Mussel Mortality Research Articles

Growth and Mortality of Green Mussel Perna viridis Farmed at Ambong Bay and Marudu Bay, Sabah, Malaysia

Abstract Asian green mussel is commercially farmed in tidal waters in several enclosed bays in Sabah, Malaysia. In this study, two areas on the west coast of Sabah – Ambong Bay and Marudu Bay – were selected for the monitoring of the growth and mortality rates of green mussels farmed in suspension raft. Both growth and survival rates were then correlated with physicochemical parameters (dissolved oxygen, pH, salinity, temperature, water transparency, chlorophyll-a), nutrients (phosphate (PO4 3-), ammonia (NH3-N), nitrate (NO3-N) and nitrite (NO2-N) and condition index of mussel from each study site, as well as between the study sites. The twelve-month growth study (September 2017 to August 2018) was started with a total of 180 mussel specimens (90 at each site). The initial size (mean) of the mussel seed used was 47.7 ± 3.5 mm and 51.1 ± 3.9 mm for Marudu Bay and Ambong Bay, respectively. Mussels in Marudu Bay attained mean size of 73.47 ± 11.05 mm (SGR 0.17% ± 0.22) compared to 64.05 ± 7.44 mm (SGR 0.11% ± 0.22) for Ambong Bay at the end of the experiment. The cumulative mortality rates were 9.2% ± 4.9 and 55.5% ± 30.0 for Marudu Bay and Ambong Bay, respectively. The Pearson correlation indicated a significant positive relationship between mortality and water transparency (r = 0.684, p<0.01). There was a significant negative relationship between ammonia in seawater and mussel mortality (r = -0.561, p<0.01), as well as significant negative relationships between nitrate and growth (r = -0.480, p<0.05) and mortality (r = -0.460, p<0.05), as indicated by Spearman’s Rank-order Correlation analysis. Overall, the growth performance of green mussels farmed in Marudu Bay was better than in Ambong Bay, however, the mortality of mussels in Ambong Bay was higher.

Emerging pathogens caused disease and mortality in freshwater mussels,Hyriopsis cumingii,in China

In August 2019, massive mortalities of cultured Hyriopsis cumingii occurred in a local shellfish breeding experimental base in Chongqing, China. In this report, we identified 14 isolates from cultured mussels in mortality events and demonstrated that these strains caused mortality from 9% to 97%. Significant mussel mortality (p < .05) was observed in association with most isolates after bacterial challenge; however, considerable variation in the virulence of different isolates was noted, with survival ranging from 3% to 88% after 7 days. The most virulent strains (HOP3 and GL1) in haemolymph and tissues were identified as Stenotrophomonas maltophilia and Aeromonas veronii based on biochemical and 16S rDNA sequence analysis. The LD50 values of HOP3 and GL1 were found to be 2.45 × 106 CFU/g (body weight) and 2.24 × 105 CFU/g (body weight) respectively. Histopathology of H. cumingii infected with S. maltophilia and A. veronii displayed degeneration of muscle fibres and gill necrosis. We tested the antibiotic susceptibility of the two isolates and found that HOP3 was multidrug-resistant. Growth optimization study revealed that both isolates remained surviving across wide environmental ranges. Our data lay the foundation for the pathogenic mechanism of mussel bacterial diseases, and they also suggest that effective control measures are worth further investigation.

First detection of Francisella halioticida in mussels Mytilus spp. experiencing mortalities in France.

This note describes the first detection of the bacteria Francisella halioticida in mussels Mytilus spp. from locations in Normandy and northern Brittany (France) experiencing high mussel mortalities, while it was not detected in the Bay of St Brieuc (northern Brittany), an area which was not affected by abnormal mussel mortality. The distribution of the bacteria in mussels seems to be restricted to inflammatory granulomas as observed in Yesso scallops Mizuhopecten yessoensis from Canada and Japan. F. halioticida has been identified as being involved in mass (>80%) mortality of abalones Haliotis gigantea in Japan and high (up to 40%) mortality of Yesso scallops Mizuhopecten yessoensis in Canada as well as in lesions reducing marketability of Yesso scallops in Japan. The impact of this bacterium on the health of mussels needs to be investigated in future research, especially since the cause of high mussel mortalities that have been occurring in France for the past few years is still undetermined.

