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Plastic pollution has become a significant environmental concern, with profound consequences for ecosystems worldwide, particularly for marine systems. Our study introduces 'plastiskin', a newly identified plastic pollution type encrusting intertidal organisms. Found on mussels and macroalgae, 'plastiskin' was composed of polypropylene and polyethylene. In mussels, the presence of 'plastiskin' was correlated with the absence of living endoliths in areas of the shells it covered, indicating a detrimental impact on the symbiotic endolithic community residing within mussel shells. In addition, we examined the potential negative effects of 'plastiskin' on the photosynthetic efficiency of macroalgae, however, these findings were inconclusive, stressing the need for further studies with larger sample sizes. Our baseline observations may serve as a groundwork for further investigation into the spatial distribution, temporal persistence, and ecological ramifications of 'plastiskin'. potential incorporation of 'plastiskin' as a new marine debris category into management and monitoring frameworks warrants serious consideration., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024. Published by Elsevier Ltd.)

Recent advances in genetic manipulation such as triploid breeding and artificial selection, have rapidly emerged as valuable hatchery methodologies for enhancing seafood stocks. The Pacific oyster Magallana gigas is a leading aquaculture species worldwide and key ecosystem engineer that has received particular attention in this field of science. In light of the growing recognition of the ecological effects of intraspecific variation, oyster polyploids provide a valuable opportunity to assess whether intraspecific diversity affects physiological responses to environmental stressors. While the responses of diploid and triploid oysters to climate change have been extensively investigated, research on their sensitivity to environmental pollution remains scarce. Here, we assess whether genotypic (i.e., ploidy) variation within Magallana gigas affects physiological responses to microplastic pollution. We show that diploid and triploid M. gigas have similar clearance rates and ingest similar amounts of microplastics under laboratory-controlled condition. In addition, they exhibited similar heart rates after prolonged exposure to microplastic leachates. Our findings suggest that intraspecific variations within M. gigas ploidy does not affect oyster responses to microplastic pollution. However, regardless of ploidy, our work highlights significant adverse effects of microplastic leachates on the heart rate of M. gigas and provides evidence of microplastic ingestion in the laboratory., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Heatwaves are increasingly severe and frequent, posing significant threats to ecosystems and human well-being. Characterised by high thermal variability, intertidal communities are particularly vulnerable to heat stress. Microbial endolithic communities that are found in marine calcifying organisms have been shown to induce shell erosion that alters shell surface colour, lowering body temperatures and increasing survival rates. Here, we investigate how the symbiotic relationship between endolithic microbes and the blue intertidal mussel Mytilus edulis mitigates thermal stress during the unprecedented 2022 atmospheric heatwave in the English Channel. Microbial infestation of the shell significantly enhanced mussel survival, particularly higher on the shore where thermal stress was greater. Using data from biomimetic temperature loggers, we predicted the expected thermal buffer and observed differences up to 3.2 °C between individuals with and without symbionts under the known conditions of the heat wave-induced mortality event. The ecological implications extend beyond individual mussels, affecting the reef-building capacity of mussels, with potential cascading effects for local biodiversity, carbon sequestration, and coastal defence. These findings emphasize the importance of understanding small-scale biotic interactions during extreme climate events and provide insights into the dynamic nature of the endolith-mussel symbiosis along a parasitic-mutualistic continuum influenced by abiotic factors., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Self-organized spatial patterns are increasingly recognized for their contribution to ecosystem functioning. They can improve the ecosystem's ability to respond to perturbation and thus increase its resilience to environmental stress. Plastic pollution has now emerged as major threat to aquatic and terrestrial biota. Under laboratory conditions, we tested whether plastic leachates from pellets collected in the intertidal can impair small-scale, spatial self-organization and byssal threads production of intertidal mussels and whether the effect varied depending on where the pellets come from. Specifically, leachates originating from plastic pellets collected from relatively pristine and polluted areas respectively impaired and inhibited the ability of mussels to self-organize at small-scale and to produce byssal threads compared to control conditions (i.e., seawater without leaching solution). Limitations to natural self-organizing processes and threads formation may translate to a declined capacity of natural ecosystems to avoid tipping points and to a reduced restoration success of disturbed ecosystems., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Plastic pollution and ongoing climatic changes exert considerable pressure on coastal ecosystems. Unravelling the combined effects of these two threats is essential to management and conservation actions to reduce the overall environmental risks. We assessed the capacity of a coastal ecosystem engineer, the blue mussel Mytilus edulis , to cope with various levels of aerial heat stress (20, 25, 30 and 35°C) after an exposure to substances leached from beached and virgin low-density polyethylene pellets. Our results revealed a significant interaction between temperature and plastic leachates on mussel survival rates. Specifically, microplastic leachates had no effect on mussel survival at 20, 25 and 30°C. In turn, mussel survival rates significantly decreased at 35°C, and this decrease was even more significant following an exposure to leachates from beached pellets; these pellets had a higher concentration of additives compared to the virgin ones, potentially causing a bioenergetic imbalance. Our results stress the importance of adopting integrated approaches combining the effects of multiple environmental threats on key marine species to understand and mitigate their potential synergistic effects on ecosystem dynamics and resilience in the face of the changing environment.

