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.     

The distribution of molluscs on littoral rocks in Ceylon,with notes on their ecology

Molluscs inhabiting littoral rocks were collected from 43 stations around Ceylon, and 57 species belonging to 20 families were identified. An account is given of the nature and distrbution of littoral rocks around the island, and the stations examined. On each shore, the distribution of the molluscs was studied in relation to the broad zonation of the shores themselves and observations were made on plant and animal associations of each speices. The limits of the broad zones of the shores were defined with reference to biological indicators. Analysis of the data collected led to a classification of the rocky shores into three groups, each characterized by particular combinations of mollusc species and distinct patterns of shore zonation. Some molluscs were found to be distributed over the whole island, others to be common to two groups of shores only, and still others to be confined to a single shore type. An attempt has been made to explain the presence or absence of species on various groups of shores, and to explain discontinuities in distribution on the basis of environmental factors such as the nature of the substratum, degree of wave exposure, and weed cover. A key, based primarily on external features, has been devised for the identification of the species listed.     

Functional identity and functional structure change through succession in a rocky intertidal marine herbivore assemblage

Despite the great interest in characterizing the functional structure and resilience of functional groups in natural communities, few studies have examined in which way the roles and relationships of coexisting species change during community succession, a fundamental and natural process that follows the release of new resources in terrestrial and aquatic ecosystems. Variation in algal traits that characterize different phases and stages of community succession on rocky shores are likely to influence the magnitude, direction of effects, and the level of redundancy and complementarity in the diverse assemblage of herbivores. Two separate field experiments were conducted to quantify per capita and population effects and the functional relationship (i.e., redundancy or complementarity) of four herbivore species found in central Chile during early and late algal succession. The first experiment examined grazer effects on the colonization and establishment of early-succession algal species. The second experiment examined effects on the late-successional, dominant corticated alga Mazzaella laminarioides. Complementary laboratory experiments with all species and under natural environmental conditions allowed us to further characterize the collective effects of these species. We found that, during early community succession, all herbivore species had similar effects on the ephemeral algae, ulvoids, but only during the phase of colonization. Once these algae were established, only a subset of the species was able to control their abundance. During late succession, only the keyhole limpet Fissurella crassa could control corticated Mazzaella. The functional relationships among these species changed dramatically from redundant effects on ephemeral algae during early colonization, to a more complementary role on established early-successional algae, to a dominant (i.e., keystone) effect on late succession. This study highlights that functional relationship within consumer assemblages can vary at different phases and times of community succession. Differentiation in herbivore roles emphasizes the need to evaluate consumer's impacts through different times of community succession, and through experimental manipulations to make even broad predictions about the resilience or vulnerability of diverse intertidal assemblages to human disturbances.     

Biomass, diversity and production of rocky shore macroalgae at two nutrient enrichment and wave action levels

The littoral zone of temperate rocky shores is normally dominated by perennial macroalgae (e.g. Fucus, Ascophyllum, Laminaria), but nutrient enrichment and/or permanently decreased wave action may lead to structural community changes from dominance of perennials to increased amounts of annual opportunistic species (mainly green algae). Macroalgal biomass, diversity and production as well as relationships between the two latter were studied using Solbergstrand’s rocky shore mesocosms in SE Norway in connection with a long-term experimental manipulation of nutrient addition and wave action (high and low levels of both factors applied in a crossed way to eight outdoor basins). After more than 2 years of experimental treatment, the total standing stock of macroalgae was larger in low nutrient than in high nutrient treatments as well as in high wave compared to low wave treatments (in autumn only). For macroalgal functional groups, bushy and filamentous brown and filamentous red algae were generally favoured by low nutrient concentrations, while annual filamentous and sheet-like green algae were stimulated by the nutrient enrichment. There was only one significant interaction between nutrient enrichment and wave action (for brown filamentous algae in autumn) and also only one significant main effect of the wave treatment (for bushy brown algae in autumn). Surprisingly, the high nutrient treatments supported a higher diversity of macroalgae, whereas the low nutrient treatments generally showed higher production rates. Moreover, significantly negative correlations were found between macroalgal diversity and primary productivity in both summer and autumn. This study shows that it is the biological components of the communities subjected to external forcing (nutrient addition or decreased wave action) that regulate production and this contradicts the common misperception that resource production in natural systems simply can be fast-forwarded by fertilization. The negative relationships between diversity and productivity, although a consequence of unexpected results for diversity and production, are also novel and hint towards species identities having more important functional consequences than general species dominance patterns and the amount of species per se. These results also emphasise the context dependency of findings within the field of biodiversity and ecosystem functioning.     

