Mortality,movement and behaviour of native mussels during a planned water‐level drawdown in the Upper Mississippi River

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Predator‐induced morphological defences in two invasive dreissenid mussels: implications for species replacement

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Valve movement of the freshwater mussel Anodonta anatina: a reciprocal transplant experiment between lake and river

The effects of source and caging on the valve movements of the freshwater unionid mussel (Anodonta anatina) were studied in a reciprocal transplant experiment between a lake and its outflow. Caged mussels were moved and compared with those remaining in their natural environment on the lake or river bottom. At both sites, the mussels from the study site and the transplanted mussels from the opposite site were monitored simultaneously. In river the averaged weighted valve openness was higher and the number of valve movements was lower than in the lake. The mussels monitored in the lake exhibited a diurnal rhythm of valve movements which differed between the lake-bottom and the caged animals. Caging was found to increase valve openness. On the other hand, little variation appeared in valve openness between caged and bottom animals in the river, where diurnal rhythms were almost nonexistent. In the river the valve movements were more variable in respect to time than in the lake.     

Effects of artificial epibionts on byssogenesis,attachment strength,and movement in two size classes of the blue mussel,Mytilus edulis

Blue mussels (Mytilus edulis) can alter the strength of byssal attachment and move between and within mussel aggregations on wave‐swept shores, but this movement ability may be limited by epibiont fouling. We quantified the effects of artificial epibiont fouling on the production of byssal threads, attachment strength, and movement in two size classes of blue mussels. In a factorial experiment, large epibiont‐covered mussels produced more functional byssal threads (i.e., those continuous from animal to substrate) after 24 h than large unfouled and small fouled mussels, but not more than small unfouled mussels. Small unfouled mussels formed and released more byssus bundles compared to any other treatment group, which indicates increased movement. Conversely, epibiont fouling resulted in decreased numbers of byssus bundles shed, and therefore reduced movement in small mussels. Epibiont‐covered mussels started producing byssal threads sooner than unfouled mussels, while small mussels began producing byssal threads earlier compared to large mussels. Mean attachment strength from both size classes increased by 9.5% when mussels were artificially fouled, and large mussels had a 34% stronger attachment compared to small mussels. On the other hand, a 2.3% decrease in attachment strength was found with increasing byssus bundles shed. Our results suggest that fouling by artificial epibionts influences byssal thread production and attachment strength in large mussels, whereas epibionts on small mussels impact their ability to move. Mussels are able to respond rapidly to fouling, which carries implications for the dynamics of mussel beds in their intertidal and subtidal habitats, especially in relation to movement of mussels within and among aggregations.     

Behaviour and physiology are linked in the responses of freshwater mussels to drought

1. Droughts have become prevalent in the south‐eastern U.S.A. and are predicted to become more common in the future. Drought conditions have particularly strong negative effects on sessile aquatic organisms with limited dispersal ability. This study explored the linkages between physiological tolerance, behavioural response and survival of three species of freshwater mussels subjected to drought. 2. To assess physiological tolerance, we measured survival under desiccation at 25, 35 and 45 °C in the laboratory. To assess behavioural responses, we tracked horizontal and vertical movement of mussels in a drying stream reach. 3. Uniomerus tetralasmus showed the greatest desiccation tolerance, Lampsilis straminea was intermediate, and Pyganodon grandis had the lowest tolerance at all temperatures. 4. In the drying stream reach, U. tetralasmus showed little horizontal movement and quickly became stranded. The other two species tracked the receding water. It was not until the pool had been reduced to c. 10% of its original size that ≥50% of P. grandis and L. straminea became stranded. 5. Uniomerus tetralasmus and L. straminea burrowed in response to becoming stranded; however, both species burrowed only shallowly (3–4 cm), presumably because burrowing to greater depths did not convey increasing thermal refuge. Pyganodon grandis rarely burrowed. No P. grandis survived the 15‐week drought, while 45% of L. straminea and 77% of U. tetralasmus survived. 6. Three strategies emerged for freshwater mussels to cope with drought conditions: tracking (intolerant), track then burrow (semi‐tolerant) and burrowing (tolerant). Results suggest that drought poses the greatest threat to intolerant trackers, while tolerant burrowers are the most resistant to drought conditions.     

