Interactive effects of wave exposure and tidal height on population structure of the mussel Perna perna

The influence of wave exposure and of tidal height on mussel (Perna perna Linnaeus) population structure (size, density, biomass and adult/juvenile correlations) was examined at 18 sites along the south coast of South Africa. Sites were classified as exposed or sheltered prior to sampling, without reference to the biota, on the basis of aspect, topography and wave regime. A single set of samples was collected from each site during three spring tide cycles. Adult mussels on these shores almost always attach directly to the rocks, and layering of mussels is virtually absent. Shore height always had a strong influence on population structure, but exposure had significant effects only lower on the shore, and almost exclusively on mussel sizes. Principal component analysis (PCA), based on size distribution data for each population, revealed a general upshore decrease in the modal size of the adult cohort. The effects of exposure on size distribution, however, varied with tidal height. PCA separated exposed zones, with larger mussels, from sheltered zones on the low-shore. Farther upshore the two shore types were increasingly confounded. The maximum size of mussels showed a similar pattern, with significant differences (ANOVA, p < 0.05) between exposed and sheltered sites only on the low- and mid-shores. Density was calculated from randomly placed quadrats (i.e. not necessarily from areas of 100% cover) and showed a different pattern. Adult (>15 mm) densities decreased up the shore, with low-, mid- and high-shore zones being significantly different from one another (ANOVA, p < 0.0001; followed by multiple range tests). However, exposure had no significant effect on density, nor was there a significant interaction with zone. Recruit (<15 mm) densities were positively correlated with adult (>15 mm) densities in all zones and for both exposure regimes ( p < 0.05 in all cases), but there was considerable variability and extremely low predictability in these relationships (r ² generally <0.2). Predictability tended to be greater towards the high-shore, where adults were more clumped. As with density, biomass was not affected by exposure, but decreased upshore as mean size and density decreased. A reduction in the influence of exposure farther upshore may be caused by greater emersion overriding the effects of exposure. The presence of free space within mussel beds and significant correlations between recruit and adult densities suggest that these mussel populations are recruit limited. Received: 7 January 2000 / Accepted: 6 July 2000     

Influence of wave exposure on South African mussel beds and their associated infaunal communities

This study investigates effects of wave exposure on beds of the mussel Mytilus galloprovincialis and the infaunal communities associated with them. Nine sites of varying wave exposure were sampled near Groenrivier, on the west coast of South Africa. Wave exposure was measured using a Palumbi device, and exposure ranged from 7.17×10³ to 18.49×10³ N m^–2. Percentage mussel cover at each site was measured using 50×50 cm quadrats, and three 10×10 cm samples were removed from the mussel beds in the mid-intertidal zones of each site for examination of infauna. Percentage mussel cover, mean mussel length, mussel biomass and mussel bed depth all peaked at intermediate exposures, declining towards both the most sheltered and most exposed sites. Infaunal species diversity and richness both showed the reverse trend, peaking at the most sheltered and most exposed sites, and declining at intermediate exposures. Neither infaunal abundance, nor biomass, was significantly correlated to wave exposure, although abundance was highest at sheltered sites and biomass greatest at the most exposed site, which was dominated by limpets and large robust polychaetes.Communicated by J.P. Thorpe, Port Erin     

A comparison of two cartographic exposure methods using Fucus vesiculosus as an indicator