Intertidal assemblages on artificial and natural habitats in marinas on the north-west coast of Italy

Following the progressive expansion of human populations, the number of artificial habitats is increasing in shallow waters of urbanised coastal areas. The comparison of assemblages between natural and artificial habitats is necessary to determine whether there is real loss and fragmentation of natural habitats. This study investigates the changes in intertidal assemblages caused by marinas for tourists, along exposed rocky shores on the north-west coast of Italy. Marinas, being made of transplanted boulders and by internal wave-sheltered seawalls, introduce different types of artificial habitats in a relatively small area. Intertidal assemblages on breakwaters, seawalls and adjacent rocky shores were compared at three locations, thousands of metres apart. To assess the generality of patterns through time, natural and artificial habitats were sampled at three different times, over a period of about 2 years. Data were analysed by means of multivariate and univariate analyses to test the hypotheses that assemblages and abundances of single taxa differed among habitats, consistently among locations and times of sampling. Furthermore, the variability of assemblages at the scales of tens of centimetres and metres was compared among habitats. Assemblages on seawalls were largely distinct from those on rocky shores or breakwaters. Seawalls, which supported a smaller number of species than breakwaters and rocky shores, were dominated by encrusting algae and lacked common species such as Rissoella verrucosa and Patella rustica. The abundance of main-space occupiers did not differ between breakwaters and rocky shores, but there were differences in variability of assemblages at both the spatial scales investigated. This study provides evidence for differences between intertidal assemblages supported by artificial habitats at marinas and those on adjacent rocky shores. Differences in habitat-structure (and/or wave-exposure in the case of seawalls) could explain the occurrence of distinct intertidal assemblages. Despite the nature and magnitude of these differences, varied according to the type of artificial habitat considered, neither breakwaters nor seawalls could be considered surrogates of rocky shores.Communicated by R. Cattaneo-Vietti, Genova     

Differences in kelp morphology between wave sheltered and exposed localities: morphologically plastic or fixed traits?

The ability of algae to change the shape of their thallus in response to the environment may be of functional and ecological importance to the alga, with many species of macroalgae exhibiting a great range of morphological variation across wave exposure gradients. However, differences in morphology detected between sheltered and exposed environments cannot determine whether such differences represent plastic responses to the local environment or whether morphology is genetically fixed. This study tested for differences in the morphology of the common kelp, Ecklonia radiata, between wave sheltered and exposed environments, and reciprocally transplanted juveniles to distinguish the nature of such differences (i.e. plastic vs fixed traits). Differences between exposure environments were consistent with known effects of exposure (i.e. a wide, thin thallus at sheltered sites and a narrow, thick thallus with a thick stipe at exposed sites). The reciprocal transplant experiment confirmed that morphological plasticity was the mechanism enabling this alga to display different patterns in morphology between exposure environments. Individuals transplanted to the exposed environment underwent a rapid and extreme response in morphology, which was not apparent in individuals transplanted to the sheltered environment that responded more slowly. These results suggest that stressors typical of sheltered environments (i.e. diffusion stress) may not be as influential (if at all) compared to stressors typical of exposed environments (i.e. breakage, dislodgement) in differentiating morphological characters between exposure environments.     

Effects of patch shape in intertidal algal mosaics: roles of area,perimeter and distance from edge

Shape is considered an important attribute of patches, but little attention has been paid to its influence on the structure and dynamics of rocky shore assemblages. Effects of patch shape were investigated from September 1997 to June 1998 in algal assemblages on intertidal shores at Cape Banks, south of Sydney (NSW, Australia). Three experiments were established, in order to separately investigate the importance of three different characteristics of patch shape, namely area, perimeter and distance of the central zone from the edge. In each experiment, responses for sets of replicate patches that differed on average for only one shape characteristic were compared. Patches of different shapes were produced by scraping off all resident organisms, and subsequent colonisation was recorded. The range of shapes naturally occurring at the study sites was also measured over 1 year, and the shapes of plots used in the experiments were chosen to cover this range of natural variability. Shapes of natural patches within assemblages were very variable. Consistent differences were observed in patch size, geometry and characteristics of the substratum among sheltered and exposed shores and among different times of the year, suggesting that different disturbances influence the dynamics of assemblages at these sites. The results of the three experiments showed that shape may have important effects on patch colonisation. Area, perimeter and distance of the central zone from the edge significantly affected growth of algae in experimental clearings. Responses to different patch characteristics were species specific. Effects of different attributes of patch shape also varied over time. This spatial and temporal variability of responses suggests that colonisation processes of patches of different shapes are complex, probably involving effects that can produce contrasting results. Results are relevant to the design and interpretation of field experiments, as they suggest that the shape of plots may affect the perception of patterns and processes in intertidal assemblages.Communicated by R. Cattaneo-Vietti, Genova     

