Bioinvasion threatens the genetic integrity of native diversity and a natural hybrid zone: smooth‐shelled blue mussels (Mytilus spp.) in the Strait of Magellan

Smooth‐shelled blue mussels of the Mytilus edulis species complex are widely distributed bivalve molluscs whose introductions threaten native marine biodiversity (non‐indigenous species – NIS). The aim of the present study was to identify the species and hybrids of Mytilus present in the Magellan Region (southern Chile). Results indicate that three mussel species of the Mytilus edulis complex are found in the region – M. edulis, M. chilensis (or the Southern Hemisphere lineage of Mytilus galloprovincialis), and M. galloprovincialis of Northern Hemisphere origin. For the first time, alleles of the introduced M. trossulus are reported from the Southern Hemisphere. In the Strait of Magellan the native Pacific blue mussel, Mytilus chilensis and the native Atlantic blue mussel, Mytilus edulis, meet and mix at a natural hybrid zone (about 125 km in length). This is the first record of a natural Mytilus hybrid zone in the Southern Hemisphere and is also the first record of the co‐occurrence of genes from all four Mytilus species in any one region. These results contribute to the knowledge of the biodiversity and delimitation of mussel species in southern South America, and highlight how introduced species may threaten the genetic integrity of native species through hybridization and introgression.     

Three species of Mytilus and their hybrids identified in a Scottish Loch: natives, relicts and invaders?

Three species of the mussel, Mytilus, occur in the North Atlantic region, M. edulis, M. galloprovincialis and M. trossulus, and hybrid zones are present where their distributions overlap. M. edulis is a native species in the UK. M. galloprovincialis originated in the Mediterranean and its distribution extends northwards along the Atlantic seaboard to Scotland. Baltic Sea mussels have a M. trossulus ancestry but are highly introgressed by M. edulis. In recent decades, farming of mussels on long-line rope culture systems has been introduced into Scotland. On farms in Loch Etive, a form of mussel with a fragile shell and a different shape to either M. edulis or M. galloprovincialis has been increasing in frequency over recent years. Samples of fragile shelled, normal strong shelled and intermediate mussel types were sampled from two farms in 2006 and compared with samples of M. edulis, M. galloprovincialis and M. trossulus from other sources where their species identity is well established. Abundance relative to depth, shell strength, condition index and shell morphology were analysed together with 5 allozyme loci and one nuclear DNA genetic marker (Me 15/16). The fragile shelled mussels, and many of those classed as intermediate, were identified as a mixture of M. trossulus and M. trossulus x M. edulis hybrids. This identification was strongly supported by both morphological and genetic data and is the first record of the presence of M. trossulus in UK waters. M. trossulus in Loch Etive are most likely to be a post-glacial relict population restricted to the low salinity area of the Loch that has recently increased in abundance due to commercial mussel growing activity. In addition, individual mussels of all three species and their hybrids were detected amongst Loch Etive mussels. This is the first genetic demonstration of all three species and their hybrids occurring together in one location in the Atlantic region and provides a unique opportunity to study the processes of speciation, divergence, and introgression in the genus Mytilus.     

Mytilus galloprovincialis Lmk. in Southern Africa

The Mediterranean mussel, Mytilus galloprovincialis Lmk. has been identified on the west coast of southern Africa using morphological and biochemical-genetic comparisons with samples of “pure” M. edulis L. from Denmark and M. galloprovincialis from the Mediterranean coast of Spain. Our results show that the southern African Mytilus more closely resembles M. galloprovincialis in anterior adductor, posterior adductor and hinge size. It also has a more ventrally located maximal shell width like the Mediterranean mussel. The electrophoretic examination of 23 proteins showed that many alleles diagnostic for M. galloprovincialis appeared in the southern African Mytilus. Nei's genetic distance between the southern African Mytilus and M. galloprovincialis was 0.010 ± 0.004 and the average genetic distance between these samples and M. edulis was 0.162 ± 0.077.The presence of M. galloprovincialis in southern Africa could be explained by biogeographic dispersal along the west coast of Africa during Pleistocene cooling, or by a recent inadvertant or intentional introduction by man. To test these two hypotheses, we examined ≈750 mussel shells collected from Koi-san middens and ≈350 shells from a raised-beach deposit all located on the west coast of Cape Province, South Africa. The ages of these deposits range from 1500 yr to 120000 yr and pre-date the arrival of Europeans to the Cape. This examination did not reveal any shells of Mytilus. We, therefore, conclude that the introduction of M. galloprovincialis to southern Africa was a recent event. We suggest that, since the first confirmed reference to Mytilus in southern Africa was made from a 1972 collection, the introduction occurred within the last two decades.     