Morphological variation and vertical distribution of Fucus vesiculosus were quantified at several sites in the Finnish archipelago (Baltic Sea). F. vesiculosus samples were obtained from skerries at geographical distances of 1 km or more (large scale) and at intervals of ca 100 m around a single island (small scale). The results were examined in relation to wave exposure, calculated by Baardseth and effective fetch cartographic methods. Despite the fact that the exposure indices were calculated differently they correlated strongly. Vegetative morphological characteristics of F. vesiculosus illustrate the morphological differences both within and between exposure gradients. The tallest and widest F. vesiculosus plants were found at the sheltered end of the large-scale exposure gradient. Those from equally sheltered sites of the island were smaller in all respects. Thus, the trend from small narrow plants to large wide sheltered plants was expressed differently over the different geographical scales. Consequently localities with similar exposure indices may have morphologically different F. vesiculosus populations. Shores with similar cartographic exposure indices can be different in nature. Underwater topography and shore locations, either close to the mainland or at the outermost sites of the archipelago, affect the exposure. Although a sheltered shore is indicated, the sublittoral zone may be quite exposed to the movements of water. In contrast, in an open shore environment underwater rocks, boulders and shallow water areas can provide sheltered habitats. The depth range of the F. vesiculosus belt exhibited two distinctive patterns. At sheltered sites, around islands in the outermost reaches of the archipelago F. vesiculosus can grow to a maximum depth of 5 m. In exposed habitats the belt becomes narrower, reaching a maximum depth of 3 m. Closer to the mainland F. vesiculosus is found at exposed sites to a maximum depth of 5 m; the depth range at sheltered sites is narrower, only reaching depths of 2 m or less. In conclusion, the changes in plant morphology and in the vertical belt distribution are similar to each other along both gradients at the exposed ends of the wave action spectrum; however, the two gradients diverge at the sheltered ends of the spectrum. Received: 10 August 1998 / Accepted: 11 January 1999     

Genetic structure of hybrid mussel populations in the west of Ireland: two hypotheses revisited

In Ireland, mussels on exposed rocky shores constitute an interbreeding mixture of two forms of mussels, the blue mussel, Mytilus edulis, and the Mediterranean mussel, M. galloprovincialis. Results from an Irish study in the 1980s, using partially diagnostic allozyme markers, indicated that mussels higher up the shore were more galloprovincialis-like than those lower down. In this study we set out to test two hypotheses: (a) recruits arriving on the shore are composed of genetically distinct cohorts that settle preferentially at different levels on the shore, and maintain genetic distinctiveness into adulthood; (b) recruits are genetically homogeneous, but once settled they diverge genetically over time, due to within-habitat site specific-selection. The diagnostic Me 15/16 DNA marker was used to analyse the genetic composition of newly-settled spat recruiting to artificial substrates, which were placed at two-week intervals from May–October 2002, on the mid- and low shore areas of two exposed sites in Galway Bay. Adult mussels were also collected on each sampling date. Results did not support the preferential settlement hypothesis, i.e., the genetic composition of primary settlers (≤ 500 μm) was similar between tidal heights and shores. Neither was there evidence of post settlement selective mortality, as adults were genetically similar to settling spat. In spat and adults the frequency of the M. galloprovincialis allele was high (0.56–0.80), due to high frequencies of M. galloprovincialis (> 37%) and hybrid (> 33%) genotypes, and correspondingly low frequencies of the M. edulis genotype (< 11%). Adult mussels from a nearby sheltered estuarine site, while significantly different to exposed shore mussels, still had low frequencies of the M. edulis genotype (< 17%), indicating no apparent advantage for the genotype in this environment. There are indications that the genetic composition of mussels may be changing on the Atlantic coasts of Ireland.     

Abundance,population structure and claw morphology of the semi-terrestrial crab <Emphasis Type="Italic">Pachygrapsus marmoratus</Emphasis> (Fabricius, 1787) on shores of differing wave exposure