Distribution of algae on tropical rocky shores: spatial and temporal patterns of non-coralline encrusting algae in Hong Kong

Encrusting algae have been described as dominant space occupying species on rocky shores around the world. Despite their abundance, however, most studies classify species under generic names (e.g. Ralfsia sp.) or as a functional group (e.g. encrusting algae), thereby underestimating the number of species present and their ecological importance. Studies on six rocky shores of varying exposure, in Hong Kong, recorded eight common species of encrusting algae. The greatest abundance of encrusting algae was recorded on shores of intermediate exposure, where four distinct zonation bands could be identified; a cyanobacterial Kyrtuthrix-Zone in the upper midlittoral, a Bare-Zone below this, a Mixed-Zone in the lower midlittoral and a Coralline-Zone in the infralittoral fringe. Abundance declined on shores of greater and lower exposure to wave action, where bivalves and barnacles were competitively dominant. Certain species were found in greater abundance on exposed shores (e.g. Dermocarpa sp. and Hildenbrandia occidentalis), while others preferred more sheltered shores (e.g. Hildenbrandia prototypus and Kyrtuthrix maculans). With the exception of some cyanobacterial crusts, the abundance of encrusting algae was always greatest towards the low shore, an area of decreased physical stress and increased herbivore density. Zonation patterns showed seasonal variation associated with the monsoonal climate of Hong Kong. Most species increased in abundance during the cool season, while during the summer months the cover and vertical extent of encrusting algae decreased in relation to summer temperatures, although K. maculans increased in abundance during the summer. On Hong Kong shores, encrusting algae have a high species richness and exhibit within-functional group spatial and temporal variation which is mediated by herbivory and seasonal, physical stresses.     

Feeding capabilities and limitation of herbivorous molluscs: A functional group approach

The susceptibility of an alga to an herbivorous mollusc depends, in part, upon the size and toughness of the plant relative to the feeding ability of the mollusc. In this study, algae are subdivided into seven functional groups based on these and other physiological characteristics. Herbivorous prosobranchs and chitons are subdivided into four functional groups based on the structure of their feeding apparatus. Distinct patterns in the diets of these molluscs are evident when feeding data, based on these functional groups, are examined. Most herbivorous mollusc species eat algal forms that are either minute (i.e., micro- and filamentous algae) or very large and expansive (kelp-like or crustose algae). Algae of intermediate size (erect forms 1- to 10-cm tall) are eaten to a lesser extent, possibly because they are too large to be rasped from the substratum and too small for most herbivores to occupy. Herbivorous archaeogastropods (excluding limpets) and mesogastropods tend to eat filamentous and microscopic algal forms predominantly, whereas limpets and chitons feed on large, leathery and crustose algae. These dietary differences reflect functional differences in the feeding apparatus of these herbivore groups. Radulae of herbivorous mesogastropods function like rakes and can ingest larger, tougher algae than can radulae of nonlimpet archaeogastropods. The latter function more like brooms by sweeping the substratum broadly, but exerting little force. Limpets and chitons have superior excavating abilities because their radulae have: robust buccal muscles surrounding them, a reduced number of points of contact on the substratum, and minerally hardened teeth. The feeding apparatus of chitons is most versatile since it possesses features found in all herbivorous gastropod functional groups, and thus, it can sweep and excavate simultaneously. This functional group approach suggests various hypotheses concerning algal community structure, plant/herbivore and herbivore/herbivore interactions, the relative importance of structural defenses in algae, and the evolution of specialized grazers. These hypotheses are examined using data from published accounts.     