Development of a real-time PCR assay for detection of Mytilus species specific alleles: Application to a sampling survey in Scotland

Shellfish aquaculture is a growing industry in Scotland, dominated by the production of the mussel Mytilus edulis, the native species. Recently the discovery of Mytilus galloprovincialis and Mytilus trossulus together with M. edulis and all 3 hybrids in cultivation in some Scottish sea lochs led to questions regarding the distribution of mussel species in Scotland. The establishment of an extensive sampling survey, involving the collection of mussels at 34 intertidal sites and 10 marinas around Scotland, motivated the development of a high-throughput method for identification of Mytilus alleles from samples. Three Taqman®-MGB probes and one set of primers were designed, based on the previously described Me 15/16 primers targeting the adhesive protein gene sequence, and samples were screened for the presence of M. edulis, M. galloprovincialis and M. trossulus alleles using real-time PCR. Mytilus edulis alleles were identified in samples from all 44 sites. Mytilus galloprovincialis alleles were found together with M. edulis alleles extensively in northern parts of the west and east coasts. Mytilus trossulus alleles were identified in samples from 6 sites in the west and south-west of Scotland. Because M. trossulus is generally undesirable in cultivation and therefore preventing the geographical spread of this species across Scotland is considered beneficial by the shellfish aquaculture industry, these 6 samples were further analysed for genotype frequencies using conventional PCR. Although distribution of the non-native species M. galloprovincialis and M. trossulus have proven to be more widespread than previously thought, there is no evidence from our study of either M. trossulus or M. galloprovincialis acting as an invasive species in Scotland. The real-time PCR method developed in this study has proven to be a rapid and effective tool for the identification of M. edulis, M. galloprovincialis and M. trossulus alleles from samples and should prove useful in future surveys, ecological or aquaculture management related studies in both unispecific and mixed species areas of these species.     

Species status and population structure of mussels (Mollusca: Bivalvia: Mytilus spp.) in the Wadden Sea of Lower Saxony (Germany)

Three species of mussel (genus Mytilus) occur in Europe: M. edulis (Linnaeus 1758), M. galloprovincialis (Lamarck 1819) and M. trossulus (Gould, Boston Society of Natural History 3: 343?C348, 1850). Although these species are indigenous to the North Sea, the Mediterranean and the Baltic Sea, respectively, they form an extended patchy species complex along the coasts of Europe (??the Mytilus edulis complex??) and are able to hybridize where their distributions overlap. Recent studies examining the taxonomic status and genetic composition of Mytilus populations in the Netherlands and the British Isles have revealed introgressive hybridization processes within this species complex, with hints of an invasion of nonindigenous M. galloprovincialis into the North Sea. Furthermore, an extensive international mussel fishery industry in Europe (i.e., Great Britain, the Netherlands, Denmark, and Germany) is also in discussion for a possibly anthropogenically induced bioinvasion of nonindigenous Mytilus traits into the Wadden Sea area. Although it is assumed that the Wadden Sea of Germany comprises M. edulis only, this has never been confirmed in a molecular genetic study. To assess the situation for the Wadden Sea of Lower Saxony, we conducted the first molecular study of the Mytilus genus in the region. Taxonomic identification of 504 mussels from 13 intertidal mussel banks using the nDNA marker Me15/16 revealed a population composition of 99% M. edulis and 1% M. edulis X M. galloprovincialis hybrids. Hence, the Wadden Sea population is unaffected by range expansion of nonindigenous Mytilus traits. The genetic structure of the M. edulis populations was investigated using the phylogenetic and population genetics analyses of the mitochondrial DNA cytochrome-c-oxidase subunit I (COI) and the first variable domain of the control region (VD1), which were sequenced for >120 female individuals. These results showed a heterogeneous, panmictic population due to unrestricted gene flow. This can be attributed to extensive larval dispersal linked to the tidal circulation system in the back barrier basins of the Wadden Sea.     

