Do mussel patches provide a refuge for algae from grazing gastropods?

On rocky shores, cover of macroalgae is often greater growingepibiotically on mussels compared to algae growing directlyattached to rock. A survey of two shores on the east coast ofIreland confirmed that mussel beds contained greater percentagealgal cover and more diverse algal assemblages compared to thoseon rock. The reasons for this difference are not clear. It hasbeen suggested that mussel beds provide a refuge for algae fromgrazing gastropods. Surprisingly, we found no evidence to supportthis. Using wax discs, gastropod grazing patterns were foundto be similar within the mussel beds as on rock. The musselbeds do not appear to provide a refuge for algae from grazingactivity at this scale and we suggest other possible mechanismsfor the prevalence of epibiotic algal cover on mussels. Intertidalgrazers may in fact affect the epibiotic algae on mussels andthereby affect indirectly the persistence of mussel beds. (Received 14 May 2007; accepted 20 October 2007)     

Impacts of sewage outfalls on rocky shores: Incorporating scale,biotic assemblage structure and variability into monitoring tools

Coastal systems, such as rocky shores, are among the most heavily anthropogenically-impacted marine ecosystems and are also among the most productive in terms of ecosystem functioning. One of the greatest impacts on coastal ecosystems is nutrient enrichment from human activities such as agricultural run-off and discharge of sewage. The aim of this study was to identify and characterise potential effects of sewage discharges on the biotic diversity of rocky shores and to test current tools for assessing the ecological status of rocky shores in line with the EU Water Framework Directive (WFD). A sampling strategy was designed to test for effects of sewage outfalls on rocky shore assemblages on the east coast of Ireland and to identify the scale of the putative impact. In addition, a separate sampling programme based on the Reduced algal Species List (RSL), the current WFD monitoring tool for rocky shores in Ireland and the UK, was also completed by identifying algae and measuring percent cover in replicate samples on rocky shores during Summer. There was no detectable effect of sewage outfalls on benthic taxon diversity or assemblage structure. However, spatial variability of assemblages was greater at sites proximal or adjacent to sewage outfalls compared to shores without sewage outfalls present. Results based on the RSL, show that algal assemblages were not affected by the presence of sewage outfalls, except when classed into functional groups when variability was greater at the sites with sewage outfalls. A key finding of both surveys, was the prevalence of spatial and temporal variation of assemblages. It is recommended that future metrics of ecological status are based on quantified sampling designs, incorporate changes in variability of assemblages (indicative of community stability), consider shifts in assemblage structure and include both benthic fauna and flora to assess the status of rocky shores.     

The shifting balance of littoral predator-prey interaction in regimes of hydrodynamic stress

Above lowshore levels of wave-beaten rocky shores, desiccation from tidal exposure and hydrodynamics stresses from wave action are thought to create refuges from predation, allowing concentrations of sedentary prey such as mussel beds. Underwater time-lapse photography on rocky shores in Southern California revealed that dense aggregations of spiny lobsters prey on mussels during nocturnal high tides. In contradiction of the refuge hypothesis, the densest aggregations occurred on midshore levels of the most wave-exposed site, a semi-protected site showed intermediate densities, and a protected site showed only sparse numbers of lobsters. On wave-beaten shores, the lobsters' high mobility and rapid prey handling allowed them to exploit intertidal prey in the brief period at extreme high tide, when both desiccation and hydrodynamic stresses were at a minimum. The spatial differences in lobster densities were, however, positively related to the recruitment rates of juvenile mussels, the preferred prey. A field experiment demonstrated that predation by lobsters within a mussel bed affects the age/size structure of the bed without changing primary percent coverage. Therefore, concentrations of adult prey on some wave-swept sites appear to result from elevated rates of prey recruitment that surpass rates of predation, rather than absolute refuges from predation.     

