Temporal and spatial drivers of the structure of macroinvertebrate assemblages associated with Laminaria hyperborea detritus in the northeast Atlantic.

Kelp forests occur on more than a quarter of the world's coastlines, serving as foundation species supporting high levels of biodiversity. They are also a major source of organic matter in coastal ecosystems, with the majority of primary production released and exported as detritus. Kelp detritus also provides food and shelter for macroinvertebrates, which comprise important components of inshore food-webs. Hitherto, research on kelp detritus-associated macroinvertebrate assemblages remains relatively limited. We quantified spatiotemporal variability in the structure of detritus-associated macroinvertebrate assemblages within Laminaria hyperborea forests and evaluated the influence of putative drivers of the observed variability in assemblages across eight study sites within four regions of the United Kingdom in May and September 2015. We documented 5167 individuals from 106 taxa with Malacostraca, Gastropoda, Isopoda and Bivalvia the most abundant groups sampled. Assemblage structure varied across months, sites, and regions, with highest richness in September compared to May. Many taxa were unique to individual regions, with few documented in all regions. Finally, key drivers of assemblage structure included detritus tissue nitrogen content, depth, sea surface temperature, light intensity, as well as L. hyperborea canopy density and canopy biomass. Despite their dynamic composition and transient existence, accumulations of L. hyperborea detritus represent valuable repositories of biodiversity and represent an additional kelp forest component which influences secondary productivity, and potentially kelp forest food-web dynamics.

The influence of environmental context on community composition in artificial rockpools associated with seawalls.

Artificial structures have become widespread features of coastal marine environments, and will likely proliferate further over the coming decades. These constitute new hard substrata in the marine environment which provide a fundamentally different habitat than natural shores. Eco-engineering solutions aim to ameliorate these differences by combining ecological knowledge and engineering criteria in the construction and modification of artificial substrata. Vertipools™ are artificial bolt-on rockpools intended for deployment on seawalls, where they have been shown to provide biodiversity benefits. In this study, a total of 32 Vertipools were retrofitted on eight seawalls in different environmental contexts (estuarine vs marine and urban vs rural) along the Irish Sea coastline, and were exposed to the environment for a period of two years. After two years, there were no differences in species richness, species-abundance distributions, diversity, or community composition between the specific environmental contexts examined here. Site-level variation was significant, and communities on Vertipools deployed in marine contexts were more variable in general than those in estuarine contexts. Community composition differed significantly between structural sections of the Vertipools, indicating that different sections provide specific microhabitats for colonisation. This study indicates that Vertipools provide biodiversity benefits in a variety of environmental contexts, and therefore are broadly viable as an eco-engineering solution.

Multiscale Spatial Variability and Stability in the Structure and Diversity of Bacterial Communities Associated with the Kelp Eisenia cokeri in Peru.

Ecological communities are structured by a range of processes that operate over a range of spatial scales. While our understanding of such biodiversity patterns in macro-communities is well studied, our understanding at the microbial level is still lacking. Bacteria can be free living or associated with host eukaryotes, forming part of a wider "microbiome," which is fundamental for host performance and health. For habitat forming foundation-species, host-bacteria relationships likely play disproportionate roles in mediating processes for the wider ecosystem. Here, