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There has been an increase in commercial-scale kelp cultivation in Europe, with fouling of cultivated kelp fronds presenting a major challenge to the growth and development of the industry. The presence of epibionts decreases productivity and impacts the commercial value of the crop. Several abiotic and biotic factors may influence the occurrence and degree of fouling of wild and cultivated fronds. Using a commercial kelp farm on the SW coast of Ireland, we studied the development of fouling communities on cultivated Alaria esculenta fronds over 2 typical growing seasons. The predictability of community development was assessed by comparing mean occurrence-day. Hypotheses that depth, kelp biomass, position within the farm and the hydrodynamic environment affect the fouling communities were tested using species richness and community composition. Artificial kelp mimics were used to test whether local frond density could affect the fouling communities. Species richness increased over time during both years, and species composition was consistent over years with early successional communities converging into later communities (no significant differences between June 2014 and June 2015 communities, ANOSIM; R = -0.184, p > 0.05). The timing of species occurrences was predictable across years for all shared species. Variations in biomass, depth and position within the farm had no significant effect on species richness and composition. Results from artificial kelp mimics suggest possible hydrodynamic effects. The ability to understand succession and the timing of occurrences of fouling organisms and predict their arrival has significant benefits for the seaweed cultivation industry.

The continued development of the aquaculture industry is contributing to the proliferation of artificial structures in the marine environment. Observations of seaweed farms (infrastructure and biomass) suggest they act as a habitat for associated species. Seaweed farms differ from other forms of artificial infrastructure as the material deployed already has marine organisms (i.e. culture species) growing on it. This 'priming' of ropes with juvenile sporophytes may affect future development of communities by facilitating colonizing species or suppressing competitors. We call this process 'ecological priming': the provision of a biological platform that influences the successional development of specific communities. The communities that developed on ropes primed with Alaria esculenta individuals were compared with unprimed ropes to assess the ecological priming effect, at a commercial kelp farm in south-west Ireland. Species richness increased over two cultivation seasons and species composition was consistent between years, with distinct communities developing on primed and unprimed treatments. Timing of species occurrence on primed ropes was predictable with no predictable pattern occurring on unprimed ropes. Multivariate tests indicated distinct communities between treatments, with suppression of other algal species and potential facilitation of some species that have a particular association with A. esculenta on primed ropes. Communities from primed holdfasts contained a lower diversity of algal species compared with unprimed communities. Cultivated kelp holdfasts represent a habitat for distinct assemblages that reflect ecological priming of the substratum. [ABSTRACT FROM AUTHOR]

In degraded urban habitats, nature-based solutions aim to enhance ecosystem functioning and service provision. Bivalves are increasingly reintroduced to urban environments to enhance water quality through biofiltration, yet their long-term sustainability remains uncertain. Following the restoration of the disused South Docks in Liverpool in the 1980s, natural colonization of mussels rapidly improved dock-basin water quality and supported diverse taxa, including other filter feeders. While the initial colonization phase has been well documented, there has been limited published research since the mid-1990s, despite ongoing routine water quality monitoring. Here, we assessed the long-term persistence of mussel populations, their associated biodiversity, and physico-chemical parameters of the water in Queens and Albert Docks by comparing historical (1980s to 1990s) and contemporary data from follow-up surveys (2012,2022). Following an initial period of poor water quality (high contamination and turbidity, low oxygen), the natural colonization of mussels from Albert Dock in 1988 extended throughout the South Docks. By the mid-1990s, the environment of the South Docks and its mussel populations had stabilized. The dock walls were dominated by mussels which provided important complex secondary substrate for invertebrates and macroalgae. Surveys conducted in 2012 and 2022 confirmed the continued dominance of mussels and estimates of mussel biofiltration rates confirm that mussels are continuing to contribute to maintaining water quality. A decline in salinity was observed in both docks in 2022, with evidence of recovery. While these ecosystems appear relatively stable, careful management of the hydrological regime is crucial to ensuring the persistence of mussels and resilient ecosystem service provision through biofiltration., 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.)

The success of non-native species (NNS) invasions depends on patterns of dispersal and connectivity, which underpin genetic diversity, population establishment and growth. In the marine environment, both global environmental change and increasing anthropogenic activity can alter hydrodynamic patterns, leading to significant inter-annual variability in dispersal pathways. Despite this, multi-generational dispersal is rarely explicitly considered in attempts to understand NNS spread or in the design of management interventions. Here, we present a novel approach to quantifying species spread that considers range expansion and network formation across time using the non-native Pacific oyster, Magallana gigas (Thunberg 1793), as a model. We combined biophysical modelling, dynamic patch occupancy models, consideration of environmental factors, and graph network theory to model multi-generational dispersal in northwest Europe over 13 generations. Results revealed that M. gigas has a capacity for rapid range expansion through the creation of an ecological network of dispersal pathways that remains stable through time. Maximum network size was achieved in four generations, after which connectivity patterns remained temporally stable. Multi-generational connectivity could therefore be divided into two periods: network growth (2000-2003) and network stability (2004-2012). Our study is the first to examine how dispersal trajectories affect the temporal stability of ecological networks across biogeographic scales, and provides an approach for the assignment of site-based prioritisation of non-native species management at different stages of the invasion timeline. More broadly, the framework we present can be applied to other fields (e.g. Marine Protected Area design, management of threatened species and species range expansion due to climate change) as a means of characterising and defining ecological network structure, functioning and stability., Competing Interests: Declaration of competing interest Authors declare no known 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., (Crown Copyright © 2024. Published by Elsevier B.V. All rights reserved.)

