Synergistic effects of ultraviolet radiation and conditions at low tide on egg masses of limpets (<Emphasis Type="Italic">Benhamina obliquata</Emphasis> and <Emphasis Type="Italic">Siphonaria australis</Emphasis>) in New Zealand

The effects of ultraviolet radiation (UVR), desiccation and conditions in tidal pools on embryonic survival were examined for two common pulmonate limpets that lay intertidal benthic egg masses on rocky shores in New Zealand: Benhamina obliquata and Siphonaria australis. Field surveys and manipulative experiments were conducted between December 2006 and September 2007 in the Wellington region of New Zealand (41°17′S, 174°47′E). Egg mass deposition sites in the field were species-specific: B. obliquata deposited eggs primarily in shaded crevices, whereas S. australis predominantly deposited egg masses in the sun and in tidal pools. For both species, however, embryonic mortality was greater in egg masses that had been in full sun compared to shade. For S. australis, there was also high mortality in egg masses in tidal pools or desiccated compared to those that remained submerged in flowing seawater at low tide. In outdoor experiments, embryonic mortality was also always greatest for egg masses exposed to full sun, and lowest for those in shaded treatments. Mortality was also higher if egg masses were in simulated tidal pools, and for S. australis, if desiccated, compared to those submerged in flowing seawater. Periods of particularly sunny conditions with high temperatures also resulted in higher overall mortality. Finally, egg masses of both species that were initially deposited in the shade had greater mortality in response to subsequent UV exposure compared to egg masses initially deposited in full sun. Results from this study suggest that the egg masses of these two species are highly vulnerable to UVR, as well as other intertidal stressors. Embryos of both of these species may be at risk of high mortality particularly during summer when extreme conditions of UV intensity and high temperature coincide with low tide cycles. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

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.     

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     

The role of colonization in determining spatial patterns of Proscoloplos bondi sp. nov. (Orbiniidae: Annelida) in coralline algal turf

We describe Proscoloplos bondi sp. nov., a new species of orbinid polychaete found in coralline algal turf on rocky intertidal shores near Sydney, Australia. We used field sampling to describe P. bondi's patterns of spatial and temporal variation on a number of shores. P. bondi was significantly more abundant in low- than in mid-shore areas (ANOVA, P<0.05), but local populations were extremely variable in space and time. Despite extensive sampling, we did not find evidence of sexual reproduction in P. bondi, but commonly observed worms with regenerating heads and tails, which indicated the importance of asexual reproduction. We, therefore, tested the hypothesis that adult colonization contributes to its patterns of abundance. First, patches of artificial turf were placed near to and away from natural coralline turf and sampled after 2, 4 and 12 months. Generally, populations established themselves more quickly in patches of new habitat directly adjacent to coralline turf than in patches that were isolated. Second, patches of turf were deployed for 2-week periods in low- and mid-shore areas (repeated four times). Proscoloplos bondi were found significantly more often in patches in low-shore areas than in mid-shore areas and at some sites more than others (ANOVA, P<0.05), demonstrating the importance of adult colonization in determining patterns of abundance. Overall, the limited dispersal from spatially and temporally ephemeral populations of P. bondi combined with the patchiness of coralline turfs suggest that metapopulation dynamics may be important in the persistence of P. bondi on rocky shores.Communicated by G.F. Humphrey, Sydney     

Influence of occupant microhabitat on the composition of encrusting communities on gastropod shells

The exploitation of microhabitats is widely considered to increase biodiversity in marine ecosystems. Although intertidal hermit crabs and gastropods may inhabit the same shell type and shore level, their microhabitat may differ depending on the state of the tide. On the south coast of Wales the hermit crab Pagurus bernhardus mainly inhabits the shells of Nucella lapillus (84%). Hermit crab shells had a significantly different encrusting community compared with live N. lapillus shells. At low tide the live gastropods were found on exposed rock surfaces whereas hermit crabs were restricted to tidal pools. Communities encrusting live gastropod shells were characterised by lower species richness and abundance compared with shells inhabited by hermit crabs (12 species found in total). A greater abundance and richness of epibionts was recorded from both shell types during the summer compared with winter. Differences in community composition between shell occupant types were attributed to microhabitats used by gastropods and hermit crabs and the associated desiccation pressures, rather than competitive interactions or shell characteristics. This contradicts earlier studies of subtidal shells where biological processes were considered more important than physical factors in controlling species abundance and richness patterns. The use of rockpool microhabitats by hermit crabs increases the biodiversity of rocky shores, as some species commonly found on hermit-crab-inhabited shells were rare in other local habitats.Communicated by J.P. Thorpe, Port Erin     

Calanoid copepod,cladoceran, and rotifer eggs in sea-bottom sediments of northern Californian coastal waters: Identification,occurrence and hatching

