Effects of environmental stress on intertidal mussels and their sea star predators

Consumer stress models of ecological theory predict that predators are more susceptible to stress than their prey. Intertidal mussels, Mytilus californianus, span a vertical stress gradient from the low zone (lower stress) to the high zone (higher thermal and desiccation stress), while their sea star predators, Pisaster ochraceus, range from the low zone only into the lower edge of the mussel zone. In summer 2003, we tested the responses of sea stars and mussels to environmental stress in an experiment conducted on the Oregon coast. Mussels were transplanted from the middle of the mussel bed to cages in the low and high edges of the mussel bed. Sea star predators were added to half of the mussel cages. Mussels and sea stars were sampled between June and August for indicators of sublethal stress. Mussel growth was measured, and tissues were collected for heat shock protein (Hsp70) analyses and histological analyses of reproduction. Sea stars were weighed, and tissues were sampled for Hsp70 analyses. Mussels in high-edge cages had higher levels of total Hsp70 and exhibited spawning activity earlier in the summer than mussels in the low-edge cages. Sea stars suffered high mortality in the high edge, and low-edge sea stars lost weight but showed no differences in Hsp70 production. These results suggest that stress in the intertidal zone affected the mobile predator more than its sessile prey, which is consistent with predictions of consumer stress models. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Impacts of sea star predation on mussel bed restoration

Predation by sea stars has the potential to cause elevated levels of mortality in reestablished populations of bivalves relative to levels of recruitment. Recent efforts to restore beds of the nearly extirpated green‐lipped mussel within the Hauraki Gulf, New Zealand, resulted in high abundances of sea stars occupying those beds and it is unknown whether predation poses a potential limitation to the success of restoration in this bivalve species. The contribution of predation by sea stars to the mortality of mussels across four experimental mussel beds over a 2‐year period was estimated using data from regular assessments of sea star abundance and an experimentally determined consumption rate of sea stars upon mussels. In addition, the potential size selection of mussels by sea stars was assessed to determine if large sea stars selected for recent settlers. Sea stars' abundance within the mussel beds grew to an average of 0.57 sea stars/m² within 9 months, remaining at similar levels throughout the study. These predominantly large sea stars were estimated to have consumed 30.1% of the mussels over a 25‐month period, representing a contribution of 40.4% of the mussel mortality. Sea stars predominantly selected for larger mussels, and their predation likely contributes little to any lack in mussel recruitment. Sea star predation is clearly a limiting factor on the survival of transplanted adult mussels and the present study highlights the need to continually assess predation risk to determine if remediation is necessary for the persistence of the restored beds.     

Non-trophic Interactions Control Benthic Producers on Intertidal Flats

The importance of positive effects of ecosystem engineers on associated communities is predicted to increase with environmental stress. However, incorporating such non-trophic interactions into ecological theory is not trivial because facilitation of associated species is conditional on both the type of engineer and the type of abiotic stress. We tested the influence of two allogenic ecosystem engineers (lugworms, Arenicola marina L. and cockles, Cerastoderma edule L.) on the main primary producers (microphytobenthos) of the tidal flats, under different abiotic stresses controlled by reefs of blue mussels (Mytilus edulis L.). We added 25,000 cockles or 2,000 lugworms to 5 × 5 m plots, both in a muddy site with high sedimentation rates located coastward of a mussel bed, and in a sandy site without mussels and characterized by high hydrodynamic stress. After a year, cockles increased algal biomass in the sandy area, but not in the mussel bed site, where high values were measured in all plots. However, lugworms did not affect algal biomass in any of the sites. Field measurements suggest that cockles outweighed negative effects of water currents in the site without mussels by locally increasing sediment stability, whereas mussels overruled the effects of cockles in the wake of the reefs through hydrodynamic stress alleviation and/or biodeposition. Our results suggest that non-trophic interactions by ecosystem engineering bivalves control primary production of intertidal areas, and that the sediment-stabilizing effect of cockles plays a crucial role where the overruling effects of mussel beds are not present.     

