Mud crab predation on ribbed mussels in salt marshes

Mud crab (Panopeus herbstii H. Milne Edwards) predation on Atlantic ribbed mussels (Geukensia demissa Dillwyn) was studied by a series of laboratory and field experiments at two sites at Morehead City, North Carolina, USA from 1987 to 1989. Tidal elevation had no effect on predation intensity, although mud crabs were active only when they were submerged. Horizontal distance from the water-marsh edge significantly affected mussel mortality in one of two winter experiments, despite the occurrence of virtually all the crabs at the marsh edge. Of the juvenile mussels attached to adult mussels, those totally buried in the sediment suffered mortality from mud crab predation at a rate not detectably different from those exposed above the surface. Juveniles attached to adult conspecifics, however, experienced significantly less mortality than those attached to oysters. Interestingly, the two groups of mussels (those attached to conspecifics and those attached to oysters) display shell morphological dimorphism. The more oblate shells of the mussels attached to oysters as compared to those attached to conspecifics might be induced by the higher predation rate. Alternatively, slimmer shells from individuals attached to conspecifics may be the result of living within physically compact mussel clumps.Please address all correspondence and requests for reprints to Dr Lin at his present address: Smithsonian Environmental Research Center, P.O. Box 28, Edgewater, Maryland 21037, USA     

How to toughen up your mussels: using mussel shell morphological plasticity to reduce predation losses

Improvements in stocking strategy and management could increase the yield of mussels that are on-grown from harvested wild seed mussel resources and thereby enhance the sustainability of this shellfishery. A field experiment was undertaken to ascertain shell characteristics (compression strength and thickness) of seed mussels grown at different shore heights, whether these characteristics changed after a period of growth under identical conditions, and if these characteristics reduced predation losses by crabs and birds. Results indicated that mussels grown at higher shore levels attained shell characteristics beneficial to predation resistance and that these were maintained after a period of growth at a lower shore level. A novel management plan for mussel cultivation was formulated from the results of this study by manipulating shore position according to the attainment of these predator resistant shell attributes and the spatial distribution of the main natural mussel predators (crabs and birds). This technique was expanded to address the mussel cultivation problem of low natural seed settlement.     

Mussels flexing their muscles: a new method for quantifying bivalve behaviour

We employed a novel technique to quantify how blue mussels Mytilus edulis react to predation risk in their environment by quantifying mussel gape using a Hall sensor attached to one shell valve reacting to a magnet attached to the other. Change in gape angle per second (CHIGA) versus gape angle plots resulted in a distribution with a boundary, which defined the maximum CHIGA of a mussel at all gape angles. CHIGA boundary plots for all individual mussels were similar in form. However, the CHIGA boundary increased in extent with mussel length (maximum CHIGA for mussel valve closures for mussels 2.98 and 79.6 mm long were −1.5 and −11°s⁻¹, respectively), showing that larger mussels opened and closed most rapidly. Mussel extract added to the seawater, a factor believed to signal predation, caused mussels to close significantly faster than otherwise (P < 0.001). This approach for assessing how mussels react to their environment indicates that mussel response to predation is graded and complex and may well indicate animal-based assessments of the trade-off between effective feeding and the likelihood of predation.     

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.     

Rates of fission, somatic growth and gonadal development of a fissiparous sea star, Allostichaster insignis, in New Zealand

