Variation in scope for growth: a test of food limitation among intertidal mussels

Wellington Harbour supports large populations of the mussels Aulacomya maoriana, Mytilus galloprovincialis and Perna canaliculus that are almost entirely absent from nearby coastal locations in Cook Strait. We calculated scope for growth (SFG) using ambient Cook Strait water over a broad temporal scale and a broad range of seston conditions to determine if negative SFG explains this phenomenon. Although all three mussel species had positive mean SFG values, variation in SFG was high and negative values often occurred: A. maoriana 19.1 J g⁻¹ h⁻¹, 43% of mussels showed negative SFG; M. galloprovincialis 1.26, 52% negative SFG; P. canaliculus 45.6, 27% negative SFG. Negative SFG was most often due to negative absorption efficiency caused by metabolic faecal loss that is characteristic of mussels feeding in environments with low seston quality. From our ecophysiology data we constructed a model to estimate SFG based on physiological responses to the narrow range of seston conditions typical of Cook Strait (Model One), and a model to estimate SFG based on physiological responses of mussels to the broad range of seston conditions typical of Wellington Harbour and Cook Strait (Model Two). We used seston data collected over an 18-month period from sites in Wellington Harbour and Cook Strait to derive 159 estimates of species-specific mussel SFG from both models. Both models produced higher estimates of SFG for mussels in the Harbour compared with those at Cook Strait sites. This was consistent with elevated particulate concentrations in the Harbour than at Cook Strait sites, and in agreement with previous studies. For Cook Strait mussels, both models produced negative estimates of net energy balance for long periods of time (several months), whereas for Harbour mussels negative SFG estimates were generally short in duration. We conclude that our short-term laboratory-based determinations of SFG and our long-term bioenergetics modelling estimates do not conclusively support the hypothesis of food limitation for three coexisting taxa of mussels in the intertidal region of Cook Strait, New Zealand. Handling editor: P. Viaroli     

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.     

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.     

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

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

Modelling count and growth data with many zeros

We discuss the problem of modelling survival/mortality and growth data that are skewed with excess zeros. This type of data is a common occurrence in biological and environmental studies. The method presented here allows us to utilize both the survival/mortality and growth data when both data sets contain a large proportion of zeros. The method consists of four stages. Firstly the original data is divided into two sets; one contains all the surviving organisms and the other all of the mortalities. Secondly we calculate the actual growth of the surviving organisms and of the mortalities. Thirdly we count the number of surviving organisms for which growth has occurred and the number where no growth occurred, and the same count procedure is carried out on the mortalities. Next we model the survival/mortality data and growth/no growth data using logistic regression, and separately model the growth data using an ordinary regression. Finally we combine the three models to estimate the expected growth for a specific set of values of the explanatory variables. If we used another statistical method that did not involve the dead mussels or the ones with no growth, some of the information provided by these mussels would be lost. However, using the method we propose, all of the data collected are used to achieve an optimal estimation of the mussel growth. A case study of survival and growth of blue mussels (Mytilus galloprovincialis) and ribbed mussels (Aulacomya atra maoriana) trans-located from their natural distribution to different depths and sites along the axis of Doubtful Sound, New Zealand, is used for illustration.     

Variability in early post-settlement mortality in intertidal mussels and the role of size at settlement

Early post-settlement mortality is difficult to measure in mobile benthic taxa such as mussels. Field studies typically employ settlement collectors that often contain empty shells of dead individuals, which are usually disregarded. Here, empty shells of pediveligers, i.e. recent settlers that died before beginning to grow as juveniles, were used to assess temporal and spatial patterns of early mortality in mussels, whether size at settlement determined mortality, and whether mortality was density-dependent. This study took place at two intertidal sites in a large harbour in central New Zealand where monthly cohorts of mussels were collected over a 2-year period. Monthly mortality varied substantially, ranging from 0% to 42.6%, but not across the two sites that were 27?km apart. Although there was no density-dependent mortality evident within cohorts, the proportion of pediveligers which were dead on collection was highly positively correlated across sites. Live settlers were on average larger than dead settlers, and the size distribution of dead settlers was shifted toward smaller sizes compared to live settlers. At the extremes, of all settlers with shell length of 260?µm or less, 72% were dead on collection. By contrast, for settlers 310?µm or greater in length, 6% were dead. Together, these results (1) suggest that in this system size at settlement may be an important determinant of very early, peri-metamorphic mortality in mussels, with smaller settlers at higher risk, and (2) demonstrate the value in evaluating dead pediveligers for assessing natural early post-settlement mortality in mussels.     

