Effects of Endolithic Parasitism on Invasive and Indigenous Mussels in a Variable Physical Environment

Biotic stress may operate in concert with physical environmental conditions to limit or facilitate invasion processes while altering competitive interactions between invaders and native species. Here, we examine how endolithic parasitism of an invasive and an indigenous mussel species acts in synergy with abiotic conditions of the habitat. Our results show that the invasive Mytilus galloprovincialis is more infested than the native Perna perna and this difference is probably due to the greater thickness of the protective outer-layer of the shell of the indigenous species. Higher abrasion due to waves on the open coast could account for dissimilarities in degree of infestation between bays and the more wave-exposed open coast. Also micro-scale variations of light affected the level of endolithic parasitism, which was more intense at non-shaded sites. The higher levels of endolithic parasitism in Mytilus mirrored greater mortality rates attributed to parasitism in this species. Condition index, attachment strength and shell strength of both species were negatively affected by the parasites suggesting an energy trade-off between the need to repair the damaged shell and the other physiological parameters. We suggest that, because it has a lower attachment strength and a thinner shell, the invasiveness of M. galloprovincialis will be limited at sun and wave exposed locations where endolithic activity, shell scouring and risk of dislodgement are high. These results underline the crucial role of physical environment in regulating biotic stress, and how these physical-biological interactions may explain site-to-site variability of competitive balances between invasive and indigenous species.     

Mechanisms and consequences of competition between an alien mussel, Mytilus galloprovincialis, and an indigenous limpet, Scutellastra argenvillei

The influence of an alien mussel Mytilus galloprovincialis (Lamarck) on an indigenous limpet Scutellastra argenvillei (Krauss) was investigated on the west coast of South Africa by comparing four situations in which limpets occupied rock patches surrounded by mussels: (1) ‘established patches’ of high-density adult limpets; (2) ‘experimentally cleared patches’ from which mussels were manually removed; (3) ‘naturally disturbed patches’ where mussels had been eliminated by wave action; and (4) ‘half-density’ patches formed by thinning established patches. Limpets in established patches were most effective in retarding settlement and lateral invasion by mussels, but all patches shrank due to mussel encroachment. As patches shrank, limpet density fell in established and naturally disturbed patches due to emigration. Limpets living on mussel beds were small, sparse and never achieved sexual maturity; 60% shifted to rock, whereas only 7% moved from rock to mussels. The limpets showed highest fidelity to established patches (79% after 12 months) and lowest fidelity in naturally disturbed patches (20%). Thinning of established patches reduced fidelity to 26%. Mussel beds did not provide a suitable alternative substratum for S. argenvillei after they displaced the limpets from rocks. Moreover, elimination of mussels by waves failed to allow S. argenvillei to re-establish dense, adult populations resembling those prior to arrival of the mussel. Adults of S. argenvillei feed collectively by trapping kelp blades. On the topographically complex mussel beds limpets cannot feed this way. In sum, on exposed shores where M. galloprovincialis achieves maximal recruitment and growth, S. argenvillei is incapable of preventing mussel encroachment and is likely to become completely displaced. Semi-exposed shores do, however, offer a refuge preventing global extinction of the limpet.     

Parasites of the shore crab Carcinus maenas (L.): implications for reproductive potential and invasion success

The European shore crab, Carcinus maenas, is one of the most successful marine invasive species. Its success has been in part attributed to the loss of parasites, rekindling an interest in host-parasite interactions and impacts on host fitness in this crab. In the present study, we investigated C. maenas populations from Europe, South Africa and Australia for parasites, and assessed their impact on the fitness of male crabs. For the shore crab, testes weight along with success in mating competition is traded off against other life-history traits. We therefore used this parameter as an indicator both for reproductive fitness and a possible resource trade-off in response to parasite infestation. In the native range, crabs infested with Sacculina carcini showed significantly lower testes weight than uninfected crabs. However, helminth parasites did not generally cause reduced testes weights. Crab populations from South Africa and Australia were either parasitized at very low prevalences, or were completely parasite free. However, no population level effect of this parasite release was reflected in testes weight. These findings do not support a severe fitness impact of helminth parasites on C. maenas, which questions the role of parasites on its population dynamics, both in the native area and for invasive success.     

