Temporal scales of variation in settlement and recruitment of the mussel Perna perna (Linnaeus, 1758)

Population dynamics of many intertidal organisms are strongly affected by the abundance and distribution of larvae arriving on the shore. In particular, not only absolute numbers of settlers but also the degree of synchronisation of settlement can have a strong influence on whether density-dependent or density-independent processes shape adult shape populations. Temporal variation in rates of settlement and recruitment of the mussel Perna perna on the south coast of South Africa was investigated using a nested spatial design at different temporal scales. Variability in settlement at spring tides was examined at two temporal scales: lunar (to investigate the effect of state of the moon on settlement) and tidal (to investigate the influence of state of the tide on mussel settlement). Recruitment over neap tides was examined at one temporal scale, fortnight (to investigate the effect of date on mussel recruitment).Strong temporal variation was evident for both settlement and recruitment, but not at all time scales. Distinct peaks of settler/recruit abundance were observed during the lunar and neap tide studies. Recruitment intensity differed over the course of the year, and pulsing of recruitment was generally synchronised among locations. However, the strength of pulsing differed dramatically among locations, giving a significant interaction between fortnight and location. The finest temporal scale, investigated in the tidal study, did not reveal a significant effect of the state of the tide on settlement. The state of the moon (new or full) was not significant as a main factor (p = 0.052), although generally more settlers arrived on the shore during new moon. Phase of the moon appeared to have an effect on settler abundances, but only when and where densities were high.     

Population dynamics of the green-lipped mussel, Perna canaliculus, at various spatial and temporal scales in northern New Zealand

Ecological processes that differentiate and maintain intertidal populations of mussels, Perna canaliculus, were studied within three sites at Ninety Mile Beach, northern New Zealand. At these three sites (Scott Point, The Bluff and Tonatona Beach), the dynamics of larval availability, primary and secondary settlement, recruitment and mortality rates were investigated at various spatial and temporal scales. (1) Mussel concentrations in seawater were variable with respect to study site and time of year, with highest abundances at the northernmost population (Scott Point) and lowest concentrations at the middle population (The Bluff). In seawater at all three sites, small mussels (< 0.25 mm in shell length) were more abundant in August 2000, while larger mussels (> 0.5 mm in shell length) were more abundant in March 2001. (2) Primary and secondary settlement patterns were investigated during short-term (daily) and long-term (monthly) settlement experiments, within quadrats that were cleared of all mussels in both the mussel bed and in adjacent algal band habitats. At all sites, primary settlement (< 0.5 mm in shell length) was high within the algal band habitat in August 1999, 2000 and 2001. Conversely, secondary settlement (> 2.0 mm in shell length) was high within cleared areas in the mussel beds in November-March 1999-2000 and 2000-2001. Abundance of mussels settling on artificial substrates placed in the intertidal did not differ greatly from comparable areas of natural substrates (bare rock or algae within cleared quadrats). (3) Recruitment and mortality rates were recorded during monthly surveys of the adult populations. Within three mussel size classes (< 24, 25-74 and > 75 mm in shell length), peak recruitment coincided with high mortality in August of the 2 years studied. However, the most dramatic turnover of the population was observed at Scott Point in both years, following a spawning event. In adjacent waters at Scott Point, large accumulations of drift algae covered (up to 100% cover) with juvenile mussels may deplete food supplies usually delivered to intertidal adult mussels, causing their demise. Mats of adult mussels were observed “peeling-off” from the rocky shore at this time of the year, making space available to the new recruits. Where nearshore algal accumulations were moderate to low, only moderate to low mussel turnovers were observed (e.g. Tonatona Beach and The Bluff).     

Oxidative stress in digestive gland and gill of the brown mussel (Perna perna) exposed to air and re-submersed