Lift as a mechanism of patch initiation in mussel beds

The mussel Mytilus californianus is the dominant competitor for space in the mid-intertidal zone of wave-swept rocky shores in the Pacific Northwest where it forms extensive tightly packed beds. The rate at which patches are formed in these beds, can play an important rôle in community ecology by controlling the establishment and persistence of fugitive species. Despite the biological importance of physical disturbance, the mechanism of patch initiation has not been adequately explained. Battering by logs can create patches, but is the predominant mechanism only on shores near active logging sites. In other areas, it has been speculated that the hydrodynamic forces associated with storm waves somehow cause patches to form. However, the forces acting along the direction of flow — drag and the acceleration reaction — are unlikely to initiate patch formation. Here, it is suggested that fluid-dynamic lift forces imposed on mussel beds by breaking waves are sufficient to dislodge individual mussels and trigger patch formation. Arguments are presented suggesting that the likelihood of dislodgment by lift is consistent with the observed rate of patch formation in the absence of log battering.     

Trophic structure of macrobenthic communities on the Portuguese coast. A review of lagoonal,estuarine and rocky littoral habitats

This work is based on a compilation and treatment of data obtained on several studies regarding the macrobenthos trophic structure, carried out in different habitats of the Portuguese coast: a lagoon, rocky subtidal shores and estuarine eelgrass beds. Macrobenthic organisms were assigned to four distinct trophic groups (filter feeders, detritivores, herbivores and carnivores). Detritivores were the dominant trophic group in soft-bottom communities. Filter feeders dominated in subtidal rocky shores while eelgrass communities were equally represented by detritivores and herbivores. Current intensity and sediment deposition are discussed as factors affecting the observed distribution. Nevertheless, different sampling methods used in these studies and the lack of information on feeding habits of some species can also influence the results obtained.     

Large scale patterns and trophic structure of southern African rocky shores: the roles of geographic variation and wave exposure

In this study we revise the biogeographic delimitation, and large-scale patterns of community structure of the intertidal rocky shores of southern Africa. We use binary (presence/absence) and per-species biomass data collected at fifteen localities and thirty-seven different rocky sites, encompassing the shores of southern Namibia, South Africa and southern Mozambique. Multivariate analyses revealed that the shores of southern Africa (south of 25°) can be divided into three main biogeographic provinces: the west coast or Namaqua province, the south coast or Agulhas province and the east coast or Natal province. The biomass structure of the intertidal rocky shores communities of southern Africa varied at a large scale, corresponding to biogeographic differences, while local-scale variation accorded with the intensity of local wave action. The average biomass of west coast communities was on average significantly greater than that of the south and east provinces. At a local scale, the community biomass on exposed shores was an order of magnitude greater than on sheltered shores, within all biogeographic provinces. Semi-exposed shores exhibited intermediate average biomass. The trophic structure of these communities varied significantly with wave action: autotrophs, filter-feeders and invertebrate predators were more prevalent on wave exposed than sheltered shores, whereas grazers were more abundant on sheltered and semi-exposed shores. Exposed shores were consistently dominated by far fewer species than semi-exposed and sheltered shores, independently of biogeographic differences. Within all biogeographic provinces semi-exposed and sheltered shores were more diverse than exposed shores. West coast intertidal communities therefore had high levels of biomass, but were consistently species-poor. Several working hypotheses that could explain these large and small-scale patterns are presented.     

Response of macroalgae and macroinvertebrates to anthropogenic disturbance gradients in rocky shores

The compliance of macroalgal and macroinvertebrate assemblages to anthropogenic disturbance gradients (e.g., nutrient enrichment) was investigated at intertidal rocky shores. Macroalgae and macroinvertebrates presented parallel behavior, both showing shifts in the communities’ structural variation along the gradients, in which an higher number of opportunistic species (and higher abundances) were found in more stressful sites (close to the disturbance source), in contrast to less disturbed sites (far from the disturbance source), which showed higher presence of more sensitive species (and higher abundance of several of them).The macroinvertebrate abundance and taxonomic composition, which are parameters required by the Water Framework Directive (WFD) to be included in tools for the ecological quality status assessment, responded to the disturbance gradient. Results suggest that the macroinvertebrate biological element might be considered an indicator of disturbance in intertidal rocky shores as good as the macroalgae, and therefore the development of a specific methodology based solely on benthic macroinvertebrates of rocky shores, presently a gap in the ecological quality status assessment for the WFD, seems feasible.     

