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Heat-induced mass mortalities involving ecosystem engineers may have long-lasting detrimental effects at the community level, eliminating the ecosystem services they provide. Intertidal mussels are ecologically and economically valuable with some populations facing unprecedented heat-induced mass mortalities. Critically, mussels are also frequently infested by endolithic parasites that modify shell albedo, hence reducing overheating and mortality rates under heat stress. Using a biophysical model, we explored the topographical and meteorological conditions under which endolithically driven thermal buffering becomes critical to survival. Based on meteorological data from a global climate analysis, we modelled body temperatures of infested and non-infested mussels over the last decade (2010-2020) at nine sites spread across c. 20 degrees of latitude. We show that thermal buffering is enhanced where and when heat stress is greatest, that is, on sun-exposed surfaces under high solar radiation and high air temperature. These results suggest that new co-evolutionary pathways are likely to open for these symbiotic organisms as climate continues to change, potentially tipping the balance of the relationship from a parasitic to a more mutualistic one. However, endolithically driven reductions in body temperatures can also occur at or below optimal temperatures, thereby reducing the host's metabolic rates and making the interplay of positive and negative effects complex. In parallel, we hindcasted body temperatures using empirical data from nearby weather stations and found that predictions were very similar with those obtained from two global climate reanalyses (i.e. NCEP-DOE Reanalysis 2 and ECMWF Reanalysis v5). This result holds great promise for modelling the distribution of terrestrial ectotherms at ecologically relevant spatiotemporal scales, as it suggests we can reasonably bypass the practical issues associated with weather stations. For intertidal ectotherms, however, the challenge will be incorporating body temperatures over the full tidal cycle. info:eu-repo/semantics/publishedVersion

Aim Biogeographic boundaries can act as either weak or strong barriers to the spread of species undergoing distributional change. Once a novel species spreads across a boundary, it can have a substantial impact on the ecosystem, for instance by competing with local species, and, over the long‐term, re‐engineer the ecosystem. Marine biogeographic regions are clearly defined on the coast of southern Africa and we tested the influence of their boundaries on the spread and spatial structure of an alien ecosystem engineer. Location Southern Africa. Taxon An invasive mussel, Mytilus galloprovincialis. Methods Records of M. galloprovincialis were compiled into a database to determine its decadal patterns of spread across multiple bioregions. Distribution and abundance (estimated using density and semi‐quantitative abundance scale [ACFOR]) of this mussel were surveyed to determine patterns in spatial structure across bioregions. In addition, we compared the size structure of populations at the eastern margins of its range with those of larger populations nearer the range centre. Results Initial breaching of biogeographic boundaries was associated with rapid spread, but other boundaries encountered decades later acted as barriers to further spread. Across >2,800 km of coast, spatial autocorrelation was observed in densities (low and mid shore levels) and in ACFOR abundances. Repeating spatial patterns in densities and ACFOR abundance were detected at scales of 120–160 km and of 400–990 km. Considerable effort was required to detect populations at the absolute eastern limits of its distribution. Main conclusion Detection of spatial structures at multiple scales suggests that ecologically determined processes regulate abundance at both intra‐bioregional and inter‐bioregional scales, which may help tease apart the historic and contemporary consequences of interspecific interactions on the structure of rocky shore communities. This study demonstrates the influence of biogeography in driving temporal patterns of spread and spatial structure on the distribution and abundance of an invasive species.

Self-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. National Research Foundation of South AfricaNational Research Foundation - South Africa [64801]; Fundacao para a Ciencia e Tecnologia (FCT-MEC, Portugal); French Ministere de l'Enseignement Superieur et de la Recherche; Hauts de France RegionRegion Hauts-de-France; European Funds for Regional Economical Development

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. info:eu-repo/semantics/publishedVersion

Mussels are ecological engineers in intertidal communities; they afect coastal species richness by increasing habitat spatial complexity, bufering against environmental extremes, and providing protection from predators. Parasitic activities of endolithic cyanobacteria on mussels weaken their shells, requiring the expenditure of energy on shell repair, with potential indirect efects on organisms that rely on mussels as habitat providers. Given the seasonality of reproduction and the need for strong attachment during winter storms, we examined the consequences of redirecting energy for shell repair to two other energetically expensive processes: reproduction and byssal attachment. We examined seasonality in the efects of cyanobacterial infestation on reproduction and attachment strength in two intertidal mussels, the indigenous Perna perna and the invasive Mytilus galloprovincialis from the south coast of South Africa, using both qualitative and quantitative analyses. Reproductive efects were examined by measuring the weight of mussel gonads and the density of eggs within each gonad for co-occurring infested and non-infested mussels, while attachment strength was measured for mussels exhibiting diferent levels of infestation. Endolithic infestation was found to afect reproduction by reducing the mass of gonads, but not the density of eggs within them. Attachment strength was closely correlated with the degree of endolithic infestation, with very infested mussels requiring much less force to detach them from the substratum than mussels with low or no infestation. Thus, endolithic infestation afected mussel ftness directly, by increasing the probably of mortality through wave dislodgement and by reducing reproductive output. info:eu-repo/semantics/publishedVersion