Autophagy as a second level protective process in conferring resistance to environmentally-induced oxidative stress

This conceptual paper addresses the role of lysosomal autophagy in cellular defense against environmentally-induced oxidative stress using a marine mollusc (the blue mussel) as an experimental model. It is proposed that augmented autophagic removal of oxidatively damaged organelles and proteins provides a second level or tier of defense against oxidative stress. Age pigment or lipofuscin is a product of oxidative attack on proteins and lipids and can accumulate in lysosomes, where it may generate further reactive oxygen species (ROS) and inhibit lysosomal function, resulting in autophagic failure. The previously observed protective role of augmented autophagy, induced by nutritional deprivation, against oxidative stress can be explained by this model, where autophagy boosts “cellular housekeeping” through enhanced removal of ROS-damaged proteins and organelles minimizing formation of potentially harmful stress/age pigment, and has been proposed as an anti-aging mechanism. Finally, the probable low level triggering of autophagy in mussels by fluctuating environmental regimes is considered as a potential protective mechanism that will contribute to resistance to environmentally induced oxidative stress. It is further conjectured that organisms making up functional ecological assemblages (communities) in fluctuating environments, where upregulation of autophagy should provide a selective advantage, may be pre-selected to be tolerant of pollutant-induced oxidative stress.

Addendum to: Moore MN, Viarengo A, Donkin P, Hawkins AJS. Autophagic and lysosomal reactions to stress in the hepatopancreas of blue mussels. Aquat Toxicol 2007; 84:80–91.     

Fouling-resistant surfaces of tropical sea stars

Qualitative evidence suggests sea stars are free of fouling organisms; however the presence of fouling-resistant surfaces of sea stars has not previously been documented. Field surveys were conducted in northern Queensland, Australia, during the wet and dry seasons and several tropical sea star species were examined for surface-associated micro- and macro-organisms. Mean bacterial abundances on seven sea star species were approximately 10(4) to 10(5) cells cm(-2) during both seasons. There were no consistent trends in bacterial abundances with season, species and aboral positions on sea star arms. No common generalist fouling organisms, such as algae, barnacles, serpulid polychaetes, bryozoans and ascidians, were found on any specimens of 12 sea star species. However, low numbers of parasitic and commensal macro-organisms were found on six sea star species. The gastropods Parvioris fulvescens, Asterolamia hians, Thyca (Granulithyca) nardoafrianti and Thyca crystallina were found exclusively on the sea stars Archaster typicus, Astropecten indicus, Nardoa pauciforis and Linckia laevigata, respectively. The shrimp Periclimenes soror was only found on Acanthaster planci, and the polychaete Ophiodromus sp. on A. typicus. The copepods Stellicola illgi and Paramolgus sp. were only found on L. laevigata and Echinaster luzonicus, respectively. As no common generalist fouling organisms were discovered, sea stars offer an excellent model to investigate the mechanisms driving fouling-resistant surfaces and the selective settlement of specialist invertebrates.     

The effects of climate change on species composition,succession and phenology: a case study

Climate change and its role in altering biological interactions and the likelihood of invasion by introduced species in marine systems have received increased attention in recent years. It is difficult to forecast how climate change will influence community function or the probability of invasion as it alters multiple marine environmental parameters including rising water temperature, lower salinity and pH. In the present study, we correlate changes in environmental parameters to shifts in species composition in a subtidal community in Newcastle, NH through comparison of two, 3‐year periods separated by 23 years (1979–1981 and 2003–2005). We observed concurrent shifts in climate related factors and in groups of organisms that dominate the marine community when comparing 1979–1981 to 2003–2005. The 1979–1981 community was dominated by perennial species (mussels and barnacles). In contrast, the 2003–2005 community was dominated by annual native and invasive tunicates (sea‐squirts). We also observed a shift in the environmental factors that characterized both communities. Dissolved inorganic nitrogen and phosphate characterized the 1979–1981 community while sea surface temperature, pH, and chlorophyll a characterized the 2003–2005 community. Elongated warmer water temperatures, through the fall and early winter months of the 2000s, extended the growing season of native organisms and facilitated local dominance of invasive species. Additionally, beta‐diversity was greater between 2003–2005 than 1979–1981 and driven by larger numbers of annual species whose life‐history characteristics (e.g., timing and magnitude of recruitment, growth and mortality) are driven by environmental parameters, particularly temperature.     