Potential use of beaver Castor fiber L., 1758 dams by the thick-shelled river mussel Unio crassus Philipsson, 1788

ABSTRACT

The objective of this paper is to draw attention to a previously unrecognised habitat for Unio crassus Philipsson, 1788 (Bivalvia: Unionidae), namely beaver dam-and-pond complexes. The construction of beaver ponds will allow the colonisation of high energy streams by providing flow refuges during periods of high near-bed shear stress. The absence of freshwater mussels (Unionidae) in mountain streams may result from steep slopes, shear stress during floods, lack of food suspended in the water column (seston), and unfavourable conditions for the settlement and growth of juvenile mussels. Transportation of parasitic larvae on host fish is the main dispersal mechanism for the mussels. A possible constraint on the colonisation of beaver complexes by freshwater mussels is that beaver dams may hinder the upstream movement of their minnow hosts. In the surveyed beaver ponds, the first specimens of U. crassus were found in 2008. Single mussels were also found in subsequent years. It is uncertain whether the occurrence of U. crassus depends on the periodic support of the population from the main river, but the presence of beaver ponds is certain to have given it the opportunity of colonising new areas.     

Effects of storage temperature on oxygen depletion in intra-valve water and survival of the Mediterranean mussel Mytilus galloprovincialis Lmk

Mussels are commonly air‐stored during transportation and as a result suffer from anoxia. In this study, storage temperature effects on the viability and characteristics of the released intra‐valve water of mussels were examined. Mussels kept at 20°C released all of their intra‐valve water within approximately 60 h and died within 4 days; oxygen concentration in the intra‐valve water dropped below the detectable level. In contrast, mussels kept at 0 and 5°C released 8.8% and 12% of their intra‐valve water, respectively. The oxygen concentration in this water remained stable at about 3–4 mg L⁻¹ until hour 72 of exposure to air and all mussels survived (5°C). Mussels immersed in seawater over‐saturated with oxygen (35 mg L⁻¹) did not show any uptake of the surplus oxygen into their intra‐valve water.     

Effect of anthropogenic feeding regimes on activity rhythms of laboratory mussels exposed to natural light

Anthropogenic disturbance may affect animal behaviour and should generally be minimised. We examined how anthropogenic disturbance (24 h food deprivation) affected circadian rhythms in laboratory mussels Mytilus edulis exposed to natural light in the absence of tides. Repeated measures data were collected on mussel gape angle, exhalant pumping and valve adduction using a Hall sensor system over eight consecutive 24 h periods when exposed to two feeding conditions after 24 h food deprivation. Mussels (fed once per day at either midday or midnight) exposed to natural light showed a clear day–night rhythm with increased nocturnal activity: significantly greater gape angle, increased exhalant pumping and had significantly higher valve adduction rates. However, circadian rhythms were less clear directly after anthropogenic food deprivation, in terms of the circadian rhythm in gape angle becoming significantly more apparent over the following days. Unlike mussels fed at midnight, those fed at midday displayed no significant change in gape angle from the hour before to the hour after they were fed, i.e. mussels given food at midday reacted to this food less than mussels fed at midnight. We suggest that independent of feeding time, laboratory mussels exposed to natural light and free from anthropogenic disturbance increase feeding activity at night because their circadian rhythms are strongly influenced by light levels. This study emphasises that the behaviour of animals in the laboratory and in the wild can be altered by anthropogenic disturbances such as vibrations caused by experimental setups and artificial illumination at night.     