Worldwide, microplastic pollution has numerous negative implications for marine biota, exacerbating the effects of other forms of global anthropogenic disturbance. Mounting evidence shows that microplastics (MPs) not only cause physical damage through their ingestion, but also act as vectors for hazardous compounds by leaching absorbed and adsorbed chemicals. Research on the effects of plastic pollution has, however, largely assumed that species respond uniformly, while ignoring intraspecific diversity (i.e., variation within a single species). We investigated the effects of plastic leachates derived from factory-fresh (virgin) and beached microplastics on the behavioural responses of two genetic lineages of the Mediterranean mussel Mytilus galloprovincialis. Through laboratory behavioural experiments, we found that during exposure to leachates from beached microplastics (beached MPLs), Atlantic specimens moved significantly less than Mediterranean individuals in terms of both (i) proportion of individuals responding through movement and (ii) net and gross distances crawled. In contrast, no significant intraspecific differences were observed in the behaviour of either adults or recruits when exposed to MPLs from virgin microplastics (virgin MPLs). Additionally, the reception of cues from three amino acids (L-cysteine, proline and L-leucine) at increasing concentrations (10 -5  M to 10 -3  M in charcoal-filtered seawater) was tested by electrophysiological analysis using mussels exposed to beached MPLs or control seawater. We found significant intraspecific differences in response to 10 -3  M L-cysteine (regardless of treatment) and 10 -4  M L-cysteine (in mussels exposed to beached MPLs) and to 10 -3  M proline (in mussels exposed to beached MPLs) and 10 -5  M L-leucine. Our study suggests that intraspecific variation in a marine mussel may prompt different responses to plastic pollution, potentially triggered by local adaptation and physiological variability between lineages. Our work highlights the importance of assessing the effects of intraspecific variation, especially in environmental sentinel species as this level of diversity could modulate responses to plastic pollution., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Interspecific and intraspecific diversity are essential components of biodiversity with far-reaching implications for ecosystem function and service provision. Importantly, genotypic and phenotypic variation within a species can affect responses to anthropogenic pressures more than interspecific diversity. We investigated the effects of interspecific and intraspecific diversity on microplastic ingestion by two coexisting mussel species in South Africa, Mytilus galloprovincialis and Perna perna, the latter occurring as two genetic lineages. We found significantly higher microplastic abundance in M. galloprovincialis (0.54 ± 0.56 MP items g -1 WW) than P. perna (0.16 ± 0.21 MP items g -1 WW), but no difference between P. perna lineages. Microbeads were the predominant microplastic (76 % in P. perna, 99 % in M. galloprovincialis) and polyethylene the prevalent polymer. Interspecific differences in microplastic abundance varied across locations, suggesting diverse sources of contamination. We suggest that microplastic ingestion can be species-specific even in organisms that coexist and play similar functional roles within ecosystems., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Both individual and collective anti-predator behaviours are essential for the survival of many species. This is particularly true for ecosystem engineers such as intertidal mussels, which through their collective behaviour create novel habitats for a range of organisms and biodiversity hotspots. However, contaminants may disrupt these behaviours and consequently indirectly affect exposure to predation risk at the population level. Among these, plastic litter is a major and ubiquitous contaminant of the marine environment. Here, we assessed the impact of microplastic (MP) leachates of the most produced plastic polymer, polypropylene (PlasticsEurope, 2022), at a high but locally relevant concentration (i.e. ca. 12 g L -1 ) on the collective behaviours and anti-predator responses of both small and large Mytilus edulis mussels. Indeed, in contrast to large mussels, small ones reacted to MP leachates, showing a taxis towards conspecifics and stronger aggregations. All mussels reacted to the chemical cues of the predatory crab, Hemigrapsus sanguineus, but with two different collective anti-predator behaviours. Small mussels only showed a taxis towards conspecifics when exposed to predator cues. This response was also found in large ones with a tendency to form more strongly bound aggregations and a considerable reduced activity, i.e. they significantly delayed their time to start to form aggregations and decreased their gross distance. These anti-predator behaviours were respectively inhibited and impaired in small and large mussels by MP leachates. The observed collective behavioural changes may reduce individual fitness by enhancing predation risk, particularly in small mussels that are the crab H. sanguineus's favourite preys. Given the key role of mussels as ecosystem engineers, our observations suggest that plastic pollution may have implication on M. edulis at the species level, but also enhancing a cascading effect towards a higher level of organisation such as population, community and ultimately structure and function of intertidal ecosystem., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

The impact of climate change on biodiversity has stimulated the need to understand environmental stress responses, particularly for ecosystem engineers whose responses to climate affect large numbers of associated organisms. Distinct species differ substantially in their resilience to thermal stress but there are also within-species variations in thermal tolerance for which the molecular mechanisms underpinning such variation remain largely unclear. Intertidal mussels are well-known for their role as ecosystem engineers. First, we exposed two genetic lineages of the intertidal mussel Perna perna to heat stress treatments in air and water. Next, we ran a high throughput RNA sequencing experiment to identify differences in gene expression between the thermally resilient eastern lineage and the thermally sensitive western lineage. We highlight different thermal tolerances that concord with their distributional ranges. Critically, we also identified lineage-specific patterns of gene expression under heat stress and revealed intraspecific differences in the underlying transcriptional pathways in response to warmer temperatures that are potentially linked to the within-species differences in thermal tolerance. Beyond the species, we show how unravelling within-species variability in mechanistic responses to heat stress promotes a better understanding of global evolutionary trajectories of the species as a whole in response to changing climate., (© 2023. The Author(s).)

Microbial symbionts have strong potential to mediate responses to climate change. Such modulation may be particularly important in the case of hosts that modify the physical habitat structure. By transforming the habitats, ecosystem engineers alter resource availability and modulate environmental conditions which, in turn, indirectly shape the community associated with that habitat. Endolithic cyanobacteria are known to reduce the body temperatures of infested mussels and here, we assessed whether the thermal benefits of endoliths on the intertidal reef-building mussel Mytilus galloprovincialis extends to the invertebrate community utilising mussel beds as habitat. Artificial reefs of biomimetic mussels either colonised or not colonised by microbial endoliths were used to test whether infauna species (the limpet Patella vulgata, the snail Littorina littorea and mussel recruits) in a mussel bed with symbionts experience lower body temperatures than those within a bed composed of mussels without symbionts. We found that infaunal individuals benefitted from being surrounded by mussels with symbionts, an effect that may be particularly critical during intense heat stress. Indirect effects of biotic interactions, complicate our understanding of community and ecosystem responses to climate change, especially in cases involving ecosystem engineers, and accounting for them will improve our predictions., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Our understanding of how anthropogenic stressors such as climate change and plastic pollution interact with biodiversity is being widened to include diversity below the species level, i.e., intraspecific variation. The emerging appreciation of the key ecological importance of intraspecific diversity and its potential loss in the Anthropocene, further highlights the need to assess the relative importance of intraspecific versus interspecific diversity. One such issue is whether a species responds as a homogenous whole to plastic pollution. Using manipulative field transplant experiments and laboratory-controlled hydrodynamic simulations, we assessed the relative effects of intraspecific and interspecific diversity on microplastic trapping in coastal biogenic habitats dominated by two key bioengineers, the brown intertidal macroalgae Fucus vesiculosus and F. guiryi. At the individual level, northern morphotypes of F. guiryi trapped more microplastics than southern individuals, and F. vesiculosus trapped more microplastics than F. guiryi. Canopy density varied among species, however, leading to reversed patterns of microplastic accumulation, with F. guiryi canopies accumulating more microplastics than those of F. vesiculosus, while no differences were observed between the canopies of F. guiryi morphotypes. We emphasize the importance of assessing the effects of intraspecific variation which, along with other crucial factors such as canopy density, flow velocity and polymer composition, modulates the extent of microplastic accumulation in coastal biogenic habitats. Our findings indicate that a realistic estimation of plastic accumulation in biogenic habitats requires an understanding of within- and between-species traits at both the individual and population levels., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Plastic is one of the most ubiquitous sources of both contamination and pollution of the Anthropocene, and accumulates virtually everywhere on the planet. As such, plastic threatens the environment, the economy and human well-being globally. The related potential threats have been identified as a major global conservation issue and a key research priority. As a consequence, plastic pollution has become one of the most prolific fields of research in research areas including chemistry, physics, oceanography, biology, ecology, ecotoxicology, molecular biology, sociology, economy, conservation, management, and even politics. In this context, one may legitimately expect plastic pollution research to be highly interdisciplinary. However, using the emerging topic of microplastic and nanoplastic leachate (i.e., the desorption of molecules that are adsorbed onto the surface of a polymer and/or absorbed into the polymer matrix in the absence of plastic ingestion) in the ocean as a case study, we argue that this is still far from being the case. Instead, we highlight that plastic pollution research rather seems to remain structured in mostly isolated monodisciplinary studies. A plethora of analytical methods are now available to qualify and quantify plastic monomers, polymers and the related additives. We nevertheless show though a survey of the literature that most studies addressing the effects of leachates on marine organisms essentially still lack of a quantitative assessment of the chemical nature and content of both plastic items and their leachates. In the context of the ever-increasing research effort devoted to assess the biological and ecological effects of plastic waste, we subsequently argue that the lack of a true interdisciplinary approach is likely to hamper the development of this research field. We finally introduce a roadmap for future research which has to evolve through the development of a sound and systematic ability to chemically define what we biologically compare., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022. Published by Elsevier B.V.)