In Ireland, mussels on exposed rocky shores constitute an interbreeding mixture of two forms of mussels,Mytilus edulis L. and the Mediterranean musselM. galloprovincialis Lmk. This paper presents an in-depth analysis, carried out between October 1984 and December 1986, of genetic variability at two partially diagnostic loci,Odh andEst-D, in two exposed-shore populations ofMytilus spp. in the west of Ireland. Significant differences at theOdh locus were observed in the genetic composition of adult mussels from different tidal levels. These differences were repeatable whether one was analysing replicate samples at a single point in time, samples collected at different points in time, i.e., in different years, or samples collected from different shores. Mussels recruiting to artificial substrates set out for a period of one month at different tidal levels at one of these sites were also observed to be genetically different; mussels higher up the shore exhibited higher frequencies of those alleles characteristically at high frequency inM. galloprovincialis for both theOdh andEst-D loci. Hence, the genetic differences observed in adult mussels are much more exaggerated in juveniles and are already apparent within the first month of benthic life. Possible reasons for the observed microgeographic differentiation are discussed. It is concluded that the observed genetic differences between mussels at different tidal levels arise either in the pelagic/attachment stage or very shortly after settlement.     

Seasonal variation in utilization of biogenic microhabitats by littorinid snails on tropical rocky shores

Mobile species may actively seek refuge from stressful conditions in biogenic habitats on rocky shores. In Hong Kong, the upper intertidal zone is extremely stressful, especially in summer when organisms are emersed for long periods in hot desiccating conditions. As a result, many species migrate downshore between winter and summer to reduce these stressful conditions. The littorinids Echinolittorina malaccana and E. vidua, for example, are found on open rock surfaces high on the shore in winter but the majority migrate downshore in summer to the same tidal height as a common barnacle, Tetraclita japonica. In the laboratory, where environmental conditions could be controlled to approximate those occurring on the shore, we tested whether the downshore migration allowed littorinids to select barnacles as biogenic habitats to reduce stress and if this behaviour varied between seasons. In summer, littorinids demonstrated a strong active preference for the barnacles, which was not observed in the cool winter conditions, when animals were found on open rock surfaces even when barnacles were present. Littorinids, therefore, only actively select biogenic habitats during the summer in Hong Kong when they migrate downshore, suggesting that such habitats may play an important, temporal, role in mitigating environmental stress on tropical 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 comparison of spatial learning and memory capabilities in intertidal gobies

For the majority of animals, the ability to orient in familiar locations is a fundamental part of life, and spatial memory allows individuals to remember key locations such as food patches, shelter, mating sites or areas regularly occupied by predators. This study determined if gobies collected from rocky platforms and sandy beaches differ in their ability to learn and memorise the locations of tide pools in a simulated rocky intertidal zone. Intertidal rock pool gobies show acute homing abilities and, therefore, should be expected to display superior learning and memory capabilities. In contrast, it is unlikely that natural selection would favour similar learning skills in sandy shore fishes because they simply shift back and forth with the tides. The learning abilities of gobies were tested using small replica rock platforms, containing four tide pools that retained varying depths of water at simulated low tide. Gobies were categorised as having learnt the task if they were able to consistently locate the tide pool that retained the most water at simulated low tide as the pool with the most favourable conditions. Rock pool species were able to locate the deepest pool to wait out low tide for ~95 % of the trials, while species from sandy shores were found in the deepest pool ~10 % of trials. Despite repeated stranding, sandy shore fish continued to follow the tide out. We propose that rock pool species memorised the location of rock pools during simulated high tide enabling them to relocate the best refuge for low tide.     

Characterizing temperate rocky shores from the perspective of an early juvenile snail: the main threats to survival of newly hatched Nucella emarginata

Mortality factors most likely to constitute substantial selective pressures for early juvenile gastropods on temperate rocky shores were identified by examining the vulnerability of hatchlings of an intertidal snail, Nucella emarginata, to heat stress, desiccation, and predation in 1992 and 1993. The highest temperature of substrata measured at tidal heights colonized by N. emarginata in Barkley Sound, British Columbia, Canada, was 28.5°C. This temperature was not lethal to hatchlings in laboratory tests. In laboratory and field desiccation experiments, all hatchlings died within 6 h of emersion. Early juveniles could not survive direct exposure to even moderate drying conditions for the duration of a low tide. Hence, intertidal microhabitats which dry up even for short periods during low tides would prove lethal. Of 45 intertidal animal species to which hatchlings were exposed in the laboratory, small decapod crustaceans were the only organisms to cause substantial hatchling mortality. Of these, Pagurus hirsutiusculus and Hemigrapsus nudus were by far the most abundant in the field, and are probably the only important predators of early juvenile N. emarginata at most sites. Total predator densities in the field were as high as 438 individuals m^–2, suggesting that predation pressure may be intense. Desiccation and predation by decapod crustanceans appear to be the most significant threats to early juvenile N. emarginata. These factors commonly occur on most temperate rocky shores and undoubtedly constitute major selective agent influencing population parameters and shaping life-history strategies and early juvenile traits of intertidal invertebrates.     

