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.     

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.     

Distribution and abundance of co-occurring chthamalid barnacles Chthamalus montagui and Chthamalus stellatus (Crustacea,Cirripedia) on the southwest coast of Portugal

Two co-occurring chthamalid barnacle species occur on the rocky shores of continental Portugal: Chthamalus stellatus (Poli) and Chthamalus montaguiSouthward. In the present study, patterns of distribution and abundance (density and percentage cover), both vertically (tidal level) and horizontally (wave action), were studied for each species on headlands (more wave action) and not on headlands (less wave action) and at two spatial scales (shore and site) and three tidal levels (low, mid and upper). Chthamalus montagui was significantly more abundant than C. stellatus in both degrees of wave action and at all tidal levels. Despite this great difference in abundance and considering each species individually, C. montagui was more abundant at mid tide-level, while C. stellatus was more abundant at the low tide-level, and C. montagui was less abundant on headlands, while C. stellatus was more abundant on headlands. Small-scale variability (between sites, tens of meters) of abundance of each species was detected, while differences on a larger scale (between shores, tens of kilometres) were not significant.     

The vertical distribution of Chthamalus montagui and Chthamalus stellatus (Crustacea,Cirripedia) in two areas of the NW Mediterranean Sea

The Mediterranean Sea is characterised by a small tidal range (0.3–1 m). Despite this, intertidal communities are well established and their upper limits often extend above mean high water level. Organisms living in the intertidal region and in the supralittoral zone rely on both tides and wave action to perform their biological functions. Lack of food, desiccation and predation are common stresses in such a harsh environment. The present study deals with the vertical distribution of two species of intertidal barnacles, Chthamalus montagui Southward and Chthamalus stellatus (Poli), which are the main constituents of the barnacle belt along Mediterranean rocky shores. Previous work, carried out in the Atlantic, showed that where the distribution ranges of the two Chthamalus species overlap, C. montagui is more common in the upper barnacle zone while C. stellatus is dominant lower down. The main aims of our study are: (1) to establish if there is a relationship between position and extension of the barnacle belt on the shore and tidal range and/or wave exposure, (2) to test the hypothesis that in the study areas C. montagui is more abundant than C. stellatus high on the shore, and that the pattern is reversed lower down. Barnacle populations were monitored in summer 1998 in the Gulf of Genoa (Ligurian Sea) and in the Gulf of Trieste (North-Adriatic Sea). The two areas differ in tidal range and hydrodynamism, the former presenting quite strong wave action and a tidal range of 30 cm, the latter having limited wave action and 1 m tidal range. Three shores were randomly selected in each gulf and two transects on each shore. Counts of barnacles in 10 ^* 10 cm quadrats were done at different shore heights along each transect. The data was subjected to analysis of variance. Results showed that a more pronounced hydrodynamic regime corresponded to a shift of the barnacle belt towards the higher shore (Gulf of Genoa), while in more sheltered areas (Gulf of Trieste), the barnacle distribution was confined to the intertidal region. The relative spatial distribution of C. montagui and C. stellatus within the barnacle belt varied locally, even between transects on the same shore, and this obscured the distribution pattern along the vertical gradient. Nevertheless, it was still possible to conclude that at mid and high shore in Genoa, C. stellatus was more abundant than C. montagui, while in Trieste the pattern was reversed.     

Spatial and temporal patterns in the recruitment of the intertidal barnacle Chthamalus malayensis Pilsbry (Crustacea: Cirripedia) on the equatorial shores of Peninsular Malaysia and Singapore

Spatial and temporal variation in the recruitment of the intertidal barnacle Chthamalus malayensis was examined over one year (September 2003-August 2004) on the equatorial shores of southern Peninsular Malaysia and Singapore. A nested sampling design was applied for the first time on these shores, over three spatial scales - hundreds of kilometres, kilometres to tens of kilometres and tens of metres - and temporal variation was determined through monthly sampling of recruits. Shores within 2° north of the equator on the East and West coasts of southern Peninsular Malaysia and the southern coast of Singapore were selected as study sites. Generally, all three coasts recruited throughout the year, with varying intensities. There was a clear regional pattern, where the largest number of recruits was found on the East Coast of Malaysia, and the least in Singapore. Analyses of variance (ANOVA) showed that variability occurred on the smallest scale, but only in nine of the twelve months examined, consequently resulting in significant temporal and spatial interaction. Calculated variance components indicated that small-scale variation accounted for most of the overall variability. The potential causes of the spatial and temporal patterns of C. malayensis recruitment, and implications on tropical-temperate comparisons will be discussed.     