In response to ongoing coastal urbanization, it is critical to develop effective methods to improve the biodiversity and ecological sustainability of artificial shorelines. Enhancing the topographic complexity of coastal infrastructure through the mimicry of natural substrata may facilitate the establishment of ecosystem engineering species and associated biogenic habitat formation. However, interactions between ecosystem engineers and their substratum are likely determined by organismal size and resource needs, thus making responses to topography highly scale-dependent. Here, we assessed the topographic properties (rugosity, surface area, micro-surface orientations) that underpin the abundance and distribution of two ecosystem engineers (fucoids, limpets) across six spatial scales (1-500 mm). Furthermore, we assessed the 'biogenic' rugosity created by barnacle matrices across fine scales (1-20 mm). Field surveys and 3D scanning, conducted across natural and artificial substrata, showed major effects of rugosity and associated topographic variables on ecosystem engineer assemblages and spatial occupancy, while additional abiotic environmental factors (compass direction, wave exposure) and biotic associations only had weak influences. Natural substrata exhibited ≤67 % higher rugosity than artificial ones. Fucoid-covered patches were predominantly associated with high-rugosity substrata and horizontal micro-surfaces, while homescars of limpets (≥15 mm shell length) predominated on smoother substratum patches. Barnacle-driven rugosity homogenized substrata at scales ≤10 mm. Our findings suggest that scale-dependent rugosity is a key driver of fucoid habitat formation and limpet habitat use, with wider eco-engineering applications for mimicking ecologically impactful topography on coastal infrastructure., 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 Author(s). Published by Elsevier B.V. All rights reserved.)

Topographic complexity is often considered to be closely associated with habitat complexity and niche diversity; however, complex topography per se does not imply habitat suitability. Rather, ecologically suitable habitats may emerge if topographic features interact with environmental factors and thereby alter their surrounding microenvironment to the benefit of local organisms (e.g., resource provisioning, stress mitigation). Topography may thus act as a key modulator of abiotic stressors and biotic pressures, particularly in environmentally challenging intertidal systems. Here, we review how topography can alter microhabitat conditions with respect to four resources required by intertidal organisms: a source of energy (light, suspended food particles, prey, detritus), water (hydration, buffering of light, temperature and hydrodynamics), shelter (temperature, wave exposure, predation), and habitat space (substratum area, propagule settlement, movement). We synthesize mechanisms and quantitative findings of how environmental factors can be altered through topography and suggest an organism-centered 'form-follows-ecological-function' approach to designing multifunctional marine infrastructure., 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.)