Mortality of marine mussels Mytilus edulis and M. galloprovincialis: systematic literature review of risk factors and recommendations for future research

AbstractThe aim of this study was to summarise the literature reporting the risk factors for mortality in the mussel species Mytilus edulis and Mytilus galloprovincialis in order to identify potential science‐based solutions to prevent or mitigate mussel mortality outbreaks. We followed the PRISMA methodology: Preferred Reporting Items for Systematic Reviews and Meta‐Analyses. The studied corpus of 91 publications (114 studies) was highly heterogeneous with respect to the methodological approaches used to define or estimate mussel mortality and the related putative risk factors. Results showed that the mortality risk of both mussel species M. edulis and M. galloprovincialis varied across the seasons, increased with an elevated seawater temperature above a thermal threshold of 20 and 24°C, respectively, decreased by protecting mussels from predation, and was associated with the presence of pathogens in M. edulis. For M. galloprovincialis, using mussel spat from the same area where the farming is carried out and farming them together with another mussel species appears to reduce the mortality risk. However, for M. edulis, this could be achieved by using pure crosses and in particular mussel spat having a selected genotype. For wild bed conservation, sand accumulation and anthropogenic sedimentation should be minimised. Our analysis showed that current approaches to this research topic are limited and are unlikely to yield actionable evidence to identify mussel mortality prevention or mitigation strategies. Therefore, recommendations are offered to increase the ability of future eco‐epidemiological research to identify multiple exposures associated with mussel mortality, underpinned by standardised efforts and cooperative initiatives.

Correlation between intermediary metabolism, Hsp gene expression, and oxidative stress-related proteins in long-term thermal-stressed Mytilus galloprovincialis.

Long-term exposure of Mytilus galloprovincialis to temperatures beyond 26°C triggers mussel mortality. The present study aimed to integratively illustrate the correlation between intermediary metabolism, hsp gene expression, and oxidative stress-related proteins in long-term thermally stressed Mytilus galloprovincialis and whether they are affected by thermal stress magnitude and duration. We accordingly evaluated the gene expression profiles, in the posterior adductor muscle (PAM) and the mantle, concerning heat shock protein 70 and 90 (hsp70 and hsp90), and the antioxidant defense indicators Mn-SOD, Cu/Zn-SOD, catalase, glutathione S-transferase, and the metallothioneins mt-10 and mt-20. Moreover, we determined antioxidant enzyme activities, oxidative stress through lipid peroxidation, and activities of intermediary metabolism enzymes. The pattern of changes in relative mRNA expression levels indicate that mussels are able to sense thermal stress even when exposed to 22°C and before mussel mortality is initiated. Data indicate a close correlation between the magnitude and duration of thermal stress with lipid peroxidation levels and changes in the activity of antioxidant enzymes and the enzymes of intermediary metabolism. The gene expression and increase in the activities of antioxidant enzymes support a scenario, according to which exposure to 24°C might trigger reactive oxygen species (ROS) production, which is closely correlated with anaerobic metabolism under hypometabolic conditions. Increase and maintenance of oxidative stress in conjunction with energy balance disturbance seem to trigger mussel mortality after long-term exposure at temperatures beyond 26°C. Eventually, in the context of preparation for oxidative stress, certain hypotheses and models are suggested, integrating the several steps of cellular stress response.

The effect of experimental drill fluid on the vital activities of the bivalve mollusk Mytilus edulis L.