Mussels Perna perna were exposed to air for 24 h showing a clear increase in the levels of lipid peroxidation and oxidative DNA damage, measured as 8-oxo-7,8-dihydro-2′-deoxyguanosine (8-oxodGuo). The levels of lipid peroxidation increased both in the digestive gland and gills, while oxidative DNA damage increased only in the gills. After the 24 h of air exposure, mussels were re-submersed for a period of 3 h, leading values to return to a pre-aerial exposure levels. Control animals were kept immersed during the whole period. Several antioxidant and complementary enzymes (superoxide dismutase (SOD), catalase (CAT), glutathione peroxidase (GPx), glutathione reductase (GR), glucose-6-phosphate dehydrogenase (G6PDH), glutathione S-transferase (GST) and the levels of total glutathione (Total GSH) were assayed in a second set of experiments where one group of mussels were exposed to air for 18 h and other to 1 h re-submersion after 18 h aerial exposure. Only a 52% increase in the glutathione S-transferase activity was observed in the digestive gland, which remained elevated to about 40% after 1 h re-submersion, showing that defense systems can be modulated even during oxygen deprivation in P. perna. The DNA and lipid oxidative damage observed after aerial exposure indicates that mussels face an oxidative challenge, and are able to counteract such an “insult” as values of lipid peroxidation and DNA damage returned to control values after 3 h re-submersion.     

Cryopreservation of Greenshell™ mussel (Perna canaliculus) trochophore larvae

The Greenshell™ mussel (Perna canaliculus) is the main shellfish species farmed in New Zealand. The aim of this study was to evaluate the effects of cryoprotectant concentration, loading and unloading strategy as well as freezing and thawing method in order to develop a protocol for cryopreservation of trochophore larvae (16–20 h old). Toxicity tests showed that levels of 10–15% ethylene glycol (EG) were not toxic to larvae and could be loaded and unloaded in a single step. Through cryopreservation experiments, we designed a cryopreservation protocol that enabled 40–60% of trochophores to develop to D-larvae when normalized to controls. The protocol involved: holding at 0 °C for 5 min, then cooling at 1 °C min⁻¹ to −10 °C, holding for a further 5 min, then cooling at 0.5 °C min⁻¹ to −35 °C followed by a 5 min hold and then plunging into liquid nitrogen. A final larval rearing experiment of 18 days was conducted to assess the ability of these frozen larvae to develop further. Results showed that only 2.8% of the frozen trochophores were able to develop to competent pediveligers.     

Respiratory responses to temperature and hypoxia in the nonindigenous Brown Mussel, Perna perna (Bivalvia: Mytilidae), from the Gulf of Mexico

The respiratory responses to increasing temperature and progressive hypoxia were examined relative to temperature acclimation in the nonindigenous, brown mussel, Perna perna (Mytilidae) from the Gulf of Mexico. When oxygen uptake rate (V?_O2) was recorded at near full air O₂ saturation, rate-temperature curves for Texas specimens of P. perna were sigmoidal, V?_O2 generally increasing with increasing temperature but becoming suppressed as temperatures approached 10 and 30 °C, corresponding closely to this species' incipient thermal limits. At each tested temperature, V?_O2 did not differ among individuals acclimated to 15, 20, or 25 °C. Lack of thermal acclimation was also reflected in acclimatory Q₁₀ values>1.0 (range=1.34-2.14) recorded across acclimation groups at test temperatures equivalent to acclimation temperature. Low acute respiratory Q₁₀ values in all acclimation groups across 15-20 °C indicated a limited capacity for thermal regulation of V?_O2 within this temperature range. The ability of P. perna to regulate O₂ uptake with progressive hypoxia was temperature-dependent, increasing from poor O₂ regulation at 10 °C to good regulation at 30 °C. The O₂ regulatory ability of P. perna and other open-water mytilids in declining O₂ concentrations does not greatly differ from that of estuarine heterodont bivalves, suggesting that it is not a major factor preventing open-water species, such as P. perna, from invading estuarine environments. However, P. perna's inability to regulate O₂ uptake at temperatures>25 °C combined with its relatively low upper thermal limit of 30 °C will likely prevent it from establishing permanent estuarine populations on Gulf of Mexico shores.     

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

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

First observations of predation by New Zealand Greenshell mussels (Perna canaliculus) on zooplankton