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.     

Genetics and taxonomy of Chilean smooth-shelled mussels, Mytilus spp. (Bivalvia: Mytilidae)

It has been previously established that native smooth-shelled mussels in southern South America possess close evolutionary affinities with Northern-Hemisphere Mytilus edulis L. 1758 (McDonald et al. (1991) [5]). This result has since been challenged by authors claiming that Chilean mussels should be considered a local subspecies of M. galloprovincialis Lmk. 1819. Moreover, morphological, physiological, ecotoxicological and molecular genetic studies on Chilean smooth-shelled mussels still frequently refer to ‘M. chilensis’ Hupé 1854, even though the previous discovery of alien M. galloprovincialis and considerable heterogeneity in shell morphology among samples collected along the Chilean shores raise concerns that different Mytilus spp. species might have been included under ‘M. chilensis’. Here we reviewed the molecular and morphological data available on smooth-shelled mussels from Chile in an attempt to clarify both their genetic composition and their taxonomic status. Using multivariate analysis on sample × allozyme-frequency matrices, we confirmed the widespread occurrence of the Southern-Hemisphere form of M. edulis along the shores from the North Patagonia region of Chile to the southern tip of the South American continent. The populations sampled in southern central Chile showed some evidence of slight introgression from Southern-Hemisphere M. galloprovincialis. Morphological characterization of a sample from Dichato in southern central Chile was consistent with its previous genetic identification as Mediterranean M. galloprovincialis. The occurrence of Southern-Hemisphere M. galloprovincialis in Punta Arenas at the southern tip of the South American continent was also reported. Southern-Hemisphere M. edulis, including native Chilean smooth-shelled Mytilus, should be assigned subspecific rank and named M. edulis platensis d’Orbigny 1846.     

Oceanographic Conditions Limit the Spread of a Marine Invader along Southern African Shores

Invasive species can affect the function and structure of natural ecological communities, hence understanding and predicting their potential for spreading is a major ecological challenge. Once established in a new region, the spread of invasive species is largely controlled by their dispersal capacity, local environmental conditions and species interactions. The mussel Mytilus galloprovincialis is native to the Mediterranean and is the most successful marine invader in southern Africa. Its distribution there has expanded rapidly and extensively since the 1970s, however, over the last decade its spread has ceased. In this study, we coupled broad scale field surveys, Ecological Niche Modelling (ENM) and Lagrangian Particle Simulations (LPS) to assess the current invaded distribution of M. galloprovincialis in southern Africa and to evaluate what prevents further spread of this species. Results showed that all environmentally suitable habitats in southern Africa have been occupied by the species. This includes rocky shores between Rocky Point in Namibia and East London in South Africa (approx. 2800 km) and these limits coincide with the steep transitions between cool-temperate and subtropical-warmer climates, on both west and southeast African coasts. On the west coast, simulations of drifting larvae almost entirely followed the northward and offshore direction of the Benguela current, creating a clear dispersal barrier by advecting larvae away from the coast. On the southeast coast, nearshore currents give larvae the potential to move eastwards, against the prevalent Agulhas current and beyond the present distributional limit, however environmental conditions prevent the establishment of the species. The transition between the cooler and warmer water regimes is therefore the main factor limiting the northern spread on the southeast coast; however, biotic interactions with native fauna may also play an important role.     

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.     

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.     

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.     

Facilitation and competition between invasive and indigenous mussels over a gradient of physical stress