The massive contamination of the environment by plastics is an increasing global scientific and societal concern. Knowing whether and how these pollutants affect the behaviour of keystone species is essential to identify environmental risks effectively. Here, we focus on the effect of plastic leachates on the behavioural response of the common blue mussel Mytilus edulis, an ecosystem engineer responsible for the creation of biogenic structures that modify the environment and provide numerous ecosystem functions and services. Specifically, we assess the effect of virgin polypropylene beads on mussels' chemotactic (i.e. a directional movement in response to a chemical stimulus) and chemokinetic (i.e. a non-directional change in movement properties such as speed, distance travelled or turning frequency in response to a chemical stimulus) responses to different chemical cues (i.e. conspecifics, injured conspecifics and a predator, the crab Hemigrapsus sanguineus). In the presence of predator cues, individual mussels reduced both their gross distance and speed, changes interpreted here as an avoidance behaviour. When exposed to polypropylene leachates, mussels moved less compared to control conditions, regardless of the cues tested. Additionally, in presence of crab cues with plastic leachates, mussels significantly changed the direction of movement suggesting a leachate-induced loss of their negative chemotaxis response. Taken together, our results indicate that the behavioural response of M. edulis is cue-specific and that its anti-predator behaviour as well as its mobility are impaired when exposed to microplastic leachates, potentially affecting the functioning of the ecosystem that the species supports., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Background: The order and orientation of genes encoded by animal mitogenomes are typically conserved, although there is increasing evidence of multiple rearrangements among mollusks. The mitogenome from a Brazilian brown mussel (hereafter named B1) classified as Perna perna Linnaeus, 1758 and assembled from Illumina short-length reads revealed an unusual gene order very different from other congeneric species. Previous mitogenomic analyses based on the Brazilian specimen and other Mytilidae suggested the polyphyly of the genus Perna ., Methods: To confirm the proposed gene rearrangements, we sequenced a second Brazilian P. perna specimen using the "primer-walking" method and performed the assembly using as reference Perna canaliculus . This time-consuming sequencing method is highly effective when assessing gene order because it relies on sequentially-determined, overlapping fragments. We also sequenced the mitogenomes of eastern and southwestern South African P. perna lineages to analyze the existence of putative intraspecific gene order changes as the two lineages show overlapping distributions but do not exhibit a sister relationship., Results: The three P. perna mitogenomes sequenced in this study exhibit the same gene order as the reference. CREx, a software that heuristically determines rearrangement scenarios, identified numerous gene order changes between B1 and our P. perna mitogenomes, rejecting the previously proposed gene order for the species. Our results validate the monophyly of the genus Perna and indicate a misidentification of B1., Competing Interests: Rita Castilho is an Academic Editor for PeerJ., (© 2022 Cunha et al.)

Plastic contamination has major effects on biodiversity, enhancing the consequences of other forms of global anthropogenic disturbance such as climate change and habitat fragmentation. Despite this and the recognised importance of intraspecific diversity, we still know relatively little about how plastic pollution affects diversity below the species level. Here, we assessed the effects of intraspecific variation in a habitat forming species (the Mediterranean mussel Mytilus galloprovincialis) on the trapping and ingestion of microplastics. We focused on symbiont-induced phenotypic variation in mussel beds. Using fractal analysis, we measured an increase in the complexity of mussel bed surfaces by ca. 15% caused by phototropic shell-degrading endoliths. By simulating high tide flow conditions and incoming waves, we found that symbionts significantly increased microplastic accumulation in mussel beds. This likely reflects deceleration of near-bed flow velocities, creation of turbulence in the bottom boundary layer and consequently increased particle retention. This effect was not constant at high tide, with no effect of infestation on retention at the base of the mussel bed under mid and high flow conditions and reduced microplastic trapping on the surface of mussel shells. Nevertheless, under natural conditions, the ingestion and trapping of microplastic were higher by the mussels comprising beds with symbionts than those in beds without symbionts. Given the dependency of many species on mussel biogenic habitats, there is an increased risk of plastics moving up the food chain in mussel beds infested by symbiotic endoliths. Our results highlight how the effects of within-species phenotypic diversity may influence the consequences of rising levels of plastic pollution., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Recruitment of mussels is a complex process with the successful arrival of individuals hinging on the availability of suitable habitats. We examined the effects of adult mussels as settlement habitat and the degree to which the suitability of habitat they offer is species-specific by comparing the recruitment success of intertidal mussels. We hypothesised that mussel recruitment and early growth are dictated by the quality of habitat offered by conspecifics adults. We used a unique experimental arena on the south coast of South Africa, where Mytilus galloprovincialis and two lineages of Perna perna co-exist. Treatments were based on the translocation of individuals of M. galloprovincialis , western- and eastern lineage of P . perna to a single site, where artificial beds were created and sampled monthly over one year. Recruit's number, their sizes and growth were greater within beds of the western lineage of Perna than eastern lineage or Mytilus beds. The results clearly demonstrate that the quality of settlement habitat offered by adult beds differs among adult lineages/species and affects rates of settlement and the early growth of recruits. This effect extends to the intraspecific level; we found greater differences in density and growth of recruits between lineages of Perna than between either lineage and M. galloprovincialis ., Supplementary Information: The online version contains supplementary material available at 10.1007/s10750-022-04994-7., Competing Interests: Conflict of interestThe authors declare that there are no competing interests., (© The Author(s), under exclusive licence to Springer Nature Switzerland AG 2022, Springer Nature or its licensor holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.)

