Effect of periphyton grazing by Hydrobia ulvae on the growth of Zostera noltii on a tidal flat in the Dutch Wadden Sea

The decrease of the intertidal seagrass Zostera noltii in the Dutch Wadden Sea may have been the result of enhanced periphyton load due to eutrophication. Decrease of this seagrass species coincided with an increase in the mudsnail Hydrobia ulvae. Feeding of this mudsnail on periphyton may have partly counteracted an increase of periphyton biomass. Exclosure experiments on seagrass stands in the Dutch Wadden Sea in 1987 showed that density of periphyton on leaves of Z. noltii decreased significantly with increasing density of grazers. An increased density of mudsnails significantly enhanced the density and biomass of seagrass, in particular of the below ground parts. Since this seagrass species survives winter in temperate climate zones mainly by means of rhizomes, grazing may also influence the seagrass dynamics in the long term. Results of the experiment were in agreement with field observations on coinciding low densities of mudsnails and high densities of fouling of seagrass stands on the tidal flats of western Wadden Sea in the late 1970s.     

Clonal architecture in an intertidal bed of the dwarf eelgrass <Emphasis Type="Italic">Zostera noltii</Emphasis> in the Northern Wadden Sea: persistence through extreme physical perturbation and the importance of a seed bank

Genotypic structure and temporal dynamics of the dwarf seagrass, Zostera noltii, were studied in an intertidal meadow that has persisted since prior to 1936 near the Wadden Sea island of Sylt. Samples were collected from two 10 × 10 m plots separated by 250 m from May 2002 to June 2005 and from four 1 × 1 m plots from June 2003 to September 2004. All the samples were genotyped with nine microsatellite loci. No genotypes were shared between the plots separated by 250 m. Genetic diversity was higher in the Wadden Sea than in the other regions of its geographic range. The average clone size (genets) (SD) in the two plots was 1.38 (0.26) and 1.46 (0.4) m2, respectively, with a range up to 9 m2 and <20% persisted for >4 years. A high genetic and genotypic diversity was maintained by annual recruitment of seedlings despite a dramatic decrease in ramet density that coincided with the severe heat stress event of 2003. Fine-scale (1 m2) analysis suggested that extensive loss of seagrass cover precluded space competition among the genets, while a persistent seed bank prevented local extinction. Long-term persistence of Z. noltii meadows in the intertidal Wadden Sea was achieved by high genet turnover and frequent seedling recruitment from a seed bank, in contrast to the low diversity observed in large and long-living clones of Z. noltii and other seagrasses in subtidal habitats.     

The spatiotemporal expression pattern of trypsinogen and bile salt-activated lipase during the larval development of red porgy (<Emphasis Type="Italic">Pagrus pagrus</Emphasis>, Pisces,Sparidae)

A restoration programme was introduced in the Mondego Estuary (Portugal) to recover seagrass beds of Zostera noltii endangered by eutrophication. A long-term survey of 10 years was used to assess the development of the processes involved, focusing one of the key species (Cyathura carinata, Isopoda). The mitigation measures implemented since 1998 (nutrient loading reduction, freshwater circulation improvement and seagrass bed protection) enhanced water quality and seagrass recovery, thus preventing the development of macroalgal blooms. C. carinata was resilient to the occurrence of floods and macroalgal blooms, although both events caused dispersion of individuals. This isopod was not much influenced by the changes occurring in the estuary, showing an unalterable population structure during the entire study period. After 1998, its density and biomass became more stable at an inner unvegetated sand flat area, where this isopod was most abundant; its population slightly increased in a bare mud flat at the middle section of the estuary; but it could not establish successfully in a downstream Z. noltii bed, contrarily to other common estuarine species. Apart from other unknown reasons, the disrupted balanced between trematodes and their hosts, caused by the eutrophication processes, may have an important role in the discontinuity of C. carinata at the Z. noltii bed. If the intertidal areas become fully restored to the original seagrass coverage, high prevalence and intensity trematodes may prevent this isopod and other crustaceans from recovering within the intervened areas, by enhancing host mortality and recruitment failure. In order to avoid this kind of situation, it may be necessary to survey the levels of parasite infestation within the target hosts and safeguard areas where crustaceans present healthy populations.     

Are Wadden Sea tidal systems with a higher tidal range more resilient against sea level rise?