Comparative studies on the biology of nerita atramentosa Reeve, bembicium nanum (Lamarck) and cellana tramoserica (Sowerby) (gastropoda: Prosobranchia) in S.E. Australia

Populations of three coexisting intertidal gastropods, Nerita atramentosa Reeve, Bembicium nanum (Lamarck) and Cellana tramoserica (Sowerby), were sampled from a shore in Botany Bay, New South Wales, from July 1972 to September 1973. The recruitment and growth rates of each species were analysed from size frequency distributions. Mortality of each age cohort, and longevity, were estimated from analyses of the densities of Nerita atramentosa and Bembicium nanum.Nerita atramentosa showed no significant mortality during the first two years on the shore, but high mortality (at an instantaneous rate of 0.084 deaths/individual/month) after reaching the age of reproductive maturity, which was 20 months from settlement and at a mean shell-length of 13.5 mm. No growth could be detected after reproductive maturity was reached. Longevity of N. atramentosa was estimated as 3–5.5 years. Bembicium nanum juveniles showed higher mortality (at an instantaneous rate of 0.233 deaths/individual/month) than reproductively mature animals (0.060 deaths/individual/month). Reproductive maturity was reached at a mean shell-breadth of 11.0mm, i.e., about ten months after settlement on the shore. Longevity was estimated as from 4–8 years. Cellana tramoserica showed different growth patterns depending on the time of settlement.The three species showed different patterns of growth and life history relating to variable recruitment (which was demonstrated in all three species) and different rates of mortality of age cohorts.     

Isolated and combined impacts of blue mussels (Mytilus edulis) and barnacles (Balanus improvisus) on structure and diversity of a fouling community

Effects of two presumably dominant competitors, the blue mussel Mytilus edulis and the barnacle Balanus improvisus on recruitment, population dynamics and community structure on hard substrata were experimentally investigated in the subtidal Kiel Fjord, Western Baltic. The hypothesis that blue mussels and/or barnacles are local dominants and strongly influence succession and community structure was tested by monitoring succession in the presence and absence of simulated predation on either or both species. Manipulations included blue mussel removal, barnacle removal, combined blue mussel and barnacle removal, as well as a control treatment for natural (non-manipulated) succession. In the second part of the experiment, recovery from the treatments was monitored over 1 year.During the manipulative phase of the experiment, blue mussels had a negative effect on recruitment of species, whereas barnacles had no significant effect. Even so, a negative synergistic effect of blue mussels and barnacles was detected. Calculation of species richness and diversity H′ (Shannon Index) showed a negative synergistic effect of blue mussels and barnacles on community structure. Additionally, diversity H′ was negatively affected by the dominant competitor M. edulis. These effects were also detectable in the ANOSIM-Analysis. The non-manipulative phase of the experiment brought about a drastic loss of diversity and species richness. Blue mussels dominated all four communities. Barnacles were the only other species still being able to coexist with mussels. Effects of simulated predation disappeared fast.Thus, in the absence of predation on blue mussels, M. edulis within a few months dominates available space, and diversity of the benthic community is low. In contrast, when mussel dominance is controlled by specific predators, more species may persist and diversity remains high.     