The carbon footprint of flying overseas to conferences, meetings, and workshops to share and build knowledge has been increasingly questioned over the last two decades, especially in environmental and climate sciences, due to the related colossal carbon emissions. Here, we infer the value of scientific meetings through the number of publications produced either directly or indirectly after attending a scientific conference, symposium, or workshop (i.e., the conference-related production) and the number of publications produced per meeting (i.e., the conference-related productivity) as proxies for the academic value of these meetings, and relate them to both the number of meetings attended and the related carbon emissions. We show that conference-related production and productivity, respectively, increase and decay with the number of meetings attended, and noticeably that the less productive people exhibit the largest carbon footprint. Taken together, our results imply that a twofold decrease in the carbon footprint F CO 2 of a given scientist would result in a twofold increase in productivity through a fivefold decrease in the number of meeting attended. In light of these figures, we call for both the implementation of objective and quantitative criteria related to the optimum number of conferences to attend in an effort to maximize scientific productivity while minimizing the related carbon footprint, and the development of a rationale to minimize the carbon emission related to scientific activities., Competing Interests: The authors have no conflicts of interest., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Temperature extremes are predicted to intensify with climate change. These extremes are rapidly emerging as a powerful driver of species distributional changes with the capacity to disrupt the functioning and provision of services of entire ecosystems, particularly when they challenge ecosystem engineers. The subsequent search for a robust framework to forecast the consequences of these changes mostly ignores within-species variation in thermal sensitivity. Such variation can be intrinsic, but can also reflect species interactions. Intertidal mussels are important ecosystem engineers that host symbiotic endoliths in their shells. These endoliths unexpectedly act as conditionally beneficial parasites that enhance the host's resistance to intense heat stress. To understand how this relationship may be altered under environmental change, we examined the conditions under which it becomes advantageous by reducing body temperature. We deployed biomimetic sensors (robomussels), built using shells of mussels (Mytilus galloprovincialis) that were or were not infested by endoliths, at nine European locations spanning a temperature gradient across 22°of latitude (Orkney, Scotland to the Algarve, Portugal). Daily wind speed and solar radiation explained the maximum variation in the difference in temperature between infested and non-infested robomussels; the largest difference occurred under low wind speed and high solar radiation. From the robomussel data, we inferred body temperature differences between infested and non-infested mussels during known heatwaves that induced mass mortality of the mussel Mytilus edulis along the coast of the English Channel in summer 2018 to quantify the thermal advantage of endolith infestation during temperature extremes. Under these conditions, endoliths provided thermal buffering of between 1.7°C and 4.8°C. Our results strongly suggest that sustainability of intertidal mussel beds will increasingly depend on the thermal buffering provided by endoliths. More generally, this work shows that biomimetic models indicate that within-species thermal sensitivity to global warming can be modulated by species interactions, using an intertidal host-symbiont relationship as an example., (© 2021 John Wiley & Sons Ltd.)

AbstractSelf-organized, regular spatial patterns emerging from local interactions among individuals enhance the ability of ecosystems to respond to environmental disturbances. Mussels self-organize to form large, regularly patterned biogenic structures that modify the biotic and abiotic environment and provide numerous ecosystem functions and services. We used two mussel species that form monospecific and mixed beds to investigate how species-specific behavior affects self-organization and resistance to wave stress. Perna perna has strong attachment but low motility, while Mytilus galloprovincialis shows the reverse. At low density, the less motile P. perna has limited spatial self-organization compared with M. galloprovincialis , while when coexisting, the two species formed random spatial patterns. At high density, the two species self-organized in similar ways, while when coexisting, patterns were less strong. Spatial pattern formations significantly shaped resistance to hydrodynamic stress. At low density, P. perna beds with strong attachment and M. galloprovincialis beds with strong spatial organization showed higher retention rates than mixed beds. At high density, the presence of strongly attached P. perna significantly increased retention in mixed and P. perna beds compared with M. galloprovincialis beds. Our study emphasizes the importance of the interplay of species-specific behaviors to spatial self-organization and stress tolerance in natural communities.

Plastic pollution is ubiquitous with increasing recognition of its direct effects on species' fitness. Little is known, however, about its more subtle effects, including the influence of plastic pollution on the morphological, functional and behavioral traits of organisms that are central to their ability to withstand disturbances. Among the least obvious but most pernicious forms of plastic-associated pollution are the chemicals that leach from microplastics. Here, we investigate how such leachates influence species' traits by assessing functional trait compensation across four species of intertidal mussels, through investigations of byssal thread production, movement and aggregation behavior for mussels held in natural seawater or seawater contaminated by microplastic leachates. We found no evidence for compensation of functional traits, but for each species, microplastic leachates reinforced one trait while others remained unaffected. Two species (Perna perna and Mytilus galloprovincialis), were characterized by a resistance strategy to disturbance; they produced more byssal threads in microplastic leachate seawater than in control seawater, while motility and aggregation remained essentially unaffected. In contrast, the other two species (M. edulis and Choromytilus meridionalis), showed a resilience strategy to disturbance through increased motility and aggregation in leachate seawater, while byssal thread production remained unaffected. These results suggest that the competitive abilities of intertidal mussels may be related to their sensitivity to microplastic leachates or other chemical disturbance. Importantly, the trait strengthening observed will affect the ability of these mussels to form spatially patterned beds, with implications for their quality as autogenic ecological engineers or foundation species. Thus, our findings have implications for the ability of mussel beds to tolerate disturbance, and hence for central ecosystem services, such as their ability to support biodiversity and enhance secondary and tertiary production. The results suggest that an inconspicuous aspect of plastic pollution has the potential to influence other communities and ecosystems in powerful ways., (© 2020 by the Ecological Society of America.)

