations exhibited large and highly significant heterozygote deficiencies. The reason for this inter-specific difference in population structure is unknown. If it is the result of selection, this suggests that selection acts differentially upon the two species, because demographic attributes and reproductive biology are very similar in the two species. For both M. galloprovincialis and P. canaliculus, significant levels of population genetic heterogeneity were recorded among three locations separated by only 8 to 12 km. Neither species exhibited shell length-dependent genetic variation at the LAP locus, suggesting that for these two species the LAP polymorphism is not associated with variation in shell length. Received: 30 December 1996 / Accepted: 6 January 1997     

Condition cycle of the green-lipped mussel Perna canaliculus in New Zealand

The condition of the green-lipped mussel Perna canaliculus (Gmelin) from 7 sites around the northern half of New Zealand was determined monthly between December 1973 and February 1975. Seven separate estimates of condition were calculated for each sample; 4 condition indices, 2 percentage glycogen analyses and 1 estimate of percentage solids. All showed a similar annual cycle, with minimum values in winter (June/ August) rising to peak levels in spring (October/ December). The 7 sites were divided into 3 groups, (i) a central New Zealand group, characterised by high levels of condition index (CI) throughout most of the year >10 CI_weight for 8 to 11 months, average to high percentage glycogen (annualmeans 25 to 35% dry weight), and high percentage solids (annual means > 20% dry weight); (ii) an intertidal group, having low levels of CI (<10 CI_weight for 10 to 11 months), low percentage glycogen(mean <20%), and low percentage solids (mean <18%); and (iii) a northern group, showing intermediate values for each of the estimates of condition. Difficulties in defining the causes of changes in the condition cycle are discussed. There is an inverse relationship between CI and mussel size, resulting from changes in body proportions. An inverse relationship is also suggested between CI and water temperature. A precise index, $$CI_{weight} = \frac{{100 \times dry meat weight}}{{whole{\text{ weight - shell weight}}}}$$ is recommended for biological studies on mussels and a rapid simple index, $$CI_{commercial} = \frac{{100 \times wet meat weight}}{{whole{\text{ (live) weight}}}}$$ for use in the field and in mussel farming practice.     

Combining heat-transfer and energy budget models to predict thermal stress in Mediterranean intertidal mussels

Recent studies have emphasised that organisms can experience physiological stress well within their geographic range limits. Developing methods for mechanistically predicting the presence, absence and physiological performance of organisms is therefore important because of the ongoing effects of climate change. In this study, we merged a biophysical–ecological (BE) model that estimates the aquatic (high tide) and aerial (low tide) body temperatures of Mytilus galloprovincialis with a Dynamic Energy Budget (DEB) model to predict growth, reproduction and mortality of this Mediterranean mussel in both intertidal and subtidal environments. Using weather and chlorophyll-a data from three Mediterranean sites along the Italian coasts, we show that predictions of sublethal and lethal (acute) stress can potentially explain the observed distribution (both presence and absence) of mussels in the intertidal and subtidal zones, and the maximum size of animals in the subtidal zones. Importantly, our results suggest that different mechanisms limit the intertidal distribution of mussels, and that these mechanisms do not follow a simple latitudinal gradient. At the northernmost site (Palermo), M. galloprovincialis appears to be excluded from the intertidal zone due to persistent exposure to lethal aerial temperatures, whereas at the southernmost sites (Porto Empedocle and Lampedusa) sublethal stress is the most important driver of mussel intertidal distribution. Our predictions provide a set of hypotheses for future work on the role of climate change in limiting intertidal distribution of mussels in the Mediterranean.     