Prey selection and the functional response of sea stars (Asterias vulgaris Verrill) and rock crabs (Cancer irroratus Say) preying on juvenile sea scallops (Placopecten magellanicus (Gmelin)) and blue mussels (Mytilus edulis Linnaeus)

Predators in nature include an array of prey types in their diet, and often select certain types over others. We examined (i) prey selection by sea stars (Asterias vulgaris) and rock crabs (Cancer irroratus) when offered two prey types, juvenile sea scallops (Placopecten magellanicus) and blue mussels (Mytilus edulis), and (ii) the effect of prey density on predation, prey selection, and component behaviours. We quantified predation rates, behavioural components (proportion of time spent searching for prey, encounter probabilities) and various prey characteristics (shell strength, energy content per prey, handling time per prey) to identify mechanisms underlying predation patterns and to assess the contribution of active and passive prey selection to observed selection of prey. Sea stars strongly selected mussels over scallops, resulting from both active and passive selection. Active selection was associated with the probability of attack upon encounter; it was higher on mussels than on scallops. The probability of capture upon attack, associated with passive selection, was higher for mussels than for scallops, since mussels can not swim to escape predators. Sea stars consumed few scallops when mussels were present, and so did not have a functional response on scallops (the target prey). Rock crabs exhibited prey switching: they selected mussels when scallop density was very low, did not select a certain prey type when scallop density was intermediate, and selected scallops when scallop density was high relative to mussel density. The interplay between encounter rate (associated with passive selection) and probability of consumption upon capture (associated with both active and passive selection) explained observed selection by crabs. Scallops were encountered by crabs relatively more often and/or mussels less often than expected from random movements of animals at all scallop densities. However, the probability of consumption varied with scallop density: it was lower for scallops than mussels at low and intermediate scallop densities, but tended to be higher for scallops than mussels at high scallop densities. When mussels were absent, crabs did not have a functional response on scallops, but rather were at the plateau of the response. When mussels were present with scallops at relatively low density, crabs exhibited a type II functional response on scallops. Our results have implications for the provision of protective refuges for species of interest (i.e., scallops) released onto the sea bed, such as in population enhancement operations and bottom aquaculture.     

Experimental determination of predation intensity in an intertidal predator guild: dominant versus subordinate prey

Theoretical and empirical ecologists have long acknowledged that information about the intensity or strength of the interaction between species is crucial for an understanding of community dynamics. In communities in which predation is an important structuring process, and some predator species are commercially exploited, quantitative estimates of predation by different predator species within a guild are necessary to make even the simplest recommendations about conservation and resource management. Here, we evaluated per capita and population level components of predation intensity of three intertidal predators that feed on monospecific stands of barnacles and mussels at wave exposed sites in the rocky intertidal zone of central Chile. These prey species represent the two most distinctive stages of the mid-intertidal seascape, with mussels being competitively dominant. Our results showed that the commercially exploited gastropod Concholepas concholepas and the sea star Heliaster helianthus have similarly large per capita and population effects on the competitively dominant mussel Perumytilus purpuratus . Their per capita (by average size individual) and population effects on mussels were more than two orders of magnitude larger than those of Acanthocyclus gayi crabs and likely even larger than the effect of other predator species in this system (other crabs, whelks, birds, fish). The overall pattern of predation on barnacles was similar to that on mussels, but some differences occurred in the way different components of predation intensity were distributed across predator species. Despite the roughly similar pattern of population predation intensity between prey species, the expected consequences for the prey population, and hence the rest of the community, were acutely different for mussels and barnacles.     