The interactions between invasive exotic and indigenous species can have profound harmful effects on the recipient community; however, not all such interactions are negative. Facilitation is increasingly recognised as important in shaping natural communities and is believed to vary under different conditions. Earlier studies have shown that the indigenous intertidal mussel Perna perna initially facilitates survival of the invasive Mytilus galloprovincialis in the low mussel zone by providing protection against waves, but later excludes M. galloprovincialis through interference competition for space. Here, we examined interactions between these species in the mid and upper mussel zones, moving mussels to experimental plots in different combinations of densities and species. Mussels were left on the shore for more than a year and treatment effects on mortality, shell length and condition were compared. In the high zone, treatment had no effects and P. perna showed greater mortality than M. galloprovincialis, indicating that its exclusion from the high shore is due to emersion stress. In the mid zone, treatment had no significant effects on M. galloprovincialis, but multiple comparisons among treatments involving P. perna showed that facilitation occurred. P. perna survived better at higher densities, but survived even better when mixed with the physiologically more tolerant M. galloprovincialis. Length data indicated both inter- and intraspecific competition for P. perna in the mid zone. Whereas facilitation occurs strongly in the low zone (P. perna facilitates M. galloprovincialis) and weakly in the mid zone (M. galloprovincialis facilitates P. perna), the lack of facilitation in the high zone suggests that the probability of facilitation is not linearly linked to increasing physical stress. Instead it is likely to be hump shaped: relatively unimportant under conditions that are benign for a particular species, significant under more severe conditions, and overridden by physical stress under very harsh conditions.     

Mytilus galloprovincialis (Mollusca: Bivalvia) in a warm-temperate South African estuarine embayment

The distribution, population structure, condition and gonad indices, and the attachment strengths of the invasive Mediterranean mussel Mytilus galloprovincialis are described from the Knysna Estuary, South Africa. Mussels were found in the rocky intertidal at the mouth of the estuary and on all man-made hard substrata as far as 12?km from the mouth in 2013. However, mussel beds were found only in the lower reach of the estuary, where mussel densities were as high as 120 per 0.01?m². When compared to mussels from the embayment regions of the estuary, those at the estuary mouth, where wave action was high, were generally smaller, had significantly lower condition and gonad indices, and greater attachment strength. This suggests that mussels within the sheltered regions of the estuary invest more energy in growth and reproduction, and that this is one reason for their success as an invasive species in the Knysna estuarine embayment.     

Zebra Mussel Infestation of Unionid Bivalves (Unionidae) in North America

SYNOPSIS. In 1989, zebra mussels received national attentionin North America when they reached densities exceeding 750,000/m2in a water withdrawal facility along the shore of western LakeErie of the Laurentian Great Lakes. Although water withdrawalproblems caused by zebra mussels have been of immediate concern,ecological impacts attributed to mussels are likely to be themore important long-term issue for surface waters in North America.To date, the epizoic colonization (i.e., infestation) of unionidbivalve mollusks by zebra mussels has caused the most directand severe ecological impact. Infestation of and resulting impactscaused by zebra mussels on unionids in the Great Lakes beganin 1988. By 1990, mortality of unionids was occurring at somelocations; by 1991, extant populations of unionids in westernLake Erie were nearly extirpated; by 1992, unionid populationsin the southern half of Lake St. Clair were extirpated; by 1993,unionids in widely separated geographic areas of the Great Lakesand the Mississippi River showed high mortality due to musselinfestation. All infested unionid species in the Great Lakes(23) have become infested and exhibited mortality within twoto four years after heavy infestation began. Data indicate thatmean zebra mussel densities >5,000–6,000/m² and infestationintensities >100-200/unionid in the presence of heavy zebramussel recruitment results in near total mortality of unionids.At present, all unionid species in rivers, streams, and akesthat sympatrically occur with zebra mussels have been infestedand, in many locations, negatively impacted by zebra mussels.We do not know the potential consequences of infestation onthe 297 unionid species found in North America, but believezebra mussels pose an immediate threat to the abundance anddiversity of unionids.     

Contrasting spatial heterogeneity of sessile organisms within mussel (Perna perna L.) beds in relation to topographic variability

