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.     

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.     

How wave exposure, group size and habitat complexity influence foraging and population densities in fishes of the genus Halichoeres (Perciformes: Labridae) on tropical rocky shores

Wave exposure and habitat complexity have been used to explain variations in the distribution patterns and behavior of many reef fishes. This study analyzed the influence of both factors on density and foraging activity, and the influence of group size on foraging in three species of the genus Halichoeres (Labridae) on tropical rocky shores. It was shown that initial phases (IP) and terminal phases (TP) Halichoeres, were influenced by wave exposure, although foraging in Halichoeres brasiliensis TP was not influenced by wave exposure. The IP in all three species were positively associated with rugosity and algal cover while the TP was positively associated with depth. Habitat complexity also influences foraging in these species. Group size influenced foraging activity, except in H. brasiliensis TP. We also found variations in microhabitat patches used for foraging between species and differences in the stomach contents between species and phases, showing that functional roles change in response to ontogenetic shifts.     

Differential meiospore size and tolerance of ultraviolet light stress within and among kelp species along a depth gradient

 Kelp are differentially stratified along a gradient of UV exposure (as a function of water depth). The role of ultraviolet light in seaweed zonation has not been fully explored. This study found a significant meiospore size difference within and among the kelp species examined: Pterygophora californica Ruprecht (high to mid-subtidal), Macrocystis integrifolia Bory (high to mid-subtidal), Laminaria groenlandica Rosenvinge (high-subtidal), Alaria marginata Postels and Ruprecht (low-intertidal) and Hedophyllum sessile Setchell (mid-intertidal). This size difference was correlated with the depth distribution of adult plants, with the largest meiospores originating from shallow-dwelling adult kelp exposed to high UV light. Under UV stress in the laboratory, meiospores from adults growing in high-UV environments displayed greater germination and survival rates than the progeny of adult kelp occupying lower-UV environments. This suggested that in Barkley Sound, British Columbia, Canada, differential tolerance to UV (possibly determined by meiospore size) may limit the upper settlement position of kelp species and individuals. Tolerance to UV may be an important determinant of kelp zonation on rocky shores. Received: 16 June 1998 / Accepted: 6 January 2000     

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.     

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.     

Zonation of the interstitial polychaeteStygocapitella subterranea (Stygocapitellidae) in European sandy beaches

The zonation ofStygocapitella subterranea Knoellner (Polychaeta, Stygocapitellidae) in sandy beaches is compared for several localities along the European coasts. In tidal beaches the species occupies a zone near average high tide level (AHTL), whereas in non-tidal beaches it is found at a fairly long distance from the water line. The regularities observed inS. subterranea indicate that the interstitial fauna may prove well suited for the establishment of a general scheme of zonation in sandy beaches analogous to that for rocky shores proposed byStephenson andStephenson (1949).     

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.     

Demography and phenology of the intertidal green algaCodium setchellii: The enigma of local scarcity on sand-influenced rocky shores

Demographic and phenological attributes that characterize rare and scarce species have not been extensively investigated for marine benthic organisms. From 1984 to 1987, 1 investigated the population ecology of the low intertidal, psammophytic green algaCodium setchellii Gardner that forms low-density populations on sand-influenced rocky benches on N.E. Pacific shores. To address why the alga is not more common in the habitat in which it is most abundant, I quantified sand fluctuations and population dynamics ofC. setchellii at several sites along the central coast of Oregon, USA. Sand movement was highly unpredictable both spatially and temporally on a local scale. AlthoughC. setchellii became fertile in late autumn to winter, the alga and rocky surfaces were often deeply buried by sand in these seasons. Algal recruitment was also spatially and temporally variable, and mortality of small thalli was high. Thallus growth was generally slow, and the alga had the capacity to persist at a given size for long periods. On sand-influenced rocky shores, mortality of largethalli was low, with some thalli persisting for several years. In laboratory experiments,C. setchellii lost substantial biomass during 4 to 5 wk sand burials, whereas two sympatric species of psammophytic algae lost little to no biomass. The within-site scarcity ofC. setchellii may be a consequence of (1) largely unpredictable environmental conditions that reduce algal recruitment and survival, and (2) apparent lack of specialized anatomical or reproductive adaptations to persist on sand-scoured rocky shores. The alga's persistence may be due to its slow growth and long lifespan.     

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.     

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     

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.     

