High intertidal community organization on a rocky headland in Maine,USA

A mosaic patchwork of the barnacle Balanus balanoides L., the mussel Mytilus edulis L., and the alga Fucus vesiculosus L. was found in the transitional region between the mid and high intertidal zones on a rocky headland on Mount Desert Island, Maine, USA. The development of the mosaic was observed by following recruitment and survival of B. balanoides in denuded patches located at the same tidal level along a 60 m stretch of shore. Barnacle recruitment was least under canopies of F. vesiculosus and greatest in open areas kept moist at low tide by surf. Barnacle survival after settlement was least under the F. vesiculosus canopy due to the whiplash effect of the algal fronds in the surf and greatest in open areas free from competition from mussels. In open areas, early mortality was correlated with settlement density. In areas of dense settlement (60 spat cm^-2) up to 90% mortality resulted within 5 months from crowding associated with growth. In older individuals crowding produced hummocks of elongated, weakly attached barnacles which were more prone to removal by surf than uncrowded barnacles. Mussels exerted competitive dominance over barnacles for space and the presence of mussel beds prevented further barnacle recruitment. Mussels suffered extensive mortality during winter storms when surf removed dense mats of weakly attached mussels. The patchy distribution of mussels and barnacles results from irregular rock substrata producing numerous environmental patches with respect to wave exposure and drainage at low tide, and from densitydependent mortality of both mussels and barnacles which creates patches of new colonizable space within each environmental patch.     

Community succession following massive ice-scour on a rocky intertidal shore: recruitment,competition and predation during early,primary succession

The importance of species interactions and recruitment variability was examined during the first year and a half of primary succession (1988–1989) on an exposed rocky seashore near Halifax, Nova Scotia. Previous work suggested that emergent rock on these shores is normally dominated by fucoid rockweeds because predatory whelks control the sessile animal competitors, mussels and barnacles, and because herbivorous littorinids control ephemeral algal competitors. Abundances of all species except seasonal ephemeral algae were very small throughout this experiment and we found no significant effects of carnivory, herbivory, plant-animal competition or plant-plant competition. A slight facilitation of Fucus recruitment is attributed to a thin mat of ephemeral, blue-green algae. Very few other studies have directly manipulated intertidal ephemeral algae. As primary succession may be very rare in this assemblage, these results may be specific to these circumstances, but they highlight the varying importance of species interactions with variable recruitment. In particular, it appears that variations in recruitment success may be important to community structure, even when recruitment is not limited by propagule supply. The scale of the study also provides insight into successional processes occurring after the recent, extensive ice-scour of exposed seashores in this region.     

Variation and persistence of the middle rocky intertidal community of central Chile,with and without human harvesting

Humans are an important intertidal predator in central Chile. Following a five-year study we report on the effects of the exclusion of humans from the rocky intertidal at Las Cruces, central Chile. The middle intertidal of harvested and non-harvested areas diverged in species diversity and composition during the experiment. In harvested areas the middle intertidal rocky shore was dominated throughout the study by a monoculture of mussels, Perumytilus purpuratus. When humans were excluded, the middle intertidal community switched to one dominated by barnacles (predominantly Jehlius cirratus and Chthamalus scabrosus); this community has persisted for at least three years, despite the presence of forces e.g. mussel larvae, that have the potential to alter the community structure. Such changes were mediated by the muricid gastropod Concholepas concholepas, a key-stone predator. As a consequence of the above changes, the species diversity, H', (primary space occupiers) in the non-harvested area increased from H'=0 at the beginning of the study in 1983 (when the middle intertidal community was dominated by mussels), to values ca H'=2 toward the middle of the study in 1984 (which coincided with the maximum predatory impact of C. concholepas) and subsequently decreasing to ca H'=0.5 at the end of the study in 1987 (when the middle intertida community was dominated by barnacles).Please address all correspondence to J. C. Castilla, Department of Zoology, University of Washington, Seattle, Washington 98195, USA, until March 1990     

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.     