Predation on invasive zebra mussel, Dreissena polymorpha, by pumpkinseed sunfish, rusty crayfish, and round goby

The enemy release hypothesis states that invasive species are successful in their new environment because native species are not adapted to utilize the invasive. If true for predators, native predators should have lower feeding rates on the invasive species than a predator from the native range of the invasive species. We tested this hypothesis for zebra mussel (Dreissena polymorpha) by comparing handling time and predation rate on zebra mussels in the laboratory by two North American species (pumpkinseed, Lepomis gibbosus, and rusty crayfish, Orconectes rusticus) and one predator with a long evolutionary history with zebra mussels (round goby, Neogobius melanostomus). Handling time per mussel (7 mm shell length) ranged from 25 to >70 s for the three predator species. Feeding rates on attached zebra mussels were higher for round goby than the two native predators. Medium and large gobies consumed 50–67 zebra mussels attached to stones in 24 h, whereas pumpkinseed and rusty crayfish consumed <11. This supports the hypothesis that the rapid spread of zebra mussels in North America was facilitated by low predation rates from the existing native predators. At these predation rates and realistic goby abundance estimates, round goby could affect zebra mussel abundance in some lakes.     

Whole-community facilitation regulates biodiversity on Patagonian rocky shores

Background

Understanding the factors that generate and maintain biodiversity is a central goal in ecology. While positive species interactions (i.e., facilitation) have historically been underemphasized in ecological research, they are increasingly recognized as playing important roles in the evolution and maintenance of biodiversity. Dominant habitat-forming species (foundation species) buffer environmental conditions and can therefore facilitate myriad associated species. Theory predicts that facilitation will be the dominant community-structuring force under harsh environmental conditions, where organisms depend on shelter for survival and predation is diminished. Wind-swept, arid Patagonian rocky shores are one of the most desiccating intertidal rocky shores ever studied, providing an opportunity to test this theory and elucidate the context-dependency of facilitation.

Methodology/Principal Findings

Surveys across 2100 km of southern Argentinean coastline and experimental manipulations both supported theoretical predictions, with 43 out of 46 species in the animal assemblage obligated to living within the matrices of mussels for protection from potentially lethal desiccation stress and predators having no detectable impact on diversity.

Conclusions/Significance

These results provide the first experimental support of long-standing theoretical predictions and reveal that in extreme climates, maintenance of whole-community diversity can be maintained by positive interactions that ameliorate physical stress. These findings have important conservation implications and emphasize that preserving foundation species should be a priority in remediating the biodiversity consequences of global climate change.     

Oxidative stress in Perna perna and other bivalves as indicators of environmental stress in the Brazilian marine environment: Antioxidants, lipid peroxidation and DNA damage

Oxidative stress can take place in marine bivalves under a series of environmental adverse conditions. The study of different systems related to oxidative stress in these organisms can give important information about their physiological status and also about environmental health. Bivalves have been proposed as good sentinel organisms in pollution monitoring studies through the analysis of biochemical biomarkers, and most of the biomarkers analyzed are those related to oxidative stress. However, it is very important to know how other environmental factors not associated to the presence of pollutants might affect these parameters. We have studied a series of mechanisms related to oxidative stress in mussels which inhabit the Brazilian coast, especially in Perna perna species, subjected to different stress conditions, such as the exposure to different contaminants in the laboratory and in the field, the exposure of mussels to air and re-submersion, simulating the tidal oscillations, and in mussels collected at different seasons. Both oxidative damage levels and antioxidant defense systems were strongly affected by the different environmental stress. This review summarizes the data obtained in some studies carried out in bivalves from the Brazilian coast.     