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. info:eu-repo/semantics/publishedVersion

Our understanding of how anthropogenic stressors such as climate change and plastic pollution interact with biodiver-sity is being widened to include diversity below the species level, i.e., intraspecific variation. The emerging apprecia-tion 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 experi-ments and laboratory-controlled hydrodynamic simulations, we assessed the relative effects of intraspecific and inter -specific 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 den-sity 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 accu-mulation in biogenic habitats requires an understanding of within-and between-species traits at both the individual and population levels. info:eu-repo/semantics/publishedVersion

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 degrees 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 degrees C and 4.8 degrees 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. Fundacao para a Ciencia e TecnologiaPortuguese Foundation for Science and TechnologyEuropean Commission [UIDB/04326/2020]; National Research Foundation of South AfricaNational Research Foundation - South Africa [64801]; Department of Science and TechnologyDepartment of Science & Technology (India); National FoundationNational Science Foundation (NSF); South African National Research FoundationNational Research Foundation - South Africa; French Ministere de l'Enseignement Superieur et de la Recherche; Hauts de France RegionRegion Hauts-de-France; European Funds for Regional Economical Development info:eu-repo/semantics/publishedVersion

Through seafood consumption, microplastic (MP) pollution is potentially threatening human health. Commercial bivalves in particular are a cause of major concern because their filter-feeding activity directly exposes them to MP in the water column and they are then ingested by humans. Here, we provide a quantitative and qualitative baseline data on MP content in the soft tissues of three commercially important bivalves (Ruditapes decussatus, Cerastoderma spp. and Polititapes spp.) collected in Ria Formosa lagoon, southern Portugal. The abundance of MPs (items per soft tissue weight) did not significantly differ among species. On average, R. decussatus exhibited the highest MP abundance (on average, 18.4 +/- 21.9 MP items g(-1) WW), followed by Cerastoderma spp. (11.9 +/- 5.5 MP items g(-1) WW) and Polititapes spp. (10.4 +/- 10.4 MP items g(-1) WW). Overall, 88% of the MPs found were synthetic fibres, the majority of which were blue (52%). Size categories >0.1-1 mm and >1-5 mm were the most common (60% and 34% respectively). The most represented polymers were polyethylene (PE) and polystyrene (PS). The unexpectedly high number of MPs recorded in the three commercially exploited species suggests that this semi-closed lagoon system is experiencing a higher anthropogenic pressure than are open coastal systems. Portuguese Foundation for Science and Technology: PTDC/MAR-EST/3223/2014 IF/01413/2014/CP1217/CT0004 UIDB/04326/2020 SFRH/BPD/119344/2016 info:eu-repo/semantics/publishedVersion

Aim Brown mussels exhibit a trans-Atlantic distribution putatively caused by either native dispersal or artificial gene flow, likely in concert with the transport of enslaved people from Africa. Evolutionary history and demographic models of this widespread species may clarify how the present-day distribution was impacted by natural versus artificial dispersal. Particularly, dating the timing of the South American/African split may determine whether the human slave trade likely impacted the contemporary distribution of brown mussels. Location Coastal Brazil, Morocco, South Africa, and Mozambique. Taxon Perna perna (Linnaeus 1758). Methods We genotyped a total of 644 samples from 18 populations at 10 microsatellite loci. We estimated genetic structure with clustering algorithms in STRUCTURE and GENETIX. We estimated genetic distances by characterizing patterns of pair-wise F-ST using the program FSTAT, evaluating differences among and between regions via AMOVA, and testing isolation by distance in IBDWS. To estimate and date the most likely pathway by which P. perna crossed the Atlantic Ocean we used Bayesian factors from thermodynamically heated coalescent simulations in the program MIGRATE-n. Results We found no general pattern of reduced or elevated levels of genetic diversity within any region across site or locus. We identified four genetic clusters: East South Africa (ESA), West South Africa (WSA), Brazil (BR) and North Africa (MO); F-ST ranged from 0.06 to 0.11 among regions and exhibited a significant pattern of isolation by distance. Migration models indicated that P. perna dispersed from WSA to MO and from there to BR of approximately 2,000 years. Main conclusions Multiple lines of evidence suggest the Brazilian populations of P. perna have been a long-standing native population, originating from northern Africa and are unlikely a consequence of the African slave trade. Although, human introduction cannot be ruled out South American P. perna populations exhibited genetic characteristics indicative of a divergent, isolated and established population, featuring the genetic signature expected for a native population. info:eu-repo/semantics/publishedVersion