Wave action is known to influence the abundance and distribution of intertidal organisms. Wave action will also determine the duration and suitability of various foraging windows (high-tide and low-tide, day and night) for predation and can also affect predator behaviour, both directly by impeding prey handling and indirectly by influencing prey abundance. It remains uncertain whether semi-terrestrial mobile predators such as crabs which can access intertidal prey during emersion when the effects of wave action are minimal, are influenced by exposure. Here, we assessed the effect of wave action on the abundance and population structure (size and gender) of the semi-terrestrial intertidal crab Pachygrapsus marmoratus on rocky shores in Portugal. The activity of P. marmoratus with the tidal cycle on sheltered and exposed shores was established using baited pots at high-tide to examine whether there was activity during intertidal immersion and by low-tide searches. Because prey abundance varies along a wave exposure gradient on most Portuguese shores and because morphology of crab chelipeds are known to be related to diet composition, we further tested the hypothesis that predator stomach contents reflected differences in prey abundance along the horizontal gradient in wave exposure and that this would be correlated with the crab cheliped morphology. Thus, we examined phenotypic variation in P. marmoratus chelipeds across shores of differing exposure to wave action. P. marmoratus was only active during low-tide. Patterns of abundance and population structure of crabs did not vary with exposure to wave action. Stomach contents, however, varied significantly between shores of differing exposure with a higher consumption of hard-shelled prey (mussels) on exposed locations, where this type of prey is more abundant, and a higher consumption of barnacles on sheltered shores. Multivariate geometric analysis of crab claws showed that claws were significantly larger on exposed shores. There was a significant correlation between animals with larger claws and the abundance of mussels in their stomach. Variation in cheliped size may have resulted from differing food availability on sheltered and exposed shores.     

Larval supply and juvenile recruitment of coral reef fishes to marine reserves and non-reserves of the upper Florida Keys,USA

For marine organisms, decoupling between the planktonic larval stage and the benthic-associated juvenile stage can lead to variable patterns of population replenishment, which have the potential to influence the effectiveness of marine reserves. We measured spatial and temporal variability in larval supply and recruitment of fishes to coral reefs of different protection levels and tested whether protection level influenced the relationship between supply and recruitment. We sampled pre-settlement larvae and newly settled recruits from four reefs (two reserves and two non-reserves) in the Florida Keys National Marine Sanctuary, USA. Replicate point measures of larval supply over 14 months and 17 monthly measurements of recruitment varied significantly among months and sites. Sites with the same protection level had significantly different patterns of larval supply as well as larval and recruit diversity, but recruitment magnitude differed only by protection level, where densities were greater at reserves. Differences in larval supply among sites included two particularly large peaks in larval abundance at one site, possibly associated with the observed passage of small-scale oceanographic features. To examine whether relationships between larval supply and recruitment varied by protection level, we selected one species that was present in both the light trap samples and the monthly recruitment surveys. Recruitment of the bicolor damselfish Stegastes partitus was significantly and positively related to larval supply at three of the four sites thus, protection level did not influence this linkage. Since local variability among sites can lead to spatial differences in population replenishment, characterization of larval supply and recruitment to potential marine reserve sites may help to identify optimal locations in a region and contribute to more effective reserve design.     

Distribution of a marginal population of <Emphasis Type="Italic">Mytilus edulis</Emphasis>: responses to biotic and abiotic processes at different spatial scales

Physical and biological processes interact to produce pattern in nature. Pattern is scale dependent as processes generating pattern are heterogeneous in time and space. We tested some causes of variation in abundance and distribution of three marginal populations of sublittoral blue mussels, Mytilus edulis, in the non-tidal northeastern Baltic Sea. We studied the role of substrate inclination, perennial algae and siltation along local wave exposure gradients on mussel distribution over a regional salinity gradient. We found marked differences on regional scales (p < 0.001) with lower densities and biomasses of mussels with declining salinity. Along local gradients, mussel densities increased with increasing exposure (p < 0.001) and declining slope and sedimentation (p < 0.01). Site specifically, densities of blue mussels and the perennial red algae, Furcellaria lumbricalis, were positively related, results supported by a colonisation experiment. Also, young post-recruits showed significant relations to adult biomass, wave exposure, algal biomass, bottom slope and sediment cover. Findings showed that the relative importance of the determinants affecting blue mussels at the edge of their range vary with scale and are affected by the density and size structure of mussel populations. The study provides an indication of the types of factors that may be invoked as causes of spatial variation in marginal blue mussel populations and reinforces the need to consider multiple aspects when distributional patterns are assessed.     