Intertidal assemblages on seawalls and vertical rocky shores in Sydney Harbour,Australia

Abstract Understanding the ecological role of artificial structures, such as seawalls, in shallow coastal waters is necessary in order to plan sound strategies of conservation and management of natural habitats. In Sydney Harbour (NSW, Australia), about 50% of the foreshore is made of retaining seawalls This study evaluates the changes caused to natural assemblages of organisms by these structures, by comparing intertidal assemblages between seawalls and vertical rocky shores. The following hypotheses were tested: that assemblages on seawalls would differ from those on rocky shores at mid‐, but not at low‐shore levels; where assemblages differ between habitats, there would be differences in cover/abundances of widespread species; patterns would be consistent among locations and through time; the variability of assemblages at the scales of 10s of cm and metres would differ between seawalls and rocky shores at mid‐ and low‐shore levels. To test these hypotheses, assemblages on seawalls and rocky shores were sampled at three locations, at roughly 4‐monthly intervals, over a period of about 18 months. Results indicated that mid‐shore assemblages on seawalls were different from those on rocky shores, but this was not the case at low‐shore levels. Few taxa were unique to either habitat. Cover of common species of algae and sessile animals and abundances of mobile grazers were variable with few consistent patterns. Variability at the scales sampled differed between habitats and heights on the shore. Seawalls and rocky shores, in general, supported a similar suite of species, but patterns of abundance and variation differed among locations and from height to height in each habitat. The implications of these findings for the future management of seawalls are briefly considered.     

BENTHIC METABOLISM ON A SHELTERED ROCKY SHORE: ROLE OF THE CANOPY IN THE CARBON BUDGET1

While the importance of canopy‐forming algae in structuring ecosystems is recognized, their role in the carbon budget is still not well understood. To our knowledge, no measurements of rocky shores primary production and respiration under emersion periods have been carried out in situ. A benthic chamber coupled to a CO₂‐infrared gas analyzer was used to measure gross primary production and respiration on the Ascophyllum nodosum (L.) Le Jol. zone of a sheltered rocky shore in Brittany, France. Over a year of monthly measurements on the zone with and without the A. nodosum canopy showed fairly high production and respiration values for the global community as well as carbon fluxes due to the canopy that largely dominated the benthic metabolism of the zone. The strong canopy respiration relative to the primary production also suggested a high metabolic activity by microscopic heterotrophs on the surface of the alga. Both the canopy and the understory annual primary production and respiration were under the control of light and temperature seasonal variations. Finally, the range of the amount of carbon produced on the A. nodosum zone during diurnal emersions was estimated. Additional measures accounting for the day–night cycles and seasonal light variations over an entire tidal cycle are, however, necessary to establish an annual carbon budget. Such measures using the benthic chamber together with complementary techniques would allow a better understanding of the functioning of sheltered rocky shores.     

Tectonically controlled late cretaceous terrestrial to neritic deposition (Northern Calcareous Alps, Tyrol, Austria)