Background and Aims: Worldwide, invasive species are spreading through marine systems at an unprecedented rate with both positive and negative consequences for ecosystems and the biological functioning of organisms. Human activities from shipping to habitat damage and modification are known vectors of spread, although biological interactions including epibiosis are increasingly recognized as potentially important to introduction into susceptible habitats., Methods: We assessed a novel mechanism of spread - limpets as transporters of an invasive alga, Sargassum muticum, into beds of the seagrass Zostera marina - and the physiological impact of its invasion. The association of S. muticum with three limpet species and other habitats was assessed using intertidal surveys on rocky shores and snorkelling at two seagrass sites in the UK. A 4-year field study tested the effect of S. muticum on Z. marina shoot density, dry weight and phenolic compounds (caffeic and tannic acid) content, and a laboratory experiment tested the impact of S. muticum on nutrient partitioning (C/H/N/P/Si), photosynthetic efficiency (Fv/Fm) and growth of Z. marina., Results: On rocky shores 15 % of S. muticum occurrences were attached to the shells of live limpets. In seagrass beds 5 % of S. muticum occurrences were attached to the shells of dead limpets. The remainder were attached to rock, to cobblestones, to the seagrass matrix or embedded within the sand. Z. marina density and phenolics content was lower when S. muticum co-occurred with it. Over 3 years, photosynthetic responses of Z. marina to S. muticum were idiosyncratic, and S. muticum had no effect on nutrient partitioning in Z. marina., Conclusions: Our results show limpets support S. muticum as an epibiont and may act as a previously unreported transport mechanism introducing invaders into sensitive habitats. S. muticum reduced production of phenolics in Z. marina, which may weaken its defensive capabilities and facilitate proliferation of S. muticum. The effect of S. muticum on Z. marina photosynthesis requires further work but having no effect on the capacity of Z. marina to sequester nutrients suggests a degree of resilience to this invader., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Background: Marine macroalgae ('seaweeds') are a diverse and globally distributed group of photosynthetic organisms that together generate considerable primary productivity, provide an array of different habitats for other organisms, and contribute many important ecosystem functions and services. As a result of continued anthropogenic stress on marine systems, many macroalgal species and habitats face an uncertain future, risking their vital contribution to global productivity and ecosystem service provision., Scope: After briefly considering the remarkable taxonomy and ecological distribution of marine macroalgae, we review how the threats posed by a combination of anthropogenically induced stressors affect seaweed species and communities. From there we highlight five critical avenues for further research to explore (long-term monitoring, use of functional traits, focus on early ontogeny, biotic interactions and impact of marine litter on coastal vegetation)., Conclusions: Although there are considerable parallels with terrestrial vascular plant responses to the many threats posed by anthropogenic stressors, we note that the impacts of some (e.g. habitat loss) are much less keenly felt in the oceans than on land. Nevertheless, and in common with terrestrial plant communities, the impact of climate change will inevitably be the most pernicious threat to the future persistence of seaweed species, communities and service provision. While understanding macroalgal responses to simultaneous environmental stressors is inevitably a complex exercise, our attempt to highlight synergies with terrestrial systems, and provide five future research priorities to elucidate some of the important trends and mechanisms of response, may yet offer some small contribution to this goal., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Background and Aims: Marine macroalgae ('seaweeds') are critical to coastal ecosystem structure and function, but also vulnerable to the many environmental changes associated with anthropogenic climate change (ACC). The local habitat conditions underpinning observed and predicted ACC-driven changes in intertidal macroalgal communities are complex and probably site-specific and operate in addition to more commonly reported regional factors such as sea surface temperatures., Methods: We examined how the composition and functional trait expression of macroalgal communities in SW England varied with aspect (i.e. north-south orientation) at four sites with opposing Equator- (EF) and Pole-facing (PF) surfaces. Previous work at these sites had established that average annual (low tide) temperatures vary by 1.6 °C and that EF-surfaces experience six-fold more frequent extremes (i.e. >30 °C)., Key Results: PF macroalgal communities were consistently more taxon rich; 11 taxa were unique to PF habitats, with only one restricted to EF. Likewise, functional richness and dispersion were greater on PF-surfaces (dominated by algae with traits linked to rapid resource capture and utilization, but low desiccation tolerance), although differences in both taxon and functional richness were probably driven by the fact that less diverse EF-surfaces were dominated by desiccation-tolerant fucoids., Conclusions: Although we cannot disentangle the influence of temperature variation on algal ecophysiology from the indirect effects of aspect on species interactions (niche pre-emption, competition, grazing, etc.), our study system provides an excellent model for understanding how environmental variation at local scales affects community composition and functioning. By virtue of enhanced taxonomic diversity, PF-aspects supported higher functional diversity and, consequently, greater effective functional redundancy. These differences may imbue PF-aspects with resilience against environmental perturbation, but if predicted increases in global temperatures are realized, some PF-sites may shift to a depauperate, desiccation-tolerant seaweed community with a concomitant loss of functional diversity and redundancy., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Habitat complexity is widely considered an important determinant of biodiversity, and enhancing complexity can play a key role in restoring degraded habitats. However, the effects of habitat complexity on ecosystem functioning - as opposed to biodiversity and community structure - are relatively poorly understood for artificial habitats, which dominate many coastlines. With Greening of Grey Infrastructure (GGI) approaches, or eco-engineering, increasingly being applied around the globe, it is important to understand the effects that modifying habitat complexity has on both biodiversity and ecological functioning in these highly modified habitats. We assessed how manipulating physical (primary substrate) and/or biogenic habitat (bivalves) complexity on intertidal artificial substrata affected filtration rates, net and gross primary productivity (NPP and GPP, respectively) and community respiration (CR) - as well as abundance of filter feeders and macro-algae and habitat use by cryptobenthic fish across six locations in three continents. We manipulated both physical and biogenic complexity using 1) flat or ridged (2.5 cm or 5 cm) settlement tiles that were either 2) unseeded or seeded with oysters or mussels. Across all locations, increasing physical and biogenic complexity (5 cm seeded tiles) had a significant effect on most ecological functioning variables, increasing overall filtration rates and community respiration of the assemblages on tiles but decreasing productivity (both GPP and NPP) across all locations. There were no overall effects of increasing either type of habitat complexity on cryptobenthic fish MaxN, total time in frame or macro-algal cover. Within each location, there were marked differences in the effects of habitat complexity. In Hobart, we found higher filtration, filter feeder biomass and community respiration on 5 cm tiles compared to flat tiles. However, at this location, both macro-algae cover and GPP decreased with increasing physical complexity. Similarly in Dublin, filtration, filter feeder biomass and community respiration were higher on 5 cm tiles compared to less complex tiles. In Sydney, filtration and filter feeder biomass were higher on seeded than unseeded tiles, and fish MaxN was higher on 5 cm tiles compared to flat tiles. On unseeded tiles in Sydney, filter feeder biomass also increased with increasing physical complexity. Our findings suggest that GGI solutions via increased habitat complexity are likely to have trade-offs among potentially desired functions, such as productivity and filtration rates, and variable effects on cryptobenthic fish communities. Importantly, our results show that the effects of GGI practices can vary markedly according to the environmental context and therefore should not be blindly and uniformly applied across the globe., 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. We only found two fish cryptobenthic species in Arraial do Cabo (Scartella cristata and Parablennius pilicornis,Table S17) and one in Ravenna (Parablennius incognitus,Table S18). The most abundant species in Arraial do Cabo, Brazil, was the molly miller blenny Scartella cristata (Table S17). Contrary to Sydney, there were no effects of complexity (physical or biogenic) on MaxN in either Arraial do Cabo (Fig. 5b–Table 5; Table S14) or Ravenna (Fig. 5c–Table 5; Table S14). Similar to Sydney, there was no effect of physical or biogenic complexity on the total time in frame (Fig. 5e and f; Table 5; Table S15)., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Thousands of artificial ('human-made') structures are present in the marine environment, many at or approaching end-of-life and requiring urgent decisions regarding their decommissioning. No consensus has been reached on which decommissioning option(s) result in optimal environmental and societal outcomes, in part, owing to a paucity of evidence from real-world decommissioning case studies. To address this significant challenge, we asked a worldwide panel of scientists to provide their expert opinion. They were asked to identify and characterise the ecosystem effects of artificial structures in the sea, their causes and consequences, and to identify which, if any, should be retained following decommissioning. Experts considered that most of the pressures driving ecological and societal effects from marine artificial structures (MAS) were of medium severity, occur frequently, and are dependent on spatial scale with local-scale effects of greater magnitude than regional effects. The duration of many effects following decommissioning were considered to be relatively short, in the order of days. Overall, environmental effects of structures were considered marginally undesirable, while societal effects marginally desirable. Experts therefore indicated that any decision to leave MAS in place at end-of-life to be more beneficial to society than the natural environment. However, some individual environmental effects were considered desirable and worthy of retention, especially in certain geographic locations, where structures can support improved trophic linkages, increases in tourism, habitat provision, and population size, and provide stability in population dynamics. The expert analysis consensus that the effects of MAS are both negative and positive for the environment and society, gives no strong support for policy change whether removal or retention is favoured until further empirical evidence is available to justify change to the status quo. The combination of desirable and undesirable effects associated with MAS present a significant challenge for policy- and decision-makers in their justification to implement decommissioning options. Decisions may need to be decided on a case-by-case basis accounting for the trade-off in costs and benefits at a local level., Competing Interests: Declaration of competing interest The authors declare the following financial interests/personal relationships which may be considered as potential competing interests:Antony Knights reports financial support was provided by Natural Environment Research Council. Anaelle Lemasson reports financial support was provided by Natural Environment Research Council. Paul Somerfield reports financial support was provided by Natural Environment Research Council., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Switching from fossil fuels to renewable energy is key to international energy transition efforts and the move toward net zero. For many nations, this requires decommissioning of hundreds of oil and gas infrastructure in the marine environment. Current international, regional and national legislation largely dictates that structures must be completely removed at end-of-life although, increasingly, alternative decommissioning options are being promoted and implemented. Yet, a paucity of real-world case studies describing the impacts of decommissioning on the environment make decision-making with respect to which option(s) might be optimal for meeting international and regional strategic environmental targets challenging. To address this gap, we draw together international expertise and judgment from marine environmental scientists on marine artificial structures as an alternative source of evidence that explores how different decommissioning options might ameliorate pressures that drive environmental status toward (or away) from environmental objectives. Synthesis reveals that for 37 United Nations and Oslo-Paris Commissions (OSPAR) global and regional environmental targets, experts consider repurposing or abandoning individual structures, or abandoning multiple structures across a region, as the options that would most strongly contribute toward targets. This collective view suggests complete removal may not be best for the environment or society. However, different decommissioning options act in different ways and make variable contributions toward environmental targets, such that policy makers and managers would likely need to prioritise some targets over others considering political, social, economic, and ecological contexts. Current policy may not result in optimal outcomes for the environment or society., (Copyright © 2023 The Authors. Published by Elsevier Ltd.. All rights reserved.)