Accelerated sea level rise may have serious implications for the Wadden Sea ecosystem in its present state. If sediment accumulation rates on the extensive intertidal flats stay behind sea level rise, the flats will eventually submerge. Drowning of the flats has negative consequences for nature conservation and for coastal risk management. Based upon an evaluation of steady state relations for Wadden Sea tidal basins, Hofstede (Zeitschrift für Geomorphologie 59(3): 377-391, 2015) postulated that the capacity of these basins to balance sea level rise by accumulation on intertidal flats seems positively related to mean tidal range. In the present study, morphodynamical simulations with a numerical model were performed for two tidal basins in the German Wadden Sea to verify the empirically established hypothesis. The following conclusions are established. Larger mean tidal range improves the capacity of Wadden Sea tidal basins to balance sea level rise. Wadden Sea intertidal flats are effective sediment sinks and seem quite resilient against (higher rates of) sea level rise. Finally, subtidal gullies may constitute a significant sediment source for accumulation on intertidal flats in response to sea level rise. With respect to the limited comparability of the two investigated tidal systems, morphodynamical modelling of all Wadden Sea tidal systems should be conducted.     

Leaf epifauna of the seagrass Thalassia testudinum

The abundance, composition and trophic relationships of metazoan leaf epifauna of the marine angiosperm Thalassia testudinum König were studied in Barbados, West Indies. Approximately 90 species from 11 phyla consisted chiefly of nematodes, harpacticoid copepods, crustacean nauplii, ostracods, and turbellarians. Epiflora- and detritus-feeders dominated the epifauna. Increasing leaf epiphytism was accompanied by faunal changes, most notably increased nematode, harpacticoid and polychaete density. Faunal composition was very similar to that of the temperate seagrass analogue Zostera marina.     

Multivariate analysis of the bentho-demersal ichthyofauna along soft bottoms of the Eastern Atlantic: comparison between unvegetated substrates,seagrass meadows and sandy bottoms beneath sea-cage fish farms

By means of multivariate techniques, we studied: (1) the differences in the structure of bentho-demersal, non-cryptic, fish assemblages associated with unvegetated sandy substrates, vegetated meadows constituted by the seagrass Cymodocea nodosa and the bottoms under the influence of sea-cage fish farms; as well as (2) the persistence of these patterns with regard to different scales of spatial variability, across three islands of the Canarian Archipelago (Central East Atlantic). Our sampling strategy (involving three islands, with five locations per island, and two sites within each location) detected significant changes in the composition and structure of the fish assemblages between the three habitats. Fish assemblages associated with the unvegetated and vegetated bottoms were similar among the surveyed islands. In contrast, we observed a significant inter-island variability in the fish populations associated with the sea-cage fish farms. The presence of the sea-cage fish farms increased the overall fish abundance (184.8±49.8 ind 100 m⁻²) as compared to both the vegetated (38.8±9.7 ind 100 m⁻²) and unvegetated habitats (1.1±0.4 ind 100 m⁻²). Differences within and between the habitats were found to be associated with the relative abundance of a few fish species. The most abundant species were Xyrichthys novacula in the unvegetated bottoms and Diplodus annularis, Spondyliosoma cantharus and Mullus surmuletus on the seagrass meadows. Finally, we recorded an increase in the abundance of Heteroconger longissimus, Trachinus draco and Pagellus acarne in the bottoms beneath the sea-cage fish farms. These species, in addition to a group of large benthic chondrichthyes, were responsible for the differences between islands in the composition and structure of the demersal ichthyofauna beneath the sea-cage fish farms.     

Reintroduction of eelgrass (Zostera marina) in the Dutch Wadden Sea; review of research and suggestions for management measures

Eelgrass (Zostera marina L.) in the Dutch Wadden Sea historically covered an area varying from 65–150 km² in the eulittoral as well as the sublittoral zones. At present, this area comprises less than 1 km² eulittoral eelgrass stands, with an associated decrease in habitat diversity. The causes for this decline are presumably connected with the ‘wasting disease’ and the closure of the former Zuiderzee in the early 1930s resulting in increased tidal range and increased currents. After a slight recovery of the eelgrass populations on the intertidal flats a definite decline started in the early 1970s, possibly connected to increased turbidity. The present water quality and turbidity do not negatively influence eelgrass growth up to a depth of at least 0.6m below Mean Sea Level. Based on mesocosm experiments and field experiments it is concluded that re-establishment of eelgrass should be possible in sheltered bays and on unexposed tidal flats. The most suitable depths for a reintroduction are those between 0 and 20–40 cm below mean sea level.     