A comparative study on intertidal faunal biodiversity of selected beaches of Mumbai coast

Comparative study has been done to examine the biodiversity and ecological status of the intertidal region of Tata Institute of Fundamental Research (TIFR), Bandstand and National Centre for Performing Arts (NCPA) rocky beaches in Mumbai, West coast of India. A total of 50 species of intertidal organisms were recorded from these shores. Shannon and Simpson's diversity index, Margalefs richness index and Pielou's evenness index indicated different level of ecological state of the shore in different months. Dendrograms and 2-D non metric MDS ordination from Bray-Curtis similarity matrix of occurrence of intertidal organisms from these sites showed highest similarity and combination pattern of occurrence between Nerita oryzarum and Planaxis sulcatus in TIFR and Bandstand shore. Nerita oryzarum and Tactarius malaccanus at NCPA shore. Abundance/biomass comparison (ABC) method of determining level of disturbance also pointed towards the polluted status of these shores. Study concludes that though these beaches are highly disturbed due to anthropogenic activities, they still support a rich intertidal biodiversity which need immediate attention for protection and conservation.     

Biomonitoring of heavy metal (Cd,Cu, Pb,and Zn) concentrations in the west intertidal area of Peninsular Malaysia by using Nerita lineata

Snails, Nerita lineata, were collected from 15 sites along the west intertidal area of Peninsular Malaysia from December 2005 until April 2006. The concentrations of heavy metals (Cd, Cu, Pb, and Zn) were determined in the total soft tissues, operculums, and shells of the snails. Different patterns of heavy metal distributions were found in the different tissues (shell, operculums, and soft tissues) as well as spatial variations of heavy metal concentrations in the snails. This shows that the distribution of metals in the shells and the total soft tissues of N. lineata were not similar which could be due to different rates of metal accumulation, excretion, and sequestration. Since N. lineata is abundant on the rocky shores, below jetties and mangrove trees along the west intertidal area of Peninsular Malaysia and accumulate heavy metals, the snails are therefore potential biomonitors of heavy metal contamination for the west intertidal area of Peninsular Malaysia.     

Genetic structure of hybrid mussel populations in the west of Ireland: two hypotheses revisited

In Ireland, mussels on exposed rocky shores constitute an interbreeding mixture of two forms of mussels, the blue mussel, Mytilus edulis, and the Mediterranean mussel, M. galloprovincialis. Results from an Irish study in the 1980s, using partially diagnostic allozyme markers, indicated that mussels higher up the shore were more galloprovincialis-like than those lower down. In this study we set out to test two hypotheses: (a) recruits arriving on the shore are composed of genetically distinct cohorts that settle preferentially at different levels on the shore, and maintain genetic distinctiveness into adulthood; (b) recruits are genetically homogeneous, but once settled they diverge genetically over time, due to within-habitat site specific-selection. The diagnostic Me 15/16 DNA marker was used to analyse the genetic composition of newly-settled spat recruiting to artificial substrates, which were placed at two-week intervals from May–October 2002, on the mid- and low shore areas of two exposed sites in Galway Bay. Adult mussels were also collected on each sampling date. Results did not support the preferential settlement hypothesis, i.e., the genetic composition of primary settlers (≤ 500 μm) was similar between tidal heights and shores. Neither was there evidence of post settlement selective mortality, as adults were genetically similar to settling spat. In spat and adults the frequency of the M. galloprovincialis allele was high (0.56–0.80), due to high frequencies of M. galloprovincialis (> 37%) and hybrid (> 33%) genotypes, and correspondingly low frequencies of the M. edulis genotype (< 11%). Adult mussels from a nearby sheltered estuarine site, while significantly different to exposed shore mussels, still had low frequencies of the M. edulis genotype (< 17%), indicating no apparent advantage for the genotype in this environment. There are indications that the genetic composition of mussels may be changing on the Atlantic coasts of Ireland.     