Quantification of echinoderms (Echinodermata) on Georges Bank,and the potential influence of marine protected areas on these populations

The spatiotemporal variation of the distribution of echinoderms in the Georges Bank ecosystem was examined from 2005 to 2012. Density and abundance of representatives from echinoderm classes (brittle stars, sand dollars, sea stars, and sea urchins) were estimated using a drop camera video survey of the benthos in areas open and closed to fish trawling. The influence of closed areas on these echinoderm populations relative to a suite of key environmental and biotic factors was evaluated using Canonical Correspondence Analysis (CCA). While marine protected areas appeared to influence the distribution of sand dollars and sea stars, the distribution of brittle stars and sea urchins seemed to be defined mainly by preferred habitat conditions. According to the CCA, depth, sediment stability, temperature, predator abundance, and management area were the most important factors explaining this echinoderm assemblage. On Georges Bank, echinoderms dominate the benthic biota and are present in a variety of habitats. They can alter marine communities and are preferred prey and main predators for several commercially targeted species. The detailed information presented here (on the scale of kilometers) on abundance and spatial distribution of these populations is thus valuable towards the implementation of ecosystem‐based fisheries management.     

Oxidative stress in Perna perna and other bivalves as indicators of environmental stress in the Brazilian marine environment: antioxidants, lipid peroxidation and DNA damage

Oxidative stress can take place in marine bivalves under a series of environmental adverse conditions. The study of different systems related to oxidative stress in these organisms can give important information about their physiological status and also about environmental health. Bivalves have been proposed as good sentinel organisms in pollution monitoring studies through the analysis of biochemical biomarkers, and most of the biomarkers analyzed are those related to oxidative stress. However, it is very important to know how other environmental factors not associated to the presence of pollutants might affect these parameters. We have studied a series of mechanisms related to oxidative stress in mussels which inhabit the Brazilian coast, especially in Perna perna species, subjected to different stress conditions, such as the exposure to different contaminants in the laboratory and in the field, the exposure of mussels to air and re-submersion, simulating the tidal oscillations, and in mussels collected at different seasons. Both oxidative damage levels and antioxidant defense systems were strongly affected by the different environmental stress. This review summarizes the data obtained in some studies carried out in bivalves from the Brazilian coast.     

Salinity as a constraint affecting food and habitat choice of mussel-feeding diving ducks

Observations of freshwater drinking in Eiders feeding primarily on mussels led us to hypothesize that the highly saline sea water enclosed in mussels could cause salt-related dehydration problems in the ducks, since they consume entire mussels. The proportion of sea water increases with increasing mussel size. Smaller duck species are more sensitive to the higher salt content of larger mussels than are larger ducks; however, salt stress may be avoided by feeding in habitats with lower salinity, by feeding on less salty food items, by utilizing smaller mussels, by drinking fresh water, or by employing low energy foraging techniques. A possible evolutionary strategy for solving the salt problem might be to increase body mass, enabling ducks to utilize larger mussels without passing an upper salt consumption limit. At the same time, foraging on larger mussels is more economical. Although large size facilitates the utilization of brackish and marine environments, it may be selected against in ducks breeding in fresh water, where fish competition may reduce optimal food item size. In conclusion, salinity is an important habitat barrier in both breeding and overwintering diving ducks, but there are various ways of crossing this barrier. To understand better how ducks utilize their habitats, however, it is necessary to measure habitat salinity levels and the size of both ducks and their preferred and less-preferred food types.     

Interactive effects of food availability and aerial body temperature on the survival of two intertidal Mytilus species

Intertidal organisms must episodically contend with the rigors of both the terrestrial and the marine environments. While body temperatures during high tide are driven primarily by water temperature, aerial body temperatures are driven by multiple environmental factors such that temperature of an organism during low tide is usually quite different from air temperature. Thus, whereas decades of research have investigated the effects of water temperature on intertidal species, considerably less is known about the physiological impacts of temperature during aerial exposure at low tide, especially with regard to the interaction of aerial body temperature with other stressors. We examined the interactive effects of aerial body temperature and food supply on the survival of two intertidal blue mussels, Mytilus galloprovincialis and Mytilus trossulus. Survival was monitored for nine weeks using a simulated tidal cycle, with two levels of food and three levels of aerial body temperature (30, 25, and 20 °C). Decreased food supply significantly reduced the survival of mussels, but only under the 30 °C treatment. In the other two thermal regimes there were no significant effect of food on survival. When aerial body temperatures are high, food availability may have a greater effect on intertidal organisms. Decreases in ocean productivity have been linked to increased in ocean temperatures, thus intertidal organisms may become more susceptible to thermal stress as climates shift.     