Local and meso-scale patterns of recruitment and abundance of two intertidal crab species that compete for refuges

Interference competition for limited habitat or refuges is known to produce density-dependent mortality and generate patterns of micro-habitat distribution. While in mobile species the outcome of interference at a local scale can usually be determined from differences in body size and behavior, the population-level consequences of such interactions vary depending on rates of settlement and recruitment at a site, which are not directly correlated to local reproductive success. Previous experimental studies in central Chile demonstrated that interference competition for refuges is the primary factor driving microhabitat segregation between the predatory crabs Acanthocyclus gayi and Acanthocyclus hassleri, with the latter species monopolizing galleries inside mussel beds and excluding A. gayi to rock crevices. Between April 2001 and March 2006 we quantified monthly recruitment rates in artificial collectors at 17 sites over 900 km of the central coast of Chile. Results show that recruitment rates of A. hassleri are almost two orders of magnitude lower than those of A. gayi, and that they are tightly and positively correlated among sites across the region, suggesting that at scales of kilometers larval stages of these species are affected by similar oceanographic processes. Total crab densities per site were also positively correlated between species and strongly associated to mussel cover, with overall low crab densities at all sites where mussel cover was lower than about 60%. At all sites with mussel cover >60%, the ratio of A. gayi to A. hassleri density progressively decreased from recruits (2.6) to juveniles (0.5) to adults (0.04), overcoming initial differences in recruitment rates. The relative success of the inferior competitor at sites with low mussel cover does not appear to provide a potential mechanism favoring regional coexistence through dispersal to other sites (“mass effects”), because their densities were lower than at sites of high mussel cover. Yet, at many sites of low mussel cover the dominant competitor is virtually absent, allowing A. gayi to attain larger population sizes at the scale of the region. Thus, the factors limiting the dominant competitor from successfully utilizing other microhabitats seem to be the most critical factor in promoting both local and regional coexistence between these species.     

Population structure,northern range limit,and recruitment variation in the intertidal limpet Lottia scabra

The northern range limit of the intertidal limpet Lottia scabra is Cape Arago, Oregon (43°N), where adult survival is excellent, the population is small (<300), and recruitment is low; the range limit may be set by limited recruitment. Between June 2012 and March 2013, 25 sites from the middle of the species range (33°N) to Cape Arago were sampled and population size frequency distributions, densities, and nearest neighbor distances were compared to the amount of rocky and sandy shore and kelp bed size. North and south of 37°N, the densities of new recruits averaged 22 and 86 m^?2, respectively. This shift was associated with the range limit of Macrocystis pyrifera kelp beds; we hypothesize that slower currents in M. pyrifera beds may limit larval dispersal leading to higher recruitment. North and south of 40°N, adult density averaged <1 and 458 m^?2, respectively, with the species absent from many sites to the north. This shift was associated with a sharp drop in the amount of rocky shoreline and an increase in uninhabitable sandy shore. Near the northern range limit, >80 % of the individuals were solitary and may be unable to spawn successfully. Recruitment at Cape Arago was infrequent and likely due to self-recruitment. This study suggests that the range limit was set by the absence of M. pyrifera and too little rocky shore leading to high larval wastage, low settlement, low population densities, and, due to an Allee effect, very small effective population sizes.     

Ecological studies on intertidal New Zealand Sphaeromatidae (Isopoda: Flabellifera)

Ten sympatric rocky shore species and 3 brackish-water species of Sphaeromatidae were studied. Rocky shores were classified in terms of substrate and exposure to wave action. Although most of the rocky-shore species occur on all shore classes, vertical and horizontal distribution in each species, as well as densities, vary with substrate and wave action. A close relationship between distribution and breeding exists in the rocky-shore species; maximum differences in the vertical distribution of juveniles and adults occur, together with the highest reproductive capacities, in the species exposed to the greatest wave action. Tolerances to temperature, salinity and desiccation are wider in the species of brackish water and less exposed rocky shores, correlating with the greater variations in these conditions where wave action is reduced. Exoskeletons and pereopods are stouter in the species of more exposed shores, except where the microhabitat affords protection from wave action. In Isocladus armatus, geographic and seasonal variations in relative frequencies of colour forms correlate with differences in environmental temperatures. Under laboratory conditions, the colour forms show differences in tolerances to temperature and salinity.This work was carried out in the Zoology Department, University of Canterbury, New Zealand; the text was completed in Zoologisches Institut der Universität Kiel, Germany.     