Can intraspecific diversity functionally supersede interspecific diversity? Recent studies have established the ecological effects of intraspecific variation on a number of ecosystem dynamics including resilience and productivity and we hypothesised that they may functionally exceed those of species diversity. We focused on a coastal ecosystem dominated by two coexisting bioengineering mussel species, one of which, Perna perna, displays two distinct phylogeographic lineages. A manipulative field experiment revealed greater habitat structural complexity and a more benign microscale environment within beds of the eastern lineage than those of the western lineage or the second species (Mytilus galloprovincialis); the latter two did not differ. Similarly, while infaunal species abundance and biomass differed significantly between the two lineages of Perna, there was no such difference between Mytilus and the western Perna lineage. The evenness and diversity of associated infaunal assemblages responded differently. Diversity differed relatively weakly between species, while evenness showed a very strong difference between conspecific lineages. Our results show that variation within a species can functionally supersede diversity between species. As the two P. perna lineages have different physiological tolerances, we expect them to react differently to environmental change. Our findings indicate that predicting the ecosystem-level consequences of climate change requires an understanding of the relative strengths of within- and between-species differences in functionality., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Anthropogenic litter is ubiquitous throughout marine ecosystems, but its abundance and distribution are driven by complex interactions of distinct environmental factors and thus can be extremely heterogeneous. Here we compare the extent of anthropogenic litter pollution at a sheltered lagoon habitat and nearby open coast sites. Monthly surveys over a period of five months showed that both the types and sources of litter always differed significantly between lagoon and open coast sites. Pollution within the lagoon was mainly land-derived and was largely made up of construction materials (70% to 95%). At open coast sites, construction materials represented a minor portion of pollution (4% to 12%) while plastics were the most abundant (82% to 95%). We show that stranded anthropogenic litter in adjacent marine habitats can differ significantly and stress the importance of sampling at appropriate spatial scales to gain realistic insights into the sources of pollution., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Understanding the spread of invasive species in many regions is difficult because surveys are rare. Here, historical records of the invasive marine mussel, Semimytilus algosus, on the shores of Angola and Namibia are synthesised to re-construct its invasive history. Since this mussel was first discovered in Namibia about 90 years ago, it has spread throughout the western coast of southern Africa. By the late 1960s, the species was well established across a range of 1005 km of coastline in southern Angola and northern Namibia. Although only coarse spatial resolution data are available since the 1990s, the distribution of S. algosus clearly increased substantially over the subsequent decades. Today, the species is distributed over 2785 km of coastline, appearing in southern Namibia in 2014, whence it spread across the border to northern South Africa in 2017, and in northern Angola in 2015. Conspicuously, its current range appears to be relatively contiguous across at least 810 km of shore in southern Angola and throughout Namibia, with isolated, spatially disjunct occurrences towards the southern and northern limits of its distribution. Despite there being few occurrence records that are unevenly distributed spatially and temporally, data for the distributional patterns of S. algosus in Angola and Namibia provide invaluable insights into how marine invasive species spread in developing regions that are infrequently monitored., Competing Interests: The authors have declared that no competing interests exist.

Plastic waste has become ubiquitous in ecosystems worldwide. Few, recent studies report evidence of coastal vegetated habitats acting as sink for plastics, yet assessments have been completed either for macro or microplastics and focussing on just one type of vegetated habitat. Here, we investigated the role of marine coastal vegetated habitats as sinks for macro (≥5 mm) and microplastics (<5 mm) through a comprehensive, multi-habitat approach. We assessed the occurrence, abundance and physical properties of macro and microplastics in the canopy and superficial sediment of two intertidal (seagrass Zostera noltei, saltmarsh Sporobolus maritimus) and two subtidal (mixed seagrass meadows of Cymodocea nodosa and Zostera marina, rhizophytic macroalga Caulerpa prolifera) habitats in the Ria Formosa lagoon (Portugal). Our results showed that coastal vegetated habitats trapped macro and microplastics in the sediment at variable degrees (1.3-17.3 macroplastics 100 m -2 , and 18.2-35.2 microplastics kg - 1 ). Macroplastics accumulated in all vegetated habitat but not in nearby unvegetated areas, yet only S. maritimus habitat presented a significant trapping effect. Microplastics occurred in the sediment of all vegetated and unvegetated areas with similar abundances and high variability. Microplastics, all of type fibre, were recorded on all canopies except for S. maritimus. Overall, the trapping capacity of microplastics in the sediment and on the canopy was higher for subtidal than for intertidal vegetated habitats. We conclude that generalizations in the trapping effect of coastal vegetated areas should be done with caution, since it may be highly variable and may depend on the plastic size, habitat and tidal position. Since these habitats support a high biodiversity, they should be included in assessments of plastic debris accumulation and impacts in coastal areas. Further research, including experimental studies, is needed to shed more light on the role of coastal vegetated habitats as plastic sinks., (Copyright © 2020 Elsevier B.V. All rights reserved.)

The 'Abundant-Centre Hypothesis' is a well-established but controversial hypothesis stating that the abundance of a species is highest at the centre of its range and decreases towards the edges, where conditions are unfavourable. As genetic diversity depends on population size, edge populations are expected to show lower intra-population genetic diversity than core populations, while showing high inter-population genetic divergence. Here, the genetic implications of the Abundant-Centre Hypothesis were tested on two coastal mussels from South Africa that disperse by means of planktonic larvae, the native Perna perna and the invasive Mytilus galloprovincialis. Genetic structure was found within P. perna, which, together with evidence from Lagrangian particle simulations, points to significant reductions in gene flow between sites. Despite this, the expected diversity pattern between centre and edge populations was not found for either species. We conclude that the genetic predictions of the Abundant-Centre Hypothesis are unlikely to be met by high-dispersal species with large population sizes, and may only become evident in species with much lower levels of connectivity.

Extreme events such as heat waves have increased in frequency and duration over the last decades. Under future climate scenarios, these discrete climatic events are expected to become even more recurrent and severe. Heat waves are particularly important on rocky intertidal shores, one of the most thermally variable and stressful habitats on the planet. Intertidal mussels, such as the blue mussel Mytilus edulis, are ecosystem engineers of global ecological and economic importance, that occasionally suffer mass mortalities. This study investigates the potential causes and consequences of a mass mortality event of M. edulis that occurred along the French coast of the eastern English Channel in summer 2018. We used an integrative, climatological and ecophysiological methodology based on three complementary approaches. We first showed that the observed mass mortality (representing 49 to 59% of the annual commercial value of local recreational and professional fisheries combined) occurred under relatively moderate heat wave conditions. This result indicates that M. edulis body temperature is controlled by non-climatic heat sources instead of climatic heat sources, as previously reported for intertidal gastropods. Using biomimetic loggers (i.e. 'robomussels'), we identified four periods of 5 to 6 consecutive days when M. edulis body temperatures consistently reached more than 30 °C, and occasionally more than 35 °C and even more than 40 °C. We subsequently reproduced these body temperature patterns in the laboratory to infer M. edulis thermal tolerance under conditions of repeated heat stress. We found that thermal tolerance consistently decreased with the number of successive daily exposures. These results are discussed in the context of an era of global change where heat events are expected to increase in intensity and frequency, especially in the eastern English Channel where the low frequency of commercially exploitable mussels already questions both their ecological and commercial sustainability.

