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.     

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     

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.     

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.     

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.     

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     

Size-dependent, spatial and temporal genetic variation at a leucine aminopeptidase (LAP) locus among blue mussel (Mytilus galloprovincialis) populations along a salinity gradient

Biochemical genetic variation at a leucine aminopeptidase (LAP) locus is related to salinity variation in several marine bivalve molluscs. This paper details an investigation of the Long Island Sound model of LAP selection (LAP genotype-dependent mortality occurring among newly settled Mytilus edulis mussels) carried out in 1997 among three M. galloprovincialis mussel populations along the salinity gradient of Wellington Harbour, New Zealand. Significant LAP genotypic heterogeneity was observed at the LAP locus between small (<25 mm shell length) and large (>25 mm shell length) M. galloprovincialis from Petone and Eastbourne (the two sites experiencing the greatest salinity variation), whereas genotypic heterogeneity was not significantly different between small and large mussels from Seatoun (the site experiencing the least salinity variation). The Lap ³ allele decreased in frequency and the Lap ⁴ allele increased in frequency at Petone and Eastbourne, whereas the Lap ³ and Lap ⁴ allele frequencies remained effectively constant at Seatoun. Both these findings are consistent with the Long Island Sound model of selection. At all three locations, the Lap ^3,3 genotype decreased in frequency from small to large mussels, whereas the Lap ^3,4 genotype increased in frequency from small to large mussels. All other LAP genotypes occurred at low frequencies (<0.10) at all three locations and showed no evidence of frequency change from small to large-size mussels nor evidence of clinal change among the three locations. These genotype frequency data possibly indicate that the Lap ^3,3 genotype is at a selective disadvantage compared to the Lap ^3,4 genotype at all three locations, and that this selective disadvantage is related to the extent of salinity variation which exists at each location. Further investigation is required before it can be determined if the Long Island Sound model of selection best describes the size-dependent and location-specific changes in LAP allele and genotype frequencies along this salinity gradient. Comparison of the population genetic structure at the LAP locus in 1995 and in 1997 revealed a profound change from heterozygote excesses to heterozygote deficiencies for all three M. galloprovincialis populations. The reason for the change is unknown, but the change indicates that population genetic structure at the LAP locus is highly variable in time, but consistent in space, among these M.␣galloprovincialis populations. Received: 5 February 1998 / Accepted: 27 May 1998     

The musselsMytilus galloprovincialis andM. trossulus on the Pacific coast of North America

Most recent authors have called the bay mussels of the Pacific coast of North AmericaMytilus edulis Linnaeus, 1758. Thirteen samples ofedulis-like mussels were collected from California, Oregon, and Alaska, USA, in 1985, 1986 and 1987. Electrophoretic evidence from wight loci indicates that these samples consist of two genetically distinct groups, neither of which is similar toM. edulis from the Atlantic Ocean. Mussels in southern California are very similar toM. galloprovincialis Lamarck, 1819 from the Mediterranean Sea; it is probable thatM. galloprovincialis was introduced accidentally to southern California. Mussels in Oregon and Alaska are similar to those from the Baltic Sea and parts of eastern Canada; the nameM. trossulus Gould, 1850 has priority for this taxon. In central and nothern California,M. galloprovincialis, M. trossulus and their hybrids co-occur. Despite the presence of hybrids betweenM. galloprovincialis andM. trossulus, the genetic integrity which they maintain across large areas of the world warrants their recognition as two distinct species.     

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.     

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.     

The Mytilus edulis species complex in southwest England: effects of hybridization and introgression upon interlocus associations and morphometric variation

Loglinear analysis of electrophoretic data from two hybrid Mytilus edulis x galloprovincialis populations in southwest England revealed non-significant associations between genotypes at four allozyme loci, each of which is partially diagnostic for differences between the two taxa. Significant non-random genotypic associations within the context of the non-significant model involved all four assayed loci equally, consistent with their occurrence in a relatively tight linkage group. Multivariate analyses were used to examine electrophoretic variation from the two hybrid populations, and morphometric variation in the hybrid populations and in four allopatric (two M. edulis and two M. galloprovincialis) populations from western Europe While the number of hybrid mussels is high at both sites (22% at Croyde, 53% at Whitsand) the two taxa have largely maintained the genetic differences which exist between them in allopatry. However, morphological differences between the taxa have been eroded for mussels within the hybrid zone, whereas these differences are quite pronounced for mussels from allopatric populations. It is proposed that each taxon within the genus maintains its genetic identity, despite high dispersal potential, widespread hybridization, and high levels of introgression, as a result of adaptation to different environments. The worlwide occurrence of all four Mytilus hybrid zones at ecotones between recognized biogeographical provinces which are characterized by differences in temperature and salinity is consistent with such an interpretation.     

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.     

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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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     

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.     

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     

A possible hybrid zone in the Mytilus edulis complex in Japan revealed by PCR markers

The adhesive protein allele of mussels collected at 13 points in Japan from Hokkaido to Kyushu was analyzed by the polymerase chain reaction using a set of primers which amplifies a part of the nonrepetitive region of the adhesive protein gene. While most mussels exhibited a 126 bp fragment, characteristic of the pure Mytilus galloprovincialis, 55 of 64 mussels sampled at Hiura and 1 of 14 mussels at Hakodate Port exhibited 168 and 126 bp fragments. Sequence analysis of the two fragments indicated that the 168 and 126 bp fragments are almost identical to previously reported sequences in M. trossulus and M. galloprovincialis, respectively. Since the frequency of heterozygous individuals in Hiura is very high, it is unlikely that they are simple hybrids. However, it is evident that mixing of genes occurred between the two species off Hokkaido. Received: 6 September 1996 / Accepted: 9 October 1996     

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     

Heterozygosity and growth in transplanted mussels

Growth comparisons were made involving mussels (Mytilus spp.) collected from five different localities in Britain in 1980–1981. Two of the localities, Mumbles, South Wales, and Bude, Southwest England, have pure populations of M. edulis and M. galloprovincialis, respectively. The other three, Whitsand Bay, Southwest England, Croyde Bay, Southwest England and Robin Hood's Bay, Northeast England, have hybrid populations with both M. edulis and M. galloprovincialis ancestry. To make growth comparisons, mussels from different populations were mixed in oyster nets and transplanted to three different localities. After periods of transplantation varying between several months and one year, growth was assessed by measuring increase in shell length or dry body weight. Starch-gel electrophoresis was used to assay variation in the transplanted mussels at three allozyme loci partially diagnostic for M. edulis and M. galloprovincialis. The results provide evidence of small growth differences between populations and between allozyme genotypes within populations. These differences accounted for no more than a few percent of the total variation in growth between mussels. Statistically significant results were obtained, but were frequently found not to be reproducible. There is no clear evidence of a growth difference between M. edulis and M. galloprovincialis. Allozyme heterozygotes appear to have growth rates intermediate between allozyme homozygotes; this study thus fails to provide evidence for overdominance with respect to growth rate.     

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.