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

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

Tidal rates of settlement of the intertidal barnacles Chthamalus stellatus and Chthamalus montagui in western Europe: the influence of the night/day cycle

Chthamalus montagui and Chthamalus stellatus are abundant barnacles in western Europe. Tidal settlement of Chthamalus in SW Ireland and SW Portugal was studied in relation to a night and day factor and at different temporal (dates) and spatial (shores and sites) scales. Based on the identifiable cyprids and metamorphs, Chthamalus settlement in SW Ireland was comprised mainly of C. stellatus but was composed of C. montagui only in SW Portugal.In SW Ireland and SW Portugal, settlement rates of Chthamalus (mean number of settlers per 25 cm²±S.E.) were higher after one day tidal cycle (597±158.7 in SW Ireland, 144±23.6 in SW Portugal) than one night tidal cycle (55±12.1 in SW Ireland, 13±2.2 in SW Portugal), but significant differences were only detected in SW Portugal. Different models were proposed for explaining this pattern related to night and day variability of the physical processes responsible for transporting cyprids to shore (1), and/or of pre-settlement behaviour (2) and/or of settlement behaviour of cyprids (3).Spatial patterns of tidal settlement of both species or at both locations seem similar with small scale variability (between sites, 5 to 30 m apart) in settlement of Chthamalus being the only spatial scale at which variability was detected in both locations.The composition of Chthamalus cohorts settling during one tidal cycle differed considerably between locations/species: settlers of C. stellatus were mainly cyprids; settlers of C. montagui during the day (when most settlement occurred) were essentially metamorphs.     

Spatial variation in the recruitment of the intertidal barnacles Chthamalus montagui Southward and Chthamalus stellatus (Poli) (Crustacea: Cirripedia) over an European scale

Spatial variation in the recruitment of the intertidal barnacles Chthamalus montagui and Chthamalus stellatus was examined over an European scale. The study was carried out using standardised protocols at a series of locations. The five locations chosen (SW Ireland, NW Spain, SW Portugal and NW and NE Italy) span a large part of the range of these species in Europe. The spatial scales were location (hundreds of kilometres) and shore (thousands of metres).Estimates of total cumulative recruitment (cyprids and metamorphs) summed over the year (April 1997 to March 1998) showed substantial variation between locations which was dependent on the species. Recruitment was highest in SW Portugal for C. montagui and in SW Ireland and NW Spain for C. stellatus. Overall recruitment of C. montagui was higher than that of C. stellatus at all locations except SW Ireland, where recruitment of the two species was not significantly different. There were significant differences among shores in each location.The recruitment period of both species varied with location, with recruitment beginning earlier further south. In general, recruitment of C. montagui and C. stellatus was recorded in 8 months in NW Spain and NE Italy, while only in 7 months in SW Ireland. Recruitment of C. montagui occurred in 10 months in SW Portugal, but no recruits of C. stellatus were found. In all locations there was at least one distinct peak of recruitment. In SW Ireland both species showed only one peak of recruitment, a month after initiation. At the more southerly Atlantic locations, as well as in the Mediterranean, two unequal peaks of recruitment were generally seen. During recruit census, the number of cyprids, in comparison to metamorphs, found at any location was very low. In SW Ireland and NW Spain cyprids of both species were found, while in SW Portugal and in the Mediterranean, only cyprids of C. montagui were found.     

Patterns of adult abundance in Chthamalus stellatus (Poli) and C. montagui Southward (Crustacea: Cirripedia) emerge during late recruitment