Very long-chain aliphatic hydrocarbons in lipids of mussels (Mytilus edulis) suspended in the water column near petroleum operations off Sable Island,Nova Scotia,Canada

Mussels (Mytilus edulis) suspended in the water column in 1994 and 1995 for the monitoring of oil drilling operations off Sable Island, Nova Scotia were examined for hydrocarbon profiles, particularly aliphatic hydrocarbons. A spring bloom of phytoplankton occurred during the 90-d suspension period in 1995. Hydrocarbons isolated from the 1995 suspended mussels showed very high concentrations of both biogenic hydrocarbons and very long-chain n-alkanes from C₂₀ to C₃₂, initially thought to be petrogenic. Both types of hydrocarbons were either not detected or were only present in trace amounts in the mussels suspended in 1994 at similar sites. The biogenic hydrocarbons in the 1995 mussels were apparently of planktonic origin, from the spring bloom, and were dominated by heneicosahexaene (21:6), followed by pristance, heptadecane, and varions monounsaturated and polyunsaturated phytenes, heptadecenes, nonadecenes and heneicosenes. They could be readily hydrogenated to yield the basic alkanes. The 1995 mussels suspended within 1 km from the oil well platform were probably slightly tainted by petrogenic hydrocarbons, as evidenced by the detection of phytane and high concentrations of total aliphatic hydrocarbons, whereas the mussels suspended 10 km from the platform showed only high concentrations of biogenic hydrocarbons and the novel long-chain n-alkanes. The occurrence of an unusual phytoplankton bloom during the suspension period severely interfered with the petroleum monitoring role of mussels by altering the mussel hydrocrbon profiles through the accumulation into and probably selective depuration of xenobiotic hydrocarbons from the mussel, tissues.J. Parsons (deceased)     

Gamete production,somatic growth and multiple-locus enzyme heterozygosity inMytilus edulis

Mussels,Mytilus edulis L., were collected from the lower shore in Stony Brook Harbor, New York, USA, in April 1984. The mussels were in spawning condition at the time of collection. Mantle weight was used as an index of fecundity and somatic tissue weight and growth rate were measured so that the relative production of gametes and somatic tissue with age could be calculated. For each individual mussel the genotypes at loci coding for five enzymes were determined. Somatic tissue production exceeds gamete production for the first two and a half years of life, but thereafter it declines and is exceeded by gamete production. There is a positive correlation between multiple-locus enzyme heterozygosity and fecundity in mussels which have grown beyond the size at which gamete production starts to exceed somatic production. There is no correlation between heterozygosity and fecundity in younger individuals. We conclude that the higher scope for growth of more heterozygous bivalves, described elsewhere, is translated into allocation of energy to somatic growth during early life and into gamete production in later life. Our observations perhaps explain why, in other studies, the correlation between growth rate and heterozygosity in bivalves is manifested most clearly when young individuals are examined.     

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.     

Effects of mussel aquaculture on the nitrogen cycle and benthic communities in Kenepuru Sound,Marlborough Sounds,New Zealand