Seasonal changes in population structure and incidence of fission were measured in intertidal and subtidal populations of Allostichaster insignis, a fissiparous sea star. Population size structure was stable over the course of the 1-year study. Sea stars in the subtidal zone attained greater maximum size (mean arm length, R = 35 mm) than those in the intertidal population (20 mm). Fission rates were greatest among small individuals (R < 20 mm). The frequency of fission ranged from 5 to 32% with peaks in early austral summer in the intertidal zone, and in autumn and winter in the subtidal zone. Sexual reproduction occurred in early spring in sea stars larger than 12 mm. The populations were heavily biased toward males. In the laboratory, A. insignis of three size classes (small, R = 9–13 mm; medium, 19–21 mm; and large, 29–31 mm) were fed mussels ad libitum or starved (not fed macroscopic food) for ∼1 year in a 3 × 2 factorial experiment. Small and medium-sized sea stars divided throughout the experiment and the ramets of most individuals regenerated sufficiently to divide again after 6–9 months. Unfed sea stars did not undergo fission (with one exception), had a higher mortality rate, and did not grow. Small, fed sea stars grew significantly faster than medium-sized or large individuals. At the end of the experiment, the pyloric caeca index (a measure of nutritional condition) was greater in fed than in unfed animals. Gonads (only testes were observed) developed in medium-sized and large, fed sea stars. Our field and laboratory results indicate that asexual reproduction in A. insignis predominantly occurs in small, well-nourished individuals. Ramets grow gradually through repeated fission and regeneration to a size (mean length of regenerating arms, R _r ∼ 20 mm) at which they begin to switch to sexual reproduction as the dominant reproductive mode.     

Prevalence, patterns, and effects of shell damage on Geukensia demissa in South Carolina estuarine habitats

Shelled molluscs frequently exhibit a record of damage on exterior surfaces that can evidence past predation attempts and may affect survival and growth. In South Carolina populations of the ribbed marsh mussel, Geukensia demissa, >90% of the individuals and up to 60% of the total shell area are damaged. A trend toward greater amounts of damage occurred on mid-marsh compared to oyster reef mussels from the barrier beach side of inlets. Shell damage effects on survivorship and shell and tissue growth were assessed seasonally during multi- and single-season field experiments. Mussels from a common mid-marsh site were divided into size classes (~50 or 70 mm), treated to create two damage levels (undamaged and damaged), and replaced within mid-marsh exclusion cages to minimize additional shell damage. In both multi- and single-season experiments increased shell damage resulted in significantly greater mortality. Linear shell growth was unaffected by increased damage, but 50 mm mussels grew twice as fast. Shell mass increased 16–50% in the multi-season and single-season winter period, but decreased 7–12% during the single-season summer period. Tissue mass significantly decreased 31–43% in 50 mm damaged mussels, but increased by 33% for 70 mm mussels in both multi-season and the single-season winter period experiments. Shell damage did reduce tissue mass 43% in 70 mm single-season summer mussels. Experimental results indicate shell damage from a simulated increase in predation can affect negatively both survival and growth of marsh mussels. Seasonal timing of shell damage and initial mussel size also influenced the effects of sublethal predation on shell and tissue growth. The previously unrecognized importance of sublethal predation and the resultant significant negative effects of shell damage on survival and growth will affect the distribution and population dynamics of G. demissa in coastal marshes and will influence the overall contribution of ribbed mussels to estuarine ecosystems.     

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.     

Population dynamics of the endangered fan mussel <Emphasis Type="Italic">Pinna nobilis</Emphasis> in a marine lake: a metapopulation matrix modeling approach

A metapopulation, time-invariant, stage-classified matrix model was developed to assess the dynamics of an important Pinna nobilis population in the marine Lake Vouliagmeni (Korinthiakos Gulf, Greece). The main aim of the study was to provide insight on the life cycle of the fan mussel and reveal influential factors for its population dynamics, with a special focus on the effect of poaching. The size of the fan mussel shell was selected as a state variable, and the model consisted of five size classes. The lake was divided in two regions, a shallow region of high (illegal) fishing mortality and high recruitment (region 1) and a deeper region of low mortality and low recruitment (region 2). The estimation of the transition matrix (stage-specific growth and mortality probabilities) was based on a tagging survey between 2005 and 2006, while independent annual surveys for abundance estimation using distance sampling techniques were utilized for the estimation of recruitment and stage-specific fertilities. The population was found to be increasing with an intrinsic rate of increase r = 0.038; however, r was not statistically different from zero. The life expectancy and expected lifetime offspring production of individuals in region 1 was markedly lower than that of individuals in region 2. Due to poaching, the life expectancy of a yearling fan mussel was less than 2.5 years in region 1, while it was almost 12 years in region 2. The highest expected annual natural mortality of fan mussels occurred on their first year of life after settlement (~43%) and greatly declined at greater sizes. Perturbation analysis revealed that the population growth rate was most sensitive to the vital rates of the larger size classes in region 2 and to fertilities corresponding to offspring that settled in the same region. The spatial distribution and abundance of the species was greatly dependent on the extent of poaching, which caused a size segregation of individuals, with small and young individuals being abundant in region 1, and larger and older individuals being restricted in region 2. If poaching ceased, the fan mussel population would be increasing with a significantly higher intrinsic rate of increase (r = 0.186), while if region 2 was also illegally exploited at the same intensity as region 1, the fan mussel population would be decreasing with r = −0.364 and would eventually collapse. The existence of refuge areas, where fan mussels may grow and reproduce, providing adjacent areas with offspring, seems crucial for the viability of local populations. Transplantation of fan mussels from high mortality areas to low mortality refuges might prove to be an effective measure to protect local populations of the species.     