Can climate change jeopardize predator control of invasive herbivore species? A case study in avocado agro-ecosystems in Spain

Climate change is one of the most important factors affecting the phenology, distribution, composition and diversity of organisms. In agricultural systems many pests and natural enemies are arthropods. As poikilotherm organisms, their body temperature is highly dependent on environmental conditions. Because higher trophic levels typically have lower tolerance to high temperatures than lower trophic levels, trends towards increasing local or regional temperatures may affect the strength of predator/prey interactions and disrupt pest control. Furthermore, increasing temperatures may create climate corridors that could facilitate the invasion and establishment of invasive species originating from warmer areas. In this study we examined the effect of environmental conditions on the dynamics of an agro-ecosystem community located in southern Spain, using field data on predator/prey dynamics and climate gathered during four consecutive years. The study system was composed of an ever-green tree species (avocado), an exotic tetranychid mite, and two native species of phytoseiid mites found in association with this new pest. We also present a climatological analysis of the temperature trend in the area of study during the last 28 years, as evidence of temperature warming occurring in the area. We found that the range of temperatures with positive per capita growth rates was much wider in prey than in predators, and that relative humidity contributed to explain the growth rate variation in predators, but not in prey. Predator and prey differences in thermal performance curves could explain why natural enemies did not respond numerically to the pest when environmental conditions were harsh.     

Seasonal variations in mercury,cadmium, lead and arsenic species in Norwegian blue mussels (Mytilus edulis L.) – Assessing the influence of biological and environmental factors

BackgroundBlue mussels (Mytilus edulis L.) can accumulate undesirable substances, including the potentially toxic elements (PTEs) cadmium (Cd), mercury, (Hg), lead (Pb), arsenic (As) and As species. In this study, the levels of PTEs and As species were determined in samples of blue mussels to assess the influence of environmental and biological factors, and evaluate the potential risk associated with blue mussels in terms of food and feed safety.MethodologyBlue mussels were collected monthly from one location in Western Norway from February 2018 to December 2018, and from April 2019 to April 2020. Samples were analyzed for PTEs using inductively coupled plasma mass spectrometry (ICP-MS), and high-performance liquid chromatography (HPLC) coupled to ICP-MS. Temperature, salinity and fluorescence (chlorophyll a) were monitored in the seawater column by STD/CTD, to assess the potential influence of these environmental factors on the PTE levels in the mussels.ResultsThe results showed seasonal variations in the PTEs, with somewhat higher concentrations in spring and winter months. Unusually high levels of total As (101.2 mg kg^-1 dw) and inorganic As (53.6 mg kg^-1 dw) were observed for some of the time points. The organic As species arsenobetaine was generally the major As species (17–82% of total As) in the mussels, but also simple methylated As species and arsenosugars were detected. Principal components analysis (PCA) did not show a consistent relationship between the environmental factors and the PTE concentrations, showing contrary results for some elements for the periods studied. The condition index (CI) could explain variations in element concentration with significant correlations for Cd (r = −0.67, p = 0.009) and Pb (r = −0.62, p = 0.02 in 2019/20 and r = −0.52, p = 0.02 in 2018), whereas the correlation between As and CI was not significant (r = 0.12 in 2018, and r = −0.06 in 2019/20). Higher concentrations of iAs and arsenosugars coincided with increased signals of chlorophyll a, suggesting that phytoplankton blooms could be a source of As in the blue mussels.ConclusionTo our knowledge, this is the first study of As species in blue mussels collected over a time period of two years, providing an insight into the natural variations of these chemical forms in mussels. In terms of mussel as food and future feed material, concentrations of Cd, Hg and Pb were below the maximum levels (MLs) established in the EU food and feed legislation. However, levels of As and iAs in mussels at some time points exceeded the MLs for As in the feed legislation, and the margin of exposure (MOE) was low if these mussels were for human consumption, highlighting the importance of determining the chemical forms of As in feed and food.     