Environmental stress,competition and dominance of Crassostrea virginica near Beaufort,North Carolina,USA

Observations and experiments were made at 2 intertidal areas near Beaufort, North Carolina, USA from May 1977 to July 1978 to determine why the oyster Crassostrea virginica dominated the community in areas protected from wave action but not in areas directly exposed to waves. Barnacles, oysters, the green alga Enteromorpha sp. and the mussel Brachiodontes exustus were the main occupiers of primary space at the mid and low intertidal levels of exposed areas. The intertidal community at the protected site consisted of a mid intertidal occupied by the barnacles Balanus amphitrite and Chthamalus fragilis, and the oyster C. virginica, and a low intertidal dominated by C. virginica. The exposed area was highly variable with high colonization and mortality for all species producing large seasonal changes in structure. The protected site remained constant throughout the year; there was no evidence of further colonization of either barnacles or oysters and mortality was very low. Recolonization experiments, selective removal of species, and growth and survival data demonstrated that C. virginica does not become dominant at exposed locations because (1) the constant wave shock at the ocean site reduces growth and increases mortality of young and adult oysters and (2) oysters are outcompeted by the mussel B. exustus. The monopolization of space by C. virginica at protected sites contrasts with studies north of Cape Hatteras where the abundance of predators produces a more diverse and heterogeneous community. Predation was unimportant in Beaufort because predators were absent at the exposed areas and the oyster drill Urosalpinx cinerea was restricted to the subtidal zone at the protected site. This absence of predators indicates a higher level of environmental stress near Beaufort compared to areas farther north.     

Population dynamics of the acorn barnacles, Tetraclita squamosa and Tetraclita japonica (Cirripedia: Balanomorpha), in Hong Kong

Tetraclita squamosa and Tetraclita japonica are common, intertidal barnacles. In Hong Kong, the population dynamics of the two species exhibited spatial and temporal variation on two semi-exposed shores. T. squamosa produced egg masses from May–June and annual settlement and recruitment occurred from June–July. In contrast, settlement and recruitment of T. japonica was sparse from March–May, but intensified in July–October. Mature gonads and egg masses were, however, only present from September–November, suggesting the larvae from the two settlement pulses originated from other locations as well as Hong Kong populations. Settlement intensity and post-recruitment mortality of the two species varied between sites, possibly due to spatial variation in free rock space, physical transport of larvae and abundance of food in the water column.Communicated by T. Ikeda, Hakodate     

Seascape-dependent subtidal-intertidal trophic linkages

In this study, we test in southern New Zealand a conceptual model of food web linkage that is seascape dependent, which can explain some of the variability in rocky shore community structure among sites and coasts. Using a comparative-experimental approach at local and distant sites we demonstrate that mobile subtidal predators (fish and crabs) can exert strong predation pressure on small mussels in the low tidal zone, but only in sites where the seascape includes subtidal reefs. On intertidal benches with adjacent subtidal reefs (+SR), 60-100% of small (5-15 mm) transplanted mussels were removed within a day from experimental tiles on the low shore when unprotected from predation, compared to fully caged controls that had approximately 100% survival over several months. In partial cages that exclude fish but not crabs, survivorship was intermediate. In contrast, on benches without subtidal reefs (-SR) 40-100% of mussels survived for months, even if unprotected. This difference is expressed in lower cover (0-60%) of mussels on rocks at +SR benches compared to -SR benches (70-99%). The central to northern west coast of the South Island is composed mostly of -SR benches, and predation on small mussels there was low and similar to the -SR benches on the east coast, whereas the +SR benches on the east coast had much greater predation. This contrasts to other studies in New Zealand that examined only predation on larger mussels by seastars and concluded that predation is strong on the west coast and weak on the east coast. Excluding large predators from low-shore areas with new recruits for a year in one +SR site showed longer-term predation effects on their abundance and cover. Short-term sampling at the east coast sites showed that mussel settlement was greater in -SR compared to +SR sites, providing some evidence that seascapes may also affect settlement. Overall, predation depended on the local seascape and ultimately affected community structure via suppression of effective recruitment rates. This study emphasizes the importance of predation on early life stages of basal species and the influence of seascapes on top-down interactions between subtidal predators and their intertidal prey.     