The aim of this study was to investigate when adult distribution patterns are established in the barnacles Chthamalus stellatus and C. montagui. Adult ‘zones’ were identified by analysing field counts of both species at mid and upper shore heights. Monthly collections of cyprids, < 1 month old metamorphs and recruits (all metamorphosed individuals older than approximately 1 month) were made for C. stellatus and C. montagui in natural barnacle beds at six shores in SW Ireland. This was carried out over one year in 1996/1997, using a hierarchical sampling design. Abundance of total recruits (0-3 months old) was compared between adult zones after the main settlement season had ended. In addition, scales of variability in 0-3 month recruitment into adult zones were compared between the species at two scales: shores (1000s of metres) and sites within shores (10s of metres). Older recruits of each species, up to 11 months of age, were also compared between adult zones.The majority of settlement (measured as attached cyprids) occurred between August and October 1996. In October, there was no effect of adult zone on the abundance of total (0-3 month) recruitment up to that point in either species. Despite this homogeneity in recruitment between adult zones, significant spatial variation was found in 0-3 month recruits of both species at both of the scales examined. In C. stellatus the amount of variation associated with the larger scale (shore) was more than twice that of sites or of the residual variation (replicates within sites). 0-3 month recruitment in C. montagui was also most variable at the scale of shores but the residual variability (between replicates within site) was of similar magnitude to that of shores. Variability in 0-3 month C. montagui recruitment was relatively low at the scale of sites.There was a small but consistent input of recruits to adult zones over 9 months of the year, complicating the assessment of when adult patterns were set-up in these species. By June 1997, characteristic patterns of adult dominance had been established at all shores. Settlement had completely ceased by this time and individual barnacles were potentially 11 months old. Neither settlement nor early recruitment are significant in determining adult zonation patterns in these species. Instead, differential mortality patterns in individuals up to the age of 11 months are implicated in determining patterns of distribution of both species.     

Isolation of highly polymorphic microsatellite markers from the intertidal barnacle Chthamalus montagui Southward

This study reports the isolation and characterization of seven highly polymorphic microsatellite loci in Chthamalus montagui (Crustacea, Cirripedia). The loci were isolated from a library constructed from genomic DNA enriched for CA repeats. The markers yielded three to 43 alleles per locus (mean 16.7) in samples averaging 49 individuals. Observed heterozygosity ranged from 0.08 to 0.58 (mean 0.39). These microsatellite loci will be valuable tools for population genetic studies of this species.     

Settlement behavior of Chthamalus anisopoma larvae largely determines the adult distribution

Summary In the northern Gulf of California the adult distribution of the intertidal barnacle species, Chthamalus anisopoma, on exposed shores is approximately between 0.0 and 2.0 m above mean low water (MLW). The species is typically absent in protected (from wave splash) areas. In this study, I examined a series of alternative hypotheses relating to the factors that could be responsible for limiting the distribution. Post-settlement factors appear to be unimportant because settlement was largely restricted to areas within the adult distribution. Two processes could account for the high correlation between settlement and adult distributions. First, hydrodynamic factors could restrict deposition of larvae to sites that coincidently were in areas in which individuals could survive to maturity. Second, larvae may choose to settle only on sites where they can survive to maturity. Of the two, the later was supported as settlement could be induced on surfaces outside the adult distribution using transplanted adult conspecifics as cues. Thus, competent larvae were present outside the adult distribution of Chthamalus zone but did not settle under normal conditions. Also, there was no evidence that pre-emption of space by other sessile species, by itself, restricted the distribution of Chthamalus. Settlement within the existing adult distribution may be an evolutionary response to increased mortality for individuals settling outside the adult distribution compared to those settling within it.     

Chthamalus stellatus (Poli) und C. depressus (Poli) in der adria

A study has been made of the ecology of Chthamalus stellatus (Poli) and C. depressus (Poli) in the region of Rovinj (Yugoslavia). The distribution has been examined at several localities of different habitat conditions particularly with respect to wave action.     

Does colonization contribute to spatial patterns of common invertebrates in coralline algal turf?

Abstract The potential of colonization to contribute to the spatial patterns of six common invertebrates in coralline algal turf was investigated on a rocky shore near Sydney, Australia. The species, which included two amphipods (Elasmopus warra, Hyale spp.), a small bivalve (Lasaea australis), a fly larva (Limonia marina), and two microgastropods (Amphithalamus incidata and Eatoniella atropurpurea), had a range of dispersal modes (larval dispersal, crawling, swimming, rafting, and passive transport). Field sampling between May 1997 and November 1999 demonstrated that the amphipods were more abundant in low‐shore areas, the fly larvae and bivalves were more abundant in mid‐shore areas, and the abundances of gastropods did not vary with tidal height. Furthermore, abundances of all species varied among patches separated by tens of metres at one time or another. To test whether rates of colonization could contribute to established patterns of abundance, habitat mimics were deployed for 2‐week periods. The supply of new individuals matched long‐term patterns of abundance at different tidal heights for E. warra and L. marina. Colonization rates also differed among patches separated by tens of metres for three of the six species. Overall, there was little evidence to suggest that common species in coralline turf are limited by colonization on local scales, regardless of their major mode of dispersal. However, the potential for colonization to determine patterns of abundance varied from species to species.     