Identification of Mytilus spp. and Pecten maximus in Irish Waters by Standard PCR of the 18S rDNA Gene and Multiplex PCR of the 16S rDNA Gene

Two molecular protocols for the identification of mussel and scallop have been developed using specific primers targeting the mitochondrial 16S ribosomal DNA gene and the nuclear 18S ribosomal DNA gene. Primers for the mitochondrial 16S ribosomal DNA gene in multiplex polymerase chain reaction (PCR) protocols yielded diagnostic DNA fragments for the mussels Mytilus edulis, Mytilus galloprovincialis, and the hybrid Mytilus edulis/galloprovincialis (335 bp), the king scallop Pecten maximus (382 bp) and the black scallop Mimachlamys varia (398 bp). DNA from the queen scallop Aequipecten opercularis showed no consistent PCR amplification of the 16S rDNA gene. Primers for the nuclear 18S rDNA gene in standard PCR protocols yielded similar-sized, diagnostic DNA fragments (approx. 190 bp) for the mussels Mytilus edulis, Mytilus galloprovincialis, and the hybrid Mytilus edulis/galloprovincialis, the king scallop Pecten maximus, the black scallop Mimachlamys varia, and the queen scallop Aequipecten opercularis. Both protocols have been tested with Mytilus spp., P. maximus, and 6 other bivalve species from a wide range of locations in Irish and European waters. Cross reaction of the specific primers with DNA template from any of the 6 other bivalve species was not observed. Rapid DNA extraction using FTA Card technology and the16S rDNA primers allowed for the detection of at least 10 mussel larvae in a subsample of natural plankton.     

Byssus attachment strength of two mytilids in mono-specific and mixed-species mussel beds

The mussel Xenostrobus securis is endemic to the brackish waters of New Zealand and Australia, but has successfully invaded the inner Galician Rías of NW Spain, where it coexists with the indigenous mussel Mytilus galloprovincialis. In this laboratory study, the plasticity of the byssus attachment strength of two mytilids was compared by manipulating substratum, salinity, and bed assembly. M. galloprovincialis showed stronger byssus detachment strength than X. securis, despite lower byssus coverage. Both species responded similarly to the substratum, with substantially lower byssus strength on methacrylate, which offered the lowest surface free energy. Byssus detachment values for M. galloprovincialis were lower at lower salinity. In mixed beds, a number of mussels moved upwards, eventually colonising the upper layers of the assemblage. This behaviour increased byssus strength but only for X. securis. X. securis is adapted to a wide spectrum of abiotic conditions, a trait that may promote its dissemination within estuarine environments.     

Intertidal assemblages associated with natural<Emphasis Type="Italic">corallina</Emphasis> turf and invasive mussel beds

There is considerable concern about conservation of biodiversity in highly disturbed and urbanized environments, although a very large proportion of biodiversity (i.e. the small and cryptic invertebrates) have been little studied in this regard. Many biogenic structures (e.g. coral reefs, mussel beds, foliose algae) provide habitat for a large number of small invertebrates. The features of these habitats to which these animals respond are complex and poorly documented. Invasive species are increasing in abundance and diversity in many disturbed estuaries, but most previous studies have concentrated on effects of invasive species on surrounding macroscopic assemblages. This study examines the assemblages of small invertebrates and algae living in natural patches of coralline turf and in patches of the invasive mussel, Mytilus galloprovincialis, on seawalls in Sydney Harbour. Although most taxa identified were common to both habitats, they were generally more abundant in turf than in the mussels, especially the more widespread and numerous taxa. Few taxa were unique to either habitat and those were generally sparse and patchy. In addition, there were relatively more smaller animals in the algal turf than in the mussels, although it is not known whether these were juveniles of adults present in both habitats, or different species. These data show that coralline turf and mussel beds do not provide similar intertidal habitat for associated assemblages and that overgrowth of natural biota by mussels may have strong indirect effects on associated assemblages. These warrant further experimental investigation, so that the effects of invasive species on local biodiversity can be better understood and managed.     

Dynamic measurements of black oystercatcher (Haematopus bachmani) predation on mussels (Mytilus californianus)

Intertidal zone mussels can face threats from a variety of predatory species during high and low tides, and they must balance the threat of predation against other needs such as feeding and aerobic respiration. Black oystercatchers (Haematopus bachmani) on the Pacific coast of North America can depend on the mussel Mytilus californianus for a substantial portion of their diet. Observations suggest that oystercatchers tend to focus on mussels beginning to gape their valves during rising tides, following periods of aerial emersion. We present detailed, autonomous field measurements of the dynamics of three such predation events in the rocky intertidal zone. We measured accelerations of up to 4 g imposed on mussels, with handling times of 115–290 s required to open the shell and remove the majority of tissue. In each case a single oystercatcher attacked a mussel that had gaped the shell valves slightly wider than its neighbors as the rising tide began to splash the mussel bed, but no other obvious characteristic of the mussels, such as body temperature or orientation, could be linked to the oystercatcher's individual prey choice.     

