Spatial structure of an intertidal molluscan assemblage on a sheltered sandy beach

The intertidal molluscan fauna on a sandy-mud beach in Newport Bay, California, USA, is divisible into two vertically distinct species assemblages which correspond to the midlittoral zone and sublittoral fringe observed on rocky coastlines and exposed sandy beaches. The species assemblage comprising the midlittoral zone is unusual, however, in that the numerically dominant species are not confined to this zone but range downward through the sublittoral fringe. The two species assemblages are vertically separated at +0.5 to +1.0 ft (+0.15 to +0.30 m; relative to 0.0 tidal datum at mean lower low water), which is higher than previously observed for the separation between the midlittoral zone and sublittoral fringe on rocky shores and exposed sandy beaches. The species composition of some of the samples at +0.5 and +1.0 ft (+0.15 and +0.30 m) was intermediate between samples higher and lower on the beach, while several samples were devoid of molluscs altogether. Additionally, the surface sedimentary environment changes in terms of an increased percent silt-clay fraction and higher sorting coefficients below +0.5 ft (+0.15 m). Since infaunal zonation is correlated with tidal height at the substrate surface, environmental factors operative at the surface are probably most important in influencing the zonation on this beach.     

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

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

Macrozoobenthic community structure in the Ría de Foz,an intertidal estuary (Galicia,Northwest Spain)

The community structure of the macrobenthic (>1 mm) fauna of an intertidal soft-bottom estuary, the Ría de Foz (Galicia, Northwest Spain) was investigated on 99 occasions during June, September and December 1984 and March 1985. The infaunal data were subjected to classification and ordination techniques. The analyses indicated that sediment characteristics (grain size, percentage silt-clay and organic-matter content) and tidal height are the most important factors governing the distribution and abundance of the intertidal communities. Three faunal assemblages were identified and examined in terms of species dominance, constancy and fidelity. In the clean sands of the outer estuary two main assemblages were distinguished; one, corresponding to the boreal LusitanianTellina community, inhabited the middle and lower tidal levels; the other, typical of a biocoenosis of sands and slightly muddy mediolittoral sands, inhabited the upper tidal level. The third benthic assemblage occupied the muddy sands and mud of the inner estuary and displayed some features in common with theCardium edule-Scrobicularia community described by Thorson.     

Description of a Tetraclita stalactifera panamensis community on a rocky intertidal Pacific shore of Panama

The midlittoral zone at Paitilla Beach, Panama, Is occupied by a group of organisms closely associated with Tetraclita stalactifera panamensis. Upon death, this cirripede provides housing for a number of species bound by common substrata and interacting with each other due to their close physical proximity; the faunistic group associated with each barnacle qualifies then, as a small, well-defined community which represents a subsample of the more comprehensive midlittoral community dominated by T. stalactifera panamensis. In patches where it is abundant, T. stalactifera panamensis occupies 28% of the surface area available. It has a relative abundance of only 1% of the specimens found in the zone, but all the other 95 species (with an average of 26, 810 individuals/m²) identified from the area are associated either with the parietal canals, or the external or internal surfaces of T. stalactifera panamensis tests. Each one of these structures represents a habitat which provides surface for attachment, shelter, or both, to 37 species of polychaetes, 32 of mollusks, 20 of crustaceans, 3 of actiniarians, 2 of pycogonids, 2 of sipunculans and an unknown number of tanaid, nemertean and turbellarian species. The single most diverse space-niche is represented by the parietal canals (H=0.783), occupied mostly by polychaetes. dead barnacles harbor an average of 131 individuals per test, which compose a fauna of higher diversity (H=0.678) than that associated with live barnacles (H=0.163). The latter contain 53 specimens per test on the average. Although there is considerable overlapping of species associated with the different T. stalactifera panamensis space-niches, mollusks are clearly more abundant on internal surfaces, polychaetes in parietal, canals; Balanus spp. on external surfaces, and anemones at the base of the parapet on the external surfaces of the tests. Over 220 species of macro-invertebrates were identified from the rocky intertidal at Paitilla. Most of the species are concentrated in the lower (157 species, 1824 specimens/m²) and midlittoral (96 species, 26, 810 specimens/m²) intertidal zones. Splash zone is depauperate (6 species, 1578 specimens/m²). Of the entire fauna identified from the intertidal, 73 species (33%) occur in more than one zone and 147 (67%) are restricted to a particular belt. This last category of zone-restricted species breaks down into 42% for the lower intertidal, 17% for the midlittoral, 6% for the intermediate zone, and 2% for the splash zone.     