Growth and reproductive rates of Littorina rudis from three contrasted shores in North Wales,UK

Data are presented on growth and reproductive rates of Littorina rudis Maton from 3 contrasted habitats: a sheltered saltmarsh, a moderately sheltered boulder shore and an exposed cliff face. Growth was fastest in L. rudis from the moderately sheltered boulder shore, followed closely by snails from the sheltered saltmarsh. Growth was much slower and the asymptotic size much less in L. rudis from the exposed cliff face, this being attributed to the reduced time available for foraging caused by longer and more erratic periods when the substratum dries out. Eggs extracted from the brood chambers of the saltmarsh L. rudis took about 72 d to hatch in the laboratory at 10°C. No seasonal trend was detected in the percentage of sexually mature snails, which fluctuated erratically round about 54 to 99% in each population. Similarly, the ratio of females: males fluctuated erratically about 1.0. The number of eggs and embryos held in the brood chamber peaked in May–June and birth rates (release of young from the brood chamber) peaked in July-August. Following a lull in August, the brood chamber contents were maintained at moderately high levels throughout the winter, whereas the birth rates declined to very low levels. These differences in the seasonal cycle of brood chamber contents and birth rates were probably caused by seasonal differences in embryological development rate and in the propensity of the young to leave the brood chamber. Fecundity increased with parental size and, because of this, the smaller, slower growing exposed-shore L. rudis had much lower potential fecundities than L. rudis from the other shores. However, the size-specific fecundity of the exposedshore L. rudis was about 1.7 times that of L. rudis on the more sheltered shores. This difference was apparently not attributable to smaller young or to a greater capacity of the brood chamber in the exposedshore L. rudis, and must therefore have been caused by higher rates of egg production. 18% of mature females from the boulder shore and 5% of those from the cliff face were oviparous, having a jelly gland in place of the brood chamber.     

Habitat modification affects recruitment of abalone in central New Zealand

The habitat experienced during early life-history stages can determine the number and quality of individuals that recruit to adult populations. In a field experiment, biogenic habitat complexity was manipulated (presence or absence of foliose macroalgae) at two depths (2–3 m and 5–6 m) and the habitat-dependent effects on recruitment of the black foot abalone (Haliotis iris) were examined at three field sites along the south coast of Wellington, New Zealand (41°20′S, 174°47′E), between July and November 2005. Recruit density (<5 weeks post-settlement) was measured on cobbles covered with crustose coralline algae. Habitats of low complexity (barrens treatments) had consistently greater densities of recruits than habitats of high complexity (algae treatments). However, recruits in algae habitats were larger, and for deep habitats, there was greater survival in algae habitats compared with barrens habitats. While depth had no significant effect on early recruit (<2 weeks post-settlement) density, late recruit (<5 weeks post-settlement) density was greater in shallow habitats, and so it seems recruit survival was greater in shallow habitats. In this experiment, algal habitat complexity had strong effects on early recruit abundance, but habitat-dependent variations in recruit growth and survival may modify initial patterns of abundance and determine recruitment to adult abalone populations.     

Mechanisms of habitat segregation between an invasive (Mytilus galloprovincialis) and an indigenous (Perna perna) mussel: adult growth and mortality