Comparative SEM Studies of Lattice Organs: Putative Sensory Structures on the Carapace of Larvae from Ascothoracida and Cirripedia (Crustacea Maxillopoda Thecostraca)

Putative sensory structures, called lattice organs, were studied with scanning electron microscopy from ascothoracid or cypris larvae representing a wide range of families of the Ascothoracida and Cirripedia. These organs, situated dorsally on the carapace were, with few exceptions, always found in two anterior and three posterior pairs. The lattice organ morphology displayed by the Ascothoracida, a seta–like structure with a terminal pore, is believed to be the most plesiomorphic condition. Within the Cirripedia lattice organ morphology varied from types resembling the Ascothoracida in the Acrothoracica and the lepadomorph Capitulum mitella, to an elongate pore field with a larger terminal pore in most Thoracica and Rhizocephala. Akentrogonid Rhizocephala seem to display the most apomorphic condition. While lattice organ morphology was generally constant at the family level, cases were seen where closely related species such as Chthamalus stellatusand Chthamalus montaguishowed minor, but clear cut differences. Lattice organs in 2 + 3 pairs are argued to represent a synapomorphy for the Ascothoracida and the Cirripedia. The results confirm that the cyprid morphology at the ultrastructural level will prove to be of high value in estimating phylogeny within the Cirripedia.     

Effects of sewage pollution on the structure of rocky shore macroinvertebrate assemblages

The urgency to find efficient indices and indicators to prevent further deterioration of coastal areas is one of the hot topics in today’s scientific publication. However, a detailed knowledge of community responses to anthropogenic impacts is essential to sustain those indices. The studies on the response of benthic community to sewage pollution on intertidal rocky shores are generally based on visual census and do not take into account the tidal levels. In order to fulfil this gap in this study: (i) the sampling was performed by destructive sampling, with all individuals identified to the species level; (ii) the sampling was done at all levels of the intertidal (sublittoral fringe, eulittoral, and littoral fringe). Sewage pollution changed the environmental variables and the abundance of macroinvertebrates, being Mytilus galloprovincialis, Melarhaphe neritoides, and Chthamalus montagui the species most responsible for the dissimilarities observed. Effects were different on the three intertidal zones: community structure changed in the sublittoral fringe; suspension-feeders abundances and species richness increased in the eulittoral; no differences were detected in the littoral fringe. Moreover, the results confirm that the presence of sewage discharges tended to benefit suspension feeders, and that the sensitive species were replaced by opportunistic ones.     

Exposure of seawalls to waves within an urban estuary: effects on intertidal assemblages

Abstract Despite being largely protected from sea swell by headlands, shores within Sydney Harbour, New South Wales, Australia are subjected to considerable wave action from wind waves and the numerous recreational and commercial boats and ships using the waterways. The aim of this study was to test whether assemblages on artificial shores, that is, seawalls, either exposed to or sheltered from waves would show similar patterns to those published for natural shores exposed to or sheltered from sea swell. Specifically, the hypotheses were tested that sheltered seawalls would have greater cover of algae, whereas exposed seawalls would have greater cover of sessile invertebrates and greater abundance of mobile invertebrates. It was found that encrusting algae had greater cover on sheltered seawalls, whereas cover of turfing algae was greater on exposed walls. Sessile filter‐feeders generally had greater cover on exposed seawalls, although a dominant space occupier, the oyster Saccostrea glomerata, showed either no difference, or greater cover on sheltered seawalls among different locations. The pattern for the dominant grazer on seawalls, the pulmonate limpet Siphonaria denticulata showed the predicted pattern low on the shore, but the opposite pattern at mid‐tidal levels. Despite some inconsistencies the results were similar to those predicted from studies on natural shores, showing that assemblages on urban structures respond to wave action in a way that is predictable from studies on natural shores. Experiments were carried out where assemblages were transplanted to sites with differing exposure to waves to determine whether differences in recruitment or post‐recruitment mortality were responsible for patterns of difference. We found that some sessile invertebrates did not survive when transplanted to sheltered seawalls, suggesting that difference in survival of organisms was responsible for observed patterns. This was not, however, found when the experiment was repeated at other locations. It is likely that processes affecting the assemblages are temporally and spatially very variable, or that more than one process interacts to cause observed patterns. Identifying such complexity requires rigorously structured sampling designs and appropriate manipulative experiments.     