Nitrogen pools and transformations and benthic communities at a Perna canaliculus farm and a nearby reference site without direct influence of marine farming in Kenepuru Sound, New Zealand, were compared on four dates between September 1982 and May 1983. The organic nitrogen pool in the top 12 cm sediment was 7.4 to 10.8 mol m^-2 at the mussel farm and 6.1 to 8.9 mol m^-2 at the reference site. The nitrate and nitrite pools were similar in both sediments, but the ammonium pool in the mussel farm sediment was about twice as high as in the reference sediment. In January, the sediment ammonium concentrations ranged from 418 nmol cm^-3 (surface) to 149 nmol cm^-3 (12 cm depth) at the mussel farm and from 86 to 112 nmol cm^-3 at the reference site. The molar C:N ratio of the sediment organic matter was 6.2 to 7.2 at the mussel farm and 7.9 to 10.0 at the reference site. The molar N:P ratio of the sediment organic matter was 4.3 to 7.2 and 3.3 to 6.1 at mussel farm and reference site, respectively. The total nitrogen mineralisation rate in the top 12 cm sediment ranged from 21.7 to 37.1 mmol m^-2 d^-1 at the mussel farm and from 8.5 to 25.0 mmol m^-2 d^-1 at the reference site. Ammonium excretion by mussels was about 4.7% (January) and 7.4% (May) of the combined nitrogen mineralisation by mussels and sediment. The sediment-denitrification rate was 0.7 to 6.1 mmol m^-2 d^-1 at the mussel farm and 0.1 to 0.9 mmol m^-2 d^-1 at the reference site. In January, 76 and 93% of the nitrate reduced in the sediments were denitrified at the mussel farm and reference site, respectively. The denitrification rate on the mussel lines (determined on detritus-covered mussels) was twice the mussel farm sediment-denitrification rate and 10 times the reference sediment-denitrification rate. Total denitrification at the mussel farm was 21% higher than at the reference site. The loss of nitrogen through mussel harvest and denitrification was 68% higher at the mussel farm. The surface layers of both sediments contained about 75 mg m^-2 chlorophyll a. Sediment phaeophytin levels were 52 mg m^-2 at the reference site and 137 mg m^-2 at the mussel farm. While the benthic infauna of the mussel-farm sediment consisted only of polychaete worms, the reference sediment contained also bivalve molluscs, brittle stars and crustaceans.     

Resource allocation in Mytilus edulis on the shore and in suspended culture

The annual cycle of carbon and nitrogen content of the flesh of wild and cultivated mussels (Mytilus edulis L.) in Killary Harbour, Ireland, was measured over two years, starting in February, 1980 and ending in November, 1981. The carbon and nitrogen contents of mussel gametes were determined and the allometry of growth of wild and cultured mussel shells was examined with respect to length, weight and organic content. The carbon and nitrogen contents of the organic fraction of the shell were determined. These data were combined with those we had previously published on growth rate, gametogenesis and the annual cycle of ash-free dry weight (AFDW) of mussels in the same locality. Estimates were made of fecundity, reproductive effort and the partitioning of carbon and nitrogen between soma, gametes and shell. In suspended culture, cumulative production after eighteen months is equal to cumulative production after six to seven years on the shore. For comparison, partitioning of carbon and nitrogen resources between soma, shell and gonad is estimated when total cumulative production by wild and cultivated mussels is approximately equal. Differences in resource allocation are considerable. Wild mussels allocate some 57% of their carbon budget and 52% of their nitrogen budget to gamete output. In culture, mussels allocate only 22% of their carbon budget and 19% of their nitrogen budget to gamete output. It is concluded that in response to a higher production rate, cultivated mussels increase allocation of resources to somatic growth.     

How to toughen up your mussels: using mussel shell morphological plasticity to reduce predation losses

Improvements in stocking strategy and management could increase the yield of mussels that are on-grown from harvested wild seed mussel resources and thereby enhance the sustainability of this shellfishery. A field experiment was undertaken to ascertain shell characteristics (compression strength and thickness) of seed mussels grown at different shore heights, whether these characteristics changed after a period of growth under identical conditions, and if these characteristics reduced predation losses by crabs and birds. Results indicated that mussels grown at higher shore levels attained shell characteristics beneficial to predation resistance and that these were maintained after a period of growth at a lower shore level. A novel management plan for mussel cultivation was formulated from the results of this study by manipulating shore position according to the attainment of these predator resistant shell attributes and the spatial distribution of the main natural mussel predators (crabs and birds). This technique was expanded to address the mussel cultivation problem of low natural seed settlement.     