The invasive mussel Mytilus galloprovincialis and the indigenous mussel Perna perna coexist intertidally on the south coast of South Africa through partial vertical habitat segregation: M. galloprovincialis dominates the upper shore and P. perna the lower shore. Recruitment patterns can explain the zonation of P. perna, but not the invasive species. We examined the role of post-recruitment interactions by measuring spatial and temporal differences in adult growth and mortality rates of the two species. Specifically, we tested the hypothesis that interspecific differences in growth and mortality reflect adult distribution patterns. The two study locations, Plettenberg Bay and Tsitsikamma, are 70 km apart with two sites (separated by 300–400 m) per location, each divided into three vertical zones. Growth was measured seasonally using different marking methods in 2001 and 2003. Cumulative adult mortality was measured through summer in 2003/2004. Both species generally grew more slowly upshore, but they showed different effects of season. For P. perna, growth was significantly reduced in winter in the low zone, but unaffected by season in the high zone. For M. galloprovincialis, growth was either unaffected by season or increased in winter, even in the high zone. Thus, growth of P. perna and M. galloprovincialis was reduced under cool winter and warm summer temperatures, respectively; and while growth was more similar between species in summer, M. galloprovincialis grew much faster than P. perna in winter. Mortality of P. perna increased upshore. For M. galloprovincialis, mortality was not zone-dependent and was significantly greater than for P. perna on the low-shore and (generally) across the shore in Tsitsikamma. Both species had higher growth and mortality rates in Plettenberg Bay than in Tsitsikamma. Thus, P. perna seems able to maintain spatial dominance on the low-shore and at certain sites because of higher mortality of M. galloprovincialis. We conclude that seasonality in growth of the two species reflects their biogeographic affinities and that coexistence is possible through pre-recruitment effects that limit the vertical distribution of P. perna and post-recruitment effects that limit M. galloprovincialis.     

Patterns of mussel recruitment in southern Africa: a caution about using artificial substrata to approximate natural recruitment

Quantifying sessile marine invertebrate recruitment often requires destructive sampling or extrapolation from artificial substrata, the latter introducing the danger of artifacts. We measured intertidal mussel recruitment into mussel beds and into brushes at three-month intervals for five years across 3,200 km of southern Africa and determined substrata effects on recruitment rate. Recruitment into mussel beds showed a strong, coast-wide gradient, with high recruitment on the West coast, diminishing on the South coast, and increasing slightly on the East coast. At scales of 10 s of km, brushes reflected natural temporal recruitment variability, with a strong significant linear correlation between recruitment into brushes and into mussel beds. However, the relationship became semi-logarithmic when comparing among locations at a scale of 100 s of km. Artificial substrata thus reflect local natural settlement well but may be a poor indicator of it when spatial scales are large, particularly when mussel bed topography is complex, or localities have very different recruitment densities.     

Wave action and competitive interaction between the invasive mussel Mytilus galloprovincialis and the indigenous Perna perna in South Africa

On the south coast of South Africa, the invasive alien mussel Mytilus galloprovincialis shows partial habitat segregation with the indigenous mussel Perna perna. P. perna predominates in the lower mussel zone and M. galloprovincialis in the upper zone, with mixed beds where the two overlap. We examined competitive interactions between these species by translocating mussels into small plots at high densities. Treatments involved different combinations of species and densities placed in each zone. Mortality was monitored regularly and at the end of each experiment, growth and condition index were measured. The experiment was attempted three times. The first two attempts were disrupted by wave action, especially winter storms, but provided information on species-specific effects of wave action. In experiment one, wave induced mortality decreased from a mean for both species of approximately 90% on the low shore to ca. 50% on the high shore, and was 15–30% lower for P. perna than M. galloprovincialis in each zone. In experiment two, M. galloprovincialis mortality was not affected by zone (Kruskal–Wallis test, P > 0.05), but was higher than P. perna mortality in the low zone (P < 0.05). P. perna survival was significantly (P < 0.05) lower on the high than mid and low zones, apparently due to the effects of greater emersion. Condition index showed a similar pattern, being lowest in the low zone for M. galloprovincialis and in the high zone for P. perna (3-way ANOVA, P < 0.05). Growth rates were fastest for both species in the low zone (Kruskal–Wallis, P < 0.05 in both cases). The third experiment was run for 12 months in the low zone only and provided evidence of intraspecific competition for P. perna and of interspecific competition. Condition was significantly greater for P. perna in all treatments (2-way ANOVA), as was growth (Kruskal–Wallis P < 0.05). Significant treatment effects indicated that P. perna had a negative effect on M. galloprovincialis survival (Kruskal–Wallis, P < 0.05). Again wave action was important; by the end of the experiment all mussels had been removed from plots stocked only with M. galloprovincialis. Thus P. perna improves survival of M. galloprovincialis on the low shore in the short term, by providing protection against wave action, but excludes it competitively in the longer term. The results show that partial habitat segregation is likely to be a permanent feature on the south coast, with M. galloprovincialis unable to dominate the low shore due to the effects of waves and competitive exclusion by P. perna. This study is the first attempt to examine the mechanisms of interaction between invasive and indigenous marine mussel species and provides evidence of the importance of environmental conditions in the mediation of this interaction.     