The effect of two drill fluids on the survival rate and linear growth rate of bivalve mollusks Mytilus edulis has been studied. The standard water-based drill fluid includes barite and carboxymethyl cellulose, which are highly toxic substances. The main components of an experimental presumably biodegradable solution are the ground brown algae Saccharina latissima, inhabiting the coast of the Barents Sea, and ferrochrome lignosulfonate, which also applies to toxic components. The minimal concentration of a standard drill fluid led to the 100% death after 9 days of exposure; the experimental drill fluid did not affect the mortality of mussels at all. The mollusks exposed to the concentration of 5 and 10 g/L of experimental drill fluid have been characterized by a high relative linear growth. An experimental drill fluid that does not contain barite and carboxymethyl cellulose may be considered safe for marine bivalves; and its components seems to be easily degradable, since some of them may be used as a food source by some detritus-feeding invertebrates

Temperature-Related Responses of an Invasive Mussel and 2 Unionid Mussels to Elevated Carbon Dioxide.

Zebra mussels (Dreissena polymorpha) have exacerbated the decline of native freshwater mussels (order Unionida) in North America since their arrival in the 1980s. Options for controlling invasive mussels, particularly in unionid mussel habitats, are limited. Previously, carbon dioxide (CO2) showed selective toxicity for zebra mussels, relative to unionids, when applied in cool water (12 °C). We first determined 96‐h lethal concentrations of CO2 at 5 and 20 °C to zebra mussels and responses of juvenile plain pocketbook (Lampsilis cardium). Next, we compared the time to lethality for zebra mussels at 5, 12, and 20 °C during exposure to partial pressure of CO2 (PCO2) values of 110 to 120 atm (1 atm = 101.325 kPa) and responses of juvenile plain pocketbook and fragile papershell (Leptodea fragilis). We found efficacious CO2 treatment regimens at each temperature that were minimally lethal to unionids. At 5 °C, plain pocketbook survived 96‐h exposure to the highest PCO2 treatment (139 atm). At 20 °C, the 96‐h lethal concentration to 10% of animals (LC10) for plain pocketbook (173 atm PCO2, 95% CI 147–198 atm) was higher than the LC99 for zebra mussels (118 atm PCO2, 95% CI 109–127 atm). Lethal time to 99% mortality (LT99) of zebra mussels in 110 to 120 atm PCO2 ranged from 100 h at 20 °C to 300 h at 5 °C. Mean survival of both plain pocketbook and fragile papershell juveniles exceeded 85% in LT99 CO2 treatments at all temperatures. Short‐term infusion of 100 to 200 atm PCO2 at a range of water temperatures could reduce biofouling by zebra mussels with limited adverse effects on unionid mussels. Environ Toxicol Chem 2020;39:1546–1557. Published 2020. This article is a U.S. Government work and is in the public domain in the USA. Environmental Toxicology and Chemistry published by Wiley Periodicals LLC on behalf of SETAC.

Quantifying the loss of filtration services following mass mortality of invasive dreissenid mussels

Freshwater mussels are ecosystem engineers that provide important ecosystem functions and services, such as bio-filtration. A method for quantifying filtration capacity in rivers was developed and applied in a case study that estimated the loss in filtration services provided by invasive alien dreissenid mussels following a mass mortality event in the river Meuse in the Netherlands. Mass mortality resulted from a sudden water level drop caused by damage to a weir during severe winter conditions. The low water level allowed an assessment of dreissenid densities on river groynes. Imagery acquired by an unmanned aerial vehicle (UAV) was used to construct 3D images of groynes and to determine surface areas inhabited by mussels. A bootstrapping approach was applied to assess the loss of filtration capacity due to the mass mortality event. The filtration capacity of dreissenids on groynes was sufficient to filter 0.3% to 12.3% of the discharge in a 25 km impounded stretch of the river Meuse during high and low discharges, respectively. On average, 3.6% of the discharge could be filtered by dreissenids on the groynes. Mortality on air exposed groynes was 100% leading to a full loss of dreissenid filtration capacity at these locations. Initial recolonization of groynes formerly exposed to air was observed 17 months after the low water event, ultimately leading to a recovery of the filtration capacity.