Observations made overseas of predation by blue mussels and zebra mussels on mesozooplankton (>200 μm) have raised concern within New Zealand that the Greenshell mussel, Perna canaliculus, which is cultured in large tonnages throughout hundreds of marine farms within the New Zealand coastal zone, could exert ecologically detrimental effects by preying on zooplankton. We conducted experiments at Clova Bay, Pelorus Sound in May 2002 to determine the rates that P. canaliculus ingests prey, up to and including the mesozooplankton size range. Single mussels from farms were incubated with seawater enriched with zooplankton (>60 μm) in gently circulated 15-l pails. Depletion of chlorophyll-a (chl-a), ciliate microzooplankton, and nauplii, copepodites, and adults of copepods was determined over 5 h, relative to controls with no mussels. Two experiments were made over consecutive days. Gut contents of these experimental mussels, and of mussels examined soon after collection from a farm, were described.Gut contents of experimental and of freshly collected mussels (standard shell length ∼90 mm) had numerous copepod parts, whole copepods and larval bivalves present. Experimental mussels cleared chl-a and ciliates from 59- to 137-l individual⁻¹ day⁻¹, respectively, averaged across the two experiments. Faster ciliate than chl-a clearance was probably caused by the high proportion (56%) of phytoplankton below the retention size for P. canaliculus (ca. 5 μm) and by faster ciliate grazing in controls than treatments. The average clearance rates of adult, copepodite, and naupliar copepod stages by mussels were 20, 31, and 49 l individual⁻¹ day⁻¹, respectively. The clearance rates of each copepod stage were not significantly different between the two experiments. Clearance of nauplii was significantly greater than of adults and copepodites, while adult and copepodite clearance rates were nearly significantly different. The mean lengths of the adult, copepodite, and naupliar copepods were 430, 265, and 165 μm, respectively. The decreasing clearance rates with increasing size and development of prey (from ciliates, through naupliar, copepodite to adult copepods), suggested that prey escape ability, related to body size and/or morphology, affected capture rates. Mussel faecal samples indicated complete digestion of the gut contents. Pseudofaecal samples showed very low rejection rates of mesozooplankton by mussels. The results are considered in context of current biophysical modelling studies of impacts of large mussel farms in New Zealand. Designs of future experiments to improve accuracy of estimates of mesozooplankton clearance rates by P. canaliculus are considered.     

Ecophysiological aspects of cardiac activity in the subtropical mussel Perna viridis (L.) (Bivalvia: Mytilidae)

Cardiac activity in the subtropical mytilid mussel Perna viridis was monitored to assess the influence of various endogenous and exogenous parameters. Temperature had a positive, significant, effect on heart rate; the Q₁₀ ranged from 2.5 to 2.6. Salinity, over a wide range (15-35‰), had no significant influence on heart rate reflecting the species' euryhalinity. Reduced oxygen tensions, either induced by emersion or seawater hypoxia, elicited rapid, significant, bradycardia. Positive inotropisms (indicative of greater heart output) were evident during seawater hypoxia and may represent an energetically advantageous mechanism of reducing heart rate while increasing stroke volume to maintain haemolymph circulation to organs associated with digestion and basal metabolism. Malnourishment for 14 days did not affect heart rate. The heart rate of larger mussels was slower than those of smaller individuals. Gender-based differences in P. viridis' heart rate were not present. No diurnal (24 h) variation in cardiac activity was evident while mussels were exposed to relatively stable, immersed conditions. Heart rate plasticity often reflected the ambient environmental conditions and such a flexible physiological response may partly explain the species' dominance in transitional, polluted harbours.     

Evidence of cannibalism and bentho-pelagic coupling within the life cycle of the mussel, Perna canaliculus

The feeding ecology of the green-lipped mussel, Perna canaliculus, was investigated within three intertidal mussel beds along Ninety Mile Beach, northern New Zealand, between August 2000 and March 2001. Adult mussels of different sizes (45-105 mm in shell length) were collected from the intertidal sites about 30 min after being submerged by the incoming tide for gut content analyses. Results of these analyses indicate that mussels consume a variety of phytoplankton, micro- and mesozooplankton, including mussel larvae and post-larvae. Cannibalism of juveniles of up to 620 μm was recorded for intertidal mussels, and conspecifics of up to 2.4 mm were found within the stomachs of additional mussels collected in August 2000 from a nearby subtidal site. For all three intertidal populations, mussel larvae and juveniles contribute about 70% of the food particle consumption during the spawning peak in August, while phytoplankton and other zooplankton constitute the majority of the food source (about 99%) in March, during gametogenesis. Larger intertidal mussels tended to have more food particles in their stomachs than smaller mussels within all three populations. Distinctive differences in food consumption among intertidal populations directly coincide with variations in total particulate matter (TPM), particulate organic matter (POM) and percent organic matter (OM) in the adjacent seawater.Separate experiments designed to test the feeding behavior of mussels feeding at different times during the incoming tide were conducted at one of the intertidal sites during August 2000 and March 2001. Results from these experiments indicate a marked shift in food consumption from bivalves to other mesozooplankton in August, and from phytoplankton to mesozooplankton in March. The observed combination of mussel predatory and grazing behavior over the incoming tide and through the year provides evidence for a strong food-web link between the benthic and pelagic life stages of this species. Furthermore, the high rate of cannibalism during some months of the year suggests that this source of food may significantly contribute to the energy budget of wild populations, with potential implications for evolutionary adaptive success.     