Mechanisms of habitat segregation between an invasive and an indigenous mussel: settlement,post-settlement mortality and recruitment

The mussel Mytilus galloprovincialis is highly invasive worldwide, but displays varying degrees of local and regional coexistence with indigenous mussels through spatial habitat segregation. We investigated the roles of settlement, post-settlement mortality, juvenile growth and recruitment in partial habitat segregation between the invasive M. galloprovincialis and the indigenous mussel Perna perna on the south coast of South Africa. We used two study locations, Plettenberg Bay and Tsitsikamma, 70 km apart, with two sites (separated by 300–400 m) per location, each divided into three vertical zones. There were no significant effects in Tsitsikamma, where daily settlement and monthly recruitment were significantly lower than in Plettenberg Bay. In Plettenberg Bay, settlement (primary and secondary) and recruitment of both species decreased upshore. Post-settlement mortality was measured over two consecutive 6-day periods during a spring tide and a neap tide. For both species mortality was low on the low-shore. High-shore mortality was consistently low for M. galloprovincialis, but increased dramatically for P. perna during spring tide. No data were obtained for growth of P. perna, but juvenile M. galloprovincialis grew more slowly farther upshore. P. perna recruited mainly in spring and summer, with a peak in summer far greater than for M. galloprovincialis. Recruitment of M. galloprovincialis was more protracted, continuing through autumn and winter. Thus local coexistence is due to a combination of pre- and post-recruitment factors differing in importance for each species. P. perna is excluded from the high-shore by recruitment failure (low settlement, high mortality). High survival and slow growth in juveniles may allow large densities of M. galloprovincialis to accumulate there, despite low settlement rates. With no differences between species in settlement or mortality on the low-shore, exclusion of M. galloprovincialis from that zone is likely to be by post-recruitment processes, possibly strengthened by periodic heavy recruitments of P. perna. At larger scales, larval retention and protracted recruitment contribute to the success of M. galloprovincialis at Plettenberg Bay, while recruitment limitation may explain why M. galloprovincialis is less successful at other sites.     

Settlement, recruitment and potential predators and competitors of juvenile echinoderms in the rocky subtidal zone

Recruitment patterns of marine invertebrates are affected both by settlement and early post-settlement events. This study examined the settlement and recruitment patterns of echinoderms at three sites in the rocky subtidal zone of Bocabec Cove, Bay of Fundy, Canada using artificial turf collectors and quadrats on the natural substrate. Potential predators were quantified at two of the sites along transects and in 1-m² quadrats. Both potential predators and competitors were quantified in 0.0625-m² quadrats. Settlement varied across sites (1.5–3 km apart) and two years of sampling (2004, 2005). The site of most potential settlement differed for the three groups of echinoderms: sea urchins (Strongylocentrotus droebachiensis), sea stars (Asterias spp.) and sea cucumber (Psolus fabricii). Settlement densities on the artificial turf collectors tended to be greater than the densities of settlers on the natural substrate. On the natural substrate, the only significant difference between densities of juveniles over time was that newly settled sea stars were found in July and were not found the following October. Large lobsters and carnivorous worms were potential predators with densities that varied between sites. Potential competitors that differed in abundance between sites were herbivorous gastropods and conspecifics for sea urchins; and carnivorous worms for sea stars. This study suggests that patterns of recruitment are either set up by patterns of settlement or by events during the first few weeks/months on the benthic substrate for these echinoderms.     