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. French Ministere de l'Enseignement Superieur et de la Recherche Hauts de France RegionRegion Hauts-de-France European Funds for Regional Economical Development Pierre Hubert Curien PESSOA Fellowship Fundacao para a Ciencia e Tecnologia (FCT-MEC, Portugal) [IF/01413/2014/CP1217/CT0004] National Research Foundation of South AfricaNational Research Foundation - South Africa [64801] South African Research Chairs Initiative (SARChI) of the Department of Science and Technology National FoundationNational Science Foundation (NSF) South African National Research Foundation (NRF)National Research Foundation - South Africa info:eu-repo/semantics/publishedVersion

Intertidal mussels are important ecosystem engineers that increase habitat complexity and provide shelter, substratum and food to associated communities. They thus play a key role in the maintenance of local biodiversity. Intertidal mussels are frequently parasitized by endolithic cyanobacteria that erode and significantly weaken their shells. We examined succession of cyanobacterial infestation in the shells of the mussels Perna perna and Mytilus galloprovincialis on the south coast of South Africa. We examined variations in species composition of the endolithic cyanobacterial community across regions of mussel shells of different ages (variation in space) and by exposing non-infested shells in field experiments for known periods (variation in time). The results on the spatial distribution of endolith species within a shell supported those from the field experiment evaluating temporal succession. Endolithic species that were early colonists of clean shells were similar to those that were found at the distal edge, the new and growing region of the shell. These species were also found near the umbo, the oldest region of the shell, but cyanobacterial communities near the umbo also included species that appeared late in time. Thus, succession when viewed in either time or space involved the accumulation, rather than the replacement of species. Species that arrived late in the succession were similar to endolithic species found near the umbo, the oldest region of the shell. Overall, the study shows clear patterns of endolithic cyanobacterial succession within mussel shells.

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. Fundacao para a Ciencia e a Tecnologia (FCT - MEC, Portugal) [IF/01413/2014/CP1217/CT0004, UIDB/Multi/04326/2020] National Research Foundation of South AfricaNational Research Foundation - South Africa [64801] info:eu-repo/semantics/publishedVersion

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.

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. (C) 2020 Elsevier B.V. All rights reserved. National Research Foundation of South AfricaNational Research Foundation - South Africa [64801] Fundacao para a Ciencia e Tecnologia (FCT-MEC, Portugal) [UID/Multi/04326/2019, IF/01413/2014/CP1217/CT0004] info:eu-repo/semantics/publishedVersion

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 (

Upwelling strongly influences the composition and dynamics of coastal communities by affecting species abundances, recruitment, dispersal and distribution. Coastal upwelling areas are key model regions to study the responses of coastal species to climate change because they are characterized by cooler water conditions and experience lower warming rates than adjacent regions, making them effective 'control' or refuge sites. This is particularly true for the benthic species of rocky shores in upwelling areas because they are sedentary, inhabit the interface between marine and terrestrial habitats, are exposed to extremely severe and variable environmental conditions and often live near their tolerance limits. We sampled roughly 2000 km of the Atlantic coast of Morocco and Western Sahara to assess the influence of upwelling cells on patterns of diversity and abundance of northern African rocky shore species. We recorded 186 taxa, providing clarification of the distribution of 141 algae and documenting nine new species records for Morocco and Western Sahara. The results emphasize the influence of upwelling on the abundance and distribution of these organisms. The contrast between non-upwelling and upwelling areas highlights the direct and indirect importance of water temperature in shaping these communities, pointing to the consequences of large-scale warming. Such warming is likely to threaten intertidal species that already live close to their thermal tolerance limits and are not buffered by the effects of upwelling. Fundacao para a Ciencia e Tecnologia (FCT-MEC, Portugal) [UID/Multi/04326/2019, IF/01413/2014/CP1217/CT0004] National Research Foundation of South AfricaNational Research Foundation - South Africa [64801] Programme Science without Borders (Conselho Nacional de Desenvolvimento Cientifico e Tecnologico, of the Ministry of Science, Technology and Innovation of Brazil) [237998/2012-2] info:eu-repo/semantics/publishedVersion

Genetic structure in biogeographical transition zones can be shaped by several factors including limited dispersal across barriers, admixture following secondary contact, differential selection, and mating incompatibility. A striking example is found in Northwest France and Northwest Spain, where the estuarine seaweed Fucus ceranoides L. exhibits sharp, regional genetic clustering. This pattern has been related to historical population fragmentation and divergence into distinct glacial refugia, followed by post-glacial expansion and secondary contact. The contemporary persistence of sharp ancient genetic breaks between nearby estuaries has been attributed to prior colonization effects (density barriers) but the effect of oceanographic barriers has not been tested. Here, through a combination of mesoscale sampling (15 consecutive populations) and population genetic data (mtIGS) in NW France, we define regional genetic disjunctions similar to those described in NW Iberia. Most importantly, using high resolution dispersal simulations for Brittany and Iberian populations, we provide evidence for a central role of contemporary hydrodynamics in maintaining genetic breaks across these two major biogeographic transition zones. Our findings further show the importance of a comprehensive understanding of oceanographic regimes in hydrodynamically complex coastal regions to explain the maintenance of sharp genetic breaks along continuously populated coastlines. Foundation for Science and Technology (FCT-MEC, Portugal) [DL57/2016/CP1361/CT0035] National Research Foundation of South AfricaNational Research Foundation - South Africa [64801] FCTPortuguese Foundation for Science and TechnologyEuropean Commission [SFRH/BPD/88935/2012, SFRH/BPD/111003/2015] Pew Marine Fellowship (USA) [SFRH/BPD/111003/2015] [BIODIVERSA/004/2015] [IF/01413/2014/CP1217/CT0004] [UID/Multi/04326/2019] info:eu-repo/semantics/publishedVersion

Marine litter is widely dispersed throughout coastal environments. Assessing the distribution and accumulation of such contaminants is crucial to understand their environmental impacts. This study presents a baseline for the monitoring of litter and microplastics in intertidal sediments along the Atlantic shores of southern Portugal and Morocco and identifies potential sources of contamination. Although variable, distribution and composition of both litter and microplastics did not follow a latitudinal pattern. Most of the litter had an undifferentiated source. Within the identifiable sources of litter, food packaging, fishing and tobacco were the most abundant, with variable contributions among sites. Over 97% of marine litter retrieved was plastic. Fragments and filaments were the most abundant categories of plastics at sites with the highest and lowest microplastic abundance respectively. Filaments were mainly made of Polypropylene (PP,50%) and Polyethylene terephthalate (PET,29%) while the predominant polymers for fragments were Polyethene (PE, 75%) and PP (25%). Fundacao para a Ciencia e Tecnologia (FCT-MEC, Portugal) [UID/Multi/04326/2019, IF/01413/2014] South African Research Chairs Initiative (SARChI) of the Department of Science and Technology [64801]

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.

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.

Ecosystems often face a complex combination of natural and anthropogenic disturbances. The assessment of anthropogenic pressures and co-occurring natural stressors is crucial to our understanding of ecosystem structures, dynamics and their conservation. The present study provides the first experimental assessment of the combined effects of trampling (anthropogenic stressor) and endolithic induced erosion (natural stressor) on two coexisting intertidal mussel species, Mytilus galloprovincialis (de Lamarck, 1819) and Perna perna (Linnaeus, 1758). Mixed beds of the two species experiencing a wide range of endolithic parasitation were exposed to increasing intensities of human trampling. Our results clearly show that endolith-infested mussels are more vulnerable to trampling than non-infested individuals. At high trampling intensities, P. perna suffered significantly lower mortalities rates than M. galloprovincialis. More than 20% of large, infested M. galloprovincialis individuals were crushed even at low trampling intensities. Unexpectedly, mortalities rates decreased with mussel size, suggesting a sheltering role of larger conspecifics. Beyond ecological interest, such findings can provide critical experimental support for conservation and management actions. Fundacao para a Ciencia e Tecnologia (FCT-MEC, Portugal) [UID/Multi/04326/2013, IF/01413/2014/CP1217/CT0004] South African Research Chairs Initiative (SARChI) of Department of Science and Technology (DST) National Research Foundation (NRF)