We examined the spatial heterogeneity in three sessile rocky shore organisms, the mussel Perna perna, the barnacle Octomeris angulosa (Sowerby) and the red alga Gelidium pristoides (Turn.) at a range of continuous local scales along horizontal transects within mid- and upper mussel beds of South African shores. We also examined the relationships between variability of organisms and topographic features (rock depressions, slope, aspect), and between mussel, barnacle and algal variability over the same scales. To estimate spatial heterogeneity, we analyzed scaling properties of semivariograms using a fractal approach. Relationships between different variables at the different scales were examined by cross-semivariograms. Spatial dependence of P. perna variability increased with spatial dependence of topographic variability, so that scaling regions of mussel and topographic distributions corresponded well. This relationship often improved with larger local scales (mussel cover increased with depressions, steeper slope and aspect towards waves), while at smaller spatial scales, variability in mussel cover was less well explained by variability in topography. The variability of the barnacle O. angulosa exhibited spatial dependence, even on topographically unstructured shores. In contrast, the distribution of the alga G. pristoides revealed high fractal dimensions, showing spatial independence on topographically unstructured shores. Algae also showed a very strong negative relationship with mussels at most local scales, and a negative relationship with barnacles in upper zones, especially at larger local scales. Barnacles may show clear spatial dependence because of hydrodynamics (at larger local scales) and the need to find a future mate in close proximity (at smaller local scales), while algae may show a strong negative relationship with mussels because of competition for space.     

Cheating the Locals: Invasive Mussels Steal and Benefit from the Cooling Effect of Indigenous Mussels

The indigenous South African mussel Perna perna gapes during periods of aerial exposure to maintain aerobic respiration. This behaviour has no effect on the body temperatures of isolated individuals, but when surrounded by conspecifics, beneficial cooling effects of gaping emerge. It is uncertain, however, whether the presence of the invasive mussel Mytilus galloprovincialis limits the ability of P. perna for collective thermoregulation. We investigated whether varying densities of P. perna and M. galloprovincialis influences the thermal properties of both natural and artificial mussel beds during periods of emersion. Using infrared thermography, body temperatures of P. perna within mixed artificial beds were shown to increase faster and reach higher temperatures than individuals in conspecific beds, indicating that the presence of M. galloprovincialis limits the group cooling effects of gaping. In contrast, body temperatures of M. galloprovincialis within mixed artificial mussel beds increased slower and exhibited lower temperatures than for individuals in beds comprised entirely of M. galloprovincialis. Interestingly, differences in bed temperatures and heating rates were largely dependent on the size of mussels, with beds comprised of larger individuals experiencing less thermal stress irrespective of species composition. The small-scale patterns of thermal stress detected within manipulated beds were not observed within naturally occurring mixed mussel beds. We propose that small-scale differences in topography, size-structure, mussel bed size and the presence of organisms encrusting the mussel shells mask the effects of gaping behaviour within natural mussel beds. Nevertheless, the results from our manipulative experiment indicate that the invasive species M. galloprovincialis steals thermal properties as well as resources from the indigenous mussel P. perna. This may have significant implications for predicting how the co-existence of these two species may change as global temperatures continue to rise.     

Population-specific responses to an invasive species

Predicting the impacts of non-native species remains a challenge. As populations of a species are genetically and phenotypically variable, the impact of non-native species on local taxa could crucially depend on population-specific traits and adaptations of both native and non-native species. Bitterling fishes are brood parasites of unionid mussels and unionid mussels produce larvae that parasitize fishes. We used common garden experiments to measure three key elements in the bitterling–mussel association among two populations of an invasive mussel (Anodonta woodiana) and four populations of European bitterling (Rhodeus amarus). The impact of the invasive mussel varied between geographically distinct R. amarus lineages and between local populations within lineages. The capacity of parasitic larvae of the invasive mussel to exploit R. amarus was higher in a Danubian than in a Baltic R. amarus lineage and in allopatric than in sympatric R. amarus populations. Maladaptive oviposition by R. amarus into A. woodiana varied among populations, with significant population-specific consequences for R. amarus recruitment. We suggest that variation in coevolutionary states may predispose different populations to divergent responses. Given that coevolutionary relationships are ubiquitous, population-specific attributes of invasive and native populations may play a critical role in the outcome of invasion. We argue for a shift from a species-centred to population-centred perspective of the impacts of invasions.     