In the marine environment, natural reef habitats are amongst the most threatened by human activities. Although reef-based ecotourism can benefit local economies, dive tourism can damage sensitive habitats. One solution to managing conflicts between the economic value of diving and its ecological threats is the deployment of artificial reefs near popular dive sites. We surveyed recreational divers to assess divers' use, preference, and perceptions of diving artificial versus natural sites. We found that more divers prefer to dive in natural than artificial habitats, with associated biodiversity the most popular reason for preferring natural habitats, and appreciating shipwrecks the most popular reason for preferring artificial ones. Despite our sample population being highly educated and experienced, predominantly European divers, only 49 % of them perceived artificial reefs as important or somewhat important for diverting pressure from sensitive natural habitats. Similarly, only 13 % of respondents exhibited preference to avoid coral reefs to protect them. These results highlight the fact that more needs to be done to educate divers about the potential importance of artificial habitats in diverting divers from natural reefs. We suggest encouraging divers to switch out a proportion of their dives in vulnerable natural sites for artificial reefs. This is not only true for coral reefs, but should be applied to other natural reef habitats that are popular with divers such as kelp forests, sponge gardens and serpulid and coralligenous reefs. We hope that this study will provide a platform to stimulate a diver-led discussion and campaign for increased uptake of artificial reef use, resulting in reduced impacts on natural reefs., 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 B.V. All rights reserved.)

Worldwide, natural habitats are being replaced by artificial structures due to urbanisation. Planning of such modifications should strive for environmental net gain that benefits biodiversity and ecosystems. Alpha (α) and gamma (γ) diversity are often used to assess 'impact' but are insensitive metrics. We test several diversity measures across two spatial scales to compare species diversity in natural and artificial habitats. We show γ-diversity indicates equivalency in biodiversity between natural and artificial habitats, but natural habitats support greater taxon (α) and functional richness. Within-site β-diversity was also greater in natural habitats, but among-site β-diversity was greater in artificial habitats, contradicting the commonly held view that urban ecosystems are more biologically homogenous than natural ecosystems. This study suggests artificial habitats may in fact provide novel habitat for biodiversity, challenges the applicability of the urban homogenisation concept and highlights a significant limitation of using just α-diversity (i.e., multiple metrics are needed and recommended) for assessing environmental net gain and attaining biodiversity conservation goals., 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.)

Non-native species are spreading at an unprecedented rate over large spatial scales, with global environmental change and growth in commerce providing novel opportunities for range expansion. Assessing the pattern and rate of spread is key to the development of strategies for safeguarding against future invasions and efficiently managing existing ones. Such assessments often depend on spatial distribution data from online repositories, which can be spatially biased, imprecise, and lacking in quantity. Here, the influence of disparities between occurrence records from online data repositories and what is known of the invasion history from peer-reviewed published literature on non-native species range expansion was evaluated using 6693 records of the Pacific oyster, Magallana gigas (Thunberg, 1793), spanning 56 years of its invasion in Europe. Two measures of spread were calculated: maximum rate of spread (distance from introduction site over time) and accumulated area (spatial expansion). Results suggest that despite discrepancies between online and peer-reviewed data sources, including a paucity of records from the early invasion history in online repositories, the use of either source does not result in significantly different estimates of spread. Our study significantly improves our understanding of the European distribution of M. gigas and suggests that a combination of short- and long-range dispersal drives range expansions. More widely, our approach provides a framework for comparison of online occurrence records and invasion histories as documented in the peer-reviewed literature, allowing critical evaluation of both data sources and improving our understanding of invasion dynamics significantly., 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 B.V. All rights reserved.)