Effects of seston variability on the clearance rate and absorption efficiency of the mussels Aulacomya maoriana, Mytilus galloprovincialis and Perna canaliculus from New Zealand

The clearance rates (CR in l g⁻¹ h⁻¹) and net absorption efficiencies (AE) of three co-occurring mytilid species were estimated to determine the effects of variation in ambient seston quantity (total particulate matter, TPM, mg l⁻¹) and quality (particulate organic matter, POM, mg l⁻¹; percent organic matter, PCOM=POM/TPM) on these physiological functions. CR_S estimates were significantly different among the species (Mytilus galloprovincialis>Perna canaliculus=Aulacomya maoriana), but were not correlated with differences in species-dependent mean body size (P. canaliculus>M. galloprovincialis>A. maoriana). AE estimates were independent of mean body weight, and did not differ among the species. For all three species, CR responded in a simple linear manner to seston organic content, either in terms of POM (for A. maoriana and M. galloprovincialis), or PCOM (for P. canaliculus). The AE response, although also of a simple linear type, was species-dependent and involved interactions of two or three seston components: TPM, POM and PCOM for A. maoriana; POM and PCOM for M. galloprovincialis; and TPM and PCOM for P. canaliculus. For all three species, seston variation explained 15-20% of the variation in CR and 52-59% of the variation in AE. Comparisons among the species indicated that two very different responses to variation in seston quantity and quality exist. First, significant differences in CR result from species-dependent differences in the magnitude of the response to seston variation. Second, species-dependent responses to variation in seston variation resulted in the similarity of the AE responses. Thus, the three species appear to have evolved different strategies for dealing with seston variation, but with the end result that their AE responses do not differ. Finally, no evidence was found of a negative association between CR and AE for any of the three species, suggesting that under the seston conditions experienced in this work, these species do not have the physiological compensatory capacity to reduce CR in order to increase AE. The importance of a comparative (i.e., multi-species) approach utilising ambient seston is emphasised by the findings of this research if we are to understand better the feeding and digestive physiologies of suspension-feeding organisms such as mussels.     

Combined effects of food quantity and quality on Cd, Cr, and Zn assimilation to the green mussel, Perna viridis

We examined the assimilation of Cd, Cr, and Zn by the green mussel Perna viridis under complicated food conditions, including combinations of different compositions and concentrations of food (diatom and sediment), and variable food quantity and quality during particle digestion. At different combinations of food composition and quantity (5 mg l⁻¹ and 20 mg⁻¹, below and above the pseudofeces production), the Cd assimilation was significantly dependent on the food composition. The Cd assimilation efficiency (AE) decreased with increasing proportions of sediments in the diets, but its assimilation was not significantly affected by food concentration. In contrast, the assimilation of Cr and Zn decreased significantly with increasing food concentrations, whereas food composition did not significantly affect their AEs. Variations in metal gut passage time accounted partially for the difference in AEs among different combinations of food composition and quantity. By changing the type of particles during metal digestion, their AEs were maintained comparably at a low particle load (1 mg l⁻¹), suggesting that variation of food quality during digestion did not affect metal assimilation. At a higher particle load (5 mg l⁻¹), variation of food type during digestion affected the AEs of Cr and Zn. An increase in food concentrations from 1 to 15 mg l⁻¹ during digestion resulted in a significant decrease in the AEs of Cr and Zn bound with either sediments or diatoms. Conversely, decreasing the food concentrations from 15 to 1 mg l⁻¹ did not affect the AEs of metals, except for Zn bound with diatoms. Overall, our results highlighted the metal-specificity in their assimilation as influenced by complicated food environments, probably caused by different metal geochemical and biological behavior in the mussels. Feeding selectivity may have a greater control on the influx rate into the mussels than metal assimilation.     