Scale-dependent associations between native freshwater mussels and invasive Corbicula

In North America there is conflicting evidence concerning whether the invasive Asian clam, Corbicula fluminea, and native mussels (Unionidae), can successfully co-exist. One reason underlying disparate conclusions may be the different spatial scales at which data have been collected. We compared the distribution and abundance of native unionid mussels and Corbicula at two spatial scales, stream reaches and 0.25 m² patches, within one biogeographic region, the Ouachita Highlands, of the south central U.S. We found that Corbicula abundance was negatively related to native mussel abundance at small spatial scales. While Corbicula densities varied widely in patches without native mussels, and in patches where mussels occurred at low abundance, Corbicula density was never high in patches where mussels were dense. We hypothesize that the likelihood of successful Corbicula invasion decreases with increasing abundance of adult native mussels. Several mechanisms may potentially drive this pattern including lack of space for Corbicula to colonize, physical displacement by actively burrowing mussels, and locally reduced food resources in patches where native mussels are feeding. In addition, Corbicula may be unable to withstand environmental bottlenecks as readily as unionids. When patch-scale density and biomass information were pooled to represent entire stream reaches, the negative relationship between native mussels and Corbicula was no longer as apparent, and there was not a significant relationship between native mussels and Corbicula. These results point to the importance of appropriate sample scale in examining potential associations between species.     

Estimation of vital population rates to assess the relative health of mussel assemblages in the Upper Mississippi River

1. Native freshwater mussels are a guild of benthic, filter feeding invertebrates that perform important ecological functions in rivers. Because of their long lifespans (30–50 years or longer), mussels are slow to respond to human-induced alterations. Thus, development of sensitive indicators of mussel population responses to river conditions and management would be beneficial. Compared to marine species, estimation of vital rates (e.g. survival, growth) in freshwater mussels has received little attention.
2. We placed passively integrated transponder tags on 578 mussels of four species (Amblema plicata, Cyclonaias pustulosa, Obliquaria reflexa, and Pleurobema sintoxia) in a well-studied mussel assemblage in a side channel of the upper Mississippi River. Growth and survival of tagged mussels were assessed annually for 4 years across core (high density) and peripheral (low density) areas of the assemblage.
3. Overall survival was highly variable, ranging from c. 15 to 90%, and was related to life history, habitat quality, and hydrologic events. Survival, which varied significantly among species and over time, was consistently higher in the dense and species-rich core of the mussel assemblage, relative to the periphery because substrates were consistently more stable in the core of the mussel bed relative to the periphery. Substrate movement during low flows was an order of magnitude lower in the core relative to the periphery, and survival was inversely related to stability of river substrates. Patterns in habitat-specific survival indicate source–sink population dynamics such that mussels in the core habitat provide recruitment to the periphery, but mussels in the periphery are subject to unsustainably low survival; additional studies to track the source of recruitment in the periphery are needed to test this hypothesis.
4. Growth rate did not vary significantly between core and peripheral areas but did vary by species. Growth rate (proportional change per year) declined with age, and was similar at mean age for A. plicata (0.016 per year), P. sintoxia (0.015 per year), and C. pustulosa (0.013 per year), but much lower for O. reflexa (0.008 per year).
5. Effective management decisions for mussels requires a better understanding of how vital rates govern populations and how they vary across a suite of physical and biological factors. Information on how population vital rates vary among species and over time gives managers another tool to understand how mussels may respond to management actions such as habitat restoration projects. Given the importance of substrate stability inferred from this study, management actions that maintain or increase substrate stability are likely to result in high quality mussel assemblages and may restore a valuable component of ecosystem function in this region.

     

Evolution of bilaterally asymmetrical larvae in freshwater mussels (Bivalvia: Unionoida: Unionidae)