Abstract: Assessing population responses to climate-related environmental change is key to understanding the adaptive potential of the species as a whole. Coralline algae are critical components of marine shallow water ecosystems where they function as important ecosystem engineers. Populations of the calcifying algae C orallina officinalis from the center (southern UK) and periphery (northern Spain) of the North Atlantic species natural distribution were selected to test for functional differentiation in thermal stress response. Physiological measurements of calcification, photosynthesis, respiration, growth rates, oxygen, and calcification evolution curves were performed using closed cell respirometry methods. Species identity was genetically confirmed via DNA barcoding. Through a common garden approach, we identified distinct vulnerability to thermal stress of central and peripheral populations. Southern populations showed a decrease in photosynthetic rate under environmental conditions of central locations, and central populations showed a decline in calcification rates under southern conditions. This shows that the two processes of calcification and photosynthesis are not as tightly coupled as previously assumed. How the species as whole will react to future climatic changes will be determined by the interplay of local environmental conditions and these distinct population adaptive traits., Open Research Badges: This article has earned an Open Materials Badge for making publicly available the components of the research methodology needed to reproduce the reported procedure and analysis. All materials are available at https://doi.pangaea.de/10.1594/PANGAEA.899568., Competing Interests: The authors declare that they have no conflict of interest.

In modern society, plastic items have become indispensable. The rapid growth of plastic production has led to an increase in the concentration of plastic waste in the environment and, consequently, wildlife has been severely affected. As wide-ranging foragers and predators, aquatic birds are ideal sentinels for monitoring changes in their environment. Plastic found in stomach contents of stranded aquatic birds collected throughout Portugal was examined. Out of the 288 birds processed, 12.9% ingested plastics. Six of the 16 species assessed showed evidence of plastic ingestion. The Lesser Black-backed Gull (18.7%) had the highest incidence while, among those that did ingest plastics, the Northern Gannet (4.8%) had the lowest. User plastics were the most common type of plastic ingested, while microplastics and off/white-clear were the most common size and colour respectively of plastics found. This study sets a first multispecies baseline for incidence of plastic ingestion by aquatic birds in Portugal., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

We report the development of nine polymorphic microsatellite markers for Corallina officinalis (Linnaeus, 1758), a calcifying intertidal red alga and important ecosystem engineer spread along the North East Atlantic. Characterization and analysis of loci were made using 15 individuals of C. officinalis from populations in Iceland and the UK. The average number of alleles per locus was 3.78 (range 2-6) and mean of gene diversity was 0.58 (range 0.38-0.77). The set of microsatellites developed here will provide a useful molecular tool for population genetic and conservation studies.

Excessive use of plastics in daily life and the inappropriate disposal of plastic products are severely affecting wildlife species in both coastal and aquatic environments. Birds are top-predators, exposed to all threats affecting their environments, making them ideal sentinel organisms for monitoring ecosystems change. We set a baseline assessment of the prevalence of marine plastic litter affecting multi-species populations of aquatic birds in southern Portugal. By examining 160 stomach contents from 8 species of aquatic birds, we show that 22.5% were affected by plastic debris. Plastic was found in Ciconia ciconia, Larus fuscus and L. michahellis. Ciconia ciconia ingested the highest amount (number of items and total mass) of plastic debris. Polydimethylsiloxane (PDMS, silicones) was the most abundant polymer and was recorded only in C. ciconia. Plastic ingestion baseline data are of crucial importance to evaluate changes through time and among regions and to define management and conservation strategies., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Oceanographic features shape the distributional and genetic patterns of marine species by interrupting or promoting connections among populations. Although general patterns commonly arise, distributional ranges and genetic structure are species-specific and do not always comply with the expected trends. By applying a multimarker genetic approach combined with Lagrangian particle simulations (LPS) we tested the hypothesis that oceanographic features along northeastern Atlantic and Mediterranean shores influence dispersal potential and genetic structure of the intertidal mussel Perna perna. Additionally, by performing environmental niche modelling we assessed the potential and realized niche of P. perna along its entire native distributional range and the environmental factors that best explain its realized distribution. Perna perna showed evidence of panmixia across >4,000 km despite several oceanographic breaking points detected by LPS. This is probably the result of a combination of life history traits, continuous habitat availability and stepping-stone dynamics. Moreover, the niche modelling framework depicted minimum sea surface temperatures (SST) as the major factor shaping P. perna distributional range limits along its native areas. Forthcoming warming SST is expected to further change these limits and allow the species to expand its range polewards though this may be accompanied by retreat from warmer areas.

Positive and negative aspects of species interactions can be context dependant and strongly affected by environmental conditions. We tested the hypothesis that, during periods of intense heat stress, parasitic phototrophic endoliths that fatally degrade mollusc shells can benefit their mussel hosts. Endolithic infestation significantly reduced body temperatures of sun-exposed mussels and, during unusually extreme heat stress, parasitised individuals suffered lower mortality rates than non-parasitised hosts. This beneficial effect was related to the white discolouration caused by the excavation activity of endoliths. Under climate warming, species relationships may be drastically realigned and conditional benefits of phototrophic endolithic parasites may become more important than the costs of infestation.

The indigenous South African mussel Perna perna gapes during periods of aerial exposure to maintain aerobic respiration. This behaviour has no effect on the body temperatures of isolated individuals, but when surrounded by conspecifics, beneficial cooling effects of gaping emerge. It is uncertain, however, whether the presence of the invasive mussel Mytilus galloprovincialis limits the ability of P. perna for collective thermoregulation. We investigated whether varying densities of P. perna and M. galloprovincialis influences the thermal properties of both natural and artificial mussel beds during periods of emersion. Using infrared thermography, body temperatures of P. perna within mixed artificial beds were shown to increase faster and reach higher temperatures than individuals in conspecific beds, indicating that the presence of M. galloprovincialis limits the group cooling effects of gaping. In contrast, body temperatures of M. galloprovincialis within mixed artificial mussel beds increased slower and exhibited lower temperatures than for individuals in beds comprised entirely of M. galloprovincialis. Interestingly, differences in bed temperatures and heating rates were largely dependent on the size of mussels, with beds comprised of larger individuals experiencing less thermal stress irrespective of species composition. The small-scale patterns of thermal stress detected within manipulated beds were not observed within naturally occurring mixed mussel beds. We propose that small-scale differences in topography, size-structure, mussel bed size and the presence of organisms encrusting the mussel shells mask the effects of gaping behaviour within natural mussel beds. Nevertheless, the results from our manipulative experiment indicate that the invasive species M. galloprovincialis steals thermal properties as well as resources from the indigenous mussel P. perna. This may have significant implications for predicting how the co-existence of these two species may change as global temperatures continue to rise.