Multivariate morphometric analysis of two varieties of Littorina saxatilis from the Swedish west coast

Shell variation in Littorina saxatilis Olivi (=L. rudis) is examined with different techniques of multivariate analysis (canonical variate-, multiple discriminant-, principal component-, and principal coordinate analysis). In bivariate projections of the analyses, two populations, spatially only 60 m apart, but from different habitats, showed a phenotypic separation without overlap. One habitat is a boulder shore, while the other is a rocky cliff. A sample, geographically and environmentally intermediate to the exposed and sheltered populations, covered completely the morphologic gap between the two former groups, indicating a clinal variation in morphology over the environmental gradient exposed — sheltered. From this it is concluded that the two phenotypically distinct populations belong to the same species.     

Aspects écologiques de la zonation intertidale sur les côtes rocheuses des îles Galapagos

Features of the intertidal zonation on the rocky shores of the Galapagos Islands are described and discussed in relation to environmental factors, in particular temperature — which varies with regional hydrology and climatology. Owing to the thermal properties of the coastal waters and to the nature of the substrate (mainly basalt), intertidal organisms are exposed to a wide range of temperatures, which fluctuate with tide, day, and season. As a result, life is mostly restricted to the lowest levels of the shore; the middle- and upper-level populations are sparse. Local variations (biofacies) resulting from water agitation are described. Biota consist of tropical, subtropical, and warm-temperate components. Tropical species dominate in sheltered areas and in middle and upper shores.

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Contribution No. 164, the Charles Darwin Foundation.     

Wind induced upwelling as a possible explanation for mass occurrences of epiphyticEctocarpus siliculosus (Phaeophyta) in the northern Baltic Proper

The quantity of epiphytic filamentous algae varies considerably in macroalgal vegetation along rocky shores of the northern Baltic Proper. The main species responsible for irregular mass occurrences is the summer annual brown algaEctocarpus siliculosus (Dillw.) Lyngb. In this study, data collected over a 3-yr monitoring period are related to hydrographical parameters. The tideless and brackish Baltic Sea is salinity stratified, and the salinity difference between surface and bottom waters can be used to indicate upwelling events. Mass occurrences developed when the salinity difference was low. These salinity differences explained 87% of the observed increase inE. siliculosus cover. We propose that the slight salinity changes themselves do not affect the growth ofE. siliculosus, but rather they can be used as indicators of short nutrient pulses. These nutrient pulses are not detected in standard water sampling, as they are utilised by both pelagic and benthic organisms within a few hours or days. If the proposed mechanism is true, changes in wind speed and direction have an important effect on the nutrient dynamics of littoral algal communities. A higher frequency of SW winds may cause mass occurrences of filamentous algae, which are often believed to indicate eutrophication of an anthropogenic origin.     

Chemoautotrophic symbiosis in the tropical clamSolemya occidentalis (Bivalvia: Protobranchia): ultrastructural and phylogenetic analysis

Patterns of algal seasonality, and their effect on the diet and feeding preferences of the herbivorous crab Grapsus albolineatus were investigated over an 18-mo period from March 1993 to August 1994 on an exposed tropical rocky shore (Hok Tsui Peninsula at Cape d' Aguilar, Hong Kong). Algal cover was greatest in the winter months, and lowest in the summer. Foliose algae such as Ulva fasciata, Porphyra suborbiculata, and Dermonema frappieri were dominant in the winter, but died off in the summer. During the hot summer months, perennial encrusting algae e.g., Ralfsia expansa, Hildenbrandia rubra, H. occidentalis, coralline crusts and the encrusting cyanobacteria Kyrtuthrix maculans, were the dominant algal species. Seasonal variation in algal abundance influenced the dietary selectivity of the herbivorous crab G. albolineatus. In the winter, the crab fed selectively on filamentous algae (e.g. Hincksia spp., Cladophora spp., Enteromorpha spp., and the cyanobacteria Lyngbya sp.). Foliose algae (e.g. U. fasciata, P. suborbiculata, Pterocladia tenuis) formed a small part of the diet, despite being the dominant species on the shore. Foliose and filamentous algae were virtually absent from the shore in the summer, and the crabs switched to feeding solely on encrusting algae. Electivity indices revealed preferences for green and brown turf species, and avoidance of foliose algae. Faecal analysis revealed that a greater proportion of the food is digested in the winter, suggesting that G. albolineatus is able to digest filamentous algae more efficiently than encrusting algae. Feeding preferences of G. albolineatus appear to be influenced by a number of factors, including the availability, digestibility and morphology of algae. The foraging behaviour and cheliped morphology of the crab also affect food choice. The monsoonal nature of Hong Kong's climate controls the diversity and abundance of intertidal algae and, therefore, indirectly influences the diet and subsequent growth and reproductive success of the herbivorous crab G. albolineatus.     