Spatial and temporal patterns of recruitment of the tunicate Ciona intestinalis on a mussel farm in Nova Scotia, Canada

Although possibly indigenous to Nova Scotia, the ascidian Ciona intestinalis has become a problem for local mussel aquaculturists. The local population ecology of the ascidian was studied by recording the depth, distribution, and timing of recruitment of C. intestinalis at four different locations (high wave exposure, low wave exposure, recently fallowed, and historically heavily fouled with tunicates) on a mussel farm at Indian Point, Nova Scotia. Animals were collected weekly from 11 collectors deployed at a depth of 4.5 m over two seasons (June-December 2003 and 2004). Recruitment occurred in two peaks during both seasons: the first peak lasted from late June to late July while the second peak lasted from early September to mid-November. Recruitment was highest in the sheltered and historically fouled location, intermediate in the fallowed site, and lowest in the site with highest wave exposure. Collectors were also deployed at different depths. Recruitment was highest at 4.5 m and to a lesser extent at 8.5 m. Little recruitment was seen at the surface (0.5 m) or in deeper water (12.5 and 16.5 m), except during the second peak in 2004. Overall, these results seem to indicate an abbreviated larval dispersal phase with young recruits settling very close to the pool of adults located on the mussel lines.     

Tidal rates of settlement of the intertidal barnacles Chthamalus stellatus and Chthamalus montagui in western Europe: the influence of the night/day cycle

Chthamalus montagui and Chthamalus stellatus are abundant barnacles in western Europe. Tidal settlement of Chthamalus in SW Ireland and SW Portugal was studied in relation to a night and day factor and at different temporal (dates) and spatial (shores and sites) scales. Based on the identifiable cyprids and metamorphs, Chthamalus settlement in SW Ireland was comprised mainly of C. stellatus but was composed of C. montagui only in SW Portugal.In SW Ireland and SW Portugal, settlement rates of Chthamalus (mean number of settlers per 25 cm²±S.E.) were higher after one day tidal cycle (597±158.7 in SW Ireland, 144±23.6 in SW Portugal) than one night tidal cycle (55±12.1 in SW Ireland, 13±2.2 in SW Portugal), but significant differences were only detected in SW Portugal. Different models were proposed for explaining this pattern related to night and day variability of the physical processes responsible for transporting cyprids to shore (1), and/or of pre-settlement behaviour (2) and/or of settlement behaviour of cyprids (3).Spatial patterns of tidal settlement of both species or at both locations seem similar with small scale variability (between sites, 5 to 30 m apart) in settlement of Chthamalus being the only spatial scale at which variability was detected in both locations.The composition of Chthamalus cohorts settling during one tidal cycle differed considerably between locations/species: settlers of C. stellatus were mainly cyprids; settlers of C. montagui during the day (when most settlement occurred) were essentially metamorphs.     

Spatial and temporal variability among marine Bacteroidetes populations in the NW Mediterranean Sea

The abundance and structure of Bacteroidetes populations at diverse temporal and spatial scales were investigated in the Northwestern Mediterranean Sea. At a temporal scale, their relative abundance exhibited a marked seasonality, since it was higher in spring and decreased in winter. Similarly, Bacteroidetes community structure encompassed three main groups (winter, spring and summer-fall), which mimicked global bacterioplankton seasonality. At the spatial scale, relative abundances were similar in all surface samples along an inshore–offshore transect, but they decreased with depth. Analysis of the community structure identified four markedly different groups mostly related to different depths. Interestingly, seasonal changes in abundance and community structure were not synchronized. Furthermore, richness was higher when Bacteroidetes were less abundant. The variability of Bacteroidetes contributions to community structure in the temporal and spatial scales was correlated with different environmental factors: day length was the most important factor at the temporal scale, and salinity at the spatial scale. The community composition in terms of phylotypes changed significantly over time and along the depth gradients, but season or depth-specific phylogenetic clusters were not identified. Delineation of coherent Bacteroidetes sub-clusters should help to uncover higher resolution patterns within Bacteroidetes, and explore associations with environmental and biological variables.     

退化草地阿尔泰针茅与狼毒种群的小尺度种间空间关联

采用草地群落学调查与点格局分析方法,在祁连山北坡选择4种退化高寒草地群落,分析了阿尔泰针茅(Stipa krylovii)与狼毒(Stellera chamaejasme)种群的大小结构、斑块特征和种间关联关系。结果表明:随天然草地退化过程延续,阿尔泰针茅由大株丛结构演变为小株丛结构,种群密度和领地面积减小、空斑面积增大、领地密度先增大后减小,狼毒种群的株丛结构和斑块特征发生了相反的变化趋势;不同退化草地阿尔泰针茅和狼毒种群的空间关联呈现负关联、正关联和不关联规律;在未退化草地和轻度退化草地、中度退化草地,阿尔泰针茅与狼毒的关联性分别由0-14 cm、0-51 cm尺度上的负相关和0-85 cm尺度上的不相关,转变为14-100 cm、51-100 cm尺度上的不相关和86-100 cm尺度上的正相关。物种个体大小结构变化,以及狼毒种群的斑块吞并、合并和阿尔泰针茅种群的斑块破碎、被分割过程,既是物种关联性发生尺度转换的先决条件,又是引起草地群落中物种地位与作用改变的关键因素。     