Effect of the substratum in the recruitment and survival of the introduced barnacle Balanus glandula (Darwin 1854) in Patagonia, Argentina

The barnacle Balanus glandula was introduced in Argentina in the 1970s, and today it dominates the high intertidal level in most Argentinean rocky shores. The aim of this work was to evaluate the effect of the type of substrata and intertidal height on a population of Balanus glandula by conducting field surveys and one-year field experiments in which we combined different substrata (hardness: hard and soft, and texture: smooth and rough) at two intertidal heights (mid and high). In natural populations, the highest density of adults and recruits occurred on soft-rough substratum and in the high intertidal. The different textures were important only on the soft substrata and high intertidal, and the density of barnacles of the soft-rough substrata was higher than soft and smooth ones. The most suitable experimental substratum was the soft-rough of the high intertidal, which had the highest recruitment, survival and final density of barnacles at the end of the experiment. In contrast, the hard and smooth of the high and middle intertidal were the least suitable in all cases. Although the recruitment of B. glandula occurred throughout the year, it was higher in the high intertidal, and it showed a recruitment peak in the winter and a second in the summer. While most studies on this barnacle investigated the effects of granite or other volcanic hard substrata, our study also focused on soft substrata. The effects of soft substrata are particularly important because soft sedimentary rocks characterise the southern Atlantic coast of South America and the presence of soft rocks appears to optimize the success of Balanus glandula.     

Effect of rock type on the recruitment and early mortality of the barnacle Chthamalus montagui

Discontinuities in distribution of sessile species due to changes in available habitat have rarely been investigated. In the English Channel, soft sedimentary rocks including chalk could potentially present a barrier to eastward range extension of the southern barnacle Chthamalus montagui. To test the effect of rock type on recruitment and mortality, prepared settlement tiles of four calcareous rocks that form important platforms close to existing limits of distribution were fixed to the shore at a site in south-west England, known to have consistently high larval settlement. Prior to fixation, the roughness parameter “R_a” and Potential Settlement Sites (PSS) index was measured. Recruitment was monitored photographically on four occasions during the settlement season and subsequent mortality for 7 months after the settlement season. By the end of the settlement season, recruitment on the hard Kimmeridge Cementstone was significantly less than on the other three rock types and attributed to low surface roughness and fewest Potential Settlement Sites (PSS), yet only 28% of variance could be attributed to PSS. Post-recruitment mortality was more variable, however after 7 months there were significant differences between rock types. Simultaneous surveys of post-recruitment mortality in natural populations showed that although recruitment on chalk can potentially be high, survivorship was relatively low. Thus rock type could be influential in setting species geographical limits.     

Roles of larval supply and behavior in determining settlement of barnacles in a temperate mangrove forest

Recruitment is often a major influence on the spatial distribution of populations of benthic marine invertebrates, but the contributions of different components of recruitment are not well known, with the added complication that the relative importance of various life-history processes may be scale-dependent. Previously, we have shown that over a large scale across a mangrove (Avicennia marina) forest in southeastern Australia, settlement of the barnacle Elminius covertus explained its patterns of recruitment, which in turn explained the distribution of adults on mangrove pneumatophores. Post-settlement mortality had little influence on this pattern. In contrast, small-scale vertical distributions of adult barnacles along individual pneumatophores were determined by the pattern of recruitment, which differed from the pattern of settlement, so post-settlement mortality determined the vertical patterns of adults.

In this study, we tested whether larval supply and/or settlement behavior influence the observed settlement patterns of E. covertus across a forest (from seaward to landward zones). We also tested whether larval supply could explain the vertical settlement patterns along the pneumatophores. A pumping system was used to collect cypris larvae from seaward, mid and landward zones of a mangrove forest and an adjacent, unvegetated shore and from three heights above the sediment surface. We also used transplantation of wooden stakes bearing microbial films and barnacle recruits between horizontal zones of the forest to determine whether settlement was influenced by these films or recruits.

Both cyprid supply and cyprid behavior were important factors in determining the patterns of settlement of E. covertus across the forest. Cyprid supply was a result of three-fold differences in immersion times of different (landward, mid and seaward) zones across the forest and a decrease in density of cyprids in the water column from the seaward zone of the forest to the landward sections. In the absence of mangroves immediately adjacent to the forest, there was no temporally consistent difference in cyprid density across the shore and even the differences in immersion time did not produce consistent differences in cyprid supply across the shore. Wooden substrata that had been immersed at seaward sections of the forest attracted consistently more settlers than substrata immersed initially at other sections of the forest and settlement could be induced beyond the normal distribution of adults of E. covertus by stakes transplanted from the seaward zone.