Invasion of an indigenous Perna perna mussel bed on the south coast of South Africa by an alien mussel Mytilus galloprovincialis and its effect on the associated fauna

The introduced mussel Mytilus galloprovincialis is progressively increasing in abundance along the south coast of South Africa. Quantitative 0.1 m² samples were collected in the mid-zone of an indigenous Perna perna mussel bed in the 1980s prior to the arrival M. galloprovincialis (12) and in the 2000s during the M. galloprovincialis invasion (16). In addition, in situ counts of M. galloprovincialis were done on eight occasions between 1993 and 2005, and in the low- and high-zones on four occasions. In the mid-zone M. galloprovincialis was absent until 1987, its mean densities were low (< 15 individuals/0.1 m²) between 1993 and 1996, but thereafter increased steadily, peaking in 2004 (at 721 individuals/0.1 m²), before declining in 2005 (331 individuals/0.1 m²). The greatest densities of M. galloprovincialis were recorded at the high-zone (1121 individuals/0.1 m²) and the smallest in the low-zone. As M. galloprovincialis numbers increased, there was an associated, but smaller decline in P. perna numbers and the overall density of mussels increased significantly (P < 0.05). No major change was recorded in the size composition of P. perna. The density of associated fauna differed significantly (P < 0.01) between sampling dates with the lowest and highest values being recorded near the ‘beginning’ (2001) and ‘end’ (2005) of the invasion period respectively. These differences were largely due to variations in the density of barnacles, and the toothed barnacle Chthamalus dentatus appeared to be the only associated faunal species that was directly affected by the M. galloprovincialis invasion, experiencing a significantly (P ≤ 0.05), but temporary decline in density and biomass values.     

Isolated and combined impacts of blue mussels (Mytilus edulis) and barnacles (Balanus improvisus) on structure and diversity of a fouling community

Effects of two presumably dominant competitors, the blue mussel Mytilus edulis and the barnacle Balanus improvisus on recruitment, population dynamics and community structure on hard substrata were experimentally investigated in the subtidal Kiel Fjord, Western Baltic. The hypothesis that blue mussels and/or barnacles are local dominants and strongly influence succession and community structure was tested by monitoring succession in the presence and absence of simulated predation on either or both species. Manipulations included blue mussel removal, barnacle removal, combined blue mussel and barnacle removal, as well as a control treatment for natural (non-manipulated) succession. In the second part of the experiment, recovery from the treatments was monitored over 1 year.During the manipulative phase of the experiment, blue mussels had a negative effect on recruitment of species, whereas barnacles had no significant effect. Even so, a negative synergistic effect of blue mussels and barnacles was detected. Calculation of species richness and diversity H′ (Shannon Index) showed a negative synergistic effect of blue mussels and barnacles on community structure. Additionally, diversity H′ was negatively affected by the dominant competitor M. edulis. These effects were also detectable in the ANOSIM-Analysis. The non-manipulative phase of the experiment brought about a drastic loss of diversity and species richness. Blue mussels dominated all four communities. Barnacles were the only other species still being able to coexist with mussels. Effects of simulated predation disappeared fast.Thus, in the absence of predation on blue mussels, M. edulis within a few months dominates available space, and diversity of the benthic community is low. In contrast, when mussel dominance is controlled by specific predators, more species may persist and diversity remains high.     

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.     

Impacts of invasive crayfish (Pacifastacus leniusculus) on endangered freshwater pearl mussels (Margaritifera laevis and M. togakushiensis) in Japan

The invasive alien crayfish Pacifastacus leniusculus is considered harmful to freshwater pearl mussels Margaritifera laevis and M. togakushiensis. It also often colonises mussel habitats in Japan. In order to test the negative effects of alien crayfish on mussels, we evaluated the predation impact of signal crayfish on freshwater pearl mussels in vitro. We tested the relationship between the survival/injury rates of mussels and crayfish predation with respect to different sizes of mussels (four classes based on shell length: 10, 30, 50 and 70 mm). Crayfish selectively fed on the flesh of the 10-mm size class mussels after breaking their shells. The shell margins of mussels in all size classes were injured by crayfish. Results also showed that crayfish particularly injured the 50-mm size class of mussels. This observation could be attributed to this mussel size being the most suitable shell size (29.56–37.73 mm in carapace length) that the crayfish can effectively hold. This study shows that the presence of invasive crayfish reduces freshwater pearl mussel populations by damaging the shell margins and/or killing the mussels. This negative impact of invasive crayfish not only decreases the mussel population but could also limit mussel recruitment, growth and reproduction.