Summary The Turonian to Santonian terrestrial to neritic succession (Lower Gosau Subgroup) in the Northern Calcareous Alps of the eastern part of the Tyrol, Austria, provides an example for deposition on a compartmentalized, narrow, microtidal to low-mesotidal, wave-dominated, mixed siliciclastic-carbonate shelf. The shelf was situated in front of a mainland with a relatively high, articulated relief, and underwent distinct changes in facies architecture mainly as a result of tectonism. The investigated succession was deposited above a deeply incised, articulated truncation surface that formed when the Eo-Alpine orogen, including the area of the future Northern Calcareous Alps, was uplifted and subaerially eroded. Distinct facies associations were deposited from (1) alluvial fans and fan deltas, (2) rivers, (3) siliciclastic lagoonal to freshwater marsh environments, (4) areally/temporally limited carbonate lagoons, (5) transgressive shores, (6) siliciclastic shelf environments, and (7) an aggrading carbonate shelf. During the Turonian to Coniacian, the combination of high rates of both subsidence and sediment accumulation, and a narrow shelf that was compartmentalized with respect to (a) morphology of the substratum, (b) fluviatile input of siliciclastics and contemporaneous input of carbonate clasts from fan deltas, (c) deposition of shallow-water carbonates, and (d) water energy and-depth gave rise to an exceptionally wide spectrum of facies as a distinguishing feature of the succession. With the exception of facies association 7, which formed only once, depositional sequences in the Turonian to Coniacian interval contain all of the facies associations 1 to 6. During Turonian to Coniacian times, the shelf was microtidal to low-mesotidal, and was dominated by waves, storm waves and storm-induced currents. In vegetated marshes, schizohaline to freshwater marl lakes existed. Transgressions occurred onto fan deltas and in association with estuaries, or in association with gravelly to rocky shores. The transgressive successions, including successions deposited from transgressive rocky carbonate shores, are overlain by regressive successions of shelf carbonates or shelf siliciclastics. Deposition of shallow-water carbonates generally occurred within lagoons and over short intervals of time. A „catch-up” succession of shelf carbonates about 100 m thick accumulated only in an area protected from siliciclastic input. In its preserved parts, the Turonian to Coniacian succession does not record deposition adjacent to major active faults. Lateral changes in thickness result mainly from onlap onto the articulated basal truncation surface. Subsidence most probably was controlled by major detachment faults outside the outcrop area, and/or was distributed over a wide area in association with secondary faults above the major detachments. During Coniacian to Early Santonian times, both the older substratum and the overlying Turonian-Coniacian succession were subaerially exposed, faulted and deeply eroded. The following Early Santonian transgression ensued with rocky carbonate shores ahead of a sandy, narrow shoreface-inner shelf environment and a deeper shelf with intermittentlydysaerobic mud. The transgression was associated with the influx of cooler and/or nutrient-rich waters, and heralds an overall deepening. Still during the Early Santonian, the deepening was interrupted by another phase of subaerial exposure. Subsequently, a short phase of shelf deposition was terminated by deepening into bathyal depths.     

Ecological interactions in the provision of habitat by urban development: whelks and engineering by oysters on artificial seawalls

Abstract Increases in human population cause increased urbanization of most habitats, including the shoreline. This has many consequences for coastal environments, in particular the trend for artificial structures, such as seawalls, to replace natural habitats. Seawalls and natural shores support many of the common intertidal species, but others important on rocky shores are absent from or rare on many seawalls. The whelk Morula marginalba Blainville is an abundant and important predator on rocky shores of south‐eastern Australia, but is infrequently recorded on artificial substrata. In Sydney Harbour, where the Sydney rock oyster (Saccostrea glomerata Gould) was locally abundant, densities of M. marginalba on some seawalls appeared to be similar to those on rocky shores and to be larger than where there were few oysters. We sampled densities and sizes of whelks in four habitats, predicting and corroborating that: (i) on seawalls with many oysters, there would be more whelks than on seawalls with few oysters; (ii) where there are many oysters, densities of whelks would be similar on seawalls and rocky shores; and (iii) whelks would be larger where oysters were abundant. Growth and survival of whelks were measured to test hypotheses from the observed differences in size and density. Survival was greater in habitats with many oysters, which could explain differences in density, but size‐specific differences in survival could not explain differences in size among habitats. On seawalls but not on rocky shores, slower growth could explain the smaller size of whelks where there were few oysters. Seawalls provide important habitat for M. marginalba, but only via their indirect effects, mediated by oysters. These interactions cannot be predicted from those on natural rocky shores. Predicting how developed areas provide suitable habitat, either in management of conservation or in assessments of potential impacts clearly depends on understanding the roles of biogenic habitats.     

