Diversity and structure of epibenthic communities across subtidal artificial hard habitats in the Bay of Cherbourg (English Channel).

To inform the performance of ecological engineering designs for artificial structures at sea, it is essential to characterise their impacts on the epibenthic communities colonising them. In this context, the present study aims to compare the community structure among natural and four different artificial hard habitats with different ages and features installed in the Bay of Cherbourg (English Channel): i) cinder blocks and ii) boulders, both installed six years prior to the study, and iii) smooth and iv) rugous concrete dykes, both installed one year prior to this study. Results showed that artificial habitats installed six years ago harboured communities with functional and taxonomic diversity characteristic of mature communities but were still different from those of natural habitat. Conversely, the two dyke habitats installed one year prior to this study presented a poorly diversified community dominated by opportunistic taxa. Furthermore, while the concrete used for the two dyke habitats presented different rugosity properties, both habitats supported similar communities, suggesting that such eco-engineering measures did not affect the settlement of early colonisers. Overall, this study highlights the need for long-term monitoring to comprehensively evaluate epibenthic colonisation of artificial structures.

Evaluation of Sub-Antarctic and Antarctic sea urchins' thermal reaction norm through righting behavior and comparison with in situ measurements.

Sea urchin's survival may depend on their capacity to recover proper orientation rapidly and effectively after inversion, enabling escape from predator and preventing desiccation. This righting behavior has been used as a repeatable and reliable indicator to assess echinoderms performance across environmental conditions, including thermal sensitivity and thermal stress. The current study aims at evaluating and comparing the thermal reaction norm for righting behavior (time for righting (TFR) and capacity to self-right) of three common sea urchins from high latitude, the Patagonian sea urchins Loxechinus albus and Pseudechinus magellanicus, and the Antarctic sea urchin Sterechinus neumayeri. In addition, to infer the ecological implications of our experiments, we compared laboratory-based and in situ TFR of these three species. We observed that populations of the Patagonian sea urchins L. albus and P. magellanicus presented similar trend of righting behavior, overly accelerating with increasing temperature (from 0 to 22°C). Little variations and high inter-individual variability were observed below 6°C in the Antarctic sea urchin TFR, and righting success strongly decreased between 7 and 11°C. For the three species, TFR was lower in in situ experiments compared to the laboratory. Overall, our results suggest that the populations of Patagonian sea urchin exhibit a wide thermal tolerance and, based on S. neumayeri's TFR, aligning with the narrow thermal tolerance of Antarctic benthos. Finally, the differences between laboratory and in situ experiments highlights the importance of considering the complexity of marine environments for future predictions.

Local flexibility in feeding behaviour and contrasting microhabitat use of an omnivore across latitudes.

As the environment is getting warmer and species are redistributed, consumers can be forced to adjust their interactions with available prey, and this could have cascading effects within food webs. To better understand the capacity for foraging flexibility, our study aimed to determine the diet variability of an ectotherm omnivore inhabiting kelp forests, the sea urchin Echinus esculentus, along its entire latitudinal distribution in the northeast Atlantic. Using a combination of gut content and stable isotope analyses, we determined the diet and trophic position of sea urchins at sites in Portugal (42° N), France (49° N), southern Norway (63° N), and northern Norway (70° N), and related these results to the local abundance and distribution of putative food items. With mean estimated trophic levels ranging from 2.4 to 4.6, omnivory and diet varied substantially within and between sites but not across latitudes. Diet composition generally reflected prey availability within epiphyte or understorey assemblages, with local affinities demonstrating that the sea urchin adjusts its foraging to match the small-scale distribution of food items. A net "preference" for epiphytic food sources was found in northern Norway, where understorey food was limited compared to other regions. We conclude that diet change may occur in response to food source redistribution at multiple spatial scales (microhabitats, sites, regions). Across these scales, the way that key consumers alter their foraging in response to food availability can have important implication for food web dynamics and ecosystem functions along current and future environmental gradients.

Non-indigenous species contribute equally to biofouling communities in international vs local ports in the Biobío region, Chile.

Growing coastal urbanization together with the intensification of maritime traffic are major processes explaining the increasing rate of biological introductions in marine environments. To investigate the link between international maritime traffic and the establishment of non-indigenous species (NIS) in coastal areas, biofouling communities in three international and three nearby local ports along 100 km of coastline in south-central Chile were compared using settlement panels and rapid assessment surveys. A larger number of NIS was observed in international ports, as expected in these 'invasion hubs'. However, despite a few environmental differences between international and local ports, the two port categories did not display significant differences regarding NIS establishment and contribution to community structure over the studied period (1.5 years). In international ports, the free space could be a limiting factor for NIS establishment. The results also suggest that local ports should be considered in NIS surveillance programs in Chile.

Canopy interactions and physical stress gradients in subtidal communities.

Species interactions are integral drivers of community structure and can change from competitive to facilitative with increasing environmental stress. In subtidal marine ecosystems, however, interactions along physical stress gradients have seldom been tested. We observed seaweed canopy interactions across depth and latitudinal gradients to test whether light and temperature stress structured interaction patterns. We also quantified interspecific and intraspecific interactions among nine subtidal canopy seaweed species across three continents to examine the general nature of interactions in subtidal systems under low consumer pressure. We reveal that positive and neutral interactions are widespread throughout global seaweed communities and the nature of interactions can change from competitive to facilitative with increasing light stress in shallow marine systems. These findings provide support for the stress gradient hypothesis within subtidal seaweed communities and highlight the importance of canopy interactions for the maintenance of subtidal marine habitats experiencing environmental stress.

Testing the physiological capacity of the mussel Mytilus chilensis to establish into the Southern Ocean.

The Southern Ocean and the Antarctic Circumpolar Current create environmental conditions that serve as an efficient barrier to prevent the colonization of non-native species (NNS) in the marine ecosystems of Antarctica. However, warming of the Southern Ocean and the increasing number of transport opportunities are reducing the physiological and physical barriers, increasing the chances of NNS arriving. The aim of this study was to determine the limits of survival of the juvenile mussels, M. chilensis, under current Antarctic conditions and those projected under climate change. These assessments were used to define the mussels potential for establishment in the Antarctic region. Experimental mussels were exposed to four treatments: -1.5 °C (Antarctic winter), 2 °C (Antarctic summer), 4 °C (Antarctic projected) and 8 °C (control) for 80 days and a combination of physiological and transcriptomics approaches were used to investigate mussel response. The molecular responses of mussels were congruent with the physiological results, revealing tolerance to Antarctic winter temperatures. However, a higher number of regulated differentially expressed gene (DEGs) were reported in mussels exposed to Antarctic winter temperatures (-1.5 °C). This tolerance was associated with the activation of the biological processes associated with apoptosis (up regulated) and both cell division and cilium assembly (down regulated). The reduced feeding rate and the negative scope for growth, for a large part of the exposure period at -1.5 °C, suggests that Antarctic winter temperatures represents an environmental barrier to M. chilensis from the Magellanic region settling in the Antarctic. Although M. chilensis are not robust to current Antarctica thermal conditions, future warming scenarios are likely to weaken these physiological barriers. These results strongly suggest that the West Antarctic Peninsula could become part of Mytilus distributional range, especially with dispersal aided by increasing maritime transport activity across the Southern Ocean.

Predator control of marine communities increases with temperature across 115 degrees of latitude.

Early naturalists suggested that predation intensity increases toward the tropics, affecting fundamental ecological and evolutionary processes by latitude, but empirical support is still limited. Several studies have measured consumption rates across latitude at large scales, with variable results. Moreover, how predation affects prey community composition at such geographic scales remains unknown. Using standardized experiments that spanned 115° of latitude, at 36 nearshore sites along both coasts of the Americas, we found that marine predators have both higher consumption rates and consistently stronger impacts on biomass and species composition of marine invertebrate communities in warmer tropical waters, likely owing to fish predators. Our results provide robust support for a temperature-dependent gradient in interaction strength and have potential implications for how marine ecosystems will respond to ocean warming.

Rapid recovery of native habitat-builders following physical disturbance on pier pilings offsets colonization of cryptogenic and non-indigenous species in a Chilean port.

Examining the effects of disturbances within marine urban communities can shed light on their assembly rules and invasion processes. The effects of physical disturbance, through the removal of dominant native habitat-builders, were investigated in the recolonization of disturbed patches and colonization of plates on pier pilings, in a Chilean port. On pilings, disturbance substantially affected community structure after 3 months, although it slowly converged across treatments after 10 months. On plates, cryptogenic and non-indigenous species richness increased with removal severity, which was not observed in natives. Opportunistic taxa took advantage of colonizing at an early successional stage, illustrating a competition-colonization trade-off, although indirect effects might be at play (e.g. trophic competition or selective predation). Recovery of the habitat-builders then occurred at the expense of cryptogenic and non-indigenous taxa. Whether natives could continue winning against increasing propagule and colonization pressures in marine urban habitats deserves further attention. The interactions between disturbance and biological invasions herein experimentally shown in situ contribute to our understanding of multiple changes imposed by marine urbanization in a growing propagule transport network.

Greater functional similarity in mobile compared to sessile assemblages colonizing artificial coastal habitats.