Dead zones enhance key fisheries species by providing predation refuge

Natural stress gradients can reduce predation intensity and increase prey abundances. Whether the harsh conditions of anthropogenic habitat degradation can similarly reduce predation intensity and structure community dynamics remains largely unexplored. Oxygen depletion in coastal waters (hypoxia) is a form of degradation that has recently emerged as one of the greatest threats to coastal ecosystems worldwide due to increased rates of eutrophication and climate change. I conducted field experiments and surveys to test whether relaxed predation could explain the paradoxically high abundance of clams that have sustained a fishery in a degraded estuary with chronic hypoxic conditions. Hypoxia reduced predation on all experimental species but enhanced the long-term survivorship of only sufficiently hypoxia-tolerant prey due to periodic extreme conditions. As a consequence, only the harvested quahog clam (Mercenaria mercenaria) thrived in hypoxic areas that were otherwise rendered dead zones with depauperate diversity and low abundances of other species. This suggests that enhanced populations of some key species may be part of a predictable nonlinear community response that sustains ecosystem services and masks overall downward trends of habitat degradation.     

Ingestion of bivalve larvae by<Emphasis Type="Italic"> Mytilus edulis</Emphasis>: experimental and field demonstrations of larviphagy in farmed blue mussels

Ingestion of bivalve larvae by Mytilus edulis was investigated. Laboratory experiments revealed that ~ 90% of bivalve larvae offered to mussels was ingested and apparently fully digested. The shell of the bivalve larvae offered no protection against digestive processes, resulting in high larval mortality once inside the stomach. Stomach content analysis (September 2001–January 2003) showed that bivalve larvae were ingested by farmed mussels year-round, with the exception of March 2002. Numbers of ingested larvae were highest in October 2001 and May 2002, which coincides with known spawning times of farmed mussels in Ireland. Mussels ingested a large size-range of bivalve larvae, suggesting that all stages of the bivalve life cycle are vulnerable to predation. It is suggested that adult bivalves routinely filter larvae from the surrounding water and that, given the high biomass of mussels present in mussel farms, filtration by adult bivalves significantly reduces numbers of bivalve larvae in nearby waters.Communicated by J.P. Thorpe, Port Erin     

Roach (<Emphasis Type="Italic">Rutilus rutilus</Emphasis>) as an important predator on blue mussel (<Emphasis Type="Italic">Mytilus edulis</Emphasis>) populations in a brackish water environment,the northern Baltic Sea