The effects of environmental changes on species distribution are generally studied at large geographical scales. However, aggregations of individuals can significantly moderate the impact of the environment at smaller, organismal scales. We focused on the intertidal macroalga Fucus guiryi and carried out field and laboratory common garden experiments to evaluate how the different individual morphologies and canopy densities typical of central and peripheral populations modify microhabitat conditions and associated levels of stress. We show that F. guiryi canopies significantly alter environmental conditions (i.e., temperature, humidity and light regimes) and mitigate the levels of stress experienced by individuals within the group. Southern algae are more branched and form denser canopies but, unexpectedly, despite these considerable differences, the mitigating effects of northern and southern canopies did not differ significantly. Microhabitat conditions beneath canopies were more stressful at marginal locations, indicating that southern populations are not more effective than northern algae at mitigating the harsher climate at the edge of the species distribution. Our findings highlight the importance of assessing structural changes in aggregating species across their distribution and relating these to local climates to understand the impact of environmental changes at scales relevant to individual organisms. Fundacao para a Ciencia e Tecnologia (FCT-MEC, Portugal) [UID/Multi/04326/2013, IF/01413/2014/CP1217/CT0004] South African Research Chairs Initiative (SARChI) of the Department of Science and Technology National Research Foundation

By altering the phenotypic properties of their hosts, endolithic parasites can modulate the engineering processes of marine ecosystem engineers. Here, we assessed the biogeographical patterns of species assemblages, prevalence and impact of endolithic parasitism in two mussel species that act as important ecosystem engineers in the southern African intertidal habitat, Perna perna and Mytilus galloprovincialis. We conducted large-scale surveys across three biogeographic regions along the South African coast: the subtropical east coast, dominated by the indigenous mussel, P. perna, the warm temperate south coast, where this species coexists with the invasive Mediterranean mussel, M. galloprovincialis, and the cool temperate west coast dominated by M. galloprovincialis. Infestation increased with mussel size, and in the case of M. galloprovincialis we found a significantly higher infestation in the cool temperate bioregion than the warm temperate region. For P. perna, the prevalence of infestation was higher on the warm temperate than the subtropical region, though the difference was marginally non-significant. On the south coast, there was no significant difference in infestation prevalence between species. Endolith-induced mortality rates through shell collapse mirrored the patterns for prevalence. For P. perna, endolith species assemblages revealed clear grouping by bioregions. Our findings indicate that biogeography affects cyanobacteria species composition, but differences between biogeographic regions in their effects are driven by environmental conditions. Agência financiadora Número do subsídio Fundacao para a Ciencia e Tecnologia (FCT-MEC, Portugal) UID/Multi/04326/2019 IF/01413/2014/CP1217/CT0004 South African Research Chairs Initiative (SARChI) of the Department of Science and Technology National Research Foundation info:eu-repo/semantics/publishedVersion

In modem 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. Fundacao para a Ciencia e a Tecnologia (FCT-MEC, Portugal) [UID/Multi/04326/2013, IF/01413/2014] South African Research Chairs Initiative (SARChI) of the Department of Science and Technology National FoundationNational Science Foundation (NSF) South African National Research Foundation (NRF)National Research Foundation - South Africa

Aim Although many studies have reported the effects of climate change on species' distributions, most of them consider each species as a physiologically homogenous unit. However, different lineages or populations inhabiting distinct bioregions within a species' distributional range can retain unique genetic diversity that could result in distinct adaptive capacities. A recent, large, climate-correlated distributional range contraction occurred at the southern edge of the intertidal macroalga Fucus vesiculosus, causing loss of genetic diversity unique to the southern clade. We tested for differential selective constraints and signs of local adaptation to thermal stress in the two genetic lineages. Location Iberian Atlantic shores. Methods We performed a series of common garden experiments and a field reciprocal transplant. Results In the laboratory, southern F. vesiculosus showed higher resilience to heat stress than northern individuals. On the southwest coast of Portugal, local individuals grew more than those transplanted from the northern range; in the north of Portugal, growth rates did not differ significantly between lineages. Conclusions We present evidence for unique adaptive traits at the retreating edge of the species' distribution that could be lost if warming trends persist. The loss of the distinct southern genetic heritage could end potential ongoing diversification or speciation processes and impoverishes the adaptive potential of the species as a whole.

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.