Geographical and seasonal variation of trace metal bioavailabilities in the Gulf of Gdansk,Baltic Sea using mussels (<Emphasis Type="Italic">Mytilus trossulus</Emphasis>) and barnacles (<Emphasis Type="Italic">Balanus improvisus</Emphasis>) as biomonitors

The barnacle Balanus improvisus and the mussel Mytilus trossulus have been used as biomonitors of the trace metals Cu, Zn, Cd, Fe, Pb, Mn and Ni at five sublittoral sites in the Gulf of Gdansk (Baltic Sea) between February 2000 and September 2001. The study has established a benchmark against which future biomonitoring programmes will be able to establish changes in local metal pollution, particularly if metal loadings in the river Vistula (draining into the Gulf) alter in the future. The study highlighted differences in trace metal bioavailabilities to both barnacles and mussels, geographically and over time. Accumulated metal concentrations of Cu, Zn, Fe, Pb and Ni, but not Cd or Mn, were correlated in the barnacles and mussels, suggesting that the bioavailabilities of the former metals to the two biomonitors were similar. The barnacles showed greater discriminatory power than the mussels as trace metal biomonitors. Concentrations of trace metals in surficial sediments (<63 m) did not correlate significantly with accumulated metal concentrations in either barnacles or mussels, indicating that sediment metal concentrations are not necessarily good proxy measures of ambient trace metal bioavailabilities to the local coastal filter feeders.Communicated by J.P. Thorpe, Port Erin     

Seaweed richness and herbivory increase rate of community recovery from disturbance

The importance of herbivores and of plant diversity for community succession and recovery from disturbance is well documented. However, few studies have assessed the relative magnitude of, or potential interactions between, these factors. To determine the combined effect of herbivory and surrounding algal species richness on the recovery of a rocky intertidal community, we conducted a 27-month field experiment assessing algal recruitment and succession in cleared patches that mimic naturally forming gaps in the ambient community. We crossed two herbivore treatments, ambient and reduced abundance, with monocultures and polycultures of the four most common algal species in a mid-high rocky intertidal zone of northern California. We found that both the presence of herbivores and high surrounding algal richness increased recovery rates, and the effect of algal richness was twice the magnitude of that of herbivores. The increased recovery rate of patches containing herbivores was due to the consumption of fast-growing, early colonist species that preempt space from perennial, late-successional species. Mechanisms linking algal richness and recovery are more numerous. In polycultures, herbivore abundance and species composition is altered, desiccation rates are lower, and propagule recruitment, survival, and growth are higher compared to monocultures, all of which could contribute the observed effect of surrounding species richness. Herbivory and species richness should jointly accelerate recovery wherever palatable species inhibit late-successional, herbivore-resistant species and recruitment and survival of new colonists is promoted by local species richness. These appear to be common features of rocky-shore seaweed, and perhaps other, communities.     

<Emphasis Type="Italic">Semibalanus</Emphasis><Emphasis Type="Italic">balanoides</Emphasis> in winter and spring: larval concentration,settlement, and substrate occupancy

This study measured the progression from pelagic larvae to juvenile barnacles, and examined whether recruitment of barnacles, Semibalanus balanoides Linnaeus, at two intertidal sites in contrasting hydrodynamic regimes was determined by pre-settlement or post-settlement processes. The two sites were 1.5 km apart in the vicinity of Woods Hole, Mass., USA. Quantitative plankton samples were taken twice weekly from December 1997 to May 1998 at a nearby site as an estimate of nearshore larval abundance. The presence of S. balanoides nauplii was noted, and cyprids were enumerated and measured. Larval settlement at the two sites [Gansett Point, Buzzards Bay (GP) and Little Harbor, Vineyard Sound (LH)] was estimated from examination of replicate settlement plates exposed for 2 or 3 days throughout the settlement season, and from replicate plots on marked rock quadrats at each site. On both plates and rocks settled cyprids and metamorphs were enumerated. Space occupancy on unmanipulated rock quadrats by all stages from cyprids to adult barnacles was also examined. Settlement occurred from 2 January to 20 May, and major settlement peaks coincided with peaks in pelagic cyprid concentration at LH, but not at GP. Space occupied by juvenile barnacles was close to zero up until late February despite substantial settlement prior to that. At LH, juvenile barnacle cover was zero at the end of the observations; all settlement failed. Almost 100% of settled cyprids failed to metamorphose within 2 days from late January to late March. Then the proportion metamorphosing increased sharply coinciding with a sudden increase of 3°C in water temperature. Observed site differences in space occupancy by juvenile barnacles suggest that while cyprid supply is a necessary condition for barnacle settlement, other factors affecting metamorphosis of settled cyprids and early juvenile mortality determine recruitment.     