Epibiosis is increasingly considered a survival strategy in space-limited environments. However, epibionts can create a new interface between its host, environment and potential predators which may alter predator-prey relationships and biological functioning. Ex-situ experiments investigated the potential costs and benefits of epibiont barnacles on mortality and feeding rate of the mussel, Mytilus edulis, and its predator, the whelk Nucella lapillus. Mussels with living epibiont barnacles suffered no mortality from whelk predation, but when barnacles were absent, mortality was ∼21% over 48 days. Further comparisons revealed the structural complexity of barnacles provided mussels with protection from whelk predation, while the presence of living barnacles increased predator-prey encounters but led to predators targeting barnacles over mussels. Feeding trials revealed feeding rate increased by ∼24% in mussels with living epibionts over mussels with dead or without epibionts, indicating potential costs of hosting epibionts. Our results show that epibionts provide important associational resistance for mussels against whelk predation but a potential cost to the mussel of hosting epibionts requiring increased energy acquisition. These findings advance our understanding of associational resistance derived from epibionts and serve to highlight the potential trade-offs affecting basibiont functioning while showing the importance of positive ecological interactions in ecosystem structure and functioning., 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.)

Distributional shifts in species ranges provide critical evidence of ecological responses to climate change. Assessments of climate-driven changes typically focus on broad-scale range shifts (e.g. poleward or upward), with ecological consequences at regional and local scales commonly overlooked. While these changes are informative for species presenting continuous geographic ranges, many species have discontinuous distributions-both natural (e.g. mountain or coastal species) or human-induced (e.g. species inhabiting fragmented landscapes)-where within-range changes can be significant. Here, we use an ecosystem engineer species (Sabellaria alveolata) with a naturally fragmented distribution as a case study to assess climate-driven changes in within-range occupancy across its entire global distribution. To this end, we applied landscape ecology metrics to outputs from species distribution modelling (SDM) in a novel unified framework. SDM predicted a 27.5% overall increase in the area of potentially suitable habitat under RCP 4.5 by 2050, which taken in isolation would have led to the classification of the species as a climate change winner. SDM further revealed that the latitudinal range is predicted to shrink because of decreased habitat suitability in the equatorward part of the range, not compensated by a poleward expansion. The use of landscape ecology metrics provided additional insights by identifying regions that are predicted to become increasingly fragmented in the future, potentially increasing extirpation risk by jeopardising metapopulation dynamics. This increased range fragmentation could have dramatic consequences for ecosystem structure and functioning. Importantly, the proposed framework-which brings together SDM and landscape metrics-can be widely used to study currently overlooked climate-driven changes in species internal range structure, without requiring detailed empirical knowledge of the modelled species. This approach represents an important advancement beyond predictive envelope approaches and could reveal itself as paramount for managers whose spatial scale of action usually ranges from local to regional., (© 2022 John Wiley & Sons Ltd.)

Marine rocky intertidal organisms are amongst those most affected by climate change with regional distributional changes observed for many species. Although often ascribed to increased sea surface temperatures, precise assessment of the local habitat conditions underpinning observed and predicted changes in community assembly is lacking. Here we examine how aspect (i.e. north-south orientation) affects intertidal community composition and how rock surface temperatures and stress responses of two dominant grazer species (Patella spp.) elucidate emergent differences. We quantified year-round temperature variation and surveyed intertidal community composition on paired natural rock gullies with Equator- (EF) and Pole-facing (PF) surfaces. We also investigated variation in limpet (Patella spp.) reproductive phenology and osmotic stress. Average annual temperatures were 0.8 °C (1.6 °C at low tide) higher, with six-fold more frequent extremes (i.e. > 30 °C) on EF than PF surfaces. Intertidal community composition varied with aspect across trophic levels with greater overall species richness, abundance of primary producers and grazers on PF-surfaces, and greater barnacle abundance on EF-surfaces. Although species richness of organisms from different biogeographical origins ('Boreal' or 'Lusitanian') did not vary, the Lusitanian limpet Patella depressa exhibited earlier reproductive development on EF-surfaces and both limpet species exhibited greater thermal stress on EF-surfaces. We argue that our study system provides a good model for understanding how temperature variation at local scales can affect community composition, as well as ecophysiological and ecological responses to climate change and so better inform and predict regional range shifts over coming decades., (Copyright © 2021. Published by Elsevier Ltd.)

A complex interplay of biotic and abiotic factors underpins the distribution of species and operates across different levels of biological organization and life history stages. Understanding ecosystem engineer reproductive traits is critical for comprehending and managing the biodiversity-rich habitats they create. Little is known about how the reproduction of the reef-forming worm, Sabellaria alveolata, varies across environmental gradients. By integrating broad-scale environmental data with in-situ physiological data in the form of biochemical traits, we identified and ranked the drivers of intraspecific reproductive trait variability (ITV). ITV was highest in locations with variable environmental conditions, subjected to fluctuating temperature and hydrodynamic conditions. Our trait selection pointed to poleward sites being the most physiologically stressful, with low numbers of irregularly shaped eggs suggesting potentially reduced reproductive success. Centre-range individuals allocated the most energy to reproduction, with the highest number of intermediate-sized eggs, whilst equatorward sites were the least physiologically stressful, thus confirming the warm-adapted nature of our model organism. Variation in total egg diameter and relative fecundity were influenced by a combination of environmental conditions, which changed depending on the trait and sampling period. An integrated approach involving biochemical and reproductive traits is essential for understanding macro-scale patterns in the face of anthropogenic-induced climate change across environmental and latitudinal gradients., (© 2021. The Author(s).)