Genetic traits in the bivalve Mytilus from Europe, with an emphasis on Arctic populations

Genetic and some ecophysiological traits of mussels collected in the European Arctic, up to their northeastern distribution limit in the Barents Sea, were studied and compared with traits of mussels from the Mediterranean, Atlantic and Baltic. The genetic traits of these populations were analysed by isoenzyme electrophoresis on seven loci in order to assess the Mytilus complex to which populations in the Arctic region belong. Ecophysiological variables, the weight-index and glycogen were analysed to assess the physiological fitness of the populations. Three distinct groups were recognised: (1) Mytilus (edulis) galloprovincialis in the Mediterranean and Spain, (2) M. (edulis) edulis along the Atlantic coast from the Netherlands northwards into Russia, and (3) the Baltic Mytilus (edulis) trossulus. The mussels from populations in the Russian Arctic all belong to the Atlantic Mytilus (edulis) edulis group. The genetic variability and ecophysiological measures indicated that the sub-Arctic White Sea mussel populations have a relatively lower performance capacity, whereas those in the Arctic at the edge of their northern distribution showed a surprisingly strong performance. Accepted: 14 June 2000     

A historical perspective of the genus Mytilus (Bivalvia: Mollusca) in New Zealand: multivariate morphometric analyses of fossil, midden and contemporary blue mussels

The taxonomic status of smooth shelled blue mussels of the genus Mytilus has received considerable attention in the last 25 years. Despite this, the situation in the southern hemisphere remains uncertain and is in need of clarification. Recent work suggests that contemporary New Zealand mussels from two cool/cold temperate locations are M. galloprovincialis. However, the distribution of Mytilus in New Zealand ranges from 35 ° to 52 ° south (～ 1800 km), meaning that large areas of the subtropical/warm temperate north and the subantarctic south remain unsampled, an important consideration when species of this genus exhibit pronounced macrogeographical differences in their distributions which are associated with environmental variables such as water temperature, salinity, wave action and ice cover. This study employed multivariate morphometric analyses of one fossil, 83 valves from middens, and 92 contemporary valves from sites spanning the distributional range of blue mussels to determine a historical and contemporary perspective of the taxonomic status of Mytilus in New Zealand. The findings indicated that all fossil and midden mussels are best regarded as M. galloprovincialis and confirmed that contemporary mussels, with one possible regional exception, are also best regarded as M. galloprovincialis. Contemporary mussels from the Bay of Islands (warm temperate/subtropical) exhibited much greater affinity to M. edulis than they did to M. galloprovincialis, indicating that mussels from this area require detailed genetic examination to determine their taxonomic status. The analyses revealed a significant difference between the fossil/midden mussels and the contemporary mussels, consistent with levels of present day differentiation among intraspecific populations and not thought to reflect any substantive temporal change between mussels of the two groups. The continuous distribution of M. galloprovincialis in New Zealand from the warm north to the subantarctic south indicates that the physiology of this species is adapted to a wide range of water temperature conditions. Therefore, the distribution of this species on a worldwide scale is unlikely to be restricted by its adaptation to warm water alone, as has previously been widely assumed. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 82 , 329–344.     

Genetic diversity of Southern hemisphere blue mussels (Bivalvia: Mytilidae) and the identification of non‐indigenous taxa

The taxonomic and evolutionary affinities of Southern hemisphere smooth‐shelled blue mussels are unclear, with studies using different marker types having identified different relationships among various geographic regions. Using an existing and a new molecular assay, the present study builds on previous work to test the distribution of blue mussels native to and introduced to the Southern hemisphere. Populations of Mytilus were sampled from New Zealand, Australia, and Chile. The nuclear‐DNA marker Me 15/16 was used to identify the taxonomic status of 484 individuals. A new restriction fragment length polymorphism (RFLP) assay was used to identify the hemisphere of origin for a subset of Mytilus galloprovincialis. The Me15/16 marker identified 478 pure M. galloprovincialis from Southern hemisphere sites and six Mytilus edulis/M. galloprovincialis hybrids from the Auckland Islands (New Zealand) and Chile. A cytoplasmic RFLP identified Northern hemisphere M. galloprovincialis in almost every Southern hemisphere region. The presence of native M. galloprovincialis at high latitudes (up to 52°S) has implications for our understanding of environmentally induced selective constraints considered to determine species distributions. Widespread occurrence of invasive Northern hemisphere blue mussels in the Southern hemisphere is documented for the first time. Identification of inter‐specific hybrids (M. edulis × M. galloprovincialis) in Chile and in the Auckland Islands (subantarctic New Zealand) illustrates that environments ranging from international ports to remote protected locations are vulnerable to bioinvasion. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 898–909.     