Urban marine ecology in southern California: the ability of riprap structures to serve as rocky intertidal habitat

Increasing human populations along marine coastlines has lead to increasing urbanization of the marine environment. Despite decades of investigations on terrestrial ecosystems, the effect of urbanization on marine life is not well understood. Riprap is the rocky rubble used to build jetties, breakwaters, and armored shorelines. Roughly 30% of the southern California shoreline supports some form of riprap, while 29% of the shoreline is natural rocky substrate. Astonishingly few studies have investigated this anthropogenic rocky habitat even though it rivals a natural habitat in area on a regional scale along a coastline that has been extensively studied. In this study, I compared the diversity and community structure of exposed rocky intertidal communities on four riprap and four natural sites in southern California. I ask the following questions: (1) does diversity or community composition differ between intertidal communities on riprap and natural rocky habitats in southern California, (2) if so, which organisms contribute to those differences, (3) which physical factors are contributing to these differences, and (4) do riprap habitats support higher abundances of invasive species than natural habitats? On average, riprap and natural rocky habitats in wave exposed environments in southern California did not differ from each other in diversity or community composition when considering the entire assemblage. However, when only mobile species were considered, they occurred in greater diversity on natural shores. These differences appear to be driven by wave exposure. The presence of invasive species was negligible in both natural and riprap habitats.     

Blue mussels, Mytilus edulis, at the edge of the range: population structure, growth and biomass along a salinity gradient in the north-eastern Baltic Sea

The blue mussel, Mytilus edulis, is the most conspicuous animal species in the northern Baltic rocky sublittoral. In the studied area the species lives at the margin of its salinity tolerance. Although dwarfed by the low-salinity conditions, blue mussels in the northern Baltic are very abundant and have a decisive role in the benthic and pelagic ecosystems. We studied abundance, size distribution, biomass and growth rate of blue mussels along a 270 km salinity gradient in the northern Baltic Sea. Samples (n=317, 1998-1999) from moderately exposed and exposed rocky shores at seven study areas were collected in the southern Archipelago Sea in the west and into the central Gulf of Finland in the east, where the species is becoming increasingly rare. The results show a marked decline in mean mussel size and biomass from the saline west to the less saline east. The growth rate also decreased with lower ambient salinity. However, abundance of small mussels was considerably higher in the central and eastern parts of the study area. Vertically, the highest biomass was recorded at intermediate depths (5 and 8 m), being lower at both shallower (3 m) and deeper bottoms (12 m). It is concluded that salinity is the foremost factor determining size structure and growth rate among populations within the area. The results suggest that predation further influences the population structure of blue mussels living at the edge of their range in the central Gulf of Finland ultimately set by their salinity tolerance.     

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.     

Cryptobenthic fishes of the “Ciclopi Islands” marine reserve (central Mediterranean Sea): assemblage composition,structure and relations with habitat features

The fish assemblage of the Ciclopi Islands marine reserve, which is located along the eastern coast of Sicily (Central Mediterranean), was unknown until our study provided insight into the cryptobenthic fish assemblage inhabiting this integral reserve zone. Visual census surveys were periodically conducted along the littoral rocky shores of Lachea, the largest island of the Ciclopi archipelago. The fish assemblage, which comprised 20 species belonging to Blenniidae, Gobiidae, Tripterygiidae, Scorpaenidae and Gobiesocidae, was dominated by Scorpaena maderensis and Trypterigion delaisi. Species richness (S) did not show significant differences between the northern, eastern, southern and western island coasts, whereas both species diversity (H) and evenness (J) were significantly higher on the western coast than on the northern coast. The effect of the island coast on fish density was not significant with regard to the whole assemblage or to single species, except for S. maderensis and T. delaisi. The pattern of variation of assemblage parameters was mainly positively correlated with microhabitat variables, especially substratum composition and type of cover. The best predictors of fish total density were bottom slope, rugosity and substratum composition. Variations in relative species density were specified by a different combination of macro- and microhabitat variables. The density pattern of S. maderensis showed a positive relationship with depth. In T. delaisi, the influence of slope was interrelated to that of exposure to waves. This species preferentially inhabited flat habitat, sheltered from the open sea and covered by small boulders, but also more-exposed rockwalls. The population densities of Tripterygion melanurus and Lipophrys trigloides were negatively affected by depth, thus reflecting their preference for the upper infralittoral. T. melanurus was associated with a more complex habitat (stones and boulders), whereas L. trigloides was mainly censused on a narrow strip of solid rock between the waterline and 1 m depth.Communicated by R. Cattaneo-Vietti, Genova     

Differences in relative abundance and size structure of the sea stars Pisaster ochraceus and Evasterias troschelii among habitat types in Puget Sound, Washington, USA