The influence of coastal upwelling on the functional structure of rocky intertidal communities

Summary Relationships between organisms at all trophic levels are influenced by the primary productivity of the ecosystem, and factors which enhance rates of primary production may modify trophic relationships and community structure. Nutrient enrichment of intertidal and nearshore waters leads to enhanced production by intertidal algae, and it was hypothesized that where rocky shores are washed by nutrient-rich upwelled waters, the intertidal communities should show a characteristic functional structure, based on the effects of enhanced primary production. Study sites were chosen on rocky shores in southern Africa, central Chile and the Canary Islands, in areas with and without coastal upwelling, and mid-shore community structure at these sites was analysed in terms of the abundance of certain functional guilds of organisms.It was found that algal cover and the biomass of herbivorous limpets supported per unit area on rocky shores were significantly greater in regions of coastal upwelling than in regions where upwelling did not occur. Ground cover by sessile filter-feeding organisms was significantly greater on shores in non-upwelled areas. However, correspondence analysis showed no functional aspect of intertidal community structure that was characteristic of coasts washed by upwelled waters. Primary reasons for this are probably the large variations in the nature of nutrient enrichment that accompanies upwelling, and in the nutrient status of non-upwelled areas. Other factors are man's exploitation of intertidal organisms and differences in the genetic origins of the intertidal species involved.     

Food supply, grazing activity and growth rate in the limpet Patella vulgata L.: a comparison between exposed and sheltered shores

The limpet Patella vulgata L. is an important microphagous grazer on intertidal rocky shores of north-west Europe, occurring across the wave exposure gradient. Groups of P. vulgata were selected at mid-tide level of two exposed shores and two sheltered, fucoid dominated shores on the Isle of Man, British Isles, and manipulated to equivalent densities and population structure. The level of grazing activity and growth rate were determined over a 1-year period. At the same time, the abundance of epilithic microalgae, measured as the concentration of chlorophyll a, was determined as an estimate of food supply. Microagal abundance showed a seasonal pattern in both exposed and sheltered conditions, with higher levels in winter compared to summer. In both seasons, the microalgal resource was more abundant on the sheltered shore studied. The level of grazing activity in P. vulgata showed a seasonal pattern on the exposed but not the sheltered shores. Averaged over the year, grazing activity on the exposed shores was over double that on sheltered shores. Thus, in sheltered conditions, food supply for limpets was high and grazing activity low; in exposed conditions, food supply was low and grazing activity high. The growth rate of P. vulgata, measured as increase in shell length, showed no significant difference between exposed and sheltered shores. Growth rate was also determined in P. vulgata at natural densities. Although the overall density declined with decreasing exposure to wave action, the density per unit area of grazeable substance was higher in shelter. In these populations, the mean growth rate was over twice as high on exposed compared to sheltered shores.     

Grazing by two species of limpets on artificial reefs in the northwest Mediterranean

The extensive presence of artificial reefs in marine coastal habitats demands a better understanding of the extent to which these structures can be considered surrogates of natural rocky shores for populations of plants and animals. The primary aim of this study was to test the hypothesis that removing limpets from the midlittoral of artificial breakwaters in the northwest Mediterranean led to changes in assemblages similar to those observed on rocky shores in the same area. Orthogonal combinations of the presence/absence of two species of limpets, P. aspera and P rustica, were produced using manual removals from June 1997 to February 1998. To test the hypothesis that the effects of limpets were variable at spatial scales comparable to those investigated on rocky shores, we repeated the experiment at two locations tens of kilometres apart, and on two reefs within each location a few kilometres apart. The results revealed strong and relatively consistent negative effects of limpets on filamentous algae, whereas interactions with other members of assemblages were complex and variable. Several taxa (Cyanophyta, encrusting and articulated coralline algae, Ralfsia and Rissoella) were abundant at one location but nearly absent at the other. This large-scale variability in patterns of distribution generated inconsistencies in the effects of limpets between locations. Within locations, several effects of P. aspera and P. rustica were observed, ranging from independent effects on some organisms, to additive or interactive effects on others. Apparently, the removal of filamentous algae by limpets resulted in positive indirect effects on Ralfsia and Rissoella. Collectively, these effects were comparable to those described for rocky shores in the northwest Mediterranean. The processes accounting for large-scale variation in grazing, however, appeared different between the natural and the artificial habitat.     

