Sand and wave induced mortality in invasive (<Emphasis Type="Italic">Mytilus galloprovincialis</Emphasis>) and indigenous (<Emphasis Type="Italic">Perna perna</Emphasis>) mussels

The ability of an invasive species to spread in a new locality depends on its interaction with the indigenous community and on variation in time and space in the environment. The Mediterranean mussel Mytilus galloprovincialis invaded the South African coast 30 years ago and it now competes and coexists with the indigenous mussel Perna perna. The two species show different tolerances to wave and sand stress, two of the main environmental factors affecting this intertidal community. P. perna is more resistant to hydrodynamic stress than M. galloprovincialis, while the invasive species is less vulnerable to sand action. Our results show that mortality rates of the two species over a period of 6 months had different timing. The indigenous species had higher mortality than M. galloprovincialis during periods of high sand accumulation in mussel beds, while the pattern reversed during winter, when wave action was high. A negative correlation between sand accumulation and attachment strength of the two mussels showed that sand not only affects mussel mortality through scouring and burial, but also weakens their attachment strength, subjecting them to a higher risk of dislodgement. Here we underline the importance of variations in time and space of environmental stress in regulating the interaction between invasive and indigenous species, and how these variations can create new competitive balances.     

Mechanisms of habitat segregation between an invasive (Mytilus galloprovincialis) and an indigenous (Perna perna) mussel: adult growth and mortality

The invasive mussel Mytilus galloprovincialis and the indigenous mussel Perna perna coexist intertidally on the south coast of South Africa through partial vertical habitat segregation: M. galloprovincialis dominates the upper shore and P. perna the lower shore. Recruitment patterns can explain the zonation of P. perna, but not the invasive species. We examined the role of post-recruitment interactions by measuring spatial and temporal differences in adult growth and mortality rates of the two species. Specifically, we tested the hypothesis that interspecific differences in growth and mortality reflect adult distribution patterns. The two study locations, Plettenberg Bay and Tsitsikamma, are 70 km apart with two sites (separated by 300–400 m) per location, each divided into three vertical zones. Growth was measured seasonally using different marking methods in 2001 and 2003. Cumulative adult mortality was measured through summer in 2003/2004. Both species generally grew more slowly upshore, but they showed different effects of season. For P. perna, growth was significantly reduced in winter in the low zone, but unaffected by season in the high zone. For M. galloprovincialis, growth was either unaffected by season or increased in winter, even in the high zone. Thus, growth of P. perna and M. galloprovincialis was reduced under cool winter and warm summer temperatures, respectively; and while growth was more similar between species in summer, M. galloprovincialis grew much faster than P. perna in winter. Mortality of P. perna increased upshore. For M. galloprovincialis, mortality was not zone-dependent and was significantly greater than for P. perna on the low-shore and (generally) across the shore in Tsitsikamma. Both species had higher growth and mortality rates in Plettenberg Bay than in Tsitsikamma. Thus, P. perna seems able to maintain spatial dominance on the low-shore and at certain sites because of higher mortality of M. galloprovincialis. We conclude that seasonality in growth of the two species reflects their biogeographic affinities and that coexistence is possible through pre-recruitment effects that limit the vertical distribution of P. perna and post-recruitment effects that limit M. galloprovincialis.     

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.     

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.     

Wave action and competitive interaction between the invasive mussel Mytilus galloprovincialis and the indigenous Perna perna in South Africa