The vertical settlement pattern could not be explained by the supply of cyprids, suggesting that larval behavior must determine the vertical settlement pattern.     

Discrimination at settlement in barnacles: Laboratory and field experiments on settlement behaviour in response to settlement‐inducing protein complexes

A major driving force to mechanistic studies of barnacle gregarious settlement is to contribute to an understanding of observed patterns of settlement in nature. In particular, how cyprids perceive adult conspecifics and how they discriminate between conspecific and allospecific barnacles are questions which have taxed researchers for nearly 50 years. The putative, active component of adult barnacles to which the cyprids respond has long been known to be a glycoprotein, referred to here as the settlement‐inducing protein complex (SIPC). The present study examines the discriminatory abilities of laboratory‐reared Balanus amphitrite and wild Semibalanus balanoides cyprids at settlement. Using a recently developed nitrocellulose membrane‐choice settlement assay, laboratory studies revealed that both species settled at a significantly higher rate on regions of membrane on which crude conspecific SIPC had been adsorbed compared to untreated regions. Moreover, when offered a choice between conspecific and allospecific SIPC, a trend to greater settlement on the conspecific regions was observed. The membrane assay was also evaluated in field trials using real‐time video footage of cyprid searching behaviour. Of 211 S. balanoides cyprids recorded during exploratory behaviour, only one settled. Exploratory behaviour was, however, clearly associated with regions of the membrane treated with either conspecific or allospecific (B. Amphitrite) SIPC compared to untreated regions. These results are generally in accord with previous reports on the discriminatory abilities of barnacle cyprids and suggest that the membrane assay may be usefully applied to field studies of settlement behaviour.     

Population structure, recruitment, and succession of the brown alga, Padina boryana Thivy (Dictyotales, Heterokontophyta), at an exposed shore of Sirinart National Park and a sheltered area of Tang Khen Bay, Phuket Province, Thailand

The brown algal genus, Padina, has a worldwide distribution in tropical and subtropical climate zones. Padina individuals are common and sometimes dominant in both the intertidal and shallow subtidal regions associated with coral reefs. We investigated the population structure and recruitment of two populations of Padina boryana at an exposed shore in Sirinart National Park (SNP) and a sheltered shore of Tang Khen Bay (TKB), Phuket Province, Thailand. From September 2005 to August 2006, the number of released spores and the height and radius of fronds were measured monthly. New recruits were recorded and monitored on manipulated permanent plots 0.25 m². The experiment was carried out monthly over a year. We found Padina recruits 1 month after the plots were cleared at both sites. There was a significant difference in percentage cover by new individuals between the two locations (P < 0.05) and also at the shore levels within the sites. At SNP, the highest recruitment was found in the uppermost zone, while at TKB high recruitment occurred at all shore levels except at 80-100 m from the shore. The factors influencing recruitment of P. boryana include wave motion, competition with other earlier successional species and sediment. The high availability of reproductive spores throughout the year, the successful recruitment and the Dictyerpa stage promote the successful establishment of Padina populations.     

Competition and facilitation between the alien mussel Mytilus galloprovincialis and indigenous species: Moderation by wave action