Data that can be used to monitor biodiversity through time are essential for conservation and management. The reef-forming worm, Sabellaria alveolata (L. 1767) is currently classed as 'Data Deficient' due to an imbalance in the spread of data on its distribution. Little is known about the distribution of this species around Ireland. Using data archaeology, we collated past and present distribution records and discovered that S. alveolata has a discontinuous distribution with large gaps between populations. Many regions lack data and should be targeted for sampling. Biodiversity surveys revealed that S. alveolata supported diverse epibiotic algal communities. Retrograding (declining) reefs supported greater infaunal diversity than prograding (growing) reefs or sand, suggesting that S. alveolata is a dynamic ecosystem engineer that has a lasting legacy effect. Similar research should be carried out for other Data Deficient species, habitats and regions. Such data are invaluable resources for management and conservation., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

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.)

Urban and periurban ocean developments impact 1.5% of the global exclusive economic zones, and the demand for ocean space and resources is increasing. As we strive for a more sustainable future, it is imperative that we better design, manage, and conserve urban ocean spaces for both humans and nature. We identify three key objectives for more sustainable urban oceans: reduction of urban pressures, protection and restoration of ocean ecosystems, and support of critical ecosystem services. We describe an array of emerging evidence-based approaches, including greening grayinfrastructure, restoring habitats, and developing biotechnologies. We then explore new economic instruments and incentives for supporting these new approaches and evaluate their feasibility in delivering these objectives. Several of these tools have the potential to help bring nature back to the urban ocean while also addressing some of the critical needs of urban societies, such as climate adaptation, seafood production, clean water, and recreation, providing both human and environmental benefits in some of our most impacted ocean spaces.

Marine heatwaves (MHWs) are an emerging threat to marine organisms that have increased in frequency and magnitude in the past decade. These extreme heating events can have differential impacts on organisms with some experiencing mortality while others survive. Here, we experimentally exposed two species of subtidal gastropod (Trochus sacellum and Astralium haematragum) to two realistic intensities of MHW to test the ability of different species to physiologically cope with extreme heating events. Extreme MHW conditions caused 100% mortality in both species within five days. While both species survived under moderate MHW conditions they showed evidence of nonadaptive metabolic depression. Both species demonstrated an inability to upregulate their metabolic rates at the higher temperatures following exposure to a MHW (i.e. reduced temperature of maximum metabolic rate; T MMR ), suggesting a lack of molecular protective responses and ongoing physiological damage. Therefore, the physiological damage endured by heatwave survivors may lessen their ability to cope with subsequent stress until fully recovered. Repairing this damage may have serious repercussions for the rate of recovery of these normally resilient species and their ability to maintain their ecological functions post MHW, especially under the predicted increasing frequency, duration and magnitude of MHWs., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Background: Under the threat of climate change populations can disperse, acclimatise or evolve in order to avoid fitness loss. In light of this, it is important to understand neutral gene flow patterns as a measure of dispersal potential, but also adaptive genetic variation as a measure of evolutionary potential. In order to assess genetic variation and how this relates to environment in the honeycomb worm (Sabellaria alveolata (L.)), a reef-building polychaete that supports high biodiversity, we carried out RAD sequencing using individuals from along its complete latitudinal range. Patterns of neutral population genetic structure were compared to larval dispersal as predicted by ocean circulation modelling, and outlier analyses and genotype-environment association tests were used to attempt to identify loci under selection in relation to local temperature data., Results: We genotyped 482 filtered SNPs, from 68 individuals across nine sites, 27 of which were identified as outliers using BAYESCAN and ARLEQUIN. All outlier loci were potentially under balancing selection, despite previous evidence of local adaptation in the system. Limited gene flow was observed among reef-sites (F ST  = 0.28 ± 0.10), in line with the low dispersal potential identified by the larval dispersal models. The North Atlantic reef emerged as a distinct population and this was linked to high local larval retention and the effect of the North Atlantic Current on dispersal., Conclusions: As an isolated population, with limited potential for natural genetic or demographic augmentation from other reefs, the North Atlantic site warrants conservation attention in order to preserve not only this species, but above all the crucial functional ecological roles that are associated with their bioconstructions. Our study highlights the utility of using seascape genomics to identify populations of conservation concern.

For much of the 20th century, the Mersey in North West England was one of the worst polluted estuaries in Europe. Water from a range of polluting industries plus domestic sewage was discharged into the Mersey Catchment and Estuary. Recovery came through a concerted clean-up campaign and tightening environmental regulations, partly driven by European Commission Directives, coupled with de-industrialisation from the 1970s onward. Recovery of oxygen levels in the Estuary led to the return of a productive ecosystem. This led to conservation designations, but also concerns about transfer of pollutants to higher trophic levels in fish, birds and humans. As part of urban renewal, ecosystems in disused dock basins were restored using mussel biofiltration and artificial de-stratification, facilitating commercial redevelopment and creation of a tourist destination. The degradation and recovery of the Mersey from peak-pollution in the mid-20th century is put in the context of wider environmental change and briefly compared to other systems to develop a hysteresis model of degradation and recovery, often to novel 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 © 2020 Elsevier Ltd. All rights reserved.)

Habitat modification coupled with the spread of non-native species (NNS) are among the top threats to marine biodiversity globally. Species are known to be transported to new locations via international shipping and secondarily spread via regional vessels and artificial structures. Rapid Assessment Surveys (RAS) combining quantitative and semi-quantitative methods compared NNS richness and assemblage composition on intertidal natural rocky shores and artificial structures in harbours in different regions along the south coast of England. Quantitative data showed that artificial habitats supported higher richness than natural habitats, while semi-quantitative data found no difference in richness among habitat types. This result was attributed to additional species found in rock pools during searches of complex microhabitats in natural habitats. Assemblages on artificial structures differed among regions, with regions and harbours with greater numbers of vessels supporting greater richness. Results highlight the importance of shipping and artificial structures for NNS introduction and spread., 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 © 2018. Published by Elsevier Ltd.)