On the south coast of South Africa, the invasive alien mussel Mytilus galloprovincialis shows partial habitat segregation with the indigenous mussel Perna perna. P. perna predominates in the lower mussel zone and M. galloprovincialis in the upper zone, with mixed beds where the two overlap. We examined competitive interactions between these species by translocating mussels into small plots at high densities. Treatments involved different combinations of species and densities placed in each zone. Mortality was monitored regularly and at the end of each experiment, growth and condition index were measured. The experiment was attempted three times. The first two attempts were disrupted by wave action, especially winter storms, but provided information on species-specific effects of wave action. In experiment one, wave induced mortality decreased from a mean for both species of approximately 90% on the low shore to ca. 50% on the high shore, and was 15–30% lower for P. perna than M. galloprovincialis in each zone. In experiment two, M. galloprovincialis mortality was not affected by zone (Kruskal–Wallis test, P > 0.05), but was higher than P. perna mortality in the low zone (P < 0.05). P. perna survival was significantly (P < 0.05) lower on the high than mid and low zones, apparently due to the effects of greater emersion. Condition index showed a similar pattern, being lowest in the low zone for M. galloprovincialis and in the high zone for P. perna (3-way ANOVA, P < 0.05). Growth rates were fastest for both species in the low zone (Kruskal–Wallis, P < 0.05 in both cases). The third experiment was run for 12 months in the low zone only and provided evidence of intraspecific competition for P. perna and of interspecific competition. Condition was significantly greater for P. perna in all treatments (2-way ANOVA), as was growth (Kruskal–Wallis P < 0.05). Significant treatment effects indicated that P. perna had a negative effect on M. galloprovincialis survival (Kruskal–Wallis, P < 0.05). Again wave action was important; by the end of the experiment all mussels had been removed from plots stocked only with M. galloprovincialis. Thus P. perna improves survival of M. galloprovincialis on the low shore in the short term, by providing protection against wave action, but excludes it competitively in the longer term. The results show that partial habitat segregation is likely to be a permanent feature on the south coast, with M. galloprovincialis unable to dominate the low shore due to the effects of waves and competitive exclusion by P. perna. This study is the first attempt to examine the mechanisms of interaction between invasive and indigenous marine mussel species and provides evidence of the importance of environmental conditions in the mediation of this interaction.     

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.     

Implications of movement behavior on mussel dislodgement: exogenous selection in a<Emphasis Type="Italic"> Mytilus</Emphasis> spp. hybrid zone

Rocky intertidal habitats often exhibit high levels of environmental heterogeneity, and the ability of organisms to move between microhabitats is likely to have a profound influence on their rates of mortality and overall fitness. Mussels within the Mytilus edulis complex are morphologically very similar, yet at sites where these species hybridize in southwest England, populations repeatedly show evidence of selection against individuals with alleles specific to M. edulis Linnaeus, in favor of those with alleles specific to M. galloprovincialis Lamarck. Differential movement rates of these two species were examined within simulated mussel beds (gravel substrate) in the winter (February) and summer (July) of 2001. M. edulis-like mussels moved more frequently and more quickly to the exterior of gravel beds than did M. galloprovincialis-like mussels. Coupled with measurements of attachment strength in the field conducted in July 2001, we used a wave force model to examine the probability of dislodgement for each species under a range of water velocities. Results suggest that by preferentially moving to the exterior of beds, M. edulis experiences higher dislodgement rates due to exposure to large hydrodynamic forces than do M. galloprovincialis. As a consequence of lower attachment strengths, M. edulis is also predicted to have higher mortality rates than M. galloprovincialis in interior portions of the bed. Thus, differential movement behavior may contribute to the differential genotype-specific mortality rates observed in the Mytilus spp. hybrid zone in southwest England, and is an example of behavior potentially modifying rates of exogenous selection in an intertidal hybrid zone.Communicated by J.P. Grassle, New Brunswick     

Spatial and temporal variation in distribution and protein ubiquitination for <Emphasis Type="Italic">Mytilus</Emphasis> congeners in the California hybrid zone

Along the west coast of North America, the invasive mussel Mytilus galloprovincialis and a native congener M. trossulus overlap in range and compete for habitat in an extensive hybrid zone along central California. The two species have been shown to exhibit differential abiotic tolerances in laboratory studies, yet little is known about how such tolerances affect spatial and temporal patterns of geographic distribution, particularly in areas of competition. We examined distributions of the two congeners and their hybrids in neighboring intertidal and subtidal habitats in Bodega Bay, CA over 2 years, and compared shell length and seasonal ubiquitin (Ub) conjugates to estimate protein turnover and physiological stress for the species at each site. The two species were spatially segregated, with M. galloprovincialis dominating the subtidal habitat, and M. trossulus constituting a majority of the intertidal mussel population. Hybrid individuals appeared in low numbers at both sites. For each habitat, there was no statistical difference between shell lengths of M. galloprovincialis and hybrids but M. trossulus mussels were statistically smaller than the other two. In regards to physiological performance, ubiquitin conjugate values showed different seasonal cycles for the two species, suggesting different periods of peak environmental stress. The highest levels of Ub-conjugated proteins were observed in winter for M. galloprovincialis and in summer for M. trossulus, consistent with the respective range edges for their distributions since Bodega Bay is near the northern range edge of the invader and the southern edge of the native species. These findings suggest that future assessments of Mytilus populations along the California coast may need to consider vertical distributions and seasonal cycles as part of monitoring and research activities.     