Temporal genetic variation in populations of the limpet Cellana grata from Hong Kong shores

Variations in the relative contributions of gene flow and spatial and temporal variation in recruitment are considered the major determinants of population genetic structure in marine organisms. Such variation can be assessed through repeated measures of the genetic structure of a species over time. To test the relative importance of these two phenomena, temporal variation in genetic composition was measured in the limpet Cellana grata, among four annual cohorts over 10 years at four rocky shores in Hong Kong. A total of 408 limpets, comprising individuals from 1998, 1999, 2006 and 2007 cohorts were screened for genetic variation using five microsatellite loci. Minor but significant genetic differentiation was detected among samples from the 1998/1999 collection (F _ST = 0.0023), but there was no significant differentiation among the 2006/2007 collection (F _ST = 0.0008). Partitioning of genetic variation among shores was also significant in 1998/1999 but not in the 2006/2007 collection, although there was no correlation between genetic and geographic distances. There was no significant difference between collections made in 1998/1999 and 2006/2007. This lack of clear structure implies a high level of gene flow, but differentiation with time may be the result of stochastic recruitment variation among shores. Estimates of effective population size were not high (599, 95% C.L. 352–11397), suggesting the potential susceptibility of the populations to genetic drift, although a significant bottleneck effect was not detected. These findings indicate that genetic structuring between populations of C. grata in space and time may result from spatio-temporal variation in recruitment, but the potential development of biologically significant differentiation is suppressed by a lack of consistency in recruitment variability and high connectivity among shores.     

Non-indigenous macroalga hosts different epiphytic assemblages to conspecific natives in southeast Australia

Codium fragile ssp. tomentosoides is one of the five most invasive macroalgae worldwide. We compared epiphytic assemblages between native (ssp. tasmanicum and ssp. novae-zelandiae combined) and non-indigenous (ssp. tomentosoides) subspecies of Codium fragile on three rocky intertidal shores in southeast Australia. Twelve species of epiphytes covered up to 20% of the surface of both native and non-indigenous Codium subspecies, but none were unique to either subspecies. For C. fragile subspecies, epiphytic cover declined from the lower (older) parts of the thallus to the branch tips. The abundance and species composition of epiphytes differed significantly between native and non-indigenous C. fragile, but differences varied among rocky shores. Results demonstrate that non-indigenous C. fragile does not play a functionally similar role to closely related native Codium subspecies. Spatial differences in epiphytic assemblages between Codium subspecies among rocky shores demonstrate that effects of non-indigenous species may be strongly location specific. Thus, our study emphasises the need to investigate variation in the effects of invaders across the regional landscape.     

Interactive effects of wave exposure and tidal height on population structure of the mussel Perna perna

The influence of wave exposure and of tidal height on mussel (Perna perna Linnaeus) population structure (size, density, biomass and adult/juvenile correlations) was examined at 18 sites along the south coast of South Africa. Sites were classified as exposed or sheltered prior to sampling, without reference to the biota, on the basis of aspect, topography and wave regime. A single set of samples was collected from each site during three spring tide cycles. Adult mussels on these shores almost always attach directly to the rocks, and layering of mussels is virtually absent. Shore height always had a strong influence on population structure, but exposure had significant effects only lower on the shore, and almost exclusively on mussel sizes. Principal component analysis (PCA), based on size distribution data for each population, revealed a general upshore decrease in the modal size of the adult cohort. The effects of exposure on size distribution, however, varied with tidal height. PCA separated exposed zones, with larger mussels, from sheltered zones on the low-shore. Farther upshore the two shore types were increasingly confounded. The maximum size of mussels showed a similar pattern, with significant differences (ANOVA, p < 0.05) between exposed and sheltered sites only on the low- and mid-shores. Density was calculated from randomly placed quadrats (i.e. not necessarily from areas of 100% cover) and showed a different pattern. Adult (>15 mm) densities decreased up the shore, with low-, mid- and high-shore zones being significantly different from one another (ANOVA, p < 0.0001; followed by multiple range tests). However, exposure had no significant effect on density, nor was there a significant interaction with zone. Recruit (<15 mm) densities were positively correlated with adult (>15 mm) densities in all zones and for both exposure regimes ( p < 0.05 in all cases), but there was considerable variability and extremely low predictability in these relationships (r ² generally <0.2). Predictability tended to be greater towards the high-shore, where adults were more clumped. As with density, biomass was not affected by exposure, but decreased upshore as mean size and density decreased. A reduction in the influence of exposure farther upshore may be caused by greater emersion overriding the effects of exposure. The presence of free space within mussel beds and significant correlations between recruit and adult densities suggest that these mussel populations are recruit limited. Received: 7 January 2000 / Accepted: 6 July 2000     

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

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

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Contribution No. 164, the Charles Darwin Foundation.     

Genetic differentiation independent of intertidal gradients in the pulmonate limpet Siphonaria kurracheensis

The intertidal limpet Siphonaria kurracheensis (Reeve, 1856) has a bimodal vertical distribution of abundance on rocky shores at Rottnest Island, Western Australia. An electrophoretic study of 5 polymorphic enzymes revealed no consistent genetic differences between adults high and low on the shore. Contrasting with this absence of a detectable genetic response to the steep environmental gradients in the intertidal zone, there were genetic differences among low-shore adults from different sites, and between adults and recruits. This genetic differentiation could be due to either localized selection or temporal variation in the genetic makeup of recruits.Communicated by G. F. Humphrey, Sydney     

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

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