Marine harbours are the focus of a diverse range of activities and subject to multiple anthropogenically induced pressures. Support for environmental management options aimed at improving degraded harbours depends on understanding the factors which influence people's perceptions of harbour environments. We used an online survey, across 12 harbours, to assess sources of variation people's perceptions of harbour health and ecological engineering. We tested the hypotheses: 1) people living near impacted harbours would consider their environment to be more unhealthy and degraded, be more concerned about the environment and supportive of and willing to pay for ecological engineering relative to those living by less impacted harbours, and 2) people with greater connectedness to the harbour would be more concerned about and have greater perceived knowledge of the environment, and be more supportive of, knowledgeable about and willing to pay for ecological engineering, than those with less connectedness. Across twelve locations, the levels of degradation and modification by artificial structures were lower and the concern and knowledge about the environment and ecological engineering were greater in the six Australasian and American than the six European and Asian harbours surveyed. We found that people's perception of harbours as healthy or degraded, but not their concern for the environment, reflected the degree to which harbours were impacted. There was a positive relationship between the percentage of shoreline modified and the extent of support for and people's willingness to pay indirect costs for ecological engineering. At the individual level, measures of connectedness to the harbour environment were good predictors of concern for and perceived knowledge about the environment but not support for and perceived knowledge about ecological engineering. To make informed decisions, it is important that people are empowered with sufficient knowledge of the environmental issues facing their harbour and ecological engineering options., (Copyright © 2018 Elsevier B.V. All rights reserved.)

The field of eco-engineering has burgeoned in recent years in response to the proliferation of artificial structures. Adding water-retaining features to seawalls has been successful in increasing biodiversity relative to the surrounding structure. Artificial rock pools may not, however, completely mimic natural rock pools. Here, we compared natural colonisation, through dispersal and recruitment, of intertidal mobile species to water-retaining flowerpots on seawalls with that into rock pools. This represents the more usual 'passive' approach to eco-engineering where features are built to enhance biodiversity and are allowed to colonise naturally, as opposed to seeding or transplanting organisms to features. While flowerpots supported some mobile species not found on the seawall, other species common on natural shores did not recruit to flowerpots. Thus, in a second experiment we tested the effectiveness of an 'active' approach through transplanting mobile organisms to flowerpots to expedite the colonisation process. For the species examined, however, most individuals did not stay in the flowerpots for more than 24 h after being transplanted. Further understanding of the processes (e.g. dispersal distances, recruitment) influencing colonisation of eco-engineered habitats is needed to effectively inform management of marine infrastructure, particularly for projects targeted at restoration rather than enhancement., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Marine ecosystems are subject to anthropogenic change at global, regional and local scales. Global drivers interact with regional- and local-scale impacts of both a chronic and acute nature. Natural fluctuations and those driven by climate change need to be understood to diagnose local- and regional-scale impacts, and to inform assessments of recovery. Three case studies are used to illustrate the need for long-term studies: (i) separation of the influence of fishing pressure from climate change on bottom fish in the English Channel; (ii) recovery of rocky shore assemblages from the Torrey Canyon oil spill in the southwest of England; (iii) interaction of climate change and chronic Tributyltin pollution affecting recovery of rocky shore populations following the Torrey Canyon oil spill. We emphasize that "baselines" or "reference states" are better viewed as envelopes that are dependent on the time window of observation. Recommendations are made for adaptive management in a rapidly changing world., (Copyright © 2017. Published by Elsevier Ltd.)

The construction of artificial structures in the marine environment is increasing globally. Eco-engineering aims to mitigate the negative ecological impacts of built infrastructure through designing structures to be multifunctional, benefiting both humans and nature. To date, the focus of eco-engineering has largely been on benefits for benthic invertebrates and algae. Here, the potential effect of eco-engineered habitats designed for benthic species on fish was investigated. Eco-engineered habitats ("flowerpots") were added to an intertidal seawall in Sydney Harbour, Australia. Responses of fish assemblages to the added habitats were quantified at two spatial scales; large (among seawalls) and small (within a seawall). Data were collected during high tide using cameras attached to the seawall to observe pelagic and benthic fish. At the larger spatial scale, herbivores, planktivores, and invertebrate predators were generally more abundant at the seawall with the added flowerpots, although results were temporally variable. At the smaller spatial scale, certain benthic species were more abundant around flowerpots than at the adjacent control areas of seawall, although there was no general pattern of differences in species density and trophic group abundance of pelagic fish between areas of the seawall with or without added habitats. Although we did not find consistent, statistically significant findings throughout our study, the field of research to improve fish habitat within human-use constraints is promising and important, although it is in its early stages (it is experimental and requires a lot of trial and error). To advance this field, it is important to document when effects were detected, and when they were not, so that others can refine the designs or scale of habitat enhancements or their study approaches (e.g., sampling protocols).

Biogenic reefs are important for habitat provision and coastal protection. Long-term datasets on the distribution and abundance of Sabellaria alveolata (L.) are available from Britain. The aim of this study was to combine historical records and contemporary data to (1) describe spatiotemporal variation in winter temperatures, (2) document short-term and long-term changes in the distribution and abundance of S. alveolata and discuss these changes in relation to extreme weather events and recent warming, and (3) assess the potential for artificial coastal defense structures to function as habitat for S. alveolata. A semi-quantitative abundance scale (ACFOR) was used to compare broadscale, long-term and interannual abundance of S. alveolata near its range edge in NW Britain. S. alveolata disappeared from the North Wales and Wirral coastlines where it had been abundant prior to the cold winter of 1962/1963. Population declines were also observed following the recent cold winters of 2009/2010 and 2010/2011. Extensive surveys in 2004 and 2012 revealed that S. alveolata had recolonized locations from which it had previously disappeared. Furthermore, it had increased in abundance at many locations, possibly in response to recent warming. S. alveolata was recorded on the majority of artificial coastal defense structures surveyed, suggesting that the proliferation of artificial coastal defense structures along this stretch of coastline may have enabled S. alveolata to spread across stretches of unsuitable natural habitat. Long-term and broadscale contextual monitoring is essential for monitoring responses of organisms to climate change. Historical data and gray literature can be invaluable sources of information. Our results support the theory that Lusitanian species are responding positively to climate warming but also that short-term extreme weather events can have potentially devastating widespread and lasting effects on organisms. Furthermore, the proliferation of coastal defense structures has implications for phylogeography, population genetics, and connectivity of coastal populations.