Seascape-dependent subtidal-intertidal trophic linkages

In this study, we test in southern New Zealand a conceptual model of food web linkage that is seascape dependent, which can explain some of the variability in rocky shore community structure among sites and coasts. Using a comparative-experimental approach at local and distant sites we demonstrate that mobile subtidal predators (fish and crabs) can exert strong predation pressure on small mussels in the low tidal zone, but only in sites where the seascape includes subtidal reefs. On intertidal benches with adjacent subtidal reefs (+SR), 60-100% of small (5-15 mm) transplanted mussels were removed within a day from experimental tiles on the low shore when unprotected from predation, compared to fully caged controls that had approximately 100% survival over several months. In partial cages that exclude fish but not crabs, survivorship was intermediate. In contrast, on benches without subtidal reefs (-SR) 40-100% of mussels survived for months, even if unprotected. This difference is expressed in lower cover (0-60%) of mussels on rocks at +SR benches compared to -SR benches (70-99%). The central to northern west coast of the South Island is composed mostly of -SR benches, and predation on small mussels there was low and similar to the -SR benches on the east coast, whereas the +SR benches on the east coast had much greater predation. This contrasts to other studies in New Zealand that examined only predation on larger mussels by seastars and concluded that predation is strong on the west coast and weak on the east coast. Excluding large predators from low-shore areas with new recruits for a year in one +SR site showed longer-term predation effects on their abundance and cover. Short-term sampling at the east coast sites showed that mussel settlement was greater in -SR compared to +SR sites, providing some evidence that seascapes may also affect settlement. Overall, predation depended on the local seascape and ultimately affected community structure via suppression of effective recruitment rates. This study emphasizes the importance of predation on early life stages of basal species and the influence of seascapes on top-down interactions between subtidal predators and their intertidal prey.     

Differences in morphology and habitat use among the native mussel <Emphasis Type="Italic">Mytilus trossulus</Emphasis>, the non-native <Emphasis Type="Italic">M. galloprovincialis,</Emphasis> and their hybrids in Puget Sound,Washington

Mytilus galloprovincialis (Mg), the Mediterranean blue mussel, is sympatric with the native M. trossulus (Mt) throughout much of the north Pacific, likely as the result of human introduction. We investigated the distribution of the two species and their hybrids (Mgt) in Puget Sound, Washington, to determine whether differences occur in habitat preference between the two species and hybrids. In addition, we investigated whether there were consistent size and shape differences between the native and introduced mussels and hybrids. Measurements of over 6,000 mussels from 30 sites, of which 1,460 were genotyped for a species-specific genetic marker, revealed that Mg and Mgt can be found throughout Puget Sound. Mg and Mgt were larger and exhibited a greater height:length ratio than Mt. Frequencies of Mg and Mgt were higher in subtidal habitats, such as docks, than on intertidal rocks, walls or pilings. Within intertidal habitats, Mg and Mgt were more frequent than Mt in the lower reaches of the intertidal. At slightly more than half the sites the frequency of the three genotypes accorded with random mating expectations suggesting no consistent barriers to gene flow between species. The standardized random sampling methods and simple morphometric identification techniques described here can be used to test whether the frequency of invasive mussels changes over time and space in Puget Sound.     

Genotype-dependent fecundity and temporal variation of spawning in hybrid mussel (Mytilus) populations

Mussel samples were collected at 4 to 6 wk intervals throughout 1987 from two hybridMytilus edulis/M. galloprovincialis populations, at Croyde Bay and Whitsand Bay, in southwest England. These were analyzed at two polymorphic loci which are diagnostic for allozyme differences which typifyM. edulis andM. galloprovincialis. Dry mantle weight as a function of shell length was determined for all individuals of each sample. Size-frequency data for the two populations was obtained in September 1987 and March 1988. For all genotypes at both sites, fecundity was a function of shell length, and in both populations the frequency ofM. galloprovincialis alleles was positively correlated with shell length. At both sites, allozyme genotype explained a significant amount of variation in mantle weight either when assessed as a main effect or when assessed as an interaction with shell length or time of collection. At Croyde,M. galloprovincialis mussels had greater estimated fecundity per unit length than theM. edulis mussels. Differences in the timing of spawning activity between theM. edulis and theM. galloprovincialis mussels were inferred, and these differences might act to reduce the amount of interbreeding at Croyde. At Whitsand, a reduced level of variability in the timing of spawning activity and fecundity between the genotypes was observed and explained by a higher degree of genetic mixing. Because theM. galloprovincialis mussels had (1) a greater estimated fecundity at any length, and (2) a greater mean length than theM. edulis mussels, the mean genotypic annual fecundity perM. galloprovincialis mussel was 2.8 times greater than an individualM. edulis mussel at Croyde, and 2.2 times greater than an individualM. edulis mussel at Whitsand. This evidence thatM. galloprovincialis mussels have an advantage in fecundity, and thus perhaps in fertility, taken together with the evidence thatM. galloprovincialis also has a higher viability, indicates directional selection in favour of theM. galloprovincialis phenotype. Because of the observed temporal stability of the population it seems likely that this selection is counterbalanced by a massive imigration ofM. edulis spat from neighbouring populations.     