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. Fundacao para a Ciencia e Tecnologia (FCT-MEC, Portugal) [UID/Multi/04326/2013, IF/01413/2014/CP1217/CT0004]; South African Research Chairs Initiative (SARChI) of the Department of Science and Technology; National Research Foundation; South African National Research Foundation (NRF); Portuguese Fundacao para a Ciencia e Tecnologia (FCT) [SFRH/BPD/85040/2012, SFRH/BPD/111003/2015] info:eu-repo/semantics/publishedVersion

Aggregations of organisms commonly benefit their members by mitigating the effects of predators and environmental stresses. Mussel aggregations also form important intertidal matrices that support associated infaunal communities, the structures of which are largely shaped by the conditions within the interstitial spaces. Intertidal mussels are frequently parasitized by phototrophic endoliths that infest the shell and have thermoregulatory effects on both solitary and aggregated mussels by changing shell albedo. A large-scale sampling was carried out May June 2016 along Portuguese and Moroccan shores to investigate a latitudinal gradient of endolithic infestation of the intertidal mussel Mytilus galloprovincialis. Endolithic infestation increased towards lower latitudes most likely as a response of greater light availability. Additionally, artificial beds of either 100% non-infested or infested biomimetic mussels were used to test whether infestation alters the temperature and humidity of the interstitial spaces within beds, and if mussels surrounded by infested mussels experience lower body temperatures than those surrounded by non-infested ones. Conditions within beds of infested mussels were significantly cooler and more humid than in non-infested beds and individuals in the centre of infested mussel beds experienced significantly lower body temperatures. Under a scenario of warming climate, endolithic infestation of mussel beds might thus represent an ecological advantage not only for M. galloprovincialis as a species but also for the associated communities. Fundacao para a Ciencia e Tecnologia (FCT-MEC, Portugal) [UID/Multi/04326/2013, IF/01413/2014/CP1217/CT0004] South African Research Chairs Initiative (SARChI) of the Department of Science and Technology National Research Foundation South African National Research Foundation (NRF) info:eu-repo/semantics/publishedVersion

Geographical discontinuities in the composition of species assemblages are often mirrored by genetic clines and local adaptation in species that extend across such biogeographic disjunction. Species distributed across such biogeographic discontinuities are thus more prone to develop genetic clines and adaptations to environmental variation. The northwest coast of France encompasses a biogeographic boundary between cold- and warm-temperate marine ecosystems. Distinct hydrographic features characterize northern and southern shores, promoting physical oceanographic breaks that can affect dispersal and connectivity between populations. Furthermore, mosaics of contrasting conditions may affect local patterns of species assemblages and potential adaptive pressures. Along this stretch of coast, the intertidal habitat is largely dominated by brown algae of the genus Fucus, key structuring species common along North Atlantic shorelines. Fucus spiralis, F. guiryi and F. vesiculosus are sister species with different mating systems. They have extensive sympatric geographical distributions and distinct but overlapping vertical distributions in the intertidal zone. We examined genetic structure and diversity across this major environmental transition zone using neutral microsatellite markers to investigate patterns of differentiation between populations of each species inhabiting the two different biogeographic regions. We found contrasting results between the dioecious species (F. vesiculosus) and the two selfing hermaphroditic lineages (F. spiralis and F. guiryi). Genetic differentiation of northern and southern populations of F. vesiculosus followed the expectation from a hypothetical oceanographic discontinuity across this biogeographical transition zone, but this was not the case for the two hermaphroditic species. The former had higher genetic diversity and structure along the distribution range investigated whereas selfing hermaphrodites had less variability and were not geographically structured. Our results suggest that genetic patterns are correlated with this biogeographical transition zone but this effect is dependent on mating system, a determining factor affecting population structure. Additionally, other factors such as intertidal elevation may also play a role in observed difference in genetic structuring. Fundacao para a Ciencia e a Tecnologia (FCT-MEC, Portugal) [CCMAR - UID/MULTI/04326/2013, EXCL/AAG-GLO/0661/2012, IF/01413/2014/CP1217/CT0004] info:eu-repo/semantics/publishedVersion

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. Portuguese National Science Foundation (FCT) [EXPL/BIA-BIC/1471/2012]; South Africa Research Chairs Initiative (SARChI) of the Department of Science and Technology info:eu-repo/semantics/publishedVersion

