Allozymes and morphometric characters of three species ofMytilus in the Northern and Southern Hemispheres

Many authors have considered the common mussels in temperate waters of the Northern and Southern Hemispheres to be a single cosmopolitan species,Mytilus edulis Linnaeus, 1758. Others have divided these mussels into several subspecies or species. Samples of mussels were collected from 36 locations in the Northern Hemisphere and nine locations in the Southern Hemisphere. Electrophoretic evidence from eight loci indicates that the Northern Hemisphere samples consist of three electrophoretically distinguishable species:M. edulis from eastern North America and western Europe;M. galloprovincialis Lamarck, 1819 from the Mediterranean Sea, western Europe, California, and eastern Asia; andM. trossulus Gould, 1850 from the Baltic Sea, eastern Canada, western North America and the Pacific coast of Siberia. Mussels from Chile, Argentina, the Falkland Islands and the Kerguelen Islands contain alleles characteristic of all three Northern Hemisphere species, but because they are most similar toM. edulis from the Northern Hemisphere, we suggest that they tentatively be included inM. edulis. These South American samples are morphologically intermediate between Northern HemisphereM. edulis andM. trossulus. Mussels from Australia and New Zealand are similar in allele frequency and morphometric characters toM. galloprovincialis from the Northern Hemisphere. FossilMytilus sp. are present in Australia, New Zealand and South America, which suggests that the Southern Hemisphere populations may be native, rather than introduced by humans. Morphometric characters were measured on samples which the allozyme data indicated contained a single species. Canonical variates analysis of the morphometric characters yields functions which distinguish among our samples of the species in the Northern Hemisphere.     

PCR-based detection of mtDNA haplotypes of native and invading mussels on the northeastern Pacific coast: latitudinal pattern of invasion

Frequencies of mitochondrial haplotypes characteristic of native Mytilus trossulus and introduced M. galloprovincialis were determined in populations along the west coast of North America from San Diego, California, to the Aleutian Islands, Alaska. We also identified the haplotypes of mussels cultured from larvae arriving in Coos Bay, Oregon, during 1988–1990 from sites in Japan in the seawater ballast of ocean-going ships. All mussels from ballast-water samples were M. galloprovincialis. We found that sampled populations north of San Francisco Bay, including Coos Bay, were entirely composed of mussels with the M. trossulus haplotype, while haplotypes of both species were present in all sites in and south of San Francisco Bay. The presence of M. trossulus in southern sites is contrary to evidence from allozyme studies, and we suggest that mtDNA introgression from M. trossulus to M. galloprovincialis may explain this discordance. This study demonstrates that, despite continued transport and release, M. galloprovincialis has not become established in northern sites. Failure to invade the north coast of North America may reflect environmental unsuitability for M. galloprovincialis. However, invasion success may be probabilistic, and the continuing release of M. galloprovincialis larvae may foreshadow a future successful invasion.     

Distribution of male and female mtDNA lineages in populations of blue mussels,Mytilus trossulus and M. galloprovincialis,along the Pacific coast of North America

Geographic variation in mitochondrial large subunit (16S) ribosomal RNA haplotypes was examined for blue mussels, Mytilus trossulus Gould, 1850 and M. galloprovincialis Lamarck, 1819, sampled from ten sites along the Pacific coast of the USA in January of 1993. Using polymerase chain reaction/restriction fragment length polymorphism (PCR/RFLP) assays we determined haplotype frequencies for both the male and female mussel mitochondrial DNA (mtDNA) lineages. Populations from Morro Bay south to San Diego, California, contained only M. galloprovincialis male and female haplotypes, while those from Arcata Bay, California, north to Port Orford, Oregon, were fixed for M. trossulus haplotypes. Populations from Monterey Bay to Bodega Bay, California, contained a mixture of M. trossulus and M. galloprovincialis haplotypes. Overall only 2 of 97 heteroplasmic individuals had a mixed M. trossulus/M. galloprovincialis mitochondrial genotype indicating that hybridization is uncommon in the populations sampled. Further, there was no evidence of extensive introgression between these mussel taxa at the level of mtDNA. This is in contrast to previously published results which suggested the significant introgression of M. trossulus haplotypes into southern populations containing primarily M. galloprovincialis nuclear genotypes. We feel the discrepancy lies in the ability of our assays to detect haplotypes corresponding to both the male and female mtDNA lineages. Potential explanations for the lack of mtDNA introgression include, low levels of backcrossing between hybrids and parental taxa, epistatic interaction between nuclear and mitochondrial genes and the breakdown of a sex-specific inheritance pattern for mtDNA in hybrids.     