Nocardiosis in Mediterranean bivalves: First detection of Nocardia crassostreae in a new host Mytilus galloprovincialis and in Ostrea edulis from the Gulf of Naples (Italy)

In this work M. galloprovincialis and O. edulis specimens were surveyed for a pathological study in the Gulf of Naples (Mediterranean sea, Campania Region, southern Italy). Clusters of Nocardia sp.-like cells were observed in histological slides. PCR amplification, sequencing and in situ hybridization were carried out in order to corroborate Nocardia species identification for both hosts. Blast results showed a 99% of maximum identity with Nocardia crassostreae sequences in Genbank. This is the first report of N. crassostreae in the new host M. galloprovincialis and, in a new area, the Mediterranean Sea.     

Competition and facilitation between the alien mussel Mytilus galloprovincialis and indigenous species: Moderation by wave action

As the alien species that most dominates space along the South African coast, the Mediterranean mussel Mytilus galloprovincialis has radically altered community composition on invaded shores. We experimentally assessed interspecific interactions between this invasive species and dominant indigenous species in conjunction with considering how wave action moderates such interactions. The density of both M. galloprovincialis and the limpet Scutellastra granularis increased with wave action. Conversely, the tube-building polycheate Gunnarea capensis was negatively affected by wave exposure, being most abundant on sheltered shores. The influence of wave action on the indigenous mussel Aulacomya ater, however, remains unclear. M. galloprovincialis outcompeted both G. capensis and A. ater at moderate to high exposure levels, whereas it had both positive and negative effects on S. granularis. It outcompeted adult limpets on primary rock space on semi-exposed and exposed shores, reducing densities of this portion of the population. However, recruitment of S. granularis was facilitated by M. galloprovincialis, as greater numbers recruited to the secondary substratum offered by mussel shells. Again this interaction intensified with wave action. Due to the extremely high density of recruits on secondary space, the net effect of M. galloprovincialis on S. granularis was positive. Thus, wave action not only influences the abundance of individual species, but also mediates both positive and negative interspecific interactions in rocky shore communities, including the impact of alien species such as M. galloprovincialis.     

Will the invasive mussel Mytilus galloprovincialis Lamarck replace the indigenous Perna perna L. on the south coast of South Africa?

The mussel Mytilus galloprovincialis is invasive worldwide, has displaced indigenous species on the west coast of South Africa and now threatens Perna perna on the south coast. We tested the hypothesis that Mytilus will replace Perna by examining changes in their distribution on shores where they co-exist. Total cover, adult density, recruit density, recruit/adult correlations and mean maximum lengths of both species were measured in 2001 at two contrasting sites (Plettenberg Bay and Tsitsikamma) 70 km apart, each including two locations 100 m apart. Cover and density were measured again in 2004. Total mussel abundance was significantly lower in Tsitsikamma, and recruit density was only 17% that of Plettenberg Bay. Abundance and cover increased upshore for Mytilus, but decreased for Perna, giving Mytilus higher adult and recruit density and total cover than Perna in the upper zones. Low shore densities of recruits and adults were similar between species but cover was lower for Mytilus, reflecting its smaller size, and presumably slower growth or higher mortality there. Thus, mechanisms excluding species differed among zones. Recruitment limitation delays invasion at Tsitsikamma and excludes Perna from the high shore, while Mytilus is excluded from the low shore by post-recruitment effects. Recruitment limitation also shapes population structure. Recruit/adult correlations were significant only where adult densities were low, and this effect was species-specific. Thus, at low densities, larvae settle or survive better near adult conspecifics. After 3 years, these patterns remained strongly evident, suggesting Mytilus will not eliminate Perna and that co-existence is possible through partial habitat segregation driven by recruitment limitation of Perna on the high shore and post-settlement effects on Mytilus on the low shore.     