Patch dynamics of mussel beds near São Sebastião (São Paulo), Brazil

In permanent quadrats on exposed and protected shores near São Sebastião (São Paulo), Brazil, changes in percent cover of mussels [Brachidontes solisianus (d'Orbigny)] and barnacles [Chthamalus stellatus bisinuatus (Pilsbury)] were followed from November 1979 through December 1982. Mussels gradually disappeared from both shores, while barnacles came to dominate the exposed, but not the protected, shore. Decreases in percent cover of mussels were related to periods of higher temperatures. Mussels were only transient dominants of the mid-intertidal zone due to lack of recruitment and high summer mortality. This is probably because the study was carried out near the northern limit of their occurrence as a dominant of mid-intertidal communities.Deceased     

Abundance of Fucus spiralis and ephemeral seaweeds in a high eulittoral zone: effects of grazers,canopy and substratum type

According to a widely accepted view of intertidal community organization in the north-west Atlantic Ocean, the abundance of perennial seaweeds is regulated by the activities of herbivores (which control ephemeral algal competitors) and carnivores (which control populations of competing filterfeeders). This concept was examined in 1987–1988 in a eulittoral belt on the Atlantic shores of Nova Scotia, Canada, dominated by a closed canopy of Fucus spiralis. A factorial design was used to test the effects of (a) grazers (there were no carnivores present, (b) adult F. spiralis canopy, and (c) substratum type on the abundance of perennial and ephemeral seaweeds. Grazers had no significant (p>0.05) effect on the density or cover of juvenile F. spiralis, but significantly reduced canopy cover of adults through the winter. The presence of grazers significantly enhanced the cover of ephemeral algae in early spring. Barnacle presence enhanced the cover of juvenile F. spiralis, but reduced the canopy of adult plants through abrasion. The most important regulator of recruitment density in F. spiralis was the presence of a canopy of conspecific adults. Canopy greatly reduced juvenile development. There was no significant relationship between the covers of perennial F. spiralis and ephemeral seaweeds. This study demonstrates that the recruitment of the perennial rockweed population examined is not dependent on the activities consumer animals. The results contrast with work in the midshore intertidal of New England, where the activities of herbivores are thought to regulate the abundances of perennial seaweeds.     

Effects of crowding and wave exposure on penis morphology of the acorn barnacle, Semibalanus balanoides

Wave action and low population density can strongly reduce the ability of sessile acorn barnacles to find mates and copulate. For Semibalanus balanoides, penis morphology varies with wave exposure and with characteristics of the mating neighborhood. Field experiments were conducted at five intertidal sites on Long Island, New York, USA from July to December 2005 to determine how wave exposure and aggregation structure influence the length, diameter, mass, and number of annulated folds of the penis. Sparsely crowded barnacles had more annulations in the penis and are inferred to have greater ability to stretch. At higher wave exposure, the diameter of the penis was greater, but the mass was not. This study identifies density of crowding as the most important cue that barnacles respond to when perceiving their mating group and details how penis morphology varies in response to wave exposure.     

Allometry and growth of the green-lipped mussel Perna canaliculus in New Zealand

Length is the most precise (and the most practical) linear measurement for predicting total weight (r>0.98 at P=0.001) in the green-lipped mussel Perna canaliculus Gmelin. The allometry varies with the environmental conditions under which the mussels grow, resulting in morphologically distinct forms of raft- and shore-grown mussels. Mussels grown intertidally are wider, less high and heavier than mussels of similar length grown in suspension. Increase in length and total weight of P. canaliculus grown in suspended cultivation was recorded at 8 experimental sites around New Zealand, during 1973–1975. Comparisons are drawn with growth on an intertidal mussel bed, where length increase was less than half that in the same period in suspension. The growth rate of mussels transferred from intertidal to suspended conditions depends on the size at transfer. Close similarity in growth rate occurred at the majority of sites in spite of a direct correlation between water temperature and length increment and substantial, differences in temperature between sites. Reasons for the uniformity are suggested. Average values for growth at sites over the northern half of New Zealand were 73 mm length (32.5 g weight) after 12 months, 113 mm (110 g) after 2 years. Growth continued throughout the year, highest growth rates corresponding to highest water temperatures. Variation due to depth was not significant. Larger mussels grew more slowly. P. canaliculus can be grown in suspended cultivation in New Zealand at a rate comparable to that in other commercial mussel-farming areas.     