Diel eclosion rhythm of a sublittoral population of the marine insect Pontomyia pacifica

Within the scope of studies on adaptation of insects to intertidal and sublittoral environments, correlations between the reproduction period of the short-lived chironomid Pontomyia pacifica Tokunaga and tidal conditions were examined at the only known Japanese location of this species in 1980. The larval habitat is situated, except for deeper tide pools in the lower midlittoral zone, mainly within the inner sublittoral zone, whose upper area is covered by Hypnea choroides, the dominating algae during the summer. The aerial adults emerge on the surface of the open sea and swarm there independent of the tidal situation above the submersed sublittoral habitat. Eclosion always started during dusk and all reproductive activities were ended within 2.5 h after sunset. The adaptation of P. pacifica (sublittoral habitat, diel eclosion after sunset with mass concentration of the adults on the water surface, no semilunar or lunar timing of the reproduction period and sinking egg masses) corresponds with those of the convergent marine chironomid Clunio balticus from Europe. On the basis of a few laboratory observations with P. pacifica and detailed experiments with C. balticus, it is supposed that the diel eclosion of P. pacifica is also controlled by an endogenous, circadian timing mechanism and the 24-h light-dark cycle as an environmental time cue.Communicated by O. Kinne, Oldendorf/Luhe     

Influence of thermal history on the response of <Emphasis Type="Italic">Montastraea annularis</Emphasis> to short-term temperature exposure

Acclimation of reef corals to environmental conditions has been related to metabolic response at large geographic scales, but regional relationships have rarely been described. Physiological responses to temperature increases of Montastraea annularis (Ellis and Solander 1786) from an inner lagoon and an outer barrier reef in the Gulf of Honduras, southern Belize, were compared in May 2003. The hypothesis that inferred differences in thermal history would result in contrasting responses to elevated temperature was tested. Ambient seawater temperatures adjacent to corals at 4–5 m depth were measured every 15 min at inner lagoon and outer barrier reef collection sites for 1 year (June 2002–May 2003). Monthly averages and 3-day running averages (warmest period, July–October 2002) of daily maximum seawater temperatures were significantly higher (by ∼0.5°C) at inner lagoon reef compared to outer barrier reef sites. M. annularis photosynthesis (P) and respiration (R) rates were measured in respirometers at six temperatures between 29°C and 35°C approximately every hour, with repeated measurements over 3 h. P and R were significantly lower across most temperature treatments for samples collected from the inner lagoon compared to outer barrier reef. Both inner and outer reef M. annularis displayed an increase in P and R with increasing temperature between 29°C and 32°C, but above 32°C P and R sharply declined. P/R ratio versus temperature showed a significant difference between the elevations of the regression lines suggesting that M. annularis from the outer barrier reefs may have been more physiologically stressed than those from the inner lagoon reefs when exposed to acute temperature changes. These results emphasize that thermal stress must be considered within the context of acclimation temperature, and that short-term exposures may have physiologically important effects on this species.     

Interlinked temporal changes in environmental conditions, chemical characteristics of sediments and macrofaunal assemblages in an estuarine intertidal sandflat (Seto Inland Sea, Japan)

Five field surveys were conducted in an estuarine intertidal sandflat of the Seto Inland Sea (Japan) between April 1994 and April 1995. Chlorophyll a, pheopigments, total organic carbon and acid-volatile sulphides (AVS) of surface and subsurface sediments, and macrofaunal assemblages were investigated in parallel at 15 stations. Monthly hydrological data of low-tide creek water adjacent to the flat were used as a complementary environmental characterisation of the study area. Strong temporal changes were found among sampling dates, most remarkably in autumn with a major increase of algal detritus and AVS, a sharp reduction in macrofaunal abundances and species richness, and a massive mortality of the clam Ruditapes philippinarum. This dystrophic event was preceded by a photoautotrophic and hypertrophic spring–summer characterized by abundant fresh (i.e., living) algal material, including microphytobenthos and macroalgae (Ulva sp.). In summer, abundant macrofaunal assemblages reached the highest biomass values (455 g wet weight m⁻² or 60.6 g ash free dry weight m⁻²), with a major contribution of filter-feeding bivalves Musculista senhousia and R. philippinarum. These are among the highest values reported in the literature for sedimentary shores. From autumn, there was a progressive recolonisation of macrofauna, initiated by few opportunistic polychaetes (e.g., Cirriformia tentaculata and Polydora sp.), apparently promoting a fast sediment recovery in winter, and followed by new bivalve recruits in the next spring. This study provides the first evidence of significant and interlinked within-year changes in chemical characteristics of sediments and macrofaunal assemblages in an estuarine intertidal flat at a small spatial scale (i.e., tens of meters). This demonstrates the high temporal variability of species–environment relations in these systems and a close relationship in seasonally driven trophodynamic processes among primary producers and benthic consumers. We conclude that a thorough parallel evaluation of the temporal changes in chemical characteristics of sediments should be taken into account in assessing the year-round distribution and changes of intertidal macrofauna, particularly in eutrophic, estuarine intertidal flats.An erratum to this article can be found at     