Suspension-feeding bivalves are organisms of major functional importance in several aquatic environments around the world. They are also important food items for many fish and benthivorous seabirds. It has commonly been thought that predation pressure on blue mussel (Mytilus edulis) populations is negligible in the Baltic Sea, owing to the scarcity of major invertebrate predators such as starfish and crabs. It has recently been shown, however, that the blue mussel is the main food item for roach (Rutilus rutilus) in the archipelago areas of the western Gulf of Finland, where this freshwater fish species has become increasingly abundant, mainly due to increased eutrophication. To quantify the influence of roach predation on blue mussel populations we measured the standing biomass and size structure of the local blue mussel population and used a bioenergetic model to estimate mussel consumption by individual roach during two consecutive summers, 1997 and 1998. The results of the model were combined with existing data on roach abundance, giving annual consumption estimates of 75–105 kg blue mussel dry weight ha^–1 in the study area, approximately two-thirds of these consumed mussels being >10 mm. This corresponds to approximately one-third of the standing population of mussels >10 mm in the area. Our results suggest that the predation effects of vertebrates on Baltic blue mussel populations are not insignificant, as previously believed. Predation by roach and other predators may have an important structuring effect on unstable blue mussel communities within the Gulf of Finland, where the species lives at the edge of its range.Communicated by M. Kühl, Helsingør     

Population dynamics of Diadema antillarum (Echinodermata: Echinoidea) following mass mortality in Panamá

In 1983, Diadema antillarum suffered mass mortality throughout the Caribbean Sea and the western Atlantic Ocean. I followed the dynamics of populations at the San Blas Islands, Panamá from April 1983 to November 1987. Density measurements indicate that populations of D. antillarum have not recovered from the die-offs that killed nearly 97% of the individuals. There was recruitment to the 1 to 1.5 cm class immediately after the mass mortality, but there has been little additional influx of juveniles since then. The low number of observable juveniles could not be attributed to elevated rates of predation on very small individuals. Rates of recruitment did not differ between reefs with artificially increased densities of D. antillarum and reefs kept free of sea urchins; thus, the lack of recruitment did not arise from absence of adults that could provide settlement cues to the larvae or protection to newly settled juveniles. Other species of sea urchins did not show a clear pattern of increase after the demise of D. antillarum. Therefore, interspecific competition directed towards D. antillarum juveniles did not increase after the mass mortality. Two reefs where Echinometra viridis, Eucidaris tribuloides and Lytechinus williamsi, were removed showed no significant differences in recruitment of D. antillarum relative to two reefs where these species were allowed to remain at their natural densities. Resident D. antillarum after the mass mortality produced gametes with the same per capita intensity and lunar synchrony as before the mass mortality. However, it is possible that the probability of fertilization of their gametes decreased because of low population density. The most likely explanation for lack of recruitment is that the reduced numbers of reproducing adults at Panamá and upstream locations resulted in levels of larval supply that were inadequate to sustain recruitment on Panamanian reefs.     

Predator-prey relationships between the rock lobster Jasus lalandii and the mussel Aulacomya ater at Robben Island on the Cape West Coast of Africa

The natural diet and mode of feeding of the rock lobster Jasus lalandii (H. Milne Edwards) was determined in a rock-lobster sanctuary near Cape Town, South Africa. Field observations were tested and confirmed by means of aquarium studies. Rock lobsters feed mainly upon ribbed mussels Aulacomya ater (Molina), which comprise the largest component of the sessile benthic fauna. Mussel remains were found as the major constituent in 97% of the rock-lobster stomachs examined. The density of rock lobsters averaged 8,100 per hectare (0.81 m^-2), while mussel biomass averaged more than 5 kg (wet whole weight) m^-2 within the same depth range (12 to 30 m). More than 80% of the mussel biomass comprised large individuals between 60 and 90 mm in length. Large rock lobsters (mainly males) were capable of feeding on all sizes of mussels, although many of these were inaccessible to predation. Smaller rock lobsters became progressively more limited in the size range of mussels which they could crack open and consume. Competition between rock lobsters for small mussels appeared to be intense, as mussels of suitable size for feeding were generally in short supply to most of the rock-lobster population. Hence, feeding and growth rates of rock lobsters are likely to be affected by the relative population densities of predator and prey. Growth rates of rock lobsters could be limited by food supplies even in areas where mussel biomass is comparatively large.     