Community Structure of Fishes and Habitat Complexity on a Tropical Rocky Shore

The relationship between the variables of reef fish community structure (fish richness, fish diversity and total number of fishes) and those of habitat complexity (total surface area, substratum diversity, topographic complexity, number of holes, percent cover of turf algae, zoanthids, millepores, massive corals, bare rock, encrusting calcareous algae, urchins, other sessile organisms and Sargassum) were examined on three different rocky shores (FA, FB and FT) at Arraial do Cabo, a tropical region located on Brazilian southeastern coast (23° S, 42° W). Fish abundance and vertical distribution were assessed by a visual census technique using strip transects. Percentage cover of benthic organisms and other substratum types were calculated by replicated transects using a chain laid down on the substratum. Topographic complexity was determined by the chain link method and number of holes were estimated by direct counts on replicated transects. More than 91 fishes belonging to 37 families were seen in all study sites during one year of visual census effort. FA and FB sites had similar fish community structure and habitat complexity characteristics, while FT showed different habitat characteristics and higher fish diversity and richness, plus total number of fishes. Vertical distribution of fishes along the rocky shores studied seemed to be predictable and was determined by factors such as feeding habits and behavior, availability of refuges and social interactions. The habitat variables that best explained the higher diversity and number of fishes observed in FT site were total surface area of rocky shores and the abundance of benthic sessile invertebrates; these conditions were typical of rocky shores more exposed to wave surge.     

Intertidal mussel beds from the South-western Atlantic show simple structure and uniform appearance: does environmental harshness explain the community?

Communities of the rocky mid-intertidal zone of the South-western Atlantic are uniform in appearance, dominated by dense monocultures of small-size mussels (Brachidontes rodriguezii and Perumytilus purpuratus). To explain this, two hypotheses have been advanced in the literature: environmental harshness due to high potential evaporation and historical contingency after the Last Glacial Maximum. In this study of Uruguayan and Argentine shores, we address the implications and predictions of these two hypotheses from a biogeographic perspective by studying the regional distribution and composition of mid-intertidal mussels. We conducted an extensive latitudinal sampling survey (21 locations, 34–54°S), along with a compilation of available information on mussel bed composition and mussel predators present along the coastline. Then we constructed latitudinal profiles of ecologically significant environmental variables with specific emphasis on potential evaporation, a proxy for desiccation stress. The results show that mussel beds are composed of two species of small mussels, which coexist over a biogeographic transition zone (40–42°S) related to sea surface water temperature. The distribution of mussels along the coastline studied is not consistent with the environmental harshness hypothesis. In addition, in the Central Patagonian zone (44–50°S), two invertebrate predators also inhabit the intertidal rocky shores. However, these localities showed higher environmental harshness (potential evaporation rate) than non-Patagonian localities. We suggest that further attention should be given to historical contingency in order to advance towards a hypothesis consistent with current knowledge on the post-glacial biogeographic history of the South-western Atlantic.     