Among anthropogenic habitats built in the marine environment, floating and non-floating structures can be colonized by distinct assemblages. However, there is little knowledge whether these differences are also reflected in the functional structure. This study compared the functional diversity of sessile and mobile invertebrate assemblages that settle over three months on floating vs. non-floating artificial habitats, in two Chilean ports. Using morphological, trophic, behavioral, and life history traits, we found differences between mobile and sessile assemblages regarding the effect of the type of habitat on the functional diversity. Compared to sessile assemblages, a greater functional similarity was observed for mobile assemblages, which suggests that their dispersal capacity enables them to balance the reduced connectivity between settlement structures. No traits, prevailing or selected in one or the other habitat type, was however clearly identified; a result warranting for further studies focusing on more advanced stages of community development.

First mussel settlement observed in Antarctica reveals the potential for future invasions.

Global biodiversity is both declining and being redistributed in response to multiple drivers characterizing the Anthropocene, including synergies between biological invasions and climate change. The Antarctic marine benthos may constitute the last biogeographic realm where barriers (oceanographic currents, climatic gradients) have not yet been broken. Here we report the successful settlement of a cohort of Mytilus cf. platensis in a shallow subtidal habitat of the South Shetland Islands in 2019, which demonstrates the ability of this species to complete its early life stages in this extreme environment. Genetic analyses and shipping records show that this observation is consistent with the dominant vectors and pathways linking southern Patagonia with the Antarctic Peninsula and demonstrates the potential for impending invasions of Antarctic ecosystems.

Habitat formation prevails over predation in influencing fouling communities.

Coastal human-made structures, such as marinas and harbors, are expanding worldwide. Species assemblages described from these artificial habitats are novel relative to natural reefs, particularly in terms of the abundance of nonindigenous species (NIS). Although these fouling assemblages are clearly distinctive, the ecosystem functioning and species interactions taking place there are little understood. For instance, large predators may influence the fouling community development either directly (feeding on sessile fauna) or indirectly (feeding on small predators associated with these assemblages). In addition, by providing refuges, habitat complexity may modify the outcome of species interactions and the extent of biotic resistance (e.g., by increasing the abundance of niche-specific competitors and predators of NIS). Using experimental settlement panels deployed in the field for 2.5 months, we tested the influence of predation (i.e., caging experiment), artificial structural complexity (i.e., mimics of turf-forming species), and their interactions (i.e., refuge effects) on the development of sessile and mobile fauna in two marinas. In addition, we tested the role of biotic complexity-arising from the habitat-forming species that grew on the panels during the trial-on the richness and abundance of mobile fauna. The effect of predation and artificial habitat complexity was negligible, regardless of assemblage status (i.e., native, cryptogenic, and nonindigenous). Conversely, habitat-forming species and associated epibionts, responsible for biotic complexity, had a significant effect on mobile invertebrates (richness, abundance, and community structure). In particular, the richness and abundance of mobile NIS were positively affected by biotic complexity, with site-dependent relationships. Altogether, our results indicate that biotic complexity prevails over artificial habitat complexity in determining the distribution of mobile species under low predation pressure. Facilitation of native and non-native species thus seems to act upon diversity and community development: This process deserves further consideration in models of biotic resistance to invasion in urban marine habitats.

Presence of the tunicate Asterocarpa humilis on ship hulls and aquaculture facilities in the coast of the Biobío Region, south central Chile.

Non-native ascidians are important members of the fouling community associated with artificial substrata and man-made structures. Being efficient fouling species, they are easily spread by human-mediated transports (e.g., with aquaculture trade and maritime transports). This is exemplified by the ascidian Asterocarpa humilis which displays a wide distribution in the Southern Hemisphere and has been recently reported in the Northern Hemisphere (NW Europe). In continental Chile, its first report dates back from 2000 for the locality of Antofagasta (23°S). Although there was no evidence about the vectors of introduction and spread, nor the source, some authors suggested maritime transport by ship hulls and aquaculture devices as putative introduction pathways and vectors. In the present study, we report for the first time the presence of A. humilis on the hull of an international ship in a commercial port in Concepción bay (36°S), south central Chile. We also found one individual associated to a seashell farm, 70 km far from Concepción bay. Further individuals were subsequently identified within Concepción bay: one juvenile settled upon international harbor pilings and a dozen individuals along aquaculture seashell longlines. For the first specimens sampled, species identification was ascertained using both morphological criteria and molecular barcoding, using the mitochondrial gene cytochrome c oxidase subunit I (COI) and a nuclear gene (ribosomal RNA 18S). The nuclear 18S gene and the mitochondrial gene COI clearly assigned the specimens to A. humilis, confirming our morphological identification. Two haplotypes were obtained with COI corresponding to haplotypes previously obtained with European and Northern Chilean specimens. The present study thus reports for the first time the presence of A. humilis in the Araucanian ecoregion, documenting the apparent expansion of this non-native tunicate in Chile over 2,000 km, spanning over three ecoregions. In addition we reveal the potential implication of the international maritime transport as a vector of spread of this species along the Eastern Pacific coast, and the putative role of aquaculture facilities in promoting local establishments of non-native tunicates.