Spatial variation in the recruitment of the intertidal barnacles Chthamalus montagui Southward and Chthamalus stellatus (Poli) (Crustacea: Cirripedia) over an European scale

Spatial variation in the recruitment of the intertidal barnacles Chthamalus montagui and Chthamalus stellatus was examined over an European scale. The study was carried out using standardised protocols at a series of locations. The five locations chosen (SW Ireland, NW Spain, SW Portugal and NW and NE Italy) span a large part of the range of these species in Europe. The spatial scales were location (hundreds of kilometres) and shore (thousands of metres).Estimates of total cumulative recruitment (cyprids and metamorphs) summed over the year (April 1997 to March 1998) showed substantial variation between locations which was dependent on the species. Recruitment was highest in SW Portugal for C. montagui and in SW Ireland and NW Spain for C. stellatus. Overall recruitment of C. montagui was higher than that of C. stellatus at all locations except SW Ireland, where recruitment of the two species was not significantly different. There were significant differences among shores in each location.The recruitment period of both species varied with location, with recruitment beginning earlier further south. In general, recruitment of C. montagui and C. stellatus was recorded in 8 months in NW Spain and NE Italy, while only in 7 months in SW Ireland. Recruitment of C. montagui occurred in 10 months in SW Portugal, but no recruits of C. stellatus were found. In all locations there was at least one distinct peak of recruitment. In SW Ireland both species showed only one peak of recruitment, a month after initiation. At the more southerly Atlantic locations, as well as in the Mediterranean, two unequal peaks of recruitment were generally seen. During recruit census, the number of cyprids, in comparison to metamorphs, found at any location was very low. In SW Ireland and NW Spain cyprids of both species were found, while in SW Portugal and in the Mediterranean, only cyprids of C. montagui were found.     

In situ enclosure experiments on the influence of cultured mussels (Perna canaliculus) on phytoplankton at times of high and low ambient nitrogen

The influence of the cultured mussel Perna canaliculus (Gmelin 1791) on the abundance of phytoplankton was investigated in Pelorus Sound, New Zealand. Four in situ enclosure experiments were undertaken, two in summer when ambient nitrogen was low, and two in winter when it was high. Each experiment had four manipulation types: added mussels; added nitrogen; both mussels and nitrogen added; and control (no additions). In summer, there was a significant increase of chlorophyll a in response to added nitrogen, indicating that the phytoplankton were nitrogen-limited. At this time, mussels caused an increase (11-17%) in phytoplankton biomass, possibly by converting particulate nitrogen to ammonium, making the nitrogen available for phytoplankton utilisation. The highest ambient chlorophyll a concentrations coincided with high ambient nitrogen in the winter. At this time, mussel grazing caused a significant decrease (5-14%) in phytoplankton concentration, indicating that within-farm depletion of phytoplankton is most likely to occur in winter. On an annual time scale, the mussels had a stabilising influence on phytoplankton biomass, reducing high ambient levels in winter and slightly increasing low levels in summer.     

Surface wettability as a determinant in the settlement of the barnacle Balanus Improvisus (DARWIN)

Several studies have shown that the initial surface wettability, is of importance in the settlement of macrofouling larvae such as barnacles, bryozoans and hydroids in the field as well as in laboratory assays. In this study we present results from laboratory assays using hydrophilic and hydrophobic polystyrene (PS) and cyprid larvae of Balanus improvisus (Darwin). The results obtained differ markedly from those reported for the barnacle Balanus amphitrite (Darwin), where a high surface wettability seemed to be preferred for settlement. Our results show that a surface with intermediary wettability (hydrophilic PS) reduced settlement by 38% as compared to surfaces of low wettability (hydrophobic PS) during an 8-day period. During the experiment, the wettability in the hydrophilic PS dishes was not significantly changed as measured by advancing contact angle with mQ water. Over an 8-day period wettability of the hydrophobic PS dishes approached that of the hydrophilic PS surfaces. We further conducted experiments with highly hydrophilic and highly hydrophobic methylsilane-treated glass surfaces with known chemistry. In this experiment, the settlement of cyprid larvae was completely inhibited by the high wettability surfaces. Contact angle measurements revealed that the wettability during the length of the experiment of the hydrophilic glass surfaces was not significantly altered. We conclude by these experiments that even an intermediate wettability can significantly affect the overall settlement success of the barnacle B. improvisus. The mechanism by which the settlement is impeded might be biologically mediated through the recognition by cyprid larvae of the molecular composition of the surface when the cyprid reverts to the settlement phase, i.e. when swimming behaviour is abandoned in favour of surface exploration, or it is mediated by physicochemical forces acting between the surface and the larval body or the larval antennules.     