Interlineage Mytilus galloprovincialis Lmk. 1819 hybridization yields inconsistent genetic outcomes in the Southern hemisphere

A Southern hemisphere lineage of the blue mussel Mytilus galloprovincialis has been diverging in allopatry from Northern hemisphere conspecifics for 0.84–1.2 million years. Secondary contact between Southern and Northern hemisphere mussels in Chile, New Zealand and Australia provides an opportunity to better understand the extent and consequences of extensive range expansion. Non-native M. galloprovincialis and hybrids, as detected from RFLP assays of nuclear and mitochondrial DNA, are present in all three countries and significant cytonuclear disequilibria exist for native homozygotes in Chile and New Zealand, non-native homozygotes in Chile and non-native heterozygotes in New Zealand. Introductions into Australia are rare events given that no pure non-native mussels were detected. Immigration from one or both taxa into the hybrid zone may underlie disequilibria in New Zealand, whilst gender-directional crossing with limited ongoing hybridization contributes to disequilibria in Chile. Hybridization dynamics do not pose a threat to the Southern lineage in Chile and Australia, but in New Zealand, introgression, continued immigration and slight hybridization gender bias towards non-native maternal parents could lead to the regional extirpation of the native lineage.     

Mytilid mussels: global habitat engineers in coastal sediments

Dense beds of mussels of the family Mytilidae occur worldwide on soft-bottoms in cold and warm temperate coastal waters and have usually been considered hot spots of biodiversity. We examined intertidal mussel beds at four distant locations around the globe with the same sampling method, to find out whether this “hot spot” designation holds universally. We studied species assemblages within the matrices of byssally interconnected mussels engineered by Mytilus edulis in the North Sea, by mixed Perumytilus purpuratus and Mytilus chilensis at the southern Chilean coast, by Musculista senhousia in the Yellow Sea and by Xenostrobus inconstans at the coast of southern Australia. In all cases, species assemblages inside mussel beds were significantly different from those outside with many species being restricted to one habitat type. However, species richness and diversity were not generally higher in mussel beds than in ambient sediments without mussels. In the North Sea (M. edulis) and at the Chilean coast (P. purpuratus, M. chilensis), mussel beds have markedly higher species numbers and diversities than surrounding sediments, but this was not the case for mussel beds in Australia (X. inconstans) and the Yellow Sea (M. senhousia) where numbers of associated species were only slightly higher and somewhat lower than in adjacent sediments, respectively. In conclusion, although soft bottom mytilid mussels generally enhance habitat heterogeneity and species diversity at the ecosystem level, mussel beds themselves are not universal centres of biodiversity, but the effects on associated species are site specific.     

Glacial history of the European marine mussels Mytilus,inferred from distribution of mitochondrial DNA lineages

Mussels of the genus Mytilus have been used to assess the circumglacial phylogeography of the intertidal zone. These mussels are representative components of the intertidal zone and have rapidly evolving mitochondrial DNA, suitable for high resolution phylogeographic analyses. In Europe, the three Mytilus species currently share mitochondrial haplotypes, owing to the cases of extensive genetic introgression. Genetic diversity of Mytilus edulis, Mytilus trossulus and Mytilus galloprovincialis was studied using a 900-bp long part of the most variable fragment of the control region from one of their two mitochondrial genomes. To this end, 985 specimens were sampled along the European coasts, at sites ranging from the Black Sea to the White Sea. The relevant DNA fragments were amplified, sequenced and analyzed. Contrary to the earlier findings, our coalescence and nested cladistics results show that only a single M. edulis glacial refugium existed in the Atlantic. Despite that, the species survived the glaciation retaining much of its diversity. Unsurprisingly, M. galloprovincialis survived in the Mediterranean Sea. In a relatively short time period, around the climatic optimum at 10 ky ago, the species underwent rapid expansion coupled with population differentiation. Following the expansion, further contemporary gene flow between populations was limited.     