Bilaterally asymmetrical glochidia (i.e. bivalved parasitic larvae bearing a large marginal appendage on a single valve) have been reported from five Asian freshwater mussel genera belonging to two separate subfamilies, the Gonideinae (i.e. Pseudodon, Solenaia, and Physunio) and Rectidentinae (i.e. Contradens and Trapezoideus). This classification requires that the bilaterally asymmetrical glochidium‐bearing mussels are not monophyletic, and suggests that this atypical larval morphology evolved twice in the same geographic region. Although homoplastic glochidium characters are known (e.g. marginal appendages and size), we hypothesized that bilaterally asymmetrical glochidia represent a novel morphological synapomorphy. We tested the monophyly of the mussels bearing bilaterally asymmetrical glochidia using a molecular matrix consisting of representatives from all six freshwater mussel families and three molecular markers (28S, 16S, and COI). Bayesian inference, maximum likelihood, and ancestral state reconstruction were employed to estimate the phylogeny and larval trait transformations. The reconstructed phylogeny rejects the monophyly of the asymmetrical glochidium‐bearing mussels and resolves two putative origins of asymmetrical glochidia; however, ancestral state reconstruction supports asymmetrical glochidia as a synapomorphy of only one supraspecific taxon of the Rectidentinae. In the Gonideinae, asymmetrical glochidia were autapomorphic of Pseudodon cambodjensis (Petit, 1865). That is, no other taxa resolved among the Gonideinae had bilaterally asymmetrical glochidia, including other Pseudodon species. We describe how the alleged intrageneric glochidial variation in Pseudodon, and in the other genera of the Gonideinae reported to have asymmetrical glochidia (i.e. Solenaia and Physunio), challenge the resolved convergence of asymmetrical glochidia. Our results are discussed in the context of freshwater mussel larval evolution, patterns in life‐history traits, and the classification of freshwater mussels generally. © 2015 The Linnean Society of London     

Zebra mussels decrease burrowing ability and growth of a native snail, <Emphasis Type="Italic">Campeloma decisum</Emphasis>

Invasive species can drive native organisms to extinction by limiting movement and accessibility to essential resources. The purpose of this study was to determine if zebra mussels (Dreissena polymorpha) affect the burrowing ability and growth rate of a native snail, Campeloma decisum. Snails with and without zebra mussels were collected from Douglas Lake, MI, and burrowing depths were studied in both the laboratory and Douglas Lake. Growth rates were calculated as the amount of shell growth from 2004 to 2005. Both the tendency of snails to burrow and the depth to which they burrowed into the substrate were significantly decreased by the presence of zebra mussels on snail shells in both laboratory and lake experiments. There was no difference in the percentage of snails that exhibited growth as a function of zebra mussel presence. However, for those snails that grew, there was a 50% higher growth rate for snails without zebra mussels compared to snails with zebra mussels. These negative effects of zebra mussels on growth and burrowing ability will likely lead to decreases in snail population densities in the future. Handling editor: S. Wellekens     

Reduced byssal thread production and movement by the intertidal mussel<Emphasis Type="Italic"> Hormomya mutabilis</Emphasis> in response to effluent from predators

Laboratory experiments were carried out to investigate byssal thread production by the intertidal mytilid mussel Hormomya mutabilis in response to effluent from the predatory crab Eriphia smithii and the starfish Coscinasterias acutispina. During the early period of the experiment, large H. mutabilis exposed to crab effluent produced a significantly smaller number of functional byssal threads than mussels in crab-free water. No significant difference in the diameter of threads produced in the two treatments was detected. The number of functional byssal threads produced by small H. mutabilis exposed to crab effluent did not differ significantly from that of mussels in crab-free water. However, small H. mutabilis exposed to crab effluent tended to discard fewer byssal bundles, that is, they shifted their attaching sites less frequently than similar mussels in crab-free water. In the presence of waterborne cues from the crab, H. mutabilis tended to reduce both the secretion of byssal threads and movement across the substratum. No significant differences in behaviour were observed between large mussels exposed to effluent from the starfish and those unexposed. The adaptive significance of the responses shown by H. mutabilis is discussed in terms of protection against predators differing in foraging behaviour. Electronic Publication     

The respiratory capacity of marine mussels (Mytilus galloprovincialis) in relation to the high temperature threshold

Thermal tolerance limits of ectotherms may result from respiratory limitations. In response to declining oxygen availability, organisms have shown to exhibit oxyregulation by enhancing ventilation and heartbeat rates. In this study we examined how this regulatory response in mussels (Mytilus) changes with increasing temperature. Experimental mussels showed extensive oxyregulation at temperatures near to their habitat temperature, but increasingly lost this capacity towards higher temperatures. At breakpoint temperature no regulation took place and respiration rates changed proportional to oxygen availability. These results revealed how thermal limitations relate to respiratory capacity of mussels.     