Understanding the extent to which neutral processes and adaptive divergence shape the spatial structure of natural populations is a major goal in evolutionary biology and is especially important for the identification of significant levels of biodiversity. Our results identified replicated habitat-specific (adaptive) phenotypic divergence in the brown macroalga Fucus vesiculosus that is independent of population (neutral) genetic structure. F. vesiculosus inhabits contiguous and contrasting marine to estuarine intertidal habitats. Combining analyses of genetic and phenotypic traits of populations living under differential selective regimes (estuaries and open coast), we investigated levels of neutral genetic differentiation and adaptive physiological responses to emersion stress. In southwest England (SW UK) and northern Iberia (N. Iberia), populations living in estuaries and marine coastal habitats were genetically characterized at six microsatellite loci. In N. Iberia, two clades with limited admixture were recovered, each including one open coast site and the adjacent estuarine location. In contrast, SW UK samples clustered according to habitat and formed three distinct groups of genotypes; one including the two open coast locations and the other two representing each of the estuarine sites. Temperature loggers revealed distinct emersion regimes that characterized each habitat type independently of the region, while water and air temperature profiles showed site-specific trends. Despite acclimation under usual conditions, trait means of emersion stress resilience showed a strong phenotypic divergence between habitats, consistent with environmental clines in exposure time observed in the different habitats. We demonstrate that neutral genetic clusters do not reflect locally adapted population units. Our results identified replicated habitat-specific (adaptive) phenotypic divergence that is independent of population (neutral) genetic structure in F. vesiculosus. The significance of such findings extends beyond the theoretical evolutionary and ecological interest of discovering parallel adaptive responses to the broader implications for conservation of intraspecific biodiversity.

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.

AimGlobal climate change has profound and diverse effects on biological diversity. Identifying present-day climate refugia is an increasingly recognized strategy for the management of biodiversity loss. Such refugia are potential safe havens that enhance environmental diversity by buffering the effects of large scale change, facilitating species persistence at regional scales and conserving unique genetic diversity. Although their ecological effects are well studied, the potential of upwelling centres to act as refugia in a scenario of climate warming remains largely unexplored. Here, we investigate whether upwelled waters act as refugia in a region heavily affected by recent climatic changes.LocationShores of south-western Iberia and north-western Africa.MethodsWe compared changes in distribution of the canopy-forming macroalga Fucus guiryi with sea surface temperature patterns in five upwelling areas and adjacent warmer regions. Population diversity and structure was inferred from genotypic analyses using nine microsatellite loci.ResultsFucus guiryi has disappeared from large expanses of non-upwelling shores, currently persisting in areas characterized by strong upwelling and reduced or non-significant rates of warming. Populations of F. guiryi were more abundant towards upwelling centres. Furthermore, the genetic characterization of populations revealed distinct genetic groups associated with each upwelling system.Main conclusionsWithin a large region of predominantly changing climate, we highlight the fundamental importance of upwelling areas as favourable, comparatively stable climates where F. guiryi has retreated and persists, preserving unique portions of the species' genetic pool.

At a proximal level, the physiological impacts of global climate change on ectothermic organisms are manifest as changes in body temperatures. Especially for plants and animals exposed to direct solar radiation, body temperatures can be substantially different from air temperatures. We deployed biomimetic sensors that approximate the thermal characteristics of intertidal mussels at 71 sites worldwide, from 1998-present. Loggers recorded temperatures at 10–30 min intervals nearly continuously at multiple intertidal elevations. Comparisons against direct measurements of mussel tissue temperature indicated errors of ~2.0–2.5 °C, during daily fluctuations that often exceeded 15°–20 °C. Geographic patterns in thermal stress based on biomimetic logger measurements were generally far more complex than anticipated based only on ‘habitat-level’ measurements of air or sea surface temperature. This unique data set provides an opportunity to link physiological measurements with spatially- and temporally-explicit field observations of body temperature.

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.

Retracing the origins of invasive species is a first critical step in identifying potential mechanisms of introduction, implementation of management strategies and forecasting the spread of the invader. Mytilus galloprovincialis is an intertidal mussel that is widely distributed in many temperate and subtropical regions. It is invasive worldwide and the most successful invasive marine species in southern Africa. Previous studies have examined genetic relationships between a few South African populations from the south-western coast and other worldwide populations, presenting evidence of a north-eastern Atlantic origin of the invasion. Here, a combination of nuclear (Me15/16 PCR-based) and mitochondrial (16S restriction fragment-length polymorphism; 16S RFLP) DNA assays was applied to infer the origin of this strong invader across its entire southern African distribution (South Africa and Namibia). The 16S RFLP confirmed the northern hemisphere as being the likely sole source of invasion. Additionally, the frequencies of haplotypes at the 16S marker and alleles at the Me15/16 locus point to north-eastern Atlantic shores as the most likely origin throughout the Namibian and South African distribution of the species.