Resilience and elasticity of intertidal communities in response to harvesting of the alien mussel Mytilus galloprovincialis

Besides the direct impacts of exploitation on target species, indirect effects on non-target species are unavoidable and find expression in changes in community structure. We quantified the effects of experimental harvesting of Mytilus galloprovincialis on intertidal communities on the South African west coast. In the mid- and low-shore, four months of harvesting at intensities greater than F = 0.3 and F = 0.6 respectively, resulted in significant changes in community composition. These changes were driven by progressively greater spatial dominance by the macroalgae Cladophora flagelliformis, Porphyra capensis and Ulva species as harvesting intensity increased. Four months after cessation of harvesting, community structure had not recovered and even areas subjected to as little as F = 0.3 supported significantly altered communities in both zones. The fact that substantial community changes were induced by even low-intensity exploitation is indicative of low resilience to harvesting. The densities and cover of the dominant taxa returned to a pre-harvest state 16 months after the cessation of harvesting, but multivariate analyses indicated that the overall community composition required 32 months for Cover Letter full recovery. Although these communities displayed elasticity within three years, it is recommended that should a M. galloprovincialis fishery be established in the region, harvesting be implemented at a maximum intensity of F = 0.3. This approach would maximise yields and protect associated intertidal communities.     

Assessing the potential benthic-pelagic coupling in episodic blue mussel (Mytilus edulis) settlement events within eelgrass (Zostera marina) communities

Coastal marine seagrass ecosystems are important nursery grounds for commercially and recreationally important species, and they serve as key settlement and recruitment sites for other species. We investigated several years (2001-2003) where episodic settlement events of blue mussels (Mytilus edulis) occurred in Barnegat Bay, NJ, USA. Population assessment indicated that blue mussels settled in eelgrass beds (Zostera marina) in late spring with peak densities exceeding 170,000 m⁻². Based on calculated filtration rates of M. edulis, we determined that for at least 53 days in 2001, the density and size distribution of M. edulis were sufficient to filter the water column volume in excess of twice a day, with maximum calculated filtration rates exceeding 8 m³ water m⁻² day⁻¹. While the settlement event in 2001 was very localized, in 2003, the settlement event was considerably more widespread throughout the bay, with maximum settling densities exceeding 175,000 individuals m⁻². Associated with these high densities, maximum calculated filtration rates exceeded 15 m³ water m⁻² day⁻¹. This filtration potential may have impeded the localized development of a brown-tide (Aureococcus anophagefferens) bloom in 2001, which occurred in other regions of the bay, but the widespread settlement event seen in 2003 may have impeded the development of any brown-tide blooms in Barnegat Bay during that summer. The decline in mussel densities throughout the summer may be a result of elevated water temperatures in this back bay, but at one site, the high settlement of M. edulis was followed by a substantial migration (>40 individuals m⁻²) of small sea stars (Asterias forbesii). In 2001, A. forbesii was a significant factor in reducing M. edulis density by the end of the summer at the Barnegat Inlet site and a community level assessment showed significant positive correlations between mussel aggregations and sea star densities (r=0.68-0.73, P<0.001). At this same site in 2003, the sea stars were again present in high densities (26 m⁻²) and were a potential mechanism for mussel decline. In other regions of the bay, sea star densities are very low, but numerous other predatory species exist, including blue crabs (Callinectes sapidus), green crabs (Carcinus maenus), spider crabs (Libinia spp.), and several Xanthid crabs. Given the high mussel densities seen in this study and the considerable predation by sea stars and other benthic predators, the benthic-pelagic coupling which these mussels provide in this system contributes to the high secondary production in these grass beds.