Stationary Pattern of a Reaction-Diffusion Mussel-Algae Model.

In this paper, we consider a reaction-diffusion mussel-algae model with state-dependent mussel mortality. This mortality involves a positive feedback term resulting from the reduction of dislodgment and predation and a negative feedback term resulting from the intraspecific competition for mussel. We first study the global stability of the nonnegative uniform steady states and then focus on the existence and nonexistence of nonconstant positive steady states. The global bifurcation of constant positive steady state is also considered. Our results suggest that the regular patterning in mussel beds may be caused by the high mobility of algae or the low diffusion of mussels.

Establishing typical values for hemocyte mortality in individual California mussels, Mytilus californianus

Hemocytes are immune cells in the hemolymph of invertebrates that play multiple roles in response to stressors; hemocyte mortality can thus serve as an indicator of overall animal health. However, previous research has often analyzed hemolymph samples pooled from several individuals, which precludes tracking individual responses to stressors over time. The ability to track individuals is important, however, because large inter-individual variation in response to stressors can confound the interpretation of pooled samples. Here, we describe protocols for analysis of inter- and intra-individual variability in hemocyte mortality across repeated hemolymph samples of California mussels, Mytilus californianus, free from typical abiotic stressors. To assess individual variability in hemocyte mortality with serial sampling, we created four groups of 15 mussels each that were repeatedly sampled four times: at baseline (time zero) and three subsequent times separated by either 24, 48, 72, or 168 h. Hemocyte mortality was assessed by fluorescence-activated cell sorting (FACS) of cells stained with propidium iodide. Our study demonstrates that hemolymph can be repeatedly sampled from individual mussels without mortality; however, there is substantial inter- and intra-individual variability in hemocyte mortality through time that is partially dependent on the sampling interval. Across repeated samples, individual mussels' hemocyte mortality had, on average, a range of ~6% and a standard deviation of ~3%, which was minimized with sampling periods ≥72 h apart. Due to this intra-individual variability, obtaining ≥2 samples from a specimen will more accurately establish an individual's baseline. Pooled-sample means were similar to individual-sample means; however, pooled samples masked the individual variation in each group. Overall, these data lay the foundation for future work exploring individual mussels' temporal responses to various stressors on a cellular level.

Tolerance of Limnoperna fortunei (Dunker, 1857) (Bivalvia: Mytilidae) to aerial exposure at different temperatures

This work evaluates the tolerance of adult Limnoperna fortunei to aerial exposure at different temperatures. The aim was to contribute knowledge about factors, such as the mussel’s capacity for survival out of water over long periods, allowing the invasion of new environments after terrestrial transport. The analysis of the results revealed that the time of exposure to air influenced mortality at different temperatures throughout the experimental period (P < 0.001). The time to reach 100% mortality at temperatures of 10°C, 20°C, and 30°C were 11, 6, and 3 days, respectively. Mussel mortality outside the water is directly related to air temperature, so the higher the temperature is, the greater the mortality. The results suggest that in the case of the land transport of structures biofouled with L. fortunei, this species can remain alive for 2 to 10 days, depending on the ambient temperature. Thus, aerial exposure may be used as a strategy to control the dispersal of the golden mussel. However, the time required for effective elimination of all individuals depends on the air temperature. The aerial exposure can also be used to periodically clean industrial systems and to eliminate the dispersion of propagules by vectors (e.g., transport of equipments/watercrafts).