Response of benthic foraminifera to heavy metal contamination in marine sediments (Sicilian coasts,Mediterranean Sea)

To examine the suitability of benthic foraminifera and their test deformations as bioindicators of pollution in coastal marine environments, we studied foraminifera and metal concentrations in 72 marine sediment samples, collected from the inner shelf along the Sicilian coast (Gulfs of Palermo and Termini) and on the south-eastern coast of Lampedusa Island. These areas are characterised by different environmental conditions. On the basis of pollution sources and foraminiferal assemblages, we recognised different zones in the Gulf of Palermo. The most polluted zones showed high metal concentrations, and low diversity of benthic foraminifera with species typical of stressed environments. By contrast, the lowest polluted zones showed a high population density and the highest percentages of epiphytes. Epiphytes were abundant where a Posidonia oceanica meadow was present and decreased in the most polluted zones. Sediments of the Gulf of Termini and Lampedusa exhibited high percentages of benthic foraminifera typical of well-oxygenated waters and low concentrations of metals, with the exception of sites located near sewage outfalls and harbour areas. Furthermore, even though deformed tests are commonly known in natural stressed environmental conditions, this study shows that in the most polluted zones, benthic foraminifera were characterised by the highest percentages of deformed individuals.     

Biotic indexes reveal the impact of harbour enlargement on benthic fauna

Environmental effects of enlargement works in Puerto Calero (Lanzarote, Canary Islands) were analysed in the sediments using abiotic variables (total hydrocarbons, polyaromatic hydrocarbons, organic matter and granulometry) and three biotic indexes (AMBI, M-AMBI and BENTIX). A before–after/control–impact (BACI) design was developed with four sampling campaigns, before (November 2004), during (March and July 2005) and after (July 2006) works. Inner stations were characterised by high concentrations of pollutants (total hydrocarbons and polyaromatic hydrocarbons). A temporal trend (2004–2006) was observed in macrofaunal assemblage structure, and thus in AMBI, M-AMBI and BENTIX indexes. Macrobenthic assemblages also mirrored the pollution gradient, with bioindicator species in inner stations (the polychaete Pseudopolydora paucibranchiata and the mollusc Abra alba), sensitive species (the sipunculid Aspidosiphon muelleri, the crustacean Pariambus typicus and the polychaete Aponuphis bilineata) in outer stations and ubiquitous species (the polychaetes Aricidea assimilis and Armandia polyophthalma) in both stations (inner and outer).     

Community development on subtidal temperate reefs: the influences of wave energy and the stochastic recruitment of a dominant kelp

Patterns of community development on subtidal rocky reefs in Marmion Lagoon, southwest Australia, were investigated with a settlement panel experiment. We tested the hypothesis that community development would differ between outer and inner reefs lines, because exposure to swell and wave energy was significantly greater on outer reefs. Following a 14-month deployment, we recorded pronounced variability between panels and sites, but did not detect any effect of wave exposure on the structure of panel assemblages. Subsequent data exploration suggested the importance of the presence of kelp recruits (Ecklonia radiata) in structuring the overall assemblage. Panel assemblages with kelp recruits were significantly different in structure to those without, principally because of greater space coverage of encrusting coralline algae and less coverage of red turfing algae, spirorbids, and bryozoans. Mature E. radiata act as ecosystem engineers in subtidal rocky reefs in southwest Australia. Our results suggested the importance of young, recruiting kelps in determining patterns of early community development on newly available hard substrata.     