Impoundments and the Decline of Freshwater Mussels: a Case Study of an Extinction Gradient

Abstract: One major factor leading to the imperilment of freshwater mussels ( Bivalvia, Unionidae) has been the large-scale impoundment of rivers. We examined the distribution and abundance of mussels at 37 sites along a 240-km length of the Little River in southeastern Oklahoma, U.S.A., which is affected by both mainstem and tributary reservoirs. We observed a mussel extinction gradient downstream from impoundments in this river: with increasing distance from the mainstem reservoir there was a gradual, linear increase in mussel species richness and abundance. Mussel species distributions were significantly nested, with only sites furthest from the impoundment containing relatively rare species. Below the confluence with the inflow from the second reservoir these same trends were apparent but much weaker, and overall mussel abundance was greatly reduced. Our results suggest that considerable stream lengths are necessary to overcome the effects of impoundment on mussel populations, and such information should be considered in conservation and management plans.     

Between-individual variation in haemocyanin concentrations in the Norway lobster<Emphasis Type="Italic"> Nephrops norvegicus</Emphasis> following exposure to hypoxia and manganese

Environmental hypoxia and the release of reduced manganese (Mn) are often combined stress factors in marine sediments. Previous investigations have shown that hypoxia induces variable responses in the concentration of the respiratory pigment haemocyanin (Hc) of crustaceans. A recent study demonstrated that the magnitude and direction of the changes was dependent on the initial level of Hc in individuals of the Norway lobster Nephrops norvegicus. The changes also took place within 24 h rather than days as suggested by previous studies. In this present study we investigated changes of Hc in individual N. norvegicus in uncontaminated sea water after exposure to and release from realistic hypoxic stress (PO ₂=6.0 kPa). Furthermore we investigated how concomitant exposure to realistic concentrations of manganese (20 mg l ^-1) modified the responses we observed. We confirmed a between-individual variation in response and a change in Hc towards an optimum level after hypoxia, but also showed that changes that took place post-hypoxic exposure occurred over a timescale similar to that of the hypoxic response itself. Manganese exposure resulted in no significant changes in Hc whereas Mn exposure combined with hypoxia resulted in a significant decrease (15%) in all individuals independent of initial Hc. N. norvegicus was thus unable to compensate for the effects of hypoxia by synthesis of Hc after exposure to naturally occurring Mn concentrations.     

Differences in relative abundance and size structure of the sea stars Pisaster ochraceus and Evasterias troschelii among habitat types in Puget Sound, Washington, USA

We surveyed patterns in the relative abundance and size structure of the sea stars Pisaster ochraceus and Evasterias troschelii in five habitat types of varying structural complexity and prey availability (sand/cobble, boulder, and rocky intertidal; pilings; and floating docks) in Puget Sound and the San Juan Islands, Washington. For both species, small sea stars were most abundant in the most structurally complex habitat type (boulder), where they occurred almost exclusively under boulders during low tide. Larger individuals became more abundant as structural complexity decreased, occurring more frequently in open habitat types (rocky shores, pilings, and docks) known to have greater abundances of prey resources. Gull foraging observations and experiments demonstrated that exposed small sea stars of both species were highly vulnerable to predation, suggesting that small sea stars require structural complexity (crevice microhabitat) as a predation refuge. Large sea stars, once attaining a size refuge from predation, appear to migrate to more exposed habitat types with more abundant food resources. These results suggest parallel ontogenetic habitat shifts in two co-occurring consumer species related to a shared predation risk at early life stages and demonstrate how the relative importance of top-down and bottom-up processes may differ with ontogeny.     

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.     

A box model of carrying capacity for suspended mussel aquaculture in Lagune de la Grande-Entrée,Iles-de-la-Madeleine,Québec