EFFECT OF MICROPLASTIC ON GREEN MUSSEL Perna viridis: EXPERIMENTAL APPROACH

Plastics become debris in the ocean that can be broken down into tiny particles of micro size (&lt;5 mm) and scattered into columns to the bottom of the water. Its very small size makes it easily accessible to various aquatic organisms, such as plankton, crustacean, and mussel. There have been many studies use mussel as a bioindicator for microplastic pollution but no one has observed the effect of microplastic exposure to mussel mortality. The aim of study was to examine the effect of microplastic on mussel mortality. Green mussels were collected from Mandalle Waters, Pangkep Regency (Pangkajene Kepualauan) then were exposed to microplastic polyethylene originating from shieving of scrub soap. The exposure was carried out for 7 days with microplastic concentrations of 0.05 (A), 0.5 (B), and 5 (C) g/L. Statistical test results showed the treatment B and C were not significantly different, but the two treatment were significantly different from treatment A. The percentage of green mussel mortality during the experiment increased with increasing microplastic concentrations with the percentage of mortality exceeding 80% at treatment C.

High mortality of mussels in northern Brittany – Evaluation of the involvement of pathogens, pathological conditions and pollutants

In 2014, a high and unusual mass mortality of mussels occurred in several important production areas along the French coasts of the Atlantic and English Channel. In the first quarter of 2016, mass mortalities hit farms on the west coast of the country once again. These heterogeneous mortality events elicited a multi-parametric study conducted during the 2017 mussel season in three sites in northern Brittany (Brest, Lannion and St. Brieuc). The objective was to assess the health status of these mussels, follow mortality and attempt to identify potential causes of the abnormal high mortality of farmed mussels in northern Brittany. Brest was the most affected site with 70% cumulative mortality, then Lannion with 40% and finally St. Brieuc with a normal value of 15%. We highlighted a temporal ‘mortality window’ that opened throughout the spring season, and concerned the sites affected by mortality of harmful parasites (including pathogenic bacteria), neoplasia, metal contamination, and tissue alterations. Likely, the combination of all these factors leads to a weakening of mussels that can cause death.

Temperature elevation and Vibrio cyclitrophicus infection reduce the diversity of haemolymph microbiome of the mussel Mytilus coruscus

Haemolymph microbiome was considered to be unique to healthy invertebrates and beneficial to the host against external pathogens, including disease resistance and maintenance of homeostasis. Here, we investigated the effects of elevated water temperature on infection of haemolymph microbiome of the hard-shelled mussel (Mytilus coruscus). Exposure to Vibrio. cyclitrophicus resulted in high mortality of mussels on day nine at 27 °C. The haemolymph was collected to determine the microbiota by 16 S rRNA gene sequencing. Exposure to waterborne V. cyclitrophicus increased the mortality of mussels that was associated with a reduction in the diversity of their microbial community. Principal coordinate analysis (PCoA) revealed that temperature was an essential factor in shaping microbial communities in mussel haemolymph. Vibrio exposure promoted the proliferation of opportunistic pathogens (e.g., Arcobacter and Francisella) at a lower temperature. A high abundance of Vibrio present in live and dead mussels, at 27 °C might contribute greatly to mortality, as indicated by linear discriminant analysis effect size (LEfSe). These data suggested that the dynamics of microbial community have unique biomarker species in mussel haemolymph that could be used as health indicators. An elevated temperature may reduce the ability of bacterial elimination function against infection in mussel haemolymph.

Control of mussel Mytilus galloprovincialis Lamarck fouling in water-cooling systems using plasma discharge.

To prevent marine macrofouling, the anti-fouling effect of liquid discharge on mussels Mytilus galloprovincialis Lamarck was investigated in a simulated water-cooling system. The effects of input energy, mussel distance from discharge center, continuous discharge time, and discharge energy distribution mode on mussel response (death or detachment) were systematically studied. The results showed that excellent anti-fouling effects could be achieved by increasing input energy, but the detachment rate and mortality of mussels decreased sharply when the mussels were farther away from the discharge center. Low frequency discharge for a long, continuous time and multiple stimuli at long intervals improved the anti-fouling effect. Shock waves are the most likely cause of mussel eradication, and the threshold values of peak pressure to prevent mussel settlement and to cause death were 0.02 MPa and 0.05 MPa, respectively.