Potential effects on zooplankton from species shifts in planktivorous mussels: a field experiment in the St Lawrence River

1. Suspension feeding by bivalves exceeds that by other planktivores in many North American rivers, and food webs may be altered substantially by differences in feeding patterns between native unionid mussels and invading dreissenid mussels. 2. We conducted an experiment comparing zooplanktivory by one unionid and two dreissenid species that addressed several primary questions. Is benthic planktivory important in this river? Has this linkage been altered substantially by dreissenids? Do the two dreissenid species differ in planktivory, and is this ecologically important if quagga mussels extend their geographical range? 3. Our 12‐day experiment consisted of controls (no mussels) and treatments with unionid (Elliptio complanata), quagga (Dreissena bugensis) or zebra (D. polymorpha) mussels in 3500‐L, 80‐μm mesh enclosures placed in a slackwater area of the St Lawrence River. 4. The density of the most abundant calanoid copepod Eurytemora affinis increased in the presence of dreissenids, probably as an indirect food web response. By day 12, a cumulative effect was shown by the most overwhelmingly abundant rotifer, Polyarthra, whose density declined dramatically in dreissenid enclosures compared with control and unionid enclosures. Rotifer densities in unionid enclosures were not different from controls, nor were dreissenid treatments different from each other. The effects on rotifers were probably from predation, as Chl‐a did not vary among treatments. 5. We conclude that benthic‐pelagic coupling via planktivory is important in slackwater areas. Dreissenids have strengthened this linkage, but range extension of quaggas should not appreciably alter effects produced by a similar biomass of zebra mussels.     

Significant fish predation on zebra mussels Dreissena polymorpha in Lake Champlain,U.S.A.

The objectives of this study were to: (1) determine which fishes were consistently eating zebra mussels Dreissena polymorpha in Lake Champlain and document their feeding behaviour and (2) quantify the diet composition of the fish predators that were found to consume zebra mussels. From 2002 to 2005, freshwater drum Aplodinotus grunniens, pumpkinseed Lepomis gibbosus, yellow perch Perca flavescens and rock bass Ambloplites rupestris all consumed zebra mussels at varying frequencies and amounts. Aplodinotusgrunniens and L. gibbosus chewed clumps of zebra mussels, expelling shells, whereas P. flavescens and A. rupestris swallowed small individuals whole. Lepomis gibbosus consumed zebra mussels at the highest frequency (65–89% of prey consumed) and zebra mussels comprised a large part of this fish’s diet (up to 40% by dry mass). Zebra mussels were also an important component of the diet of A. grunniens (up to 59% of the diet by dry mass, 40–63% frequency of consumption). The percentage of the diet comprising zebra mussels in P. flavescens and A. rupestris varied significantly from year to year but never exceeded 10%. Because A. grunniens and L. gibbosus crushed zebra mussels, the nutritional return from consuming zebra mussels would be similar to other prey; for P. flavescens and A. rupestris zebra mussels were only partially digested and the nutritional return would probably be low. As predation on zebra mussels is widespread and significant, it is possible that fish predators could contribute to regulating the population of zebra mussels in Lake Champlain.     

Indigenous microflora and opportunistic pathogens of the freshwater zebra mussel, Dreissena polymorpha

Freshwater fouling invertebrate zebra mussels (Dreissena polymorpha) harbor a diverse population of microorganisms in the Great Lakes of North America. Among the indigenous microorganisms, selective species are opportunistic pathogens to zebra mussels. Pathogenicity to zebra mussels by opportunistic bacteria isolated from the mussels was investigated in this study. Among the more than 30 bacteria isolated from temperature-stressed mussels, Aeromonas media, A. veronii, A. salmonicida subsp. salmonicida, and Shewanella putrefaciens are virulent pathogens to juvenile zebra mussels. Inoculation of a bacterial concentration of A. media, A. salmonicida subsp. salmonicida and S. putrefaciens at 10⁷ cells per zebra mussel resulted in 100% mortality within 5 days, and only 64.9% for A. veronii. In contrast, mortality was less than 12.3% following inoculation of a sterile phosphate buffer solution as a control. In addition, mortality was dependent on the size of the pathogen population used in inoculation and the incubation temperature, indicating the close relationship between the bacterial population and subsequent death. On the mussel tissue, a dense microbial population was evident from the moribund mussels viewed with Scanning Electron Microscope (SEM). Opportunistic bacteria invaded and destroyed the D. polymorpha tissue after 7 days of incubation when the bacterial inoculation was larger than 10⁵ per zebra mussel. Our results suggest that mussels are reservoirs of opportunistic pathogenic microorganisms to aquatic organisms and humans and a better understanding of the microbial ecology of the mussels will provide insights to the possible health hazards from these microorganisms.     