The niche-width variation hypothesis reconfirmed: Validation by genetic diversity in the sessile intertidal cirripedes Chthamalus stellatus and Euraphia depressa (Crustacea,Chthamalidae)

Genetic diversity in a pair of cirripede species, based on electrophoretic analysis of 25 gene loci, is higher for Chthamalus stellatus, the species with a broader biogeographical distribution, and hence, with a broader ecological niche, than that of Euraphia depressa. Comparing the genetic diversity within E. depressa we again report a higher genetic diversity among the specimens exposed to solar radiation (living in the wider ecological niche) versus the specimens confined to the sheltered dark environments in caves or under boulders. The indices of genetic diversity used in this study are polymorphism, heterozygosity, mean number of alleles per locus and gene diversity. We conclude that the niche-width variation hypothesis has been confirmed in this pair of species: genetic diversity is positively correlated with niche breadth.     

Settlement patterns of Chthamalus spp. at Praia da Oliveirinha (SW Portugal)

Recruitment of Chthamalus spp. is intense during spring and summer but the number of settled cypris larvae is very low when compared to the high number of Chthamalus spp. metamorphs. In the summer of 1997 at Praia da Oliveirinha (SW Portugal), two hypotheses were tested: 1) the cypris larvae of Chthamalus spp. metamorphose in one tidal cycle; and 2) the settlement of Chthamalus spp. is higher during diurnal high tide compared with nocturnal high tide. The results indicate that cypris larvae of Chthamalus spp. can metamorphose in one tidal cycle. The hypothesis of higher settlement during diurnal high tide could not be tested at the mid and upper levels of C. montagui distribution due to a significative heterogeneity of variances. At the lower level, the results support this hypothesis. In spite of these data having been collected on a single shore and on a few sampling dates, these results suggest that more studies using different spatial and temporal scales are needed to see if this pattern is general.     

Studies in the biochemistry of cirripede eggs. VII. Changes in the general biochemical composition during development of Chthamalus dentatus Krauss and Octomeris angulosa Sowerby

The carbohydrate, protein, total lipid, RNA, and DNA have been determined at four stages during the embryonic development in two chthamalids, Chthamalus dentatus Krauss and Octomeris angulosa Sowerby, from the South African coast. These values were compared with those of some cirripedes from the northern hemisphere. There is a relatively high utilization of lipids during the development as was previously found in Chthamalus montagui Southward. The relative volume specific energy content is similar to other cirripedes, but clearly higher than that of boreo-arctic balanids. The energy loss in C. dentatus is 54.4% of the initial energy content and that of Octomeris angulosa is 39.4%. This is lower than that found in Chthamalus montagui, but higher than that of all other cirripedes studied and especially the boreo-arctic species. The possibility of geographical significance of these differences is discussed.     

Habitat segregation and temporal variation in some tropical intertidal populations

Observations were made to document habitat segregation and seasonal changes in density for the most common intertidal organisms at a study site on the Pacific coast of Costa Rica. The pulmonate limpets Siphonaria gigas (Sowerby) and Siphonaria maura (Sowerby) were more abundant in the mid-high- and high-intertidal zone, whereas the keyhole limpet Fissurella virescens (Sowerby) was more abundant in the mid zone. Coiled gastropods were found in the mid- and mid-high-intertidal zone but were absent or scarce in the high zone. Barnacles and coralline algae were found throughout the intertidal zone and foliose algae were mostly present in the splash zone. Abundance of all species changed over the 2-yr period of the study. S. gigas and S. maura and the barnacle Chthamalus fissus Darwin exhibited well-defined fluctuations in density related to recruitment. No recruitment was observed for F. virescens or Tetraclita panamensis Pilsbry and populations of both species declined. Changes also occurred for coiled gastropods and were probably related to movement of adults. Coralline and foliose algae were more abundant during the rainy season than during the dry months. Far from being constant, populations on these tropical shores were highly dynamic with temporal changes rivaling those in the temperate zone.     

A six-year study of abundance and voltinism of Chironomidae (Diptera) in an Illinois cooling reservoir

The chironomid fauna of Baldwin Lake, an off-stream closed-cycle cooling reservoir for an 1800-megawatt coal-fired power plant, was sampled semi-monthly to monthly at four stations for six years, beginning five months after the reservoir was filled and ending 23 months after the third unit of the power plant went into operation. In the main basin three species dominated: Coelotanypus concinnus (Coquillett), Procladius bellus (Loew), and Tanypus stellatus Coquillett. The last did not become established until the lake was two years old, probably because temperatures were too low the first winter. Populations of all three species were initially large and sharply fluctuating, but later became smaller and less fluctuant in C. concinnus and P. bellus, due, probably, to more intense predation on them. Only in T. stellatus were changes in voltinism clearly evident: it changed from bivoltine to tri- and quatrivoltine as degree-days of heat increased. Photoperiod also affected its life cycle, as evidenced by synchronous development of summer generations each year. Temperature and perhaps food supply affected development time of its overwintering generation. All three species tolerated a temperature of 38 °C; their populations were kept low in the discharge channel by the current, not the temperature. Six species of Chironominae were sporadically abundant in the discharge channel.     