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.     

A mathematical model of competition and colonization in a community of marine benthic algae

A mathematical model has been constructed for the algal community on the rocky shores of a Norwegian fjord. We report here on the studies of competition and colonization along a vertical transect from the upper intertidal to the sublittoral habiats. Results on species abundance and distribution (patterns of zonation) and time to reach maturity have been compared to observations both in the fjord area and in other rocky shore areas.Competition coefficients for the algae were inferred from plant morphology and shown to be in agreement with observations of algal abundance and their zone-forming ability. Competition restricts the distribution of the species, especially at the lower elevations, but does not alter their relative position. However, increasing uniform competition prolongs the time in which zone-forming can occur, and it also decreases the overall biomass which an area can sustain. Colonization by a single species may create transient stages in community development of the same order of magnitude as algae longevity, and probably also alters the zonation pattern to some degree.The simulation results indicate that the large-scale algal distribution pattern in the Hardangerfjord area results from global stability of the rocky shore community.     

Community succession following massive ice-scour on a rocky intertidal shore: recruitment,competition and predation during early,primary succession

The importance of species interactions and recruitment variability was examined during the first year and a half of primary succession (1988–1989) on an exposed rocky seashore near Halifax, Nova Scotia. Previous work suggested that emergent rock on these shores is normally dominated by fucoid rockweeds because predatory whelks control the sessile animal competitors, mussels and barnacles, and because herbivorous littorinids control ephemeral algal competitors. Abundances of all species except seasonal ephemeral algae were very small throughout this experiment and we found no significant effects of carnivory, herbivory, plant-animal competition or plant-plant competition. A slight facilitation of Fucus recruitment is attributed to a thin mat of ephemeral, blue-green algae. Very few other studies have directly manipulated intertidal ephemeral algae. As primary succession may be very rare in this assemblage, these results may be specific to these circumstances, but they highlight the varying importance of species interactions with variable recruitment. In particular, it appears that variations in recruitment success may be important to community structure, even when recruitment is not limited by propagule supply. The scale of the study also provides insight into successional processes occurring after the recent, extensive ice-scour of exposed seashores in this region.     

Food availability and diet composition of three coexisting Mediterranean limpets (Patella spp.)

The gut contents of three intertidal patellid limpets were analysed by collecting foraging specimens on a breakwater on the Tyrrhenian coast (central Italy) between May 1988 and October 1989. The three species coexist there showing a different, but partially overlapping zonation: Patella aspera dominates the infralittoral fringe; the majority of P. caerulea inhabits the lower midlittoral, while P. rustica is most abundant in the upper midlittoral. The algae present on slivers of substratum over which each limpet collected was moving were identified. Moreover, floristic surveys were made along the shore in order to characterize the algal cover of the different zonal belts. The floristic study revealed that the basic elements of algal communities typical of western Mediterranean rocky shores are present in the study area. The algae found on the slivers under the foraging limpets were generally representative of the algal community typical of the same zone. There was a marked difference between the diets of P. rustica and P. aspera due to the fact that the first species forages on a few low lying epilithic and endolithic Cyanophyceae, while P. aspera feeds on a large number of species belonging to all the main algal classes and life forms considered, including frondose epilithics and epiphytics. The diet of P. caerulea resembles that of P. aspera in algal heterogeneity, but is dominated by Cyanophyceae as in P. rustica. A detailed analysis of the differences between gut contents of each limpet species and the relative slivers showed an obvious general correspondence, but revealed also that the diets of the three species do not completely reflect the availability of algae. These findings suggest that the basic diet segregation mechanism between the three populations is their zonal separation. However, the difference in gut contents of heterospecific limpets foraging in the same zone suggests the existence of supplementary morphy-functional or behavioural mechanisms for diet segregation between the three species.     