Spatial patterns of macroalgal abundance in relation to eutrophication

Based on a large monitoring data set from Danish coastal waters we tested the hypotheses: (1) The vertical pattern of algal abundance is regulated by exposure in shallow water and by light limitation towards deeper water, resulting in a bell-shaped abundance curve, with peak abundance located deeper at more exposed sites, (2) in deeper water, total algal abundance and abundance of perennial algae decrease along a eutrophication gradient while (3) abundance and relative abundance of opportunists increase. The vertical pattern of algal abundance showed a peak at intermediate water depths which was located deeper in more exposed areas and thus confirmed our first hypothesis. For algae growing from depths of peak abundance and deeper, the study demonstrated that total algal abundance and abundance of perennials and opportunists at given depths decreased significantly along a eutrophication gradient and the relationships had high explanatory power (R ² = 0.53–0.73). These results confirmed our second hypothesis. By contrast, the relative abundance of opportunists responded solely to salinity and was largest in the most brackish areas, in contradiction to hypothesis three. The lack of coupling between eutrophication and relative abundance of opportunists arises because both opportunists and the entire algal community were light limited and their ratio therefore relatively insensitive to changing water clarity. The analyses indicated that algal abundance initially responded slowly to increasing eutrophication but showed a more marked response at TN concentrations of 35–40 μM. However, the existence of possible threshold nutrient levels demands further analyses.     

Phenology and demography of a marine specialist herbivore: Placida dendritica (Gastropoda: Opisthobranchia) on the central coast of Oregon

The occurrence of small specialist herbivores in marine rocky intertidal communities has been largely ignored. From 1984 to 1989, the phenology and demography of the common oligophagous ascoglossan opisthobranch Placida dendritica A.& H. was documented at several sites along the central coast of Oregon, USA. P. dendritica was found on the low intertidal green algae Codium setchellii Gardn., C. fragile (Sur.) Har. and Bryopsis corticulans Setch. during spring and summer. Ascoglossan abundance on Codium spp. varied spatially and temporally. Peak density usually occurred in May or June. Maximum ascoglossan live mass was 4 to 6 mg on Codium spp. and 100 mg on B. corticulans. Larval recruitment of P. dendritica to algal hosts was continuous during spring and summer. Rapid recruitment (200 to 400 sea slugs thallus^-1mo^-1) of newly metamorphosed ascoglossans to C. setchellii in wave-protected areas suggests that high densities of competent larvae occurred in nearshore waters and/or that larvae were efficient at locating algal hosts. Ascoglossan growth was rapid, with less than one month elapsing from larval settlement to adult sexual maturity. Therefore, populations of P. dendritica were composed of many overlapping generations. Movement of adult ascoglossans among thalli of C. setchellii was rapid. Continuous larval settlement and adult movement enabled P. dendritica not only to locate and colonize its algal hosts quickly but also to maintain a persistent association with algal hosts during spring and summer.     

Diatoms of the microphytobenthic community in a tropical intertidal sand flat influenced by monsoons: spatial and temporal variations

Seasonal variations in the microphytobenthic diatom community were investigated in an intertidal sand flat of a tropical marine environment influenced by monsoons. Cores of sediments were collected along the beach gradient: low tide, mid tide and high tide zone up to a depth of 15 cm.. Diatom abundance was lowest during the monsoons and highest during the post-monsoons and the early pre-monsoon season throughout the intertidal transect. Diatom diversity was highest at the mid tide, followed by the high and low tide zones. Diatoms were viable up to a depth of 15 cm throughout the intertidal transect. The diatom community included the pennates, the permanent residents of this area, centric genera, which lead an attached mode of life and also some planktonic genera, brought in from ambient waters. Among the pennates, Navicula and Amphora were the dominant genera whereas in the case of centrics, Thalassiosira dominated the community throughout the intertidal transect down to a 15 cm depth. . Grain size fractions, which served as predictors of some diatom genera changed with tidal zones. The effect of winds on the resuspension of the pennate diatoms was evident only at the low tide zone down to a depth of 5 cm . Chlorophyll a concentration proved to be a good predictor of both pennate and centric diatom abundance at the low tide zone down to a depth of 10 cm and at the mid tide zone down to a depth of 5 cm.. However, even though chlorophyll a concentrations failed to reveal any positive correlation with the diatom abundance at both the deeper sediment layers and the high tide zone, the fact that viable cells were present at these areas reveal that the diatoms adopt survival strategies, contributing significantly to the carbon budgets of such unstable habitats.     