Annual winter water level drawdowns limit shallow‐water mussel densities in small lakes

Abstract Regulated water level fluctuations alter the physical, chemical, and biological environments in lakes. However, few studies have measured the effects of repeated annual winter drawdowns on freshwater mussel populations (Bivalvia: Unionida), and it is unknown whether drawdowns permanently constrain mussel populations to deeper depths or are resilient to the annual disturbance. We quantified mussel densities during normal water levels and their mortality after the initiation of drawdown in lakes with annual winter (December–March) water level drawdown regimes that have existed for several decades. We used systematic quadrat sampling and visual snorkel surveys to estimate the number of live mussels at the surface and buried, and shell length in six drawdown and three control lakes at exposed (0.5‐m) and unexposed (1‐m) depths during winter drawdown. We also estimated mortality of mussels along the exposed lakebed for nine drawdown lakes. Study lakes were dominated by Elliptio complanata and Pyganodon cataracta. During normal water levels, mussel densities were much lower in drawdown lakes than control lakes at 0.5‐m depths. When mussels were present in drawdown lakes at 0.5‐m depths, they were often burrowed in the substrate and smaller than buried mussels in control lakes. At unexposed 1‐m depths, mussel densities were primarily explained by geographic region (western versus central Massachusetts), and found higher densities in drawdown lakes compared to control lakes. Mussel mortality after a single drawdown event in nine drawdown lakes was not correlated with drawdown magnitude or rate. Concordance of shell length‐frequency distributions between dead and buried living mussels in drawdown lakes suggests annual mortality of colonising individuals, which are typically small and presumably of young cohorts. Annual winter drawdowns still limit densities of E. complanata and P. cataracta in shallow‐water habitats in the following autumn (September–October) after 4–5 months of potential recovery, and hence constrain their distribution in lakes exposed to drawdowns. The development of sustainable water level management practices will be essential to minimise impacts to native mussel populations.

A novel paramyxean parasite, Marteilia tapetis sp. nov. (Cercozoa) infecting the digestive gland of Manila clam Ruditapes philippinarum from the southeast coast of Korea.

Paramyxean parasites in the genus Marteilia deteriorate digestive tissues of the host organisms, resulting in mortality of oysters, cockles, and mussels. Most reports of infection by Marteilia spp. are from Europe, while a new species of Marteilia was identified recently in Japan. Here, we report a previously unidentified species in the genus Marteilia from digestive diverticula of Manila clam Ruditapes philippinarum from the south coast of Korea. Prevalence of the parasite was low, 0.5-3.3% in the study sites. We characterized this species using light and transmission electron microscopy (TEM), and analyzed the 18S rDNA sequence. Light microscopy revealed the sporulation process from uninucleated stage to spore in the epithelial tissues of the digestive gland. TEM revealed that the parasites produced four secondary cells containing four tri-cellular spores. An electron-dense haplosporosome-like structure and striated inclusions were evident in the spore and the primary cells, respectively, while refringent granules were rarely observed in the secondary cells. Phylogenetic analyses of the 18S rDNA sequence placed this isolate in the genus Marteilia, although it is not identical to other known species in the genus. Based on morphological and molecular characters, we describe this species as Marteilia tapetis sp. nov., the second Marteilia species reported parasitizing Manila clams in Asian waters.

Use of a hydrodynamic model for the management of water renovation in a coastal system