Invasive species can affect the function and structure of natural ecological communities, hence understanding and predicting their potential for spreading is a major ecological challenge. Once established in a new region, the spread of invasive species is largely controlled by their dispersal capacity, local environmental conditions and species interactions. The mussel Mytilus galloprovincialis is native to the Mediterranean and is the most successful marine invader in southern Africa. Its distribution there has expanded rapidly and extensively since the 1970s, however, over the last decade its spread has ceased. In this study, we coupled broad scale field surveys, Ecological Niche Modelling (ENM) and Lagrangian Particle Simulations (LPS) to assess the current invaded distribution of M. galloprovincialis in southern Africa and to evaluate what prevents further spread of this species. Results showed that all environmentally suitable habitats in southern Africa have been occupied by the species. This includes rocky shores between Rocky Point in Namibia and East London in South Africa (approx. 2800 km) and these limits coincide with the steep transitions between cool-temperate and subtropical-warmer climates, on both west and southeast African coasts. On the west coast, simulations of drifting larvae almost entirely followed the northward and offshore direction of the Benguela current, creating a clear dispersal barrier by advecting larvae away from the coast. On the southeast coast, nearshore currents give larvae the potential to move eastwards, against the prevalent Agulhas current and beyond the present distributional limit, however environmental conditions prevent the establishment of the species. The transition between the cooler and warmer water regimes is therefore the main factor limiting the northern spread on the southeast coast; however, biotic interactions with native fauna may also play an important role.

Background: Intraspecific variability is seen as a central component of biodiversity. We investigated genetic differentiation, contemporary patterns of demographic connectivity and intraspecific variation of adaptive behavioural traits in two lineages of an intertidal mussel (Perna perna) across a tropical/subtropical biogeographic transition., Results: Microsatellite analyses revealed clear genetic differentiation between western (temperate) and eastern (subtropical/tropical) populations, confirming divergence previously detected with mitochondrial (COI) and nuclear (ITS) markers. Gene flow between regions was predominantly east-to-west and was only moderate, with higher heterozygote deficiency where the two lineages co-occur. This can be explained by differential selection and/or oceanographic dynamics acting as a barrier to larval dispersal. Common garden experiments showed that gaping (periodic closure and opening of the shell) and attachment to the substratum differed significantly between the two lineages. Western individuals gaped more and attached less strongly to the substratum than eastern ones., Conclusions: These behavioural differences are consistent with the geographic and intertidal distributions of each lineage along sharp environmental clines, indicating their strong adaptive significance. We highlight the functional role of diversity below the species level in evolutionary trends and the need to understand this when predicting biodiversity responses to environmental change.

The accuracy of phylogenetic inference can be significantly improved by the addition of more taxa and by increasing the spatial coverage of sampling. In previous studies, the brown mussel Perna perna showed a sister-lineage relationship between eastern and western individuals contiguously distributed along the South African coastline. We used mitochondrial (COI) and nuclear (ITS) sequence data to further analyze phylogeographic patterns within P. perna. Significant expansion of the geographical coverage revealed an unexpected pattern. The western South African lineage shared the most recent common ancestor (MRCA) with specimens from Angola, Venezuela, and Namibia, whereas eastern South African specimens and Mozambique grouped together, indicating a non-sister relationship for the two South African lineages. Two plausible biogeographic scenarios to explain their origin were both supported by the hypotheses-testing analysis. One includes an Indo-Pacific origin for P. perna, dispersal into the Mediterranean and Atlantic through the Tethys seaway, followed by recent secondary contact after southward expansion of the western and eastern South African lineages. The other scenario (Out of South Africa) suggests an ancient vicariant divergence of the two lineages followed by their northward expansion. Nevertheless, the "Out of South Africa" hypothesis would require a more ancient divergence between the two lineages. Instead, our estimates indicated that they diverged very recently (310 kyr), providing a better support for an Indo-Pacific origin of the two South African lineages. The arrival of the MRCA of P. perna in Brazil was estimated at 10 [0-40] kyr. Thus, the hypothesis of a recent introduction in Brazil through hull fouling in wooden vessels involved in the transatlantic itineraries of the slave trade did not receive strong support, but given the range for this estimate, it could not be discarded. Wider geographic sampling of marine organisms shows that lineages with contiguous distributions need not share a common ancestry.

Background: Significant effects of recent global climate change have already been observed in a variety of ecosystems, with evidence for shifts in species ranges, but rarely have such consequences been related to the changes in the species genetic pool. The stretch of Atlantic coast between North Africa and North Iberia is ideal for studying the relationship between species distribution and climate change as it includes the distributional limits of a considerable number of both cold- and warm-water species.We compared temporal changes in distribution of the canopy-forming alga Fucus vesiculosus with historical sea surface temperature (SST) patterns to draw links between range shifts and contemporary climate change. Moreover, we genetically characterized with microsatellite markers previously sampled extinct and extant populations in order to estimate resulting cryptic genetic erosion., Results: Over the past 30 years, a geographic contraction of the southern range edge of this species has occurred, with a northward latitudinal shift of approximately 1,250 km. Additionally, a more restricted distributional decline was recorded in the Bay of Biscay. Coastal SST warming data over the last three decades revealed a significant increase in temperature along most of the studied coastline, averaging 0.214°C/decade. Importantly, the analysis of existing and extinct population samples clearly distinguished two genetically different groups, a northern and a southern clade. Because of the range contraction, the southern group is currently represented by very few extant populations. This southern edge range shift is thus causing the loss of a distinct component of the species genetic background., Conclusions: We reveal a climate-correlated diversity loss below the species level, a process that could render the species more vulnerable to future environmental changes and affect its evolutionary potential. This is a remarkable case of genetic uniqueness of a vanishing cryptic genetic clade (southern clade).

Background: The brown mussel Perna perna (Linnaeus, 1758) is a dominant constituent of intertidal communities and a strong invader with multiple non-native populations distributed around the world. In a previous study, two polymorphic microsatellite loci were developed and used to determine population-level genetic diversity in invasive and native P. perna populations. However, higher number of microsatellite markers are required for reliable population genetic studies.In this context, in order to understand P. perna origins and history of invasion and to compare population genetic structure in native versus invaded areas, we developed 10 polymorphic microsatellite markers., Findings: Described microsatellite markers were developed from an enriched genomic library. Analyses and characterization of loci using 20 individuals from a population in Western Sahara revealed on average 11 alleles per locus (range: 5-27) and mean gene diversity of 0.75 (range: 0.31 - 0.95). One primer pair revealed possible linkage disequilibrium while heterozygote deficiency was significant at four loci. Six of these markers cross-amplified in P. canaliculus (origin: New Zealand)., Conclusions: Developed markers will be useful in addressing a variety of questions concerning P. perna, including dispersal scales, genetic variation and population structure, in both native and invaded areas.