An ecosystem model for estimating potential shellfish culture production in sheltered coastal waters

A generic ecosystem model has been developed for estimating the potential production of shellfish culture and the effect of that cultivation on the pelagic ecosystem in sheltered coastal waters. The model describes the dynamics of a simple food web, nutrient cycling and growth of shellfish. The design of the model is closely tied to the temporal and spatial scales that are important in determining the sustainable production level for a particular embayment. The pelagic ecosystem, mussel energetics, population dynamics and hydrodynamics are coupled to allow fully dynamic predictions of the effect of the shellfish density. When applied to Beatrix Bay, an intensive culture embayment in the Pelorus Sound of New Zealand, the model successfully captured main features of the observed system behaviour. The hydrodynamic regime of the bay controls mussel growth and production. Although high fluxes of water into the bay suppress nutrient and carbon cycling signals in the system, the model simulations demonstrated that the mussel cultivation can have considerable effects on the ecosystem of the bay including food depletion and nutrient cycling. One of the most obvious effects is nutrient enhancement through mussel excretion at low cultivation densities, which promotes primary production particularly during the N-limitation period in summer. The sensitivity analysis identified uncertainty in some parameters and indicated areas for which experimental studies could lead to model improvement. The modelling exercise has established a primary predictive tool for managing mussel aquaculture of a coastal embayment to estimate relationships between the stock level and the growth rate of mussels, and the potentially achievable harvest and stocking density.     

Stylochus mediterraneus (Turbellaria: Polycladida), predator on the mussel Mytilus galloprovincialis

The polyclad Stylochus mediterraneus Galleni has been found associated with the mussel Mytilus galloprovincialis Lmk., on which it feeds. Polyclads allowed to feed freely on groups of mussels of different sizes preyed mainly on small mussels <25 mm in length. The predation rate (number of mussels eaten per no. of polyclads per no. of days) ranged between 0.07 and 0.33. The average amount of food ingested by 1 polyclad feeding on mussels 16 to 25 mm or 26 to 35 mm in length was 7.1 and 11.2 (dry weight) mg per day, respectively. Before penetrating the mussels, the worm first straddles the valves at the posterior edge of the shell and then, after having digested the posterior adductor muscle, removes and swallows the soft parts of the prey.     

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.     

Tidal height,rather than habitat selection for conspecifics,controls settlement in mussels