Habitat partitioning among four elasmobranch species in nearshore,shallow waters of a subtropical embayment in Western Australia

There is a need to explore, in an integrated and statistical manner, how the number of species, relative abundance, species composition and life-cycle stages of elasmobranchs in nearshore waters vary among habitat types and during the year. Therefore, four sites in a large marine embayment, each representing a different habitat type, were sampled at regular intervals. These sites were: (1) unvegetated, with no vegetation within at least 200 m; (2) unvegetated, immediately adjacent to sparse mangroves; (3) unvegetated, immediately adjacent to dense mangroves; and (4) vegetated, with seagrass (Posidonia australis) throughout and in the vicinity. Gill netting caught 10 shark species (5 families), 5 ray species (4 families) and 12 teleost species (10 families). Carcharhinus cautus, which contributed approximately 60% to the numbers of elasmobranchs caught, completed its life cycle in nearshore, shallow waters. Negaprion acutidens, Carcharhinus brevipinna, Carcharhinus limbatus and Rhizoprionodon acutus used these waters as a nursery area. C. cautus was caught mainly in the unvegetated sites, particularly in those near mangroves. N. acutidens was caught entirely in unvegetated sites, while R. acutus, C. brevipinna and Chiloscyllium punctatum were caught predominantly or exclusively in seagrass. The mean number of species and mean catch rate of elasmobranchs were greatest for the seagrass site and least for the unvegetated site with no vegetation within at least 200 m and were significantly less for the latter site than for the unvegetated site immediately adjacent to dense mangroves (P<0.05). The numbers of species and catch rates of elasmobranchs were significantly greater in summer and autumn than in winter (P<0.05) and, in the case of number of species, also than in spring (P<0.05). We conclude that the spatial and food resources in the nearshore, shallow waters of Shark Bay are partitioned among elasmobranch species, thus reducing the potential for competition among these species for the resources in those waters.Communicated by G.F. Humphrey, Sydney     

Spatial and temporal patterns of habitat use by Eurasian oystercatchers (<Emphasis Type="Italic">Haematopus ostralegus</Emphasis>) in the eastern Wadden Sea revealed using GPS data loggers

Detailed knowledge on species–habitat relationships is of crucial importance for the understanding of processes in marine ecosystems. Being top-predators, birds are important bio-indicators for marine systems. The aim of this study was to elucidate precise information on foraging habitat use and foraging times of oystercatchers (Haematopus ostralegus) on wide tidal flats using global positioning system (GPS) data loggers. The study was conducted to collect hints for the negative population trends in oystercatchers in the Wadden Sea. It is the first time that GPS technique has been used in a shorebird species. Although oystercatchers are known to exhibit foraging site fidelity, a number of individuals visited multiple sites. Foraging trips at night were longer, and the targeted sites were further away than those used during the day. These patterns were likely to be caused by higher risks of clutch predation by avian predators during the day that led adults to reduce their absence to defend their clutches. Our methodological approach enabled the subtle spatio-temporal patterns of habitat use to be determined on a very fine spatio-temporal scale. We suggest further potential studies using GPS data loggers that may help to reveal the reasons for the current declines in oystercatcher populations in the German Wadden Sea.     

Clonal extent,apical dominance and networking features in the phalanx angiosperm <Emphasis Type="Italic">Zostera noltii</Emphasis> Hornem.

Disaggregating seagrass meadows and studying its components separately (clones, ramets, shoots) can provide us insights on meadow dynamics and growth patterns. The clonal growth, dependent upon clonal rules may regulate and impose constraints to plant architecture and, therefore, determine how individual clones evolve into the environment. In order to investigate the relationship between clonal growth rules and clone architecture, the belowground network architecture of single-clones of the seagrass Zostera noltii was studied. Networks were traced in situ after washing out the overlying sediment, and network characteristics were measured using digital analysis: area covered by clone, total rhizome length, type of rhizomatic axes (main, secondary, tertiary, quaternary), number and length of the internodes, branching angles and branching frequencies. This approach revealed that Z. noltii is able to develop into large clones integrating up to 300 internodes, 676 cm of rhizome, 208 shoots and 4,300 cm² of plant area. Internodal length depended on both, the distance to the apical shoot (time effect) and the axes type (apical dominance effect). However, average branching angle was independent of axis type (average 58.3 ± 0.75), but varied significantly depending on the distance from the apical shoot. This average branching angle allows Z. noltii maximize the rate of centrifugal expansion, maintaining a high density in colonized areas to produce close stands but also minimizing the investment in belowground biomass and ramets overlapping. The clonal architecture of Z. noltii seems to be regulated by the interaction of both, apical dominance strength and clonal integration distance. Moreover, clonal growth rules and growth pattern seem to constrain clonality through (clonal) plant architecture regulations (i.e. branching is restricted in secondary axes, similar average branching angles regardless the axes, the higher the distance to the apex the higher the number of internodes in secondary axes, shorter internodes in secondary and tertiary axes). Future research efforts should focus on how these complex relationships between apical dominance and clonal integration interact to elucidate the temporal (seasonal) and spatial scales of both processes and the outcome at the plant architectural level.     