Abstract. In this contribution we investigate the hydrodynamic response in Alfacs Bay (Ebro Delta, NW Mediterranean Sea) to different anthropogenic modifications in freshwater flows and inner bay–open sea connections. The fresh water coming from rice field irrigation contains nutrients and pesticides and therefore affects in multiple ways the productivity and water quality of the bay. The application of a nested oceanographic circulation modelling suite within the bay provides objective information to solve water quality problems that are becoming more acute due to temperature and phytoplankton concentration peaks during the summer period when seawater may exceed 28 ∘C, leading to high rates of mussel mortality and therefore a significant impact on the local economy. The effects of different management “solutions” (like a connection channel between the inner bay and open sea) are hydrodynamically modelled in order to diminish residence times (e-flushing time) and water temperatures. The modelling system, based on the Regional Ocean Modeling System (ROMS), consists of a set of nested domains using data from CMEMS-IBI for the initial and open boundary conditions (coarser domain). One full year (2014) of simulation is used to validate the results, showing low errors with sea surface temperature (SST) and good agreement with surface currents. Finally, a set of twin numerical experiments during the summer period (when the water temperature reaches 28 ∘C) is used to analyse the effects of proposed nature-based interventions. Although these actions modify water temperature in the water column, the decrease in SST is not enough to avoid high temperatures during some days and prevent eventual mussel mortality during summer in the shallowest regions. However, the proposed management actions reveal their effectiveness in diminishing water residence times along the entire bay, thus preventing the inner areas from having poor water renewal and the corresponding ecological problems.

Dispersal and mortality of translocated thick‐shelled river mussel Unio crassus Philipsson, 1788 adults revealed by radio tracking

Abstract Freshwater mussels are one of the most endangered groups of animals. Translocation is a common conservation measure for protecting them, but the success of this procedure varies widely, and the reasons for failure are frequently unknown. The dispersal and mortality of freshwater mussels was studied using a novel method: the radio tracking of Unio crassus translocated to new sites during a reintroduction project for this threatened species in the River Biała (Poland). Individual mussels were equipped with radio transmitters (10 mussels at each of four sites during low water in autumn, and 10 mussels at each of 11 sites during the spring flood). After being released into the water, the mussels generally moved away from the bank, and then performed erratic movements, dispersing over a larger area on flat parts of the riverbed, finally congregating on steep parts of the river bed in shallow water (0.20 m) near the bank. Mussel mortality was surprisingly low: all survived the autumn trial, and 80% of the radio‐tracked adults survived the spring flood. Some of them were displaced quite far within the channel (&gt;100 m), apparently suffering no harm. The confirmed cases of mortality were a few individuals that had become stranded on gravel bars formed during the flood that were now drying out. Long‐distance displacement was more frequent in lotic or regulated parts of the channel and in the parts where the channel morphology was unstable during the flood. Radiotelemetry should be used to test the suitability of receptor sites before translocation, in order to find places where mussels will choose to stay, and to standardize mussel detectability when monitoring relocation success.

Infection by invasive parasites increases susceptibility of native hosts to secondary infection via modulation of cellular immunity.

Parasite spillover from invasive aliens to native species increases the risk of disease emergence within native biota-either by direct harm to the new host or by indirect effects like increased risks of secondary infection. One example for such a detrimental effect is the parasitic copepod Mytilicola intestinalis that infected blue mussels Mytilus edulis after being introduced into the North Sea in the early 20th century. Since 1949, the parasite was blamed for multiple mass mortalities of infested blue mussels but evidence for a direct causal involvement of M.intestinalis remained circumstantial. Here, we now examine the potential effects of primary infections by the invasive parasite on the susceptibility to secondary infections with virulent bacteria (Vibrio spp.) in a full factorial infection experiment combining parasite infection (control vs. infected) with different Vibrio infection treatments (control, bath challenge, injection) in environmental conditions that either favoured the host (ambient temperature) or the bacterium (elevated temperature). The influence of primary and secondary infections on cellular immunity (phagocytosis) and Vibrio load in the haemolymph was used to correlate these results to host survival. Our results suggest that the rate of secondary Vibrio infection is increased due to lower efficiency of the cellular immune response. As a consequence, the failure of clearing Vibrio from the haemolymph might increase mortality of mussels infected by M.intestinalis. This demonstrates that indirect effects of parasite invasions can outweigh direct effects of the infection highlighting the need for a more integrative approach to understand and predict the consequences of parasite invasions.