Expansive covers of turf-forming algae on human-dominated coast: the relative effects of increasing nutrient and sediment loads

Turf-forming algae form more extensive habitat on subtidal rock adjacent to urban than non-urban coast of South Australia. This pattern is frequently observed on the worlds temperate coasts and is variously considered to be a result of enhanced concentration of nutrients or rates of sediment deposition on urban coasts. We experimentally tested which of three components of environmental change (increased nutrients in water, increased nutrients in sediments and increased sediment deposition) best explain the expansive covers of turf-forming algae on urban coasts. All three treatments had independent and positive effects on the percentage cover of turf-forming algae. The addition of nutrients from the water column had the largest influence (²=0.55), which was more than six times greater than the effect of nutrients added to sediments (²=0.08). An increase in rate of deposition of sediments had substantial effects (²=0.35), which were about one third less than those of water-borne nutrients. Importantly, the combined effect of all three treatments caused a 77% increase in percentage cover of turf-forming algae, which is comparable to the observed difference in covers between urban and non-urban coast in South Australia (93%). These results suggest that human activities that reduce water quality in both nutrient and sediment loads account for major change observed on human-dominated coasts. Despite this knowledge, we still lack complete information on the mechanisms that switch the primary subtidal habitat from canopy-forming algae to turf-forming algae on human-dominated coasts.Communicated by M.S. Johnson, Crawley     

Wave exposure effects on population structure and recruitment in the mussel <Emphasis Type="Italic">Perna perna</Emphasis> suggest regulation primarily through availability of recruits and food,not space

Recruitment and population structure of Perna perna in low shore mussel beds were investigated over 15 months at six sites along the south coast of South Africa. Initial, subjective classification of sites as wave exposed or wave sheltered (three of each) was confirmed using the dissolution of cement blocks to measure average water flux and dynamometers for maximum wave force. Recruitment occurred throughout the year, but recruit (1–5 mm) densities were significantly higher from January to April 1996 on both shore types. Recruit densities were positively correlated with adult (>15 mm) densities for both shore types (P < 0.05) but the correlations were extremely weak (r ² < 0.06 in each case). In areas with 100% cover, adult size (mean and maximum lengths) was greater on exposed sites, but density showed the reverse and was negatively correlated with maximum wave strength (r = −0.84). Despite differences in adult densities and sizes, biomass, which is a product of the two, showed no significant difference between the two shore types (ANOVA P > 0.05). Thus wave exposure dramatically affects density, recruitment and mussel size, but not recruitment timing or biomass where there is 100% cover, and mediates a three-way interaction among food supply, larval supply and intraspecific competition for space. In contrast to shores with saturation recruitment, mussel biomass here appears to be limited by recruit supply and constraints of food, especially on sheltered shores, while density is regulated through intraspecific competition for space primarily on exposed shores and at small spatial scales.     

Spatiotemporal variations in rocky intertidal malacofauna throughout Japan in the 1970s and 1980s

Spatial and temporal variations of benthic fauna over a wide region (>1,000 km) that includes complex ocean currents have seldom been quantitatively studied. The present study revealed malacofaunal variations on Japanese rocky shores affected by the Soya, Tsushima, Oyashio, and Kuroshio currents, and in the Inland Sea. On 29 shores in seven regions in these sea areas, previous researchers have recorded the number of molluscs in quadrats in spring and summer of 1978 and 1984–1986. Reanalyses of these records revealed the following patterns. (1) Shore-to-shore variation in malacofauna was correlated with shore-to-shore geographical distance less clearly for more southern regions. This might show that in southern regions many taxa with low relative abundance are haphazardly recorded on some shores and blur the effect of shore-to-shore distance on malacofaunal dissimilarity. (2) Region-to-region variation in malacofauna was significantly correlated with region-to-region geographical distance. This suggests the influences of environments and paleontological events specific to each current surrounding each region. (3) The pattern of dissimilarity between the Inland Sea and the other sea areas greatly changed from the 1970s to 1980s. This was due in part to the following density fluctuations on some shores in the Inland Sea: increases in the non-indigenous mussel Mytilus galloprovincialis, cultured oyster Crassostrea gigas, and littorinid Littorina brevicula reportedly abundant in sheltered areas; and decreases in the littorinid Nodilittorina radiata reportedly abundant in exposed areas and muricid gastropods Thais spp. showing imposex. This suggests an increase of anthropogenic perturbations during the study period in the Inland Sea which is surrounded by large cities and ports. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.