An object-oriented model of environment–mussel aquaculture interactions and mussel carrying-capacity within Lagune de la Grande-Entrée (GEL), Iles-de-la-Madeleine, Québec, was constructed to assist in development of sustainable mussel culture in this region. A multiple box ecosystem model for GEL tied to the output of a hydrodynamic model was constructed using Simile software, which has inherent ability to represent spatial elements and specify water exchange between modelled regions. Mussel growth and other field data were used for model validation. Plackett–Burman sensitivity analysis demonstrated that a variety of bioenergetic parameters of zooplankton and phytoplankton submodels were important in model outcomes. Model results demonstrated that mussel aquaculture can be further developed throughout the lagoon. At present culture densities, phytoplankton depletion is minimal, and there is little food limitation of mussel growth. Results indicated that increased stocking density of mussels in the existing farm will lead to decreased mass per individual mussel. Depending on the location of new farm emplacement within the lagoon, implementation of new aquaculture sites either reduced mussel growth in the existing farm due to depletion of phytoplankton, or exhibited minimum negative impact on the existing farm. With development throughout GEL, an excess of phytoplankton was observed during the year in all modelled regions, even at stocking densities as high as 20 mussels m⁻³. Although mussels cultured at this density do not substantially impact the ecosystem, their growth is controlled by the flux of phytoplankton food and abundance of zooplankton competitors. This model provides an effective tool to examine expansion of shellfish farming to new areas, balancing culture location and density.     

Depression of feeding and growth rates of the seastar Evasterias troschelii during long-term exposure to the water-soluble fraction of crude oil

To test the effect of petroleum hydrocarbons on predation by the seastar Evasterias troschelii (Stimpson, 1862) on the mussel Mytilus edulis (L.), we exposed the predator with the prey to six concentrations of the water-soluble fraction (WSF) of Cook Inlet crude oil. Seastars and mussels were collected at Auke Bay, Alaska, in November 1980. During a 28 d exposure in a flow-through system, seastars were more sensitive to the WSF than mussels: the LC₅₀ for the seastars was 0.82 ppm at Day 19 and, although no mussels were exposed to WSF for more than 12 d, none died. Daily feeding rates (whether in terms of number of mussels seastar^-1 d^-1 or dry weight of mussels seastar^-1 d^-1) were significantly reduced at all concentrations above 0.12 ppm. At 0.20, 0.28 and 0.72 ppm WSF, daily feeding rates (in terms of dry weight of mussels) were, respectively, 53, 37, and 5% of the control rate; at the two highest concentrations (0.97 and 1.31 ppm WSF), the seastars did not feed. Seastars at concentrations greater than 0.12 ppm WSF grew slower than individuals from the control group and the 0.12 ppm-treatment group combined. These laboratory results show that E. troschelii is more sensitive to chronic low levels of the WSF of crude oil. The possibility that such oil pollution could reduce predation and permit M. edulis to monopolize the low intertidal zone of southern Alaska remains to be studied.     

Indirect effects of a key ecosystem engineer alter survival and growth of foundation coral species

Stegastes nigricans, a "farmerfish" that cultivates algal turf and defends territories from grazers and other intruders, can affect coral indirectly due to increased competition with farmed algal turf and/or reduced predation resulting from territorial aggression directed at corallivores. To investigate the indirect effects of this key ecosystem engineer on coral mortality and growth, we transplanted caged and exposed fragments of four coral species to patch reefs in French Polynesia on which we manipulated the presence of S. nigricans and turf, and to reefs naturally devoid of S. nigricans. Reef access was two to four times higher for herbivorous fishes, and two times higher for corallivorous fishes, when S. nigricans was removed, indicating that reef access is reduced for two important guilds of fishes when S. nigricans is present. Stegastes' territoriality indirectly benefited delicate acroporids (Montipora floweri and Acropora striata), yielding a twofold to fivefold reduction in skeletal loss due to lower predation frequencies in the presence of S. nigricans. Three corals, A. striata, M. floweri, and especially Porites australiensis, suffered mortality due to overgrowth significantly more frequently in the presence of farmed turf, but Pocillopora verrucosa did not. Algal abundance predicted the frequency of overgrowth for only A. striata and P. australiensis. M. floweri were more likely to be overgrown when exposed (uncaged) in the presence of S. nigricans, suggesting an interaction modification, in this case that initial predation increased susceptibility to competition with turf. In this community, the presence of S. nigricans may increase algal overgrowth of massive Porites by facilitating its turf competitors and simultaneously reduce predation of branching corals through territorial exclusion of corallivores. These indirect interactions may underlie previously documented community transitions from disturbance-resistant massive coral to recovering branching corals within S. nigricans territories.