Environmental heterogeneity challenges both indigenous species and invaders and can play a defining role in the dynamics of their interactions. We compare bay and open coast habitats to show how environmental heterogeneity and seasonality affect survival and physiological performances of invasive (Mytilus galloprovincialis) and indigenous (Perna perna) intertidal mussels. P. perna had significantly higher attachment strength than M. galloprovincialis. Attachment was strongly correlated with hydrodynamic stress and was lower for both species in bays. Both species had a major spawning event when wave action was weakest. In bays, there was no correlation between gonad index (GI) and attachment strength for either species, but on the open coast GI was negatively correlated with attachment. In bays, maximum GI of M. galloprovincialis was 64% higher than for P. perna, while on the open coast values did not differ between the two. Thus, on the open coast, both species invest more energy in attachment but P. perna can accommodate energetic demands of increased byssal production without altering gonad production, while M. galloprovincialis cannot. Mortality was significantly correlated to sand stress, while the correlation with wave action was very weak in bays and non-significant on the open coast probably because sand stress peaked during periods of low wave action. The success of the invader and thus the outcomes of its interaction with the indigenous species are governed by habitat-to-habitat variability. In this case the invasive species is likely to prove a weaker competitor on the more stressful and energetically demanding open coast.

Understanding population connectivity is fundamental] to ecology, and, for sedentary organisms, connectivity is achieved through larval dispersal. We tested whether coastal topography influences genetic structure in Perna Perna mussels by comparing populations inside bays and on the open coast. Higher hydrodynamic stress on the open coast produces higher mortality and thus genetic turnover. Populations on the open coast had fewer private haplotypes and less genetic endemism than those inside bays. Gene flow analysis showed that bays act as Source populations, with greater migration rates Out. of bays than into them. Differences in genetic structure on scales of 10s of kilometres show that coastal configuration strongly affects selection, larval dispersal and haplotype diversity, Rhodes University; National Research Foundation of South Africa [2069119]; Claude Harris Leon postdoctoral research fellowship Foundation

The ability of an invasive species to spread in a new locality depends on its interaction with the indigenous community and on variation in time and space in the environment. The Mediterranean mussel Mytilus galloprovincialis invaded the South African coast 30 years ago and it now competes and coexists with the indigenous mussel Perna perna. The two species show different tolerances to wave and sand stress, two of the main environmental factors affecting this intertidal community. P. perna is more resistant to hydrodynamic stress than M. galloprovincialis, while the invasive species is less vulnerable to sand action. Our results show that mortality rates of the two species over a period of 6 months had different timing. The indigenous species had higher mortality than M. galloprovincialis during periods of high sand accumulation in mussel beds, while the pattern reversed during winter, when wave action was high. A negative correlation between sand accumulation and attachment strength of the two mussels showed that sand not only affects mussel mortality through scouring and burial, but also weakens their attachment strength, subjecting them to a higher risk of dislodgement. Here we underline the importance of variations in time and space of environmental stress in regulating the interaction between invasive and indigenous species, and how these variations can create new competitive balances.

The amount of genetic structure in marine invertebrates is often thought to be negatively correlated with larval duration. However, larval retention may increase genetic structure in species with long-lived planktonic larvae, and rafting provides a means of dispersal for species that lack a larval dispersal phase. We compared genetic structure, demographic histories and levels of gene flow of regional lineages (in most cases defined by biogeographic region) of five southern African coastal invertebrates with three main types of larval development: (1) dispersal by long-lived planktonic larvae (mudprawn Upogebia africana and brown mussel Perna perna), (2) abbreviated larval development (crown crab Hymenosoma orbiculare) and (3) direct development (estuarine isopod Exosphaeroma hylecoetes and estuarine cumacean Iphinoe truncata). We hypothesized that H. orbiculare, having abbreviated larval development, would employ a strategy of larval retention, resulting in genetic structure comparable to that of the direct developers rather than the planktonic dispersers. However, regional population structure was significantly lower in all species with planktonic larvae, including H. orbiculare, than in the direct developers. Moreover, nested clade analysis identified demographic histories resulting from low levels of gene flow (isolation by distance and allopatric fragmentation) in the direct developers only, and migration rates were significantly higher in all three species having planktonic larvae than in the direct developers. We conclude that the amount of genetic structure within marine biogeographic regions strongly depends on the presence or absence of free-swimming larvae. Whether such larvae are primarily exported or retained, whether they have long or short larval duration, and whether or not they are capable of active dispersal seems to have little effect on connectivity among populations.