Identification and validation of novel SNP markers in European populations of marine Mytilus mussels

Mussels of the genus Mytilus are widespread in both northern and southern hemispheres. Mytilus taxa are very important components of marine coastal ecosystems, but are difficult to differentiate morphologically. Sequencing and Sequenom MassARRAY iPLEX genotyping technology was used to identify and verify novel SNP markers in three European taxa of Mytilus: M. edulis, M. galloprovincialis, and M. trossulus. SNPs were localized in coding and non-coding sequences of some functionally important genes. Eight SNPs located in genes of the histone family, hsp70 and p53, were discovered and applied as novel markers for Mytilus taxa on a large European scale. Five of these differentiated the M. trossulus genome, two M. galloprovincialis, and one M. edulis. Other SNPs differentiated populations within taxa. The new SNPs will be a valuable tool for population studies of European Mytilus mussels. The percentage of polymorphic SNPs ranged from 19 to 100 % in 24 samples of mussels studied. Populations from Scotland, Mecklenburg Bight, and Norway had over 90 % polymorphic loci. Most loci were in Hardy–Weinberg equilibrium in all samples except for the one from Scotland. The highest percentages of heterozygotes were observed in the Atlantic (Banyuls and Vigo) and North Sea (Tjarno and Westerschelde) populations. An excess of homozygotes was observed in samples from Scotland, Norway, and the Barents Sea. Correspondence and Structure analysis also demonstrated the great heterogeneity of these three samples.     

Origin of the antitropical distribution pattern in marine mussels (Mytilus spp.): routes and timing of transequatorial migration

Many marine species, including mussels in the Mytilus edulis species group (i.e. M. edulis L., M. galloprovincialis Lamarck, and M. trossulus Gould), have an antitropical distribution pattern, with closely related taxa occurring in high latitudes of the northern and southern hemispheres but being absent from the tropics. We tested four hypotheses to explain the timing and route of transequatorial migration by species with antitropical distributions. These hypotheses yield different predictions for the phylogenetic relationship of southern hemisphere taxa relative to their northern counter-parts. The three Mytilus species were used to test these hypotheses since they exhibit a typical antitropical distribution and representative taxa occur in both the Pacific and Atlantic. Two types of mtDNA lineages were found among populations of mussels collected from the southern hemisphere between 1988 and 1996; over 90% of the mtDNA lineages formed a distinct subclade which, on average, had 1.4% divergence from haplotypes found exclusively in northern Atlantic populations of M. galloprovincialis. These data indicate that southern hemisphere mussels arose from a migration event from the northern hemisphere during the Pleistocene via an Atlantic route. The remainder of the southern hemisphere lineages (<10%) were very closely related to mtDNA haplotypes found in both M. edulis and M. galloprovincialis in the northern hemisphere, suggesting a second, more recent migration to the southern hemisphere. There was no evidence that southern hemisphere mussels arose from Pacific populations of mussels. Received: 8 December 1998 / Accepted: 8 November 1999     

Reproductive isolation and reproductive output in two sympatric mussel species (Mytilus edulis, M. trossulus) and their hybrids from Newfoundland

The mussels Mytilus edulis L. and M. trossulus Gould are found sympatrically in most areas of Newfoundland, with a low frequency of hybrids. To assess the potential for reproductive isolation, we sampled mussels from three sites in an eastern Newfoundland Bay from May–October 1996 to determine if there were differences in the reproductive cycles of the two species and their natural hybrids. In mussels with sheil lengths of 38–42 mm, males and females with mature gametes were dominant in June for M. edulis and hybrids, while M. trossulus showed a lower frequency of individuals with mature gametes. M. trossulus and hybrids spawned over a prolonged period (from late spring to early autumn) compared with most M. edulis individuals that spawned over a period of 2–3 weeks in July. This asynchrony in spawning activity between the two species may partially explain the low frequency of hybrids found in previous studies of these mussel populations. Female and male hybrids between M. edulis and M. trossulus showed normal gonad development, ripening and spawning, providing an opportunity for the introgression of genes between the two species. M. trossulus had a higher reproductive output than M. edulis of similar shell length, while hybrids showed intermediate values of reproductive output. M. trossulus females produced smaller eggs than either M. edulis or hybrids. Differences in reproductive traits may partially explain the maintenance of the mussel hybrid zone in Newfoundland. Published online: 13 August 2002     

Mitochondrial DNA lineages in the European populations of mussels (<Emphasis Type="Italic">Mytilus</Emphasis> spp.)