As the alien species that most dominates space along the South African coast, the Mediterranean mussel Mytilus galloprovincialis has radically altered community composition on invaded shores. We experimentally assessed interspecific interactions between this invasive species and dominant indigenous species in conjunction with considering how wave action moderates such interactions. The density of both M. galloprovincialis and the limpet Scutellastra granularis increased with wave action. Conversely, the tube-building polycheate Gunnarea capensis was negatively affected by wave exposure, being most abundant on sheltered shores. The influence of wave action on the indigenous mussel Aulacomya ater, however, remains unclear. M. galloprovincialis outcompeted both G. capensis and A. ater at moderate to high exposure levels, whereas it had both positive and negative effects on S. granularis. It outcompeted adult limpets on primary rock space on semi-exposed and exposed shores, reducing densities of this portion of the population. However, recruitment of S. granularis was facilitated by M. galloprovincialis, as greater numbers recruited to the secondary substratum offered by mussel shells. Again this interaction intensified with wave action. Due to the extremely high density of recruits on secondary space, the net effect of M. galloprovincialis on S. granularis was positive. Thus, wave action not only influences the abundance of individual species, but also mediates both positive and negative interspecific interactions in rocky shore communities, including the impact of alien species such as M. galloprovincialis.     

No effect of naturally-occurring rock types on settlement or survival in the intertidal barnacle, Tesseropora rosea (Krauss)

The settlement and early survival of young barnacles, Tesseropora rosea (Krauss), on four different, naturally-occurring rock substrata were investigated in a four-factor experiment. Three factors assessed variation on three spatial scales: among shores on the New South Wales coast in eastern Australia i.e., 10 to 600 km apart; among sites within these shores i.e., 20 to 50 m apart; within these sites, i.e., < 3 m². Variation on these scales was usually significant for settlement density and proportional survival 30–60 days post-settlement. The fourth factor, different rock types, showed no consistent trends for settlement density or proportional survival. Previously observed differences in these phenomena among shores cannot, therefore, be ascribed to differences in rock substrata.     

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.     

The effect of ecdysterone on cyprids of Balanus eburneus Gould

Synthetic ecdysterone in concentrations from 0.02 to 250 · 10^?6 causes developmental abnormalities in the cyprids of Balanus eburneus Gould. These cyprids are unable to attach themselves to the substratum, but readily metamorphose into barnacles with recognizable adult shell structures; these remain encased within the cyprid valves, but live only for a short time. The possible involvement of an ecdysone system in the metamorphosis of cirripedes is discussed.     

Benthic assemblages and temperature effects on Paracentrotus lividus and Arbacia lixula larvae and settlement

Recruitment is a principal controlling factor in population dynamics of marine species. In marine invertebrates with a planktonic larval stage, such as echinoids, recruitment is assured by larval supply, settlement and juvenile survival. Larval supply and juvenile survival are affected by a wide range of factors, including temperature, presence of predators, quality and quantity of food. Echinoid larval settlement is mainly conditioned by the finding of a suitable substrate to metamorphose. The sea urchins Arbacia lixula and Paracentrotus lividus are considered key species of the Mediterranean infralittoral rocky shores. At high densities, the grazing activity of both species can produce and maintain barren grounds, a particular habitat condition characterized by extremely low cover values of erect algae with high presence of naked substrates and encrusting corallinales, poor in biodiversity and ecosystem functions. We tested the role of different settlement substrates on the metamorphosis competent larvae of the two species. Furthermore, from our larval rearing trails we were able to identify strong temperature effects on larval development of the two species. P. lividus and A. lixula larvae have been reared at 18 °C but for the second species it was necessary to use higher temperatures (22 °C) to perform settlement experiments, as in the 18 °C set all larvae died in the first week. Both species larvae have been fed Cricosphaera elongata. Metamorphosis of competent larvae has been induced using different substrates: naked stones, Lithophyllum incrustans, Stypocaulon scoparium, Corallina elongata, turf forming algae and Posidonia oceanica. For each species, two larval batches were used for settlement experiments; for each larval batch two replicates/substrates were set up. No differences in the rate of metamorphosis on any of the tested substrates were observed for P. lividus, while A. lixula showed to prefer naked stones and encrusting coralline algae Considering that A. lixula population growth may trigger barren extension on rocky shores, this may lead to a positive feedback between barren extension and A. lixula population density. Furthermore, our results suggest that the predicted rise in seawater temperature may favor A. lixula larval survival and inhibit P. lividus. Combining information on temperature tolerance with other sources of information for these species in the Mediterranean, it is possible to develop a conceptual model of the interaction between the two species and the alternative state of their habitats.     