Prey selection in <Emphasis Type="Italic">Octopus rubescens</Emphasis>: possible roles of energy budgeting and prey nutritional composition

This study explores the relationship between energy budgeting and prey choice of Octopus rubescens. Seventeen male Octopus rubescens were collected between June 2006 and August 2007 from Admiralty Bay, Washington. Prey choices made by individuals in the laboratory deviated widely from those expected from a simple optimal foraging model. O. rubescens chose the crab Hemigrapsus nudus over the clam Nuttallia obscurata as prey by a ratio of 3:1, even though prey energy content and handling times suggested that this octopus could obtain 10 times more energy intake per unit time when choosing the latter compared to the former prey species. Octopus energy budgets were similar when consuming either of the prey species except for lipid extraction efficiency that was significantly higher in octopuses consuming H. nudus. This suggests that lipid digestibility may play an important role in the prey choice of O. rubescens.     

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     

Ecology of Echinometra mathaei (Echinoidea: Echinodermata) at Diani Beach,Kenya

Studies were carried out on the inner and outer coral reefs at Diani Beach on the Kenya coast to assess the distribution, density and behaviour of Echinometra mathaei (de Blainville). Transects 1 m wide were run on the two reefs in April, June and September, 1970. Test measurements on representative samples from the animal populations on both reefs were also taken. Direct observations on specimens of E. mathaci in selected rock pools on the outer reef were made to determine their movement, gregariousness, homing and feeding behaviour. Population density was higher on the inner reef furthest from the sea at low tide than on the outer reef. On the submerged inner coral reef at low tide, E. mathaei occurred mainly exposed on the seaweeds, but, on the exposed outer reef, its main niches were crevices in rock pools and under coral ledges. Sizefrequency distributions revealed that smaller individuals occurred on the inner reef and larger ones on the outer reef. The growth rate of E. mathaei was estimated from the positions of modal values, calculated from size-frequency distributions. No gregarious or homing behaviour was observed and, once settled in a suitable crevice, E. mathaei showed little movement.     

Population dynamics of beech scale (Cryptococcus fagisuga) (Coccina,Pseudococcidae) related to physiological defence reactions of attacked beech trees (Fagus sylvatica)

Summary Changes in the density of beech scale infestation (Cryptococcus fagisuga) of a mature beech stand (Fagus sylvatica) were investigated after gap-cutting in limed and unlimed areas bordering on the gaps and in untreated areas over a 5-year period. Parallelly the concentrations of sucrose, protein amino acids, and procyanidins were examined in the inner and outer bark of non-infested beech trees and beeches infested by beech scale.Irrespective of liming in the areas bordering on the gaps the proportion of beech trees with increasing beech scale infestation was significantly higher, the proportion of beeches with diminishing infestation was significantly lower than in the untreated control areas. Trees with the same infestation tendency occurred in clusters.Physiological defence reactions in the inner and outer bark of the infested beech trees were registrated as an increase in the procyanidin content and a decrease in the protein amino acid content. A change in the outer bark—inner bark—ratio of the infested beech trees showed a transfer of compounds between the inner and outer bark. During the period of observation long-lasting shifts occurred in the pattern of compounds of the bark irrespective of actual infestation intensity.     

The effects of sand movement on intertidal seaweeds and selected invertebrates at Bound Rock,New Hampshire,USA

A seasonal study of sand movement and the benthic intertidal organisms at Bound Rock, New Hampshire, USA was conducted between November, 1973 and February, 1975. The site is subjected to irregular sand fluctuations, as well as diurnal, neapspring and major summer sand intrusions. The abundance and distribution of intertidal species was interrelated with the historical sand fluctuations at the area. For example, the lower limits of Mytilus edulis, Balanus balanoides and Porphyra umbilicalis approximated the highest summer sand elevations. Highly abraded rock surfaces in the lower intertidal zone were dominated by opportunistic annuals (e.g. Enteromorpha spp.) and perennial psammophytic or sand-loving seaweeds (e.g. Ahnfeltia plicata and Sphacelaria radicans). Overall, the intertidal seaweed populations at Bound Rock showed a lower number of perennials and fewer species than adjacent rocky shores. The low species diversity of seaweeds at the study site is attributable to unstable environmental conditions and a limited number of habitats. Even so, several psammophytic microhabitats are evident at Bound Rock depending upon the extent of sand burial and abrasion. The morphological and reproductive adaptations of several psammophytic species are discussed.Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as Scientific Contribution No. 812.Scientific Contribution No. 53 of the Jackson Estuarine Laboratory.     