A first report on coexistence and hybridization of <Emphasis Type="Italic">Mytilus trossulus</Emphasis> and <Emphasis Type="Italic">M. edulis</Emphasis> mussels in Greenland

Semi-sessile Mytilus mussels are used as indicators of climate changes, but their geographic distribution is not sufficiently known in the Arctic. The aim of this study was to investigate the taxonomic status and genetic differentiation of Mytilus populations in a Northwest Greenlandic fjord at Maarmorilik, impacted by contaminations from a former mine. In this study, mussels were collected at three sites differing in exposure to environmental factors. A total of 54 polymorphic SNPs found in the Mytilus EST and DNA sequences analyzed were successfully applied to 256 individuals. The results provided the first evidence for the existence of M. trossulus in Greenland. The mussel from M. trossulus and M. edulis taxa are shown to coexist and hybridize in the fjord. The three studied sites were found to differ significantly in the distribution of taxa with a higher prevalence of M. trossulus in the inner fjord. The identified M. edulis × M. trossulus hybrids mostly had a hybrid index score of about 0.5, indicating a similar number of alleles characteristic for M. trossulus and M. edulis. There was a low number of backcrosses between ‘pure’ taxa and hybrids. This newly discovered hybrid zone between the two taxa is unique in comparison with the Canadian populations. As Mytilus mussels in Greenland hitherto have been regarded as the one taxon M. edulis, the results have importance for biogeography and future monitoring and environmental studies.     

Microgeographic allozyme differentiation in the hybrid zone ofMytilus galloprovincialis Lmk. andM. edulis L. on the continental European coast

The EuropeanMytilus galloprovincialis Lmk. andM. edulis L. coexist and hybridize in different proportions in extended areas of the British and Atlantic French coasts.M. galloprovincialis typical allozymes seem to predominate in wave exposed areas, at high levels of attachment and in larger mussels in the British hybrid zone. Mussel samples from exposed and sheltered areas, 200 m apart, and from high and low levels of attachment were collected from a location of the French hybrid zone in 1988–92. PureM. galloprovincialis andM. edulis populations were also taken as controls. Diagnostic enzyme loci for bothMytilus (EST-D ^*, LAP-1^*, MPI^*, ODH^*) andAP-1 ^*, LAP-2^* andPGM ^* loci were studied. The frequencies of theM. galloprovincialis typical alleles were significantly greater in exposed populations than in sheltered samples (e.g. 0.729 to 0.803 vs 0.192 to 0.581 forEST-D ^*90), and at high level of attachment than at low level for the sheltered area (e.g. 0.581 vs 0.192 forEST-D ^*90). PutativeM. galloprovincialis was more abundant on the exposed coast (0.591 and 0.702) than on the sheltered shore, where it predominated at the high shore but not at the low shore location (0.371 vs 0.045). Significantly positive correlations between shell length and typicalM. galloprovincialis compound allele frequencies were found only for populations from exposed areas. Relationships between theMytilus genetic differentiation and ecological factors are discussed.     

Larval settlement can explain the adult distribution of Mytilus californianus Conrad but not of M. galloprovincialis Lamarck or M. trossulus Gould in Moss Landing, central California: Evidence from genetic identification of spat

We investigated the spatial distribution of adult and newly settled mussels (Mytilus galloprovincialis Lamarck, Mytilus trossulus Gould and Mytilus californianus Conrad) on the shore at Moss Landing, California to test the hypothesis that adult distributions are a result of settlement patterns. Adult M. californianus were most abundant on a wave-exposed rocky jetty and adults of Blue mussels (M. trossulus and M. galloprovincialis) were more abundant inside the protected Moss Landing harbor. Using taxon-specific polymerase chain reactions, we monitored recruitment during continuous 1-2 week intervals on fibrous scrubbing pads for 12 months in 2002-2003. All mussel species settled in greatest numbers on the exposed jetty, and Blue mussels settled in greater numbers there than did M. californianus. Because Blue mussels settled abundantly where their adults were rare, post-settlement mortality appeared to be the strongest influence on adult distribution. In contrast, M. californianus settled mostly in their adult habitat.     