Reintroduction of eelgrass ( Zostera marina ) in the Dutch Wadden Sea; Review of research and suggestions for management measures

Eelgras (Zostera marina L.) in the Dutch Wadden Sea historically covered an area varying from 65–150 km² in the eulittoral as well as the sublittoral zones. At present, this area comprises less than 1 km² eulittoral eelgrass stands, with an associated decrease in habitat diversity. The causes for this decline are presumably connected with the ‘wasting disease’ and the closure of the former Zuiderzee in the early 1930s resulting in increased tidal range and increased currents. After a slight recovery of the eelgrass populations on the intertidal flats a definite decline started in the early 1970s, possibly connected to increased turbidity. The present water quality and turbidity do not negatively influence eelgrass growth up to a depth of at least 0.6m below Mean Sea Level. Based on mesocosm experiments and field experiments it is concluded that re-establishment of eelgrass should be possible in sheltered bays and on unexposed tidal falts. The most suitable depths for a reintroduction are those between 0 and 20–40 cm below mean sea level.     

Habitat use pattern of tidal mud and sand flats by shorebirds (charadriiformes) wintering in southern India

Habitat use pattern and status of shorebirds in southern India were investigated in six tidal flats comprising three mudflats and three sand flats on the east coast of southern India. Totally, 7757 shorebirds belonging to 21 species were recorded during September 2000–March 2002. The birds preferred mudflats over sand flats. Density, diversity and richness of shorebirds were relatively higher in tidal flats, especially in mudflats. The shorebirds preferred mudflats over sand flats as stopover sites and sites for refueling the required energy during migration. Rhythmic changes in tidal pattern also influenced the allochthonous nutrients and prey species, which also attracted the migratory shorebirds. However, the abundance of shorebirds with reference to tidal flats *season*tide and habitats*seasons * tide had strong variations (P < 0.001). During migratory season, bird density, diversity and species richness were observed higher than in other seasons.     

Carrion-feeding on the sediment surface at nocturnal low tides by the polychaete<Emphasis Type="Italic"> Phyllodoce mucosa</Emphasis>

Harsh physical conditions in the intertidal zone are the cause of an ample amount of dead macroinvertebrates, which constitute a food source for carrion-feeders. In the European Wadden Sea, this trophic guild includes decapod crustaceans and fish when the tide is in, while during nocturnal low tides the polychaete Phyllodoce mucosa is attracted in large numbers by dead mollusks, crabs or worms on the sediment surface. Within 10 s worms emerged to the surface, crawled as far as 15 m on mucus trails towards the carcass, sucked in tissue up to one-third of their own weight, and then quickly retreated to below the surface. Abundance of P. mucosa was highest in the lower intertidal zone and winter. The seaward high abundance pattern, however, did not continue into the shallow subtidal. In summer, few were attracted during daytime or when the tide was in. However, up to 447 worms aggregated at a single crushed mussel within 20 min at dusk during low-tide exposure. This study suggests that during winter carrion-feeding is an important trophic niche on cold-temperate, intertidal mud flats occupied by a phyllodocid polychaete that is segregated in feeding time from most other scavengers and benefits from cold-sensitive benthic invertebrates.Communicated by O. Kinne, Oldendorf/Luhe     

A meiofaunal “thiobios” limited to the anaerobic sulfide system of marine sand does not exist

The spatial relationship of interstitial metazoans to sources of oxygen has been studied on a tidal flat in the Wadden Sea near Sylt (eastern part of the North Sea). Consistently, meiofauna remains in close proximity to oxygenated layers or pocket areas within the sediment. This is exemplified by the pattern of meiofaunal abundance and species composition alongside oxic burrows of the lugworm Arenicola marina L. A specific meiofauna confined to oxygen-deficient horizons of the sediment does not exist.     