By associating closely with others to form a group, an animal can benefit from a number of advantages including reduced risk of predation, amelioration of environmental conditions, and increased reproductive success, but at the price of reduced resources. Although made up of individual members, an aggregation often displays novel effects that do not manifest at the level of the individual organism. Here we show that very simple behaviour in intertidal mussels shows new effects in dense aggregations but not in isolated individuals. Perna perna and Mytilus galloprovincialis are gaping (periodic valve movement during emersion) and non-gaping mussels respectively. P. perna gaping behaviour had no effect on body temperatures of isolated individuals, while it led to increased humidity and decreased temperatures in dense groups (beds). Gaping resulted in cooler body temperatures for P. perna than M. galloprovincialis when in aggregations, while solitary individuals exhibited the highest temperatures. Gradients of increasing body temperature were detected from the center to edges of beds, but M. galloprovincialis at the edge had the same temperature as isolated individuals. Furthermore, a field study showed that during periods of severe heat stress, mortality rates of mussels within beds of the gaping P. perna were lower than those of isolated individuals or within beds of M. galloprovincialis, highlighting the determinant role of gaping on fitness and group functioning. We demonstrate that new effects of very simple individual behaviour lead to amelioration of abiotic conditions at the aggregation level and that these effects increase mussel resistance to thermal stress.

Understanding patterns of gene flow, selection and genetic diversity within and among populations is a critical element of predicting how long-term changes in environmental conditions are likely to affect species distribution. The intertidal mussel Perna perna consists of two distinct genetic lineages in South Africa, but the mechanisms maintaining these lineages remains obscure. We used regional oceanography and lineage-specific responses to environmental conditions as proxies for gene flow and local selection, respectively, to test how these mechanisms could shape population genetic structure. Laboratory experiments supported the field findings that mussels on the east coast (eastern lineage) are physiologically more tolerant of sand inundation and high temperatures than those on the south coast (western lineage). Temperature loggers mimicking mussel body temperatures revealed that mussels experience higher body temperatures during aerial exposure on the subtropical east coast than on the temperate south coast. Translocations showed that, on the east coast, the western lineage suffered higher mortality rates than local individuals, while on the south coast, mortality rates did not differ significantly between the lineages. Nearshore drogues showed remarkably little overlap between the trajectories of drifters released off the south coast and those released off the east coast. Physiological tolerances can thus explain the exclusion of western individuals from the east coast, but they cannot explain the exclusion of the eastern lineage from the south coast. In contrast, however, ocean dynamics may limit larval dispersal between the two lineages, helping to explain the absence of eastern individuals from the south coast. We emphasise the importance of a multidisciplinary approach in a macro-ecological context to understand fully the mechanisms promoting evolutionary divergence between genetic entities. Our results suggest that phylogeographic patterns of Perna perna may be maintained by a combination of local conditions and the isolating effect of the Agulhas Current that reduces gene exchange.

Gene flow among hybridizing species with incomplete reproductive barriers blurs species boundaries, while selection under heterogeneous local ecological conditions or along strong gradients may counteract this tendency. Congeneric, externally-fertilizing fucoid brown algae occur as distinct morphotypes along intertidal exposure gradients despite gene flow. Combining analyses of genetic and phenotypic traits, we investigate the potential for physiological resilience to emersion stressors to act as an isolating mechanism in the face of gene flow. Along vertical exposure gradients in the intertidal zone of Northern Portugal and Northwest France, the mid-low shore species Fucus vesiculosus, the upper shore species Fucus spiralis, and an intermediate distinctive morphotype of F. spiralis var. platycarpus were morphologically characterized. Two diagnostic microsatellite loci recovered 3 genetic clusters consistent with prior morphological assignment. Phylogenetic analysis based on single nucleotide polymorphisms in 14 protein coding regions unambiguously resolved 3 clades; sympatric F. vesiculosus, F. spiralis, and the allopatric (in southern Iberia) population of F. spiralis var. platycarpus. In contrast, the sympatric F. spiralis var. platycarpus (from Northern Portugal) was distributed across the 3 clades, strongly suggesting hybridization/introgression with both other entities. Common garden experiments showed that physiological resilience following exposure to desiccation/heat stress differed significantly between the 3 sympatric genetic taxa; consistent with their respective vertical distribution on steep environmental clines in exposure time. Phylogenetic analyses indicate that F. spiralis var. platycarpus is a distinct entity in allopatry, but that extensive gene flow occurs with both higher and lower shore species in sympatry. Experimental results suggest that strong selection on physiological traits across steep intertidal exposure gradients acts to maintain the 3 distinct genetic and morphological taxa within their preferred vertical distribution ranges. On the strength of distributional, genetic, physiological and morphological differences, we propose elevation of F. spiralis var. platycarpus from variety to species level, as F. guiryi.

Background: Environmental heterogeneity plays a major role in invasion and coexistence dynamics. Habitat segregation between introduced species and their native competitors is usually described in terms of different physiological and behavioural abilities. However little attention has been paid to the effects of behaviour in habitat partitioning among invertebrates, partially because their behavioural repertoires, especially marine benthic taxa, are extremely limited. This study investigates the effect of gaping behaviour on habitat segregation of the two dominant mussel species living in South Africa, the invasive Mytilus galloprovincialis and the indigenous Perna perna. These two species show partial habitat segregation on the south coast of South Africa, the lower and upper areas of the mussel zone are dominated by P. perna and M. galloprovincialis respectively, with overlap in the middle zone. During emergence, intertidal mussels will either keep the valves closed, minimizing water loss and undergoing anaerobic metabolism, or will periodically open the valves maintaining a more efficient aerobic metabolism but increasing the risk of desiccation., Results: Our results show that, when air exposed, the two species adopt clearly different behaviours. M. galloprovincialis keeps the shell valves closed, while P. perna periodically gapes. Gaping behaviour increased water loss in the indigenous species, and consequently the risk of desiccation. The indigenous species expressed significantly higher levels of stress protein (Hsp70) than M. galloprovincialis under field conditions and suffered significantly higher mortality rates when exposed to air in the laboratory. In general, no intra-specific differences were observed in relation to intertidal height. The absence of gaping minimises water loss but exposes the invasive species to other stresses, probably related to anoxic respiration., Conclusions: Gaping affects tolerance to desiccation, thus influencing the vertical zonation of the two species. Valve closure exposes the invasive species to higher stress and associated energy demands, but it minimizes water loss, allowing this species to dominate the upper mussel zone, where the gaping indigenous P. perna cannot survive. Thus even very simple behaviour can influence the outcome of interactions between indigenous and invasive species.