Interactions among sea otters, sea stars, and suspension-feeding invertebrates in the western Aleutian archipelago

The structuring and organizing effects of apex predators on ecosystems are becoming increasingly well documented. The enhancement of kelp forests via sea otter predation on herbivorous sea urchins is among the earliest and best known examples. This study provides evidence for direct and indirect trophic interactions among sea otters, predatory sea stars, and filter-feeding mussels (Mytilus trossulus) and barnacles (Semibalanus cariosis). In western Massacre Bay at Attu Island (173°E, 53°N), subtidal transects showed sea star body size and biomass density declined markedly between 1983 and 1994 as sea otters reinhabited this area. Mussels and barnacles translocated from the rocky intertidal zone to shallow subtidal habitats to assess loss rates from sea star predation showed lower mortality rates after the arrival of sea otters. Prey mortality rates in subtidal caged controls were consistently low and similar to those of intertidal controls in both years. These findings elucidate a trophic pathway by which sea otters can influence ecosystems separate from the well-known sea otter/sea urchin/macroalgae cascade.     

Habitat structure and complexity as determinants of biodiversity in blue mussel beds on sublittoral rocky shores

Habitat-forming, ecosystem engineer species are common in most marine systems. Still, much uncertainty exists about how individual and population-level traits of these species contribute to ecosystem processes and how engineering species jointly affect biodiversity. In this manipulative field experiment, we examined how biodiversity in marginal blue mussel beds is affected by blue mussel (1) body size, density and patch context and (2) presence of fucoid and algal structures. In the study area, bladder-wrack (Fucus vesiculosus), filamentous algae and blue mussels (Mytilus edulis) coexist at shallow depths in a variety of patch configurations and offer complex habitats with a high variability of resources. We hypothesized that complexity in terms of mussel bed structure and algal presence determines species composition and abundance. Results from the experiment were compared with macrofaunal communities found in natural populations of both engineering species. Results show that the physical structure and blue mussel patch context are important determinants for species composition and abundance. Results further show that the presence of algal structures positively affects diversity in blue mussel habitats due to increased surface availability and complexity that these algae offer. This study shows that blue mussel beds at the very margin of their distribution have an indisputable function for promoting and maintaining biodiversity and suggest that facilitative effects of habitat-modifying species are important on Baltic Sea rocky shores with fundamental importance to community structure.     

Bacterivory of a mudflat nematode community under different environmental conditions

The fate of the benthic bacterial biomass is a topic of major importance in understanding how soft-bottom environments function. Because of their high abundance, production and nutritional value, benthic bacteria may constitute an important food resource for benthic fauna. The trophic role of bacteria for a nematode community on the Brouage mudflat (Marennes-Oléron-France), dominated by three species: Chromadora macrolaima (64% of the abundance), Daptonema oxycerca (15%) and Ptycholaimellus jacobi (8%), was determined in grazing experiments using ¹⁵N pre-enriched bacteria. On intertidal flats, seasonal, tidal and circadian cycles induce strong variations in environmental conditions. Grazing experiments were performed in order to measure the effects of abiotic (temperature, salinity and luminosity) and biotic (bacterial and algal abundances) factors on assimilation rates of bacteria by nematodes. In order to assess simultaneously bacteria and algal assimilation rates, algal abundances were modified adding ¹³C pre-enriched Navicula phyllepta. Assimilation rate was significantly lower at 5°C; moreover, general trend shows a prominent temperature effect with an optimum around 30°C. Assimilation at salinity 18 was not significantly different from the assimilation at salinity 31. Assimilation was higher under light conditions than in the dark. Above 10⁹ bacteria ml⁻¹, assimilation of bacteria remained unaffected by bacterial abundance. However, assimilation of algae increased with the algal concentration. Nematode kept feeding under conditions of stress, which are typical of the surficial sediment habitat and they appeared to be principally dependent on the algal resource.