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.     

Factors influencing differential mortality within a marine mussel (Mytilus spp.) hybrid population in southwestern England: reproductive effort and parasitism

Two species of marine mussel, Mytilus edulis and M. galloprovincialis hybridize on the coasts of western Europe. Studies of hybrid mussel populations have shown that natural selection favors M. galloprovincialis-like genotypes within this hybrid zone. Many hypotheses have been proposed to explain differential mortality in these populations. This study tests two hypotheses addressing factors of mortality in a population, and describes yearly energy storage and reproductive cycles of these two species and their hybrids. No evidence was found that the two taxa have different overall levels of reproductive effort or parasite infestation. They do, however, have asynchronous spawning periods and divergent energy storage strategies. In the year of this study, 1993, the M. edulis genotypic class spawned as a group in June and July. After spawning, they built up a high level of mantle energy-storage tissues that are probably used for gametogenesis in the following winter and spring. The M. galloprovincialis genotypic group, however, spawned asynchronously, beginning in June and finishing by August, and did not build up high levels of energy-storage tissues in summer. These results add a temporal component to the interpretation of selective forces acting to shape this hybrid zone. Vulnerability of each species to mortality factors may differ because of their divergent reproductive and energy-storage cycles. Received: 15 January 1999 / Accepted: 26 July 2000     

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.     

Ecological gradients and relative abundance of native (Mytilus trossulus) and invasive (Mytilus galloprovincialis) blue mussels in the California hybrid zone

Marine communities are experiencing unprecedented rates of species homogenization due to the increasing success of invasive species, but little is known about the mechanisms that allow a species to invade and persist in a new habitat. In central California, native (Mytilus trossulus Gould 1850) and invasive (Mytilus galloprovincialis Lamarck 1819) blue mussels and their hybrids co-exist, providing an opportunity to analyze the mechanisms that determine the distributions of these taxa. Spatial and temporal variation in temperature and salinity and the relative frequencies of these mussel taxa were examined between 2000 and 2004 at four sites in San Francisco Bay and four in Monterey Bay, which were chosen for their different positions along inferred estuarine/oceanic gradients in the hybrid zone. Mussels were genetically identified as the parent species or hybrids by amplifying regions of two species-specific loci: the adhesive byssal thread protein (Glu-5′) and the internal transcribed spacer region of ribosomal DNA (ITS 1). The proportion of M. trossulus at the eight hybrid zone sites correlated negatively with average salinity (R ²=0.60) and positively with maximal temperature (R ²≥0.72), a somewhat unexpected result given what is known about the phylogeography of this species. The proportion of M. galloprovincialis showed the opposite pattern. The proportion of hybrids was correlated neither with habitat temperature nor salinity. Genotypes of mussel populations at an additional 13 sites from Coos Bay, Oregon (latitude 43.35°N) to Long Beach, California (latitude 33.72°N), sampled at various intervals between 2000 and 2004, were also determined. This survey confirmed previous reports that the hybrid zone lies between Monterey and the Cape Mendocino region (latitudes 36.63°N–40.5°N). Within Monterey and San Francisco Bays, however, the temporal comparisons (1990s vs. 2000s) revealed abrupt changes in the proportions of the two parent species and their hybrids on annual and decadal time scales. These changes indicate that the blue mussel populations are in a highly dynamic state. The survey also showed that, regardless of habitat, M. trossulus is consistently of smaller average size than either M. galloprovincialis or hybrids.