Coastal defence structures are proliferating to counter rising and stormier seas. With increasing concern about the ecological value of built environments, efforts are being made to create novel habitat to increase biodiversity. Rock pools are infrequent on artificial structures. We compared biodiversity patterns between rock pools and emergent rock and assessed the role of pool depth and substratum incline in determining patterns of biodiversity. Rock pools were more taxon rich than emergent substrata. Patterns varied with depth and incline with algal groups being more positively associated with shallow than deeper habitats. Substratum incline had little influence on colonising epibiota, with the exception of canopy algae in deeper habitats where vertical surfaces supported greater taxon richness than horizontal surfaces. The creation of artificial rock pools in built environments will have a positive effect on biodiversity. Building pools of varying depths and inclines and shore heights will provide a range of habitats, increase environmental heterogeneity, therefore creating more possible ecological niches, promoting local biodiversity., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Marine biodiversity currently faces unprecedented threats from multiple pressures arising from human activities. Global drivers such as climate change and ocean acidification interact with regional eutrophication, exploitation of commercial fish stocks and localized pressures including pollution, coastal development and the extraction of aggregates and fuel, causing alteration and degradation of habitats and communities. Segregating natural from anthropogenically induced change in marine ecosystems requires long-term, sustained observations of marine biota. In this review, we outline the history of biological recording in the coastal and shelf seas of the UK and Ireland and highlight where sustained observations have contributed new understanding of how anthropogenic activities have impacted on marine biodiversity. The contributions of sustained observations, from those collected at observatories, single station platforms and multiple-site programmes to the emergent field of multiple stressor impacts research, are discussed, along with implications for management and sustainable governance of marine resources in an era of unprecedented use of the marine environment., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)

Artificial structures can create novel habitat in the marine environment that has been associated with the spread of invasive species. They are often located in areas of high disturbance and can vary significantly in the area of free space provided for settlement of marine organisms. Whilst correlation between the amount of free space available and recruitment success has been shown in populations of several marine benthic organisms, there has been relatively little focus on invasive species, a group with the potential to reproduce in vast numbers and colonise habitats rapidly. Invasion success following different scales of disturbance was examined in the invasive acorn barnacle, Austrominiusmodestus, on a unique art installation located in Liverpool Bay. Population growth and recruitment success were examined by comparing recruitment rates within disturbance clearings of 4 different sizes and by contrasting population development with early recruitment rates over a 10 week period. Disturbed areas were rapidly recolonised and monocultures of A. modestus formed within 6 weeks. The size of patch created during disturbance had no effect on the rate of recruitment, while a linear relationship between recruit density and patch size was observed. Density-dependent processes mediated initial high recruitment resulting in population stability after 8-10 weeks, but densities continued to greatly exceed those reported in natural habitats. Given that artificial structures are likely to continue to proliferate in light of climate change projections, free-space is likely to become more available more frequently in the future supporting the expansion of fast-colonising species.

We briefly review how coastal ecosystems are responding to and being impacted by climate change, one of the greatest challenges facing society today. In adapting to rising and stormier seas associated with climate change, coastal defence structures are proliferating and becoming dominant coastal features, particularly in urbanised areas. Whilst the primary function of these structures is to protect coastal property and infrastructure, they inevitably have a significant secondary impact on the local environment and ecosystems. In this review we outline some of the negative and positive effects of these structures on physical processes, impacts on marine species, and the novel engineering approaches that have been employed to improve the ecological value of these structures in recent years. Finally we outline guidelines for an environmentally sensitive approach to design of such structures in the marine environment.

Background: Dispersal is a primary driver in shaping the future distribution of species in both terrestrial and marine systems. Physical transport by advection can regulate the distance travelled and rate of propagule supply to a habitat but post-settlement processes such as predation can decouple supply from recruitment. The effect of flow-mediated recruitment and predation on the recruitment success of an intertidal species, the eastern oyster Crassostrea virginica was evaluated in two-replicated field experiments. Two key crab species were manipulated to test predator identity effects on oyster mortality., Findings: Recruitment was ∼58% higher in high flow compared to low flow, but predation masked those differences. Predation mortality was primarily attributed to the blue crab Callinectes sapidus, whilst the mud crab Panopeus herbstii had no effect on recruit mortality. Recruit mortality from predation was high when recruit densities were high, but when recruit density was low, predation effects were not seen. Under high recruitment (supply), predation determined maximum population size and in low flow environments, recruitment success is likely determined by a combination of recruitment and resource limitation but not predation., Conclusions: Four processes are demonstrated: (1) Increases in flow rate positively affect recruitment success; (2) In high flow (recruitment) environments, resource availability is less important than predation; (3) predation is an important source of recruit mortality, but is dependent upon recruit density; and (4) recruitment and/or resource limitation is likely a major driver of population structure and functioning, modifying the interaction between predators and prey. Simultaneous testing of flow-mediated recruitment and predation was required to differentiate between the role of each process in determining population size. Our results reinforce the importance of propagule pressure, predation and post-settlement mortality as important determinants of population growth and persistence, but demonstrate that they should not be considered mutually exclusive.