Genetic Divergence Detected by ISSR Markers and Characterization of Microsatellite Regions in Mytilus Mussels

The wide distribution of microsatellites in mussels of the Mytilus edulis complex (Mytilidae) enables the analysis of inter-simple-sequence repeat (ISSR) markers. The aim of this investigation was to assess genetic differentiation in six sampling localities distributed along the European Atlantic coast to expose the potential of these markers in genetic studies requiring the detection of low polymorphism and as a source of sequences for developing microsatellite markers. We detected low genetic structuring within each member of the Mytilus edulis complex. Nei and Li distances and AMOVA clustered the individuals studied into two groups. On the basis of these results two sampling localities coming from the M. edulis × M. galloprovincialis mosaic hybrid zone in Western Europe were assigned to one species. On the other hand, mussels of a sampling locality in the Baltic Sea were not significantly different from a pure M. edulis locality supporting an extensive introgression of M. edulis in these individuals. Finally, 148 microsatellites were found in the sequences of 51 ISSR markers, and two polymorphic microsatellite markers were developed.     

Contrasting patterns of mussel abundance at neighbouring sites: does recruitment limitation explain the absence of mussels on Cook Strait (New Zealand) shores?

Wellington Harbour (New Zealand) supports large populations of mussels (Aulacomya maoriana, Mytilus galloprovincialis and Perna canaliculus), whereas these species are absent from Cook Strait shores only a few km away. The density of planktonic mussel larvae and their recruitment rates to artificial substrates were investigated at harbour (with mussels) and Cook Strait (no mussels) sites to determine if a diminished or a zero larval supply and/or settlement explains the absence of mussels from Cook Strait shores. At both locations, larvae were collected from the plankton approximately monthly between September 1998 and February 2000, and recruitment rates to artificial substrates were estimated between March 2000 and February 2001. Planktonic larval densities were almost an order of magnitude greater within the harbour than at coastal sites (mean (±S.D.) density was 982 m⁻³ (±1478) with a peak density in September 1998 of 4207 m⁻³, compared with 106 (±94) and 381 m⁻³, respectively, in March 1999). Larval recruitment at harbour sites was also significantly greater than at coastal sites (mean (±S.D.) recruitment density was 2169 m⁻² (±4207) with a peak of ca. 211,425 m⁻² in July 2000, compared with 88 m⁻² (±86) and ca. 3700 m⁻², respectively, in February 2001). It has been suggested that “bottom up” regulation of community structure, principally via a diet of particulates low in organic matter, is the explanation for the absence of suspension feeding mussels from Cook Strait sites [Helson, J. G., 2001. An investigation into the absence of mussels (Perna canalicus, Aulacomya maoriana and Mytilus galloprovincialis) from the South Coast of Wellington, New Zealand. Unpublished PhD thesis, Victoria University of Wellington, 183 pp.], but given that planktonic larval supply and recruitment rates are much reduced at coastal sites, these data may also be important in explaining the absence. Whether current levels of recruitment are sufficient to maintain an adult population is at present unknown and requires further examination.     

Gene-Flow in a Mosaic Hybrid Zone: Is Local Introgression Adaptive?

Genome-wide scans of genetic differentiation between hybridizing taxa can identify genome regions with unusual rates of introgression. Regions of high differentiation might represent barriers to gene flow, while regions of low differentiation might indicate adaptive introgression—the spread of selectively beneficial alleles between reproductively isolated genetic backgrounds. Here we conduct a scan for unusual patterns of differentiation in a mosaic hybrid zone between two mussel species, Mytilus edulis and M. galloprovincialis. One outlying locus, mac-1, showed a characteristic footprint of local introgression, with abnormally high frequency of edulis-derived alleles in a patch of M. galloprovincialis enclosed within the mosaic zone, but low frequencies outside of the zone. Further analysis of DNA sequences showed that almost all of the edulis allelic diversity had introgressed into the M. galloprovincialis background in this patch. We then used a variety of approaches to test the hypothesis that there had been adaptive introgression at mac-1. Simulations and model fitting with maximum-likelihood and approximate Bayesian computation approaches suggested that adaptive introgression could generate a “soft sweep,” which was qualitatively consistent with our data. Although the migration rate required was high, it was compatible with the functioning of an effective barrier to gene flow as revealed by demographic inferences. As such, adaptive introgression could explain both the reduced intraspecific differentiation around mac-1 and the high diversity of introgressed alleles, although a localized change in barrier strength may also be invoked. Together, our results emphasize the need to account for the complex history of secondary contacts in interpreting outlier loci.