Autophagy as a second level protective process in conferring resistance to environmentally-induced oxidative stress

This conceptual paper addresses the role of lysosomal autophagy in cellular defense against environmentally-induced oxidative stress using a marine mollusc (the blue mussel) as an experimental model. It is proposed that augmented autophagic removal of oxidatively damaged organelles and proteins provides a second level or tier of defense against oxidative stress. Age pigment or lipofuscin is a product of oxidative attack on proteins and lipids and can accumulate in lysosomes, where it may generate further reactive oxygen species (ROS) and inhibit lysosomal function, resulting in autophagic failure. The previously observed protective role of augmented autophagy, induced by nutritional deprivation, against oxidative stress can be explained by this model, where autophagy boosts "cellular housekeeping" through enhanced removal of ROS-damaged proteins and organelles minimizing formation of potentially harmful stress/age pigment, and has been proposed as an anti-aging mechanism. Finally, the probable low level triggering of autophagy in mussels by fluctuating environmental regimes is considered as a potential protective mechanism that will contribute to resistance to environmentally induced oxidative stress. It is further conjectured that organisms making up functional ecological assemblages (communities) in fluctuating environments, where upregulation of autophagy should provide a selective advantage, may be pre-selected to be tolerant of pollutant-induced oxidative stress.     

Predatory blue crabs induce byssal thread production in hooked mussels

Abstract. Blue crabs (Callinectes sapidus) prey on hooked mussels (Ischadium recurvum) growing epizoically on oyster clumps in estuaries along the Louisiana coast. In prey size‐selection experiments, blue crabs preferred small mussels (<30‐mm shell length) to larger mussels, possibly because handling time increased with mussel size. When crabs were given a choice of solitary mussels versus mussels in clumps on oysters in the laboratory, mortality was lower by 86% in clumped mussels. However, no size selection by crabs occurred with mussels in clumps, likely because smaller mussels escaped predation in crevices between larger mussels or oysters. When individuals of two size classes of mussels were exposed to water containing the scent of crabs and of mussels consumed by blue crabs, an increase in byssal thread production was induced in all mussels, but byssal thread production rate was higher for small mussels than for large mussels. We conclude that increased predation risk for small mussels has resulted in higher size‐specific production of byssal threads, and that predator‐induced production of byssal threads, which may increase clumping behavior, may reduce their risk of mortality to predatory blue crabs.     

Feeding repellence of Antarctic and sub-Antarctic benthic invertebrates against the omnivorous sea star Odontaster validus

Antarctic and sub-Antarctic benthic invertebrates are subjected to intense predation by mobile macroinvertebrates. Accordingly, chemical protection as well as other defensive mechanisms are expected to be common in organisms inhabiting these ecosystems. In order to evaluate anti-predation activities and allocation of chemical defenses within the anatomy of marine benthic Antarctic and sub-Antarctic invertebrates, 55 species were tested for feeding repellence against the sea star Odontaster validus, a common eurybathic sympatric predator. The invertebrates tested were collected from the deep waters of two poorly surveyed areas in terms of chemical ecology studies: the eastern Weddell Sea (Antarctica) and the vicinities of Bouvet Island (sub-Antarctica). Experiments were conducted at the Spanish Antarctic Base in Deception Island. In the feeding deterrence experiments, shrimp pieces were treated with crude lipophilic fractions obtained from each species, and were offered to the sea stars. A total of 29 species (53 %) from 7 different phyla (Porifera, Cnidaria, Chordata, Bryozoa, Echinodermata, Mollusca, and Annelida) showed feeding repellence against O. validus, and are therefore chemically protected against this keystone predator. Furthermore, 25 species were dissected into parts to investigate the possible allocation of defensive compounds. Some of the results obtained from these analyses support the prediction that the most exposed/vulnerable tissues concentrate chemical defenses to avoid predation against the sea stars. In summary, the results obtained in our survey support the hypothesis that deep-water Antarctic and sub-Antarctic benthic invertebrates are well protected chemically against sympatric predators, similarly to what has been reported in previous studies investigating shallow-water Antarctic species.