Settlement is a major determinant of intertidal populations. However, the energy costs of lost larvae are very high. Accordingly, arrival and attachment on suitable substrata are essential requirements for species’ survival. On the intertidal, the presence of cues left by adult or juvenile conspecifics could be vital for the successful establishment of larvae arriving on the shore. Two mussel species, the indigenous Perna perna and the invasive Mytilus galloprovincialis, co-occur on the lower eulittoral zone on the south coast of South Africa. P. perna dominates the low and M. galloprovincialis the high mussel zones, with co-existence in the mid mussel zone. This study tested the hypothesis of settlement selectivity for conspecifics in these two mussel species, to understand whether the final adult distribution of mussels on the shores is determined by active behavioural and chemical mechanisms. Preferential selection by larvae for conspecifics was tested in the field during the peak settlement period in 2004 in natural mussel beds across zones and through manipulative experiments in the mid-zone where the species co-exist. On natural beds, settlement was determined by counts of settlers attached over 48 h onto artificial collectors. Collectors were placed on beds of P. perna and M. galloprovincialis present at both high- and low-adult densities, as well as in mixed beds. On such natural beds, settlers of both species consistently favored low-zone P. perna beds. Settlement patterns over 24 h onto experimentally created mussel patches consisting of P. perna, M. galloprovincialis or the two species combined beds, set in the mixed zone, did not conform with the results of the natural beds study: settlers of both species settled with no discrimination among different patches. The results indicate that mussels, which are sedentary, lack attraction to conspecifics at settlement. This highlights the importance of tidal height in setting settlement rates, and of post-settlement events in shaping populations of these broadcast spawners.     

Hydrodynamic stress and habitat partitioning between indigenous (Perna perna) and invasive (Mytilus galloprovincialis) mussels: constraints of an evolutionary strategy

The ability of a mussel to withstand wave-generated hydrodynamic stress depends mainly on its byssal attachment strength. This study investigated causes and consequences of different attachment strengths of the two dominant mussels species on the South African south coast, the invasive Mytilus galloprovincialis and the indigenous Perna perna, which dominate the upper and the lower areas of the lower balanoid zone, respectively and co-exist in the middle area. Attachment strength of P. perna was significantly higher than that of M. galloprovincialis. Likewise solitary mussels were more strongly attached than mussels living within mussel beds (bed mussels), and in both cases this can be explained by more and thicker byssal threads. Having a wider shell, M. galloprovincialis is also subjected to higher hydrodynamic loads than P. perna. Attachment strength of both species increased from higher to lower shore, in response to a gradient of stronger wave action. The morphological features of the invasive species and its higher mortality rates during winter storms help to explain the exclusion of M. galloprovincialis from the low shore. The results are discussed in the context of the evolutionary strategy of the alien mussel, which directs most of its energy to fast growth and high reproductive output, apparently at the cost of reduced attachment strength. This raises the prediction that its invasive impact will be more pronounced at sites subject to strong but not extreme wave action.     

Scope for growth of Mytilus?galloprovincialis (Lmk., 1819) in oligotrophic coastal waters (Southern Tyrrhenian Sea, Italy)

The ‘scope for growth’ (SFG) tool was used to study the growth performance of cultivated populations of Mytilus galloprovincialis (Lmk., 1819) in an oligotrophic area of the Southern Mediterranean Sea. The study was carried out between 1993 and 1996 by using data from four seasonal oceanographic cruises and from growth experiments. Water samples were collected and analysed for total suspended matter (TSM), particulate organic carbon (POC) and nitrogen (PON), particulate lipids, proteins and carbohydrates and chloropigments. The sum of the carbon equivalents of carbohydrates, proteins and lipids is indicated as the total biopolymeric particulate organic carbon (BPC) and was converted into a unit of energy in order to calculate the SFG of a theoretical mussel of 5 cm length. In order to test the performance of mussel growth at two depths (5 and 15 m water depth), mussel body size [as ash free dry weight (AFDW)] and the actual concentrations of BPC were used to calculate the monthly SFG using the physiological energetic relationships suggested in the current literature. Data from the field cruises led us to characterise the study site as ultra-oligotrophic (annual average of chloropigment concentration approximately 0.5 μg L⁻¹). SFG calculations allowed us to identify a site where mussels grown successively were found to reach a commercial size in approximately 12 months. The good agreement obtained between energetic response and subsequent production response suggests that the available energy from particulate food could be fully available for organic production for maintaining “proportionate” growth trajectories, even in a ultra-oligotrophic system.     

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.