Marine mussels (Mytilus spp.) belong to a group of benthic species crucial to coastal ecosystems in Europe and are important for the cultivation industry. In the present study, the nuclear adhesive protein marker (Me15/16) was used for identification of Mytilus species in coastal areas, on a large geographic scale in Europe. Pure M. edulis populations were found in the White Sea and Iceland. M. edulis, M. trossulus and their hybrids were found in the Baltic Sea and the North Sea (Oosterschelde, The Netherlands). M. galloprovincialis, M. edulis and their hybrids occurred in Ireland. M. galloprovincialis populations were observed in the Sea of Azov (Black Sea), the Mediterranean and Portugal. The mitochondrial (mt) DNA coding-region ND2-COIII was studied by PCR (polymerase chain reaction) and RFLP (restriction fragment length polymorphism) assay methods. The mtDNA control region was studied by PCR. Substantial differentiation in the frequency of female haplotypes among the studied populations in Europe was observed. Despite isolation between the Mytilus taxa on a macro-geographic scale, considerable mitochondrial gene flow occurred between populations, with introgression in hybrid zones on a more local geographic scale in Europe. MtDNA of the Atlantic Iberian (Portugal) population of M. galloprovincialis was more similar to mtDNA in populations of M. galloprovincialis and M. edulis from the Atlantic coasts of the Ireland and M. edulis from the North Sea, than to M. galloprovincialis from the Mediterranean. Lower polymorphism of mtDNA in populations of the Baltic and Azov Sea mussels in comparison with other European populations was observed and can be explained by the recent history of both seas after the Pleistocene glaciation. In the M. galloprovincialis population from the Azov Sea, the presence of the male-inherited (M) genome was demonstrated for the first time by sequencing the control region and was observed at high frequency. Possible influence of mussel culture on geographic distribution of the Mytilus taxa in Europe is discussed.     

A molecular approach to the ecology of a mussel (Mytilus edulis – M. trossulus) hybrid zone on the east coast of Newfoundland, Canada

Mytilus trossulus Gould and M. edulis L. coexist in mixed populations in Atlantic Canadian waters. Although there is evidence that the two species hybridize in natural populations and that hybrids produce progeny through backcrosses, no study of the microgeographic distribution of the two forms and their hybrids has been made. Here we examine subtidal samples of mussels taken in July 1997 from two locations in eastern Newfoundland (Canada) and from wave-exposed and protected environments within each location. Shell lengths ranged from 15 to about 90 mm. Mussels were classified as pure forms or hybrids (F₁, F₂ and from backcrosses) based on four diagnostic markers, two allozyme loci (Mpi and Est-D) and two nuclear PCR-based DNA markers (ITS and Glu-5). In addition, a PCR-based mtDNA marker (COIII) was used to characterize the distribution of mtDNA mitotypes among pure and hybrid individuals. There were differences in the proportions of pure M. edulis and M. trossulus between sites and between environments at one location. M. trossulus was the predominant species at one of the two exposed sites. In all four samples, M. trossulus was also the predominant form among small individuals. The frequency of hybrids was 26% overall and did not differ among samples. Hybrids consisted mostly of backcrosses that were M. trossulus-biased among small mussels and M. edulis-biased among large ones. We conclude that both intrinsic genetic factors and extrinsic environmental factors influence the relative frequency of M. edulis, M. trossulus and their hybrids. Received: 29 June 1998 / Accepted: 4 November 1998     

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.     

Mitochondrial DNA heteroplasmy in European populations of the mussel Mytilus trossulus

Mitochondrial DNA (mtDNA) of Mytilus trossulus from the Gulf of Gdansk (southern Baltic) and M. edulis from Swansea Bay, South Wales, UK, collected in 1991, was studied by restriction-enzyme analysis. These two species were more similar to each other in haplotype frequencies than either was to M. galloprovincialis from Britain. M. trossulus resembles M. edulis in having a high frequency of heteroplasmy restricted to males. However, in contrast to M. edulis where restriction-site heteroplasmy predominates, in M. trossulus heteroplasmic individuals possess two genomes which differ in length by up to 3 kilobases.     

Hybridization between the blue mussels Mytilus galloprovincialis and M. trossulus along the Pacific coast of North America: evidence for limited introgression

Two species of blue mussel, Mytilus galloprovincialis and M. trossulus, co-occur and hybridize along the Pacific coast of North America. Using a set of polymerase chain-reaction (PCR)-based genetic markers which diagnostically identify these species, we show that they are sympatric from the Cape Mendocino region to the Monterey Peninsula in northern and central California, USA. Mussels with hybrid genotypes were detected in all populations sampled in the region of sympatry, and the frequency of hybrid genotypes in individual hybrid populations ranged from 13 to 44%. Significant frequencies of first-generation backcross genotypes were detected in two individual hybrid zone populations (Berkeley and Monterey Marina) and in the hybrid zone as a whole, indicating that the potential exists for introgression between M. galloprovincialis and M. trossulus. Despite this potential, we found no evidence of advanced introgression beyond first-generation backcrosses, suggesting that gene flow between M. galloprovincialis and M. trossulus has been quite limited. The frequency of mussels with M. trossulus and hybrid genotypes declined abruptly south of Monterey Peninsula, while the frequency of mussels with M. galloprovincialis and hybrid genotypes declined precipitously north of Cape Mendocino. These abrupt genetic discontinuities indicate that this blue mussel hybrid zone is presently positioned between two prominent coastal features and there is little, if any, export of alleles from the hybrid zone into bordering parental populations. Received: 20 August 1997 / Accepted: 26 October 1998     