Spatial and vertical patterns in the tidepool fish assemblage on the island of O`ahu

The microtides, wave regimes, and relative isolation of the Hawaiian archipelago may provide unique environmental and biogeographic effects that shape the structure of tidepool fishes. We sampled fishes across a narrow gradient at low tide from 6 sites on the island of O`ahu. We tested predictions of the hypotheses that environmental conditions (pool depth, volume, macroalgal cover, temperature, and salinity) would result in a vertically structured tidepool fish assemblage unique to basalt or limestone rocky shores. 343 fish were recorded from 40 pools, and 19 species from 10 families were identified. Tidepool fish diversity (H’: O`ahu = 2.4; Sites Average = 0.0–0.9) was typical for tropical islands, with members from Gobiidae (5 species), Blenniidae (4 species), Pomacentridae (3 species), Acanthuridae (2 species) and Kuhliidae (2 species) among the most common. Endemism (32%) was higher than other well studied assemblages yet similar to Hawaiian reef fishes (25%). Assemblage abundance varied among shores with basalt or limestone substrate, among sites, and vertically among high, mid, and low pools. In general, blenniids occurred at higher proportions on limestone shores and gobiids were more common on basalt shores. High pools were characterized by an abundance of a small sized (29.0 mm median standard length) blenniid Istiblennius zebra, while the blenniid Entomacrodus marmoratus and wrasses Thalassoma spp. were more common in low pools. Temperature was the best environmental predictor of assemblages and this relationship warrants further investigation. Our findings indicate that assemblages can vary across a narrow geographical range and intertidal shore.     

Phylogeography of the cold‐water barnacle Chthamalus challengeri in the north‐western Pacific: effect of past population expansion and contemporary gene flow

Aim Phylogeographical patterns of marine organisms in the north‐western Pacific are shaped by the interaction of past sea‐level fluctuations during glacial maxima and present‐day gene flow. This study examines whether observed population differentiation in the barnacle Chthamalus challengeri, which is endemic to the north‐western Pacific, can be explained by the interactions between historical glacial events and patterns of contemporary gene flow. Location Eleven locations in the north‐western Pacific. Methods Partial sequences of mitochondrial cytochrome c oxidase subunit I (COI), 12S, 16S and nuclear internal transcribed spacer 1 (ITS1) were obtained from 312 individuals. Parsimony haplotype networks and analysis of molecular variance (AMOVA) were used to determine whether the observed genetic structure corresponds to marine provinces (Kuroshio Current and China Sea Coastal Provinces), zoogeographical zones (oriental and Japan warm‐temperate zones) and/or potential refugial areas (Sea of Japan and East China Sea) in the north‐western Pacific. Neutrality tests, mismatch distribution analysis and Bayesian skyline plots were used to infer the demographic history of C. challengeri. Results In total, 312, 117, 182 and 250 sequences were obtained for COI, 12S, 16S and ITS1, respectively. A panmictic population was revealed, which did not conform to the ‘isolation by distance’ model. None of the a priori population groupings based on marine provinces, zoogeographical zones or potential refugial areas was associated with observed genetic patterns. Significant negative values from neutrality tests and the unimodal mismatch distribution and expansion patterns in Bayesian skyline plots for the COI and 16S data sets indicate a population expansion in the mid‐Pleistocene (c. 200 ka). Information from the fossil record suggests that there has been a northward range shift of this species from the East China Sea or the Palaeo‐Pacific coast of Japan to the Sea of Japan since the mid‐Pleistocene. Main conclusions Chthamalus challengeri has experienced a population expansion and range shift since the mid‐Pleistocene. The observed lack of population differentiation can be explained by this past population expansion and present‐day wide‐scale larval dispersal (owing to the long planktonic larval duration) across marine provinces, which have led to the successful establishment of the species in different zoogeographical zones and habitats.     

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