Spatial variation in juvenile and adult Ecklonia radiata (Laminariales) sporophytes

Juvenile and adult sporophytes of Ecklonia radiata were counted on in-shore and off-shore reefs at Marmion (Perth, Western Australia), across spatial scales of kilometres, 10s of metres and metres. The position on in-shore reefs vs. off-shore reefs did not influence recruit and adult kelp density. There was considerable site-to-site (kilometres) variation in densities of recruits but not adults. The majority of variation in both recruit and adult densities was found between quadrats separated by a few metres. There was no relationship between abundance of recruits and abundance of adult sporophytes at any of the spatial levels of investigation, suggesting a decoupling of recruitment and canopy processes. The results emphasise the patchy nature of kelp canopy distribution and suggest that relatively small-scale processes, acting on scales of a few metres, are of primary importance in generating heterogeneity in the canopy of these kelps.     

Interactions of Patella and macroalgae with settling Semibalanus balanoides (L.)

Biological interactions affecting densities of settling and newly-settled Semibalanus balanoides (L.) have been investigated by manipulative field experiments on the Isle of Man.The effects of sweeping by fucoid clumps of different species and Patella browsing have been compared on moderately-exposed shores. Patella allowed barnacle settlement by preventing growth of competing green algae, but reduced post-settlement densities. Small clumps of Fucus spiralis L., F. vesiculosus L, and F. serratus L. all reduced settlement considerably more so than limpets. F. serratus had the greatest sweeping effect.Interactions between macroalgae and Semibalanus balanoides have been investigated at all levels on sheltered shores and low down on more exposed shores. In the Fucus spiralis and F. vesiculosus zones, post-settlement numbers were higher than in adjacent areas where the canopy was removed. Barnacles did not settle readily in the Ascophyllum zone in either experimental or control areas. Settlement occurred in the upper part of the Fucus serratus zone in experimental areas where the canopy was removed but not in control areas. No settlement occurred in either treatment or control areas lower in the F. serratus zone. At all levels on the shore fucoid canopies seemed to reduce cyprid settlement, but the effect was greatest amongst F. serratus where there was total prevention. High on the shore the effect of enhanced post-settlement survival under the canopy outweighs reduction of cyprid settlement thus there are greater numbers in the controls. Competition with red algal turfs was shown to set the lower limit of the barnacle zone on a vertical pier face.     

Will the invasive mussel Mytilus galloprovincialis Lamarck replace the indigenous Perna perna L. on the south coast of South Africa?

The mussel Mytilus galloprovincialis is invasive worldwide, has displaced indigenous species on the west coast of South Africa and now threatens Perna perna on the south coast. We tested the hypothesis that Mytilus will replace Perna by examining changes in their distribution on shores where they co-exist. Total cover, adult density, recruit density, recruit/adult correlations and mean maximum lengths of both species were measured in 2001 at two contrasting sites (Plettenberg Bay and Tsitsikamma) 70 km apart, each including two locations 100 m apart. Cover and density were measured again in 2004. Total mussel abundance was significantly lower in Tsitsikamma, and recruit density was only 17% that of Plettenberg Bay. Abundance and cover increased upshore for Mytilus, but decreased for Perna, giving Mytilus higher adult and recruit density and total cover than Perna in the upper zones. Low shore densities of recruits and adults were similar between species but cover was lower for Mytilus, reflecting its smaller size, and presumably slower growth or higher mortality there. Thus, mechanisms excluding species differed among zones. Recruitment limitation delays invasion at Tsitsikamma and excludes Perna from the high shore, while Mytilus is excluded from the low shore by post-recruitment effects. Recruitment limitation also shapes population structure. Recruit/adult correlations were significant only where adult densities were low, and this effect was species-specific. Thus, at low densities, larvae settle or survive better near adult conspecifics. After 3 years, these patterns remained strongly evident, suggesting Mytilus will not eliminate Perna and that co-existence is possible through partial habitat segregation driven by recruitment limitation of Perna on the high shore and post-settlement effects on Mytilus on the low shore.