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Is habitat amount important for biodiversity in rocky shore systems? A study of South African mussel assemblages

Habitat-forming species on rocky shores are often subject to high levels of exploitation, but the effects of subsequent habitat loss and fragmentation on associated species and the ecosystem as a whole are poorly understood. In this study, the effects of habitat amount on the fauna associated with mussel beds were investigated, testing for the existence of threshold effects at small landscape scales. Specifically, the relationships between mussel or algal habitat amount and: associated biodiversity, associated macrofaunal abundance and density of mussel recruits were studied at three sites (Kidd’s Beach, Kayser’s Beach and Kini Bay) on the southern and south-eastern coasts of South Africa. Samples, including mussel-associated macrofauna, of 10 × 10 cm were taken from areas with 100 % mussel cover (Perna perna or a combination of P. perna and Mytilus galloprovincialis) at each site. The amount of habitat provided by mussels and algae surrounding the sampled areas was thereafter determined at the 4.0 m² scale. A number of significant positive relationships were found between the amount of surrounding mussel habitat and the abundances of several taxa (Anthozoa, Malacostraca and Nemertea). Likewise, there were positive relationships between the amount of surrounding algal habitat and total animal abundance as well as abundance of mussel recruits at one site, Kini Bay. In contrast, abundance of mussel recruits showed a significant negative relationship with the amount of mussel habitat at Kayser’s Beach. Significant negative relationships were also detected between the amount of mussel habitat and species richness and total abundance at Kidd’s Beach, and between amount of mussel habitat and the abundance of many taxa (Bivalvia, Gastropoda, Maxillopoda, Ophiuroidea, Polychaeta and Pycnogonida) at all three sites. No threshold effects were found, nor were significant relationships consistent across the investigated sites. The results indicate that the surrounding landscape is important in shaping the structure of communities associated with these mussel beds, with significant effects of the amount of surrounding habitat per se. The strength and the direction of habitat effects vary, however, between shores and probably with the scale of observation as well as with the studied dependent variables (e.g. diversity, abundance, mussel recruitment, species identity), indicating the complexity of the processes structuring macrofaunal communities on these shores.     

Variation in strength of attachment to the substrate explains differential mortality in hybrid mussel (Mytilus galloprovincialis and M. edulis) populations

Mussel samples were collected from a hybrid mussel (Mytilus galloprovincialis and M. edulis) Population at Croyde, southwest England, in January, March and May 1990. The strength of attachment of each mussel to the substrate was measured with a spring balance. A number of diagnostic characters were also recorded. These are shell lengh, width and height, mantle colour and genotype at two allozyme loci, esterase-D and octopine dehydrogenase. Multiple-regression analysis was used to assess the effect of the diagnostic characters on strength of attachment as dependent variable. Mussels possessing the relatively high shells and darker mantle colouration characteristic of M. galloprovincialis had higher values, on average, for strength of attachment than mussels resembling M. edulis. Phenotypically intermediate mussels had intermediate values for strength of attachment. The results suggest an adaptive difference which can account for reports of differential mortality acting in favour of M. galloprovincialiis.     

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     

Genetic variation of mitochondrial DNA in mussel (Mytilus edulis and M. galloprovincialis) populations from South West England and South Wales

Mitochondrial DNA (mtDNA) and allozyme variation were analysed in samples of mussels collected in 1984 and 1985 from four localities in South West England and one locality in South Wales, a region of Britain where the common mussel (Mytilus edulis) occurs sympatrically and hybridises with the Mediterranean mussel (M. galloprovincialis). Significant differences in mtDNA genotype frequencies for three restriction enzymes (BstEII, XbaI, and EcoRI) were observed between mussels from M. galloprovincialis populations (Padstow and Bude) and those from an M. edulis population (Swansea). Some mtDNA genotypes at high-frequency in M. galloprovincialis were not observed in M. edulis, although there was no indication that mtDNA variation provides greater overall diagnostic power than allozyme variation in distinguishing between the two forms of mussel. Construction of a phylogenetic tree of multiple mtDNA genotypes revealed small mutational distances between the genotypes characterising M. edulis and M. galloprovincialis. The results were consistent with predominant mtDNA flow from M. edulis to M. galloprovincialis. This can be explained by the dispersal of larvae to South West England from M. edulis regions to the north and east, but little dispersal in the opposite directions. Samples from two hybrid populations (Whitsand and Croyde) were analysed. mtDNA genotype frequencies at Croyde were in line with predictions made on the basis of two partially diagnostic allozyme loci (Est-D and Odh), mtDNA frequencies at Whitsand were not. Frequencies of some mtDNA genotypes at Whitsand were characteristic of M. edulis, others of M. galloprovincialis. Differential selective mortality or flow of different mtDNA genotypes and allozyme variation are proposed as possible causes of these results.     

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     

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.     

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.     

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.