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.     

Seasonal abundance,distribution and growth of postlarval and juvenile grass shrimp (Palaemonetes pugio) in a Georgia,USA, salt marsh

Postlarval and juvenile grass shrimp (Palaemonetes pugio Holthuis) ≦15 mm total length (TL) were abundant at low tide in shallow aquatic microhabitats (i.e. puddles and films of residual tidal water) in the intertidal zone of a salt marsh on Sapelo Island, Georgia, USA from 1982 to 1984. The highest concentrations of young P. pugio occurred at 190 to 200 cm above mean low water, ∼10 to 20 cm below mean high water. The intertidal distributions of young grass shrimp expanded and contracted with changes in tidal amplitude. Postlarval grass shrimp (6 to 8 mm TL) continuously recruited into the intertidal marsh population from May until October, but densities varied in a regular pattern with peaks in abundance occurring at ∼2-week intervals, corresponding to spring tide periods in the lunartidal cycle. Although present nearly year-round in the intertidal marsh, juveniles (9 to 15 mm TL) were most abundant from August to October. Apparent growth rates of individuals up to 15 mm TL averaged 0.268±0.026 (mean±95% C.I.) mm d^-1 from May to October and 0.070±0.032 mm d^-1 in November and December. Unlike larger aquatic organisms, which can forage in the emergent marsh only when it is flooded by the tide, juvenile grass shrimp have constant access to intertidal resources. Although potentially important predators in this system, the role of young P. pugio in the trophic organization of salt marsh benthic invertebrate assemblages has yet to be examined. Contribution No. 576 of the University of Georgia Marine Institute     

A diatom assemblage living below the surface of intertidal sand flats

The assemblages of species living on and in intertidal sand flats at Barnstable Harbor, Massachusetts (USA) are briefly described. An endopsammic assemblage of diatoms, mainly Amphora species, living below the surface is described for the first time in such a marine situation. Some aspects of its ecology and relationship to the other diatom assemblages living on the sediments are discussed.     

Effects of physical disturbance on nematode communities in sand and mud: a microcosm experiment

A microcosm experiment was carried out to evaluate the effects of continuous and spasmodic physical disturbance of differing frequency on the structure of nematode communities of intertidal sand and mud. There was a marked, characteristic change in abundance and diversity for both sediment types. In the sand microcosms, the majority of univariate measures of community structure, including species diversity, were lowest in the sediments subjected to a high frequency of disturbance. For the mud microcosms, most univariate measures reached their highest values in the treatments with an intermediate frequency of disturbance and were lower in treatments subjected to both higher and lower frequencies. Multivariate ordinations for both nematode assemblages showed a clear separation of undisturbed controls and disturbed treatments, but only for the muddy sediment was there a graded change in community composition with increasing frequency of disturbance. These results confirmed our a priori expectation that nematode assemblages from mobile sandy sediments would be more resilient to physical disturbance than those from sheltered muds, and these observations are considered in the context of Connell's intermediate disturbance hypothesis. Received: 7 May 1997 / Accepted: 10 October 1997     

Comparison of mussel bed community between two intertidal mytilids Septifier virgatus and Hormomya mutabilis

Mussel bed community structure of two intertidal mytilids, Septifer virgatus (Wiegmann) and Hormomya mutabilis (Gould), whose beds were contiguous vertically on a rocky intertidal shore (Wakayama Prefecture, Japan), was compared between 1982 and 1983. In the upper S. virgatus bed, crustaceans and bivalves were dominant in terms of both number of individuals and biomass. There were three barnacle species representative of the epizoans, two isopods and one amphipod as mobile fauna, and two bivalves as infauna. The lower H. mutabilis bed supported virtually no epizoans or mobile fauna. Infaunal free-ranging polychaetes and sipunculids were dominant in terms of both number of individuals and biomass. The H. mutabilis bed contained a much greater amount of sediment than did the S. virgatus bed, and the interstices among individual H. mutabilis and among their byssal threads were filled with sediment. The biomass of six of nine species dominant in the S. virgatus bed was negatively correlated with the amount of sediment. Recruitment of these faunal assemblages into artificial mussel clumps was examined in mussel enclosure experiments, and a negative sediment effect in H. mutabilis clumps was detected for one isopod and one limpet species. Factors causing the differences between two mussel bed communities are discussed, focusing on the effects of sediment.