Context-dependent effects of freshwater mussels on stream benthic communities

1. We asked whether unionid mussels influence the distribution and abundance of co‐occurring benthic algae and invertebrates. In a yearlong field enclosure experiment in a south‐central U.S. river, we examined the effects of living mussels versus sham mussels (shells filled with sand) on periphyton and invertebrates in both the surrounding sediment and on mussel shells. We also examined differences between two common unionid species, Actinonaias ligamentina (Lamarck 1819) and Amblema plicata (Say 1817). 2. Organic matter concentrations and invertebrate densities in the sediment surrounding mussels were significantly higher in treatments with live mussels than treatments with sham mussels or sediment alone. Organic matter was significantly higher in the sediment surrounding Actinonaias than that surrounding Amblema. Actinonaias was more active than Amblema and may have increased benthic organic matter through bioturbation. 3. Living mussels increased the abundance of periphyton on shells and the abundance and richness of invertebrates on shells, whereas effects of sham mussels were similar to sediment alone. Differences in the amount of periphyton growing on the shells of the two mussel species reflected differences in mussel activity and shell morphology. 4. Differences between living and sham mussel treatments indicate that biological activities of mussels provide ecosystem services to the benthic community beyond the physical habitat provided by shells alone. In treatments containing live mussels we found significant correlations between organic matter and chlorophyll a concentrations in the sediment, organic matter concentrations and invertebrate abundance in the sediment and the amount of chlorophyll a on the sediment and invertebrate abundance. There were no significant correlations among these response variables in control treatments. Thus, in addition to providing biogenic structure as habitat, mussels likely facilitate benthic invertebrates by altering the availability of resources (algae and organic matter) through nutrient excretion and biodeposition. 5. Effects of mussels on sediment and shell periphyton concentrations, organic matter concentrations and invertebrate abundance, varied seasonally, and were strongest in late summer during periods of low water volume, low flow, and high water temperature. 6. Our study demonstrates that freshwater mussels can strongly influence the co‐occurring benthic community, but that effects of mussels are context‐dependent and may vary among species.     

Evidence of phenotypic plasticity in the response of unionid mussels to turbidity

1. Increases in total suspended solids (TSS) reduce feeding and reproductive success of unionid mussels, but mussels in turbid rivers are less affected than those in clear rivers, probably due to differences in gill and palp morphology. This study was designed to determine whether the differences observed between adult mussels in populations from turbid versus clear rivers are due to phenotypic plasticity.
2. Parasitic larvae (glochidia) of Lampsilis siliquoidea (Fatmucket) obtained from a low turbidity river (TSS < 5 mg/L) were transformed on Ambloplites rupestris (Rock Bass) in the laboratory to obtain juvenile mussels for rearing under clear and turbid conditions in the laboratory (i.e. nominally 0 versus 50 mg/L suspended river sediment). Juveniles obtained were reared under these contrasting conditions until age 3–4 weeks, when they were examined in a feeding experiment under a range of TSS concentrations (0, 5, 10, 15, 20, 25 mg/L).
3. The clearance rate (volume cleared of particles per unit time) of algae by juveniles from both rearing conditions was similar for the no-TSS control and declined with increased TSS. The rate of decline was, however, lower in the group reared under turbid conditions, which is consistent with reports for adult mussels.
4. Our results indicate that differences in the clearance rate response to high TSS observed in mussels in clear and turbid rivers are probably driven in part by phenotypic plasticity. This finding provides a mechanism to explain how freshwater mussels thrive under turbid conditions as well as informs conservation efforts involving reintroduction of mussels in this highly imperilled taxon.

     

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.