Introgression and mitochondrial DNA heteroplasmy in the Baltic populations of mussels Mytilus trossulus and M. edulis

A strong clinal change in salinity occurs between the Baltic Sea and the North Sea, Atlantic Ocean, in the Danish Straits, where hybridization zone between mussels Mytilus edulis and M. trossulus has been reported. Eleven samples of mussels were studied from the Danish Straits and the inner Baltic Sea. Extensive introgression of M. edulis alleles from the North Sea into populations throughout the Baltic was ascertained for mitochondrial DNA (mtDNA) and two nuclear markers (ME15–16 and ITS). In the opposite direction, introgression of M. trossulus alleles into the M. edulis background was observed at the EFbis nuclear marker in populations from Kattegat (Danish Straits). While only M. edulis F (female) mtDNA was present in the Baltic, there were still strong differences in frequencies in the control region length variants between the Danish Straits and the inner Baltic samples, and weaker variation in coding region ND2–COIII haplotype frequencies. In the assays of the two mtDNA regions, various patterns of heteroplasmy were detected in 32% of all the studied individual mussels; this includes the presence of distinct, independently inherited M and F mitochondria in males, as well as the presence of two different distinguishable F genomes. The male-inherited M mtDNA genomes are quite common in the mussels from the Danish Straits, but very rare in males from the inner Baltic. Instead, a recombined control region variant (1r), which seems to have taken over the role of the M genome, was present in a number of specimens in the Baltic. Observations of heteroplasmy for two F genomes in some females and males confirm disruptions of the doubly uniparental inheritance mechanism in the hybrid Baltic Mytilus.     

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.     

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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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.     

Genetic population structure of a species' complex of blue mussels (Mytilus spp.)

Blue mussels representing two nominal species (Mytilus trossulus Gould, 1850 and Mytilus galloprovincialis Lamarck, 1819 were collected from 28 intertidal locations along the Pacific coast of the USA in 1990–1991 (total N=1255) and examined for variation at 15 allozyme loci. Twelve samples, mostly from a region of suspected hybridization, were analyzed for variation in seven shell characters. Principal-components analysis of allozyme data revealed three groups based on first principal-component scores, which were identified as M. trossulus, M. galloprovincialis, and hybrids. Canonical discriminant analysis of shell characters was less successful in separating mussels into discrete groups. Each location was characterized for four environmental variables: (1) temperature, (2) salinity, (3) tidal height and (4) degree of exposure to wave action, which were then used as independent variables in a series of multiple-regression analyses, with the proportions of the two species as dependent variables. Temperature and salinity had significant (P<0.05) effects on the macrogeographic distribution of the two species, whereas the effects of height in the tidal zone and degree of wave exposure were not statistically significant. Salinity was found to have a greater influence than temperature on the microgeographic distribution of the two species. M. trossulus was more abundant at locations with lower temperatures and greater salinity variation than M. galloprovincialis. The two species appear to be ecologically distinct, and their genetic integrity is at least partly the result of environmental heterogeneity.     

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.     

Calanoid copepod,cladoceran, and rotifer eggs in sea-bottom sediments of northern Californian coastal waters: Identification,occurrence and hatching

In Ireland, mussels on exposed rocky shores constitute an interbreeding mixture of two forms of mussels,Mytilus edulis L. and the Mediterranean musselM. galloprovincialis Lmk. This paper presents an in-depth analysis, carried out between October 1984 and December 1986, of genetic variability at two partially diagnostic loci,Odh andEst-D, in two exposed-shore populations ofMytilus spp. in the west of Ireland. Significant differences at theOdh locus were observed in the genetic composition of adult mussels from different tidal levels. These differences were repeatable whether one was analysing replicate samples at a single point in time, samples collected at different points in time, i.e., in different years, or samples collected from different shores. Mussels recruiting to artificial substrates set out for a period of one month at different tidal levels at one of these sites were also observed to be genetically different; mussels higher up the shore exhibited higher frequencies of those alleles characteristically at high frequency inM. galloprovincialis for both theOdh andEst-D loci. Hence, the genetic differences observed in adult mussels are much more exaggerated in juveniles and are already apparent within the first month of benthic life. Possible reasons for the observed microgeographic differentiation are discussed. It is concluded that the observed genetic differences between mussels at different tidal levels arise either in the pelagic/attachment stage or very shortly after settlement.     

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.