We surveyed patterns in the relative abundance and size structure of the sea stars Pisaster ochraceus and Evasterias troschelii in five habitat types of varying structural complexity and prey availability (sand/cobble, boulder, and rocky intertidal; pilings; and floating docks) in Puget Sound and the San Juan Islands, Washington. For both species, small sea stars were most abundant in the most structurally complex habitat type (boulder), where they occurred almost exclusively under boulders during low tide. Larger individuals became more abundant as structural complexity decreased, occurring more frequently in open habitat types (rocky shores, pilings, and docks) known to have greater abundances of prey resources. Gull foraging observations and experiments demonstrated that exposed small sea stars of both species were highly vulnerable to predation, suggesting that small sea stars require structural complexity (crevice microhabitat) as a predation refuge. Large sea stars, once attaining a size refuge from predation, appear to migrate to more exposed habitat types with more abundant food resources. These results suggest parallel ontogenetic habitat shifts in two co-occurring consumer species related to a shared predation risk at early life stages and demonstrate how the relative importance of top-down and bottom-up processes may differ with ontogeny.     

Area-specific temporal changes of species composition and species-specific range shifts in rocky-shore mollusks associated with warming Kuroshio Current

Increases of low-latitude species in various sea areas and poleward shifts of ranges of many species have been reported and linked to warming environments. To examine the generality of these trends for mollusks, we conducted 7 quadrat surveys during 1978–2006 on Pacific rocky shores in Japan (26.6–45.0°N). Results showed that the dominance of southern species increased on 11 of the 15 shores in a southern, warming sea area but on only 1 of the 6 shores in a northern area with no warming trend. The latitudes of species ranges increased on average but showed large interspecific variations which showed a weak, positive correlation with interspecific taxonomic distance. The differences of these results from the previously reported trends are discussed in relation to the unique current patterns in our study area and the phylogenetic constraints of species-specific responses to a warming environment.     

Wave exposure,swimming performance,and the structure of tropical and temperate reef fish assemblages

We examined the relationship between swimming performance, wave exposure, and the distribution patterns of labrids on temperate rocky reefs, in comparison with previous functional analyses of a tropical assemblage. Visual censuses of the distribution and abundance of labrids across two major gradients of wave exposure (depth and aspect to prevailing winds) were made at two offshore islands near Port Stephens, New South Wales, Australia. Distinct shifts in species composition and abundance were evident between high and low wave exposure habitats on temperate rocky reefs, particularly between deep and shallow habitats on exposed reef fronts. The swimming performances of temperate labrids were assessed through examination of pectoral fin shape (aspect ratio) and in situ swimming speeds. A diversity of pectoral fin morphologies was exhibited within this temperate assemblage, ranging from rounded to tapered fins (aspect ratios of 0.52 and 1.43, respectively). Fin shape was strongly correlated (Pearsons correlation 0.884, P<0.001) with swimming speed (ranging from 1.05 and 3.06 body lengths s^–1), in a relationship comparable to that observed in tropical labrids. Inter-specific differences in swimming ability provided some explanation for differences in the distribution and abundance of temperate labrids in relation to wave exposure. However, our findings suggest that although coral reef labrids appear to predominantly use high aspect-ratio fins to successfully occupy wave-exposed habitats, temperate labrids appear to be using an enhanced swimming ability through increased body size.Communicated by G.F. Humphrey, Sydney     

Amphipods and isopods in the rocky intertidal: dispersal and movements during high tide

Animals in the intertidal, both mobile and sessile, generally exhibit some zonation pattern, in which each species shows a preference for, or is confined to, some height levels. The study of zonation patterns is, however, almost exclusively based on surveys made during low tide, when many animals are relatively inactive. We studied zonation patterns of amphipods and isopods on rocky shores in southwestern Iceland, both by traditional sampling at low tide as well as by sampling during high tide. The distributional patterns seen at high tide differed significantly from that at low tide. One amphipod, Anonyx sarsi, was common around baits at all levels at high tide but absent from the intertidal at low tide. Several other species were either relatively more common or tended to be recorded higher, or in one instance, lower on the shore when the tide was in than at low tide. There was also evidence of some species changing habitats within the intertidal with the tidal cycle. Many species, however, moved little away from their respective zones occupied at low tide, and for some species, including some capable of rapid swimming, very limited mobility was indicated. We conclude that low-tide surveys of the intertidal give an incomplete picture of the community structure, and even key species may be missed in such surveys.Communicated by L. Hagermann, Helsingør