The effect of patch isolation on epifaunal colonization in two different seagrass ecosystems

The macrofaunal colonization of isolated habitats is affected by many factors, ranging from distance to the nearest source population to the dispersal mechanism of the species. We investigated the initial epifaunal colonization at two sites, one situated in the Northern Gulf of Mexico and the other in the Northern Baltic Sea. At each site, artificial seagrass units were placed at 10- and 20-m distances from a continuous seagrass meadow, as well as inside the meadow over a 5-day colonization time. With the exception of amphipods in Gulf of Mexico, patch isolation had a negative effect on colonization for the other faunal species, irrespective of the sites. This inverse colonization pattern of amphipods suggests that they are not equally sensitive to patch isolation in different regions. Our results indicate that increasing habitat isolation can have serious consequences for the community composition of seagrass epifauna. Furthermore, we emphasize the need for larger-scale latitudinal comparative studies.     

Biological and physical aspects of migration in the estuarine amphipod Gammarus zaddachi

The amphipod Gammarus zaddachi (Sexton) conducts extensive migrations along estuaries from near the limit of tidal influence in winter to more downstream reaches (where reproduction occurs) in spring. A return migration then takes place, primarily by juveniles, until the seaward areas are depopulated in winter. The present study was conducted between 1988 and 1990 in the Conwy Estuary, North Wales. This represents the first investigation on this species in a strongly tidal estuary, where the amphipods appear to migrate vertically into the water column on flood or ebb tides to control horizontal transport and to maintain preferred distributions. The timing of vertical migration seems to be largely controlled by an endogenous circatidal swimming rhythm. Phasing of peak activity relative to the time of expected high tide varies with season; upstream migrants in the autumn showed peak activity at the time of expected high tide, while in the spring at the time of downstream migration the rhythm was phase-delayed, with peak activity during the expected ebb tide. Together with the season, position along the estuary also affected the timing of peak endogenous activity; downstream migrants, originally active on the ebb tide and experimentally displaced seawards, showed a phase-advance of the rhythm relative to the time of high tide. Salinity-preference behaviour also varied between different developmental stages, with ovigerous females (downstream migrants) showing no preference between fresh and saline water, and juveniles (upstream migrants) showing a significant preference for freshwater. The interactions of endogenous rhythmicity and salinity-preference behaviour are discussed as controlling factors of migration in this species.     

Fish assemblages in seagrass beds are influenced by the proximity of mangrove forests

Mangrove forests and seagrass beds frequently occur as adjacent habitats in the temperate waters of southeastern Australia. At low tide when fish cannot occupy mangroves they might utilise adjacent habitats, including seagrass. We first sampled small fish from seagrass beds close to and far from mangroves in the Pittwater estuary, NSW, Australia. Seagrass beds close to mangroves had a greater density of fish species than beds far from mangroves (close: mean 16.0 species net⁻¹, SE 1.0; far: 13.2, 1.3; P < 0.05). In particular, juvenile fish were in greater densities near to than far from mangroves (close: 5.3, 0.4; far: 3.1, 0.4; P < 0.05). We then sampled the mangrove forests during the high tide and seagrass beds during the low tide, in beds along a continuum of distances from mangroves. Multivariate analysis showed that fish assemblages differed with distance from mangroves, and the differences were attributed to the composition of the fish assemblage (i.e. presence/absence of fish species), not the abundances of individual species. In particular, fish that utilise mangrove forests at high tide were found in greater species densities and species richness in seagrass nearer to mangroves. A negative relationship was found between the density of mangrove-utilising fish species and the distance of the bed from mangroves (R ² = 0.37, P < 0.05). This confirms the important connectivity between mangroves and seagrass for fish in temperate Australian waters.     

Seasonal variations in nitrogen-fixation (acetylene reduction) and sulphate-reduction rates in the rhizosphere of Zostera noltii: nitrogen fixation by sulphate-reducing bacteria