Colonization and reef growth on a Late Pleistocene rocky shore and abrasion platform in Western Australia

A low, rocky shoreline and attached abrasion platform of Late Pleistocene age are marked by a sharp disconformity within the Tamala Limestone Formation, exposed at Cape Burney facing the Indian Ocean near Geraldton, Western Australia. Colonization by an intertidal to shallow subtidal biota dominated by encrusting coralline red algae, oysters, and tube-dwelling worms occurs on a sandstone surface with a channeled topographic relief of 20–30 cm. The encrusting cup coral Rhizotrochus tuberculatus also is present, and this report details what is probably the first fossil occurrence of that species. The ancient rocky shore above this level retains trace fossils characteristic of a boring barnacle, probably belonging to Lithotrya. Herein named the Cape Burney sandstone, the distinctive unit on which the disconformity sits is assigned member status within the Tamala Limestone. Shell beds with a diversity of 35 species, dominated in volume by robust gastropods such as Turbo intercostalis and T. torquatus , thinly drape portions of the disconformity surface. Succeeding the shell drapes is a reef limestone with a maximum thickness of more than 2 m. The limestone is a massive accumulation of collapsed but otherwise mostly undisturbed coral fronds belonging primarily to a robust species of Acropora. Herein named the Bootenall limestone, this unit is assigned member status within the Tamala Limestone. Based on an analysis of electron spin resonance (ESR) from Acropora samples, the fringing reef developed between 120 ka and 132 ka, in the terminal stage of coastal transgression during the last interglacial period (Oxygen Isotope Substage 5e). □ Ancient rocky shores, intertidal abrasion platform, hard-substrate colonization, reef succession.     

Trends in abundance of rocky intertidal seaweeds and filter feeders across gradients of elevation,wave exposure,and ice scour in eastern Canada

On NW Atlantic rocky shores, the main basal organisms in intertidal communities are seaweeds (Ascophyllum nodosum, Fucus spp. and Chondrus crispus) and filter feeders (barnacles, Semibalanus balanoides, and mussels, Mytilus spp.). Their ecology has been extensively studied in New England (United States), but knowledge gaps exist for northern shores, which are subjected to stronger environmental stress. Therefore, we studied the above organisms on Canadian shores. We quantified the summer abundance of these seaweeds and filter feeders across full vertical (intertidal elevation) and horizontal (wave exposure and winter ice scour) environmental gradients on the Gulf of St. Lawrence and open Atlantic coasts of Nova Scotia. At the regional scale along the open Atlantic coast, seaweeds showed similar abundances in Nova Scotia than values reported for New England. However, both filter feeders were considerably less abundant in Nova Scotia. At the local scale in Nova Scotia, intense winter ice scour (which only occurs on the Gulf of St. Lawrence coast) was associated with a very low abundance of all species except barnacles. Spatial trends in Nova Scotia were similar to patterns known for certain species elsewhere, such as A. nodosum being almost restricted to sheltered habitats, regardless of elevation, and C. crispus being almost restricted to low elevations, regardless of exposure. Other trends were, however, characteristic of Nova Scotia, such as C. crispus being frequent at low elevations in exposed habitats, unlike in New England, where mussels often predominate there because of competitive advantages. In Nova Scotia, mussels were always restricted to cracks and crevices, unlike in New England, where they form extensive intertidal beds on exposed shores. The direct effects of increased environmental stress and indirect effects through altered interspecific interactions might explain the regional differences in local species distribution, which will require experimental validation. Handling editor: K. Martens     

Enhancement of habitat heterogeneity and species richness on rocky shores inundated by sand

Summary Many rocky shores are subject to periodic inundation by sand, which is often thought to reduce species richness by eliminating organisms intolerant of sand scour or sand smothering. However, regular disturbance (e.g. inundation) should promote richness by preventing the development of low diversity climax communities. A study of faunal richness on 10 regularly inundated shores showed that inundation does promote richness, but by increasing habitat heterogeneity. Some species are excluded from parts of the shore by sand, but because of the patchiness of sand deposits they are rarely excluded from the entire shore. Other species are found only on rocks associated with sand, while typically sandy shore animals occur in the sand deposits themselves. Total richness (281 species) was greater than for local noninundated shores and sandy beaches combined.