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

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

Community regulation: the relative importance of recruitment and predation intensity of an intertidal community dominant in a seascape context

Predicting the strength and context-dependency of species interactions across multiple scales is a core area in ecology. This is especially challenging in the marine environment, where populations of most predators and prey are generally open, because of their pelagic larval phase, and recruitment of both is highly variable. In this study we use a comparative-experimental approach on small and large spatial scales to test the relationship between predation intensity and prey recruitment and their relative importance in shaping populations of a dominant rocky intertidal space occupier, mussels, in the context of seascape (availability of nearby subtidal reef habitat). Predation intensity on transplanted mussels was tested inside and outside cages and recruitment was measured with standard larval settlement collectors. We found that on intertidal rocky benches with contiguous subtidal reefs in New Zealand, mussel larval recruitment is usually low but predation on recruits by subtidal consumers (fish, crabs) is intense during high tide. On nearby intertidal rocky benches with adjacent sandy subtidal habitats, larval recruitment is usually greater but subtidal predators are typically rare and predation is weaker. Multiple regression analysis showed that predation intensity accounts for most of the variability in the abundance of adult mussels compared to recruitment. This seascape-dependent, predation-recruitment relationship could scale up to explain regional community variability. We argue that community ecology models should include seascape context-dependency and its effects on recruitment and species interactions for better predictions of coastal community dynamics and structure.     

The influence of pre‐settlement and early post‐settlement processes on the adult distribution and relative dominance of two invasive mussel species

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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.     

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

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

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

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

Shell morphometrics and growth rate of the invasive bivalve mollusc Mytilus galloprovincialis in the Knysna estuarine embayment,South Africa

Shell morphometrics of the invasive mussel Mytilus galloprovincialis were compared at five sites and growth rate at four sites (in four seasons) in the Knysna estuarine embayment. Mussels from two sites (The Heads, Leisure Isle) where wave action was present had shells significantly lower for any length when compared with other more sheltered sites. There was no significant difference in shell width of mussels for any given length among sites. Mussels from The Heads had thicker shells than other sites, and those from Leisure Isle thicker shells than three other embayment sites where shells did not differ in thickness. Growth rate of mussels at two embayment sites (Thesen’s Wharf and Thesen Islands Marina) was greatest in autumn and summer whereas at The Heads and Leisure Isle there was little seasonal difference in growth rate. Growth rate of mussels at Thesen’s Wharf and Thesen Islands Marina was mainly greater in all seasons when compared with mussels at The Heads and Leisure Isle. The more rapid growth rate of mussels at the sheltered embayment sites might in part explain why M. galloprovincialis now dominates the mid- to lower intertidal on hard substrata in this region of the Knysna estuary.     

Mussel and dogwhelk distribution along the north‐west Atlantic coast: testing predictions derived from the abundant‐centre model

Aim We performed the first test of predictions from the abundant‐centre model using north‐west Atlantic coastal organisms. We tested the hypotheses that the density of intertidal mussels (Mytilus edulis and M. trossulus) and dogwhelks (Nucella lapillus) and mussel age and size would peak at an intermediate location along their distribution range. We also assessed the latitudinal variation in critical aerial exposure time. Location North‐west Atlantic coast between Newfoundland (Canada) and New York (USA), covering 1800 km of shoreline. Methods Using a nested design, we measured mussel density, age and size and dogwhelk density in 60 wave‐exposed rocky intertidal sites spread evenly in six regions. Critical aerial exposure times were determined using online data. Results Mytilus edulis peaked in abundance in Maine and was much less abundant in the other regions. Mytilus trossulus peaked in abundance in southern Nova Scotia and Maine, was less abundant in the other regions to the north, and was absent in the southernmost region (New York). Both mussel species were least abundant in a northern region (Cape Breton), although not in the northernmost region (Newfoundland). Critical aerial exposure times were negatively correlated with overall mussel density. Mussel age and size were similar among regions. Dogwhelks peaked in abundance in Maine and were much less abundant in the other regions, being positively correlated with overall mussel density across regions. Main conclusions Density data for M. edulis and N. lapillus provide limited support for an abundant‐centre pattern, while M. trossulus shows a clear ramped‐south distribution. Critical aerial exposure times suggest that physiological stress during summer and winter low tides may be lowest in Maine and southern Nova Scotia, which might partially explain mussel predominance in those regions. Winter ice scour in Cape Breton may explain the abundance trough observed there. Mussel size and age may be more limited by wave exposure at our sites (as they all face open waters) than by regional differences in environmental stress. Dogwhelks, which prey on mussels, seem to respond positively to prey density at the regional scale. Our study supports the notion that, while the abundant‐centre model is a useful starting point for research, it often represents an oversimplification of reality.     

Mytilid mussels: global habitat engineers in coastal sediments

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

Commensals of Mytilus galloprovincialis in the Black Sea: Urastoma cyprinae (Turbellaria) and Polydora ciliata (Polychaeta)

The intensity of infection of the mussel Mytilus galloprovincialis Lamarck in the Black Sea by the turbellarian Urastoma cyprinae (Graff), which lives on its gills, was found to be higher in larger hosts, reaching a maximum in mussels of 50–70 mm length. Greater numbers occured in mussels inhabiting a silty bottom than in cultivated mussels suspended above the bottom. Over the period 1982–1987, U. cyprinae was most numerous in winter and especially so in years that were colder. The spionid polychaete Polydora ciliata (Johnston) also infects M. galloprovincialis, burrowing into the shell. Young spionids of up to 1 mm length occured in mussels with a shell length of 35 mm. Numbers of this commensal reached a maximum in mussels of intermediate size.