Positive effects of the introduced green alga, Codium fragile ssp. tomentosoides, on recruitment and survival of mussels

The green macroalga, Codium fragile ssp. tomentosoides, is an important component of sheltered low-shore assemblages on breakwaters along sandy shores in the northern Adriatic Sea. Macroscopic thalli of C. fragile are not perennial, but develop from propagules and/or undifferentiated forms in early spring, when the dominant native space-occupier, the mussel Mytilus galloprovincialis, recruits. By mid-summer, rapid growth of C. fragile leads to the formation of a dense canopy. We investigated the effects of juvenile and adult thalli of C. fragile on recruitment, survival and growth of mussels. Two experiments tested the hypotheses: (1) that recruitment of mussels is greater within patches of juvenile thalli (primordia) of C. fragile than on adjacent bare surfaces; (2) that the presence of a canopy of C. fragile affects the survival and growth of mussel recruits. The number of recruits of mussels was significantly larger within clumps of primordia of C. fragile than on bare surfaces. The removal of the canopy of C. fragile affected negatively the density of mussels after 2 months from the start of the experiment, but there were no effects on the mean size of individuals, nor on the size–frequency distribution. The same trend persisted after 4 months from the start of the experiment. These results show that re-colonisation of space by mussels is enhanced by C. fragile. Given that mussels, in turn, have the potential to reduce recruitment rates of C. fragile, quick recovery of mussel beds after disturbances could be crucial for controlling the abundance of this alga on breakwaters. Results also suggest that the effects of introduced species on native assemblages can be explained only through studies encompassing different life-stages of interacting organisms.     

Population dynamics and growth of juveniles of the velvet swimming crab Necora puber (Decapoda: Portunidae)

Recruitment variability plays a critical role in determining local population densities of benthic organisms, but extreme vulnerability at the onset of juvenile life is a trait that is largely responsible for population survivorship trends. The aim of the present study was to determine the role of juvenile recruitment in the population structure of Necora puber. Juveniles of N. puber were collected from the lower intertidal of rocky shores of Plymouth Sound (southwest coast of the UK) and monthly size–frequency distribution were used to determine the dynamics and the growth of the population. The parameters of the von Bertalanffy growth function were estimated (K=0.281 year⁻¹; t ₀=0.043; C=0.103; and t _s=0.268) assuming a L _∞=105 mm. Growth was markedly seasonal and present results indicated a slower juvenile growth rate than described previously for N. puber. The recruitment period was extensive and was two times higher in 2001 than in 2000 at the start of the 1+ year, but levelled off at the end of the 1+ year class on three of the four shores studied. Instantaneous mortality as high as 5.1 year⁻¹(99.4% year⁻¹) was observed during the higher recruitment year. Early juvenile mortality appears to be density dependent and a demographic bottleneck appears to limit the number of juveniles on some shores.     

Distribution and morphological variation of low-shore algal turfs

The distribution of three functional groups of algae (filamentous, corticated terete and calcareous articulated) was investigated in low-shore, turf-forming assemblages from rocky shores in the western Mediterranean Sea (Italy). Algae were sampled along shores from three different stations. Shores were either exposed to wave action or sheltered and were characterised by various inclinations of the rocky substratum (horizontal, sloping and vertical). The hypotheses tested were: that the relative abundance of the three functional groups of algae (1) varies between sheltered and exposed shores, (2) is influenced by substratum inclination and (3) that these distributions are consistent across stations. There was no clear relationship between the morphology of turf-forming algae and shore characteristics. Covers of filamentous, corticated terete and calcareous articulated algae differed among shores and stations, but patterns were not consistent across shores with similar exposure to waves and substratum inclination. Turfs often comprised algae with different morphologies, in a combination of anchor and epiphytic species. It is suggested that functional groups of algae, as currently identified, are not effective in describing changes in distribution of algae between sheltered and exposed shores, when turf morphologies are considered. Future research on relationships between morphology and ecology of algae should consider the potential importance of positive interactions among associated species, which could override responses of individual functional groups.