Nitrogen-fixation (acetylene reduction) rates were measured over an annual cycle in meadows of the seagrass Zostera noltii Hornem in the Bassin d'Arcachon, south-west France, between March 1994 and February 1995, using both slurry and whole-core techniques. Measured rates using the slurry technique consistently overestimated those determined on whole cores, probably due to the release of labile organic carbon sources as a result of root damage during preparation of the slurries. Thus, the whole-core technique may provide a more accurate estimate of in situ activity, since disturbance of physicochemical gradients of oxygen, sulphide, nutrients and the relationship between the plant roots and the rhizosphere microflora is minimised. Rates measured by the whole-core method were 1.8- to 4-fold greater (dependent upon season) in the light than those measured during dark incubations, indicating that organic carbon diffusing from the plant roots during photosynthesis was an important factor in regulating nitrogen fixation in the rhizosphere. Additions of sodium molybdate, a specific inhibitor of sulphate-reducing bacteria (SRB) inhibited acetylene-reduction activity by >80% as measured by both the slurry and whole-core techniques throughout the year, inferring that SRB were the dominant component of the nitrogen-fixing microflora. A mutualistic relationship between Z. noltii and nitrogen-fixing SRB in the rhizosphere, based on the exchange of organic carbon and fixed nitrogen is proposed. Acetylene- and sulphate-reduction rates showed distinct summer peaks which correlated with a reduced availability of ammonium in the sediment and the annual growth cycle of Z. noltii in the basin. Overall, these data indicate that acetylene reduction (nitrogen fixation) activity in the rhizosphere of Z. noltii was regulated both by the availability of organic carbon from the plant roots and maintenance of a low NH ₄ ⁺ concentration in the vicinity of the plant roots due to efficient assimilation of NH ₄ ⁺ by Z. noltii during the growth season. Nitrogenfixation rates determined from acetylene-reduction rates measured using the whole-core technique ranged from 0.1 to 7.3 mg N m^-2d^-1, depending on season, and were calculated to contribute between 0.4 and 1.1 g N m^-2yr^-1, or 6.3 to 12% of the annual fixed nitrogen requirement of Z. noltii.     

Food sources used by sediment meiofauna in an intertidal Zostera noltii seagrass bed: a seasonal stable isotope study

In an intertidal Zostera noltii Hornem seagrass bed, food sources used by sediment meiofauna were determined seasonally by comparing stable isotope signatures (δ¹³C, δ¹⁵N) of sources with those of nematodes and copepods. Proportions of different carbon sources used by consumers were estimated using the SIAR mixing model on δ¹³C values. Contrary to δ¹⁵N values, food source mean δ¹³C values encompassed a large range, from −22.1 ‰ (suspended particulate organic matter) to −10.0 ‰ (Z. noltii roots). δ¹³C values of copepods (from −22.3 to −12.3 ‰) showed that they use many food sources (benthic and phytoplanktonic microalgae, Z. noltii matter). Nematode δ¹³C values ranged from −14.6 to −11.4 ‰, indicating a strong role of microphytobenthos and/or Z. noltii matter as carbon sources. The difference of food source uses between copepods and nematodes is discussed in light of source accessibility and availability.     

Clam predation by whelks (Busycon spp.): Experimental tests of the importance of prey size,prey density,and seagrass cover

In 57 l-m² samples within a meadow of Halodule wrightii in Bogue Sound, North Carolina, USA, densities of the clams Mercenaria mercenaria and Chione cancellata were positively associated with seagrass cover. Where seagrass was experimentally removed, marked individuals of both clam species exhibited high rates of mortality in fine sand sediments during two successive experiments spanning 13 months. In the unaltered (control) seagrass meadow, M. mercenaria density remained constant over 13 months and C. cancellata density declined at a slower rate than in the unvegetated plots. Seagrass provides these clams with a refuge from whelk (Busycon carica, B. contrarium, and B. canaliculatum) predation, the major cause of mortality and population decline in experimentally unvegetated plots. In 2 factorial field experiments in unvegetated substratum in which densities of M. mercenaria and C. cancellata were varied independently, first over 5 levels (0 X, 1/2X, 1 X, 2 X, 4 X) and subsequently over 4 levels (0 X, 1/4 X, 1 X, 4 X), there was no repeatable intra- or interspecific effect of density on percent survival, or on the rate of any mortality type. Whelk predation fell preferentially on larger size classes of both species, whereas factors which contribute to clam disappearance usually acted more intensely on smaller sizes. Experimental exclusion of large predators by caging demonstrated that even in unvegetated substratum survivorship of both clam species was high in the absence of whelks and other predators. Individuals of C. cancellata live closer to the sediment surface than those of M. mercenaria, which may explain why seagrass does not serve as effectively to protect them from whelk predation. The mechanism of whelk inhibition may depend upon sediment binding by the H. wrightii root mat, which produces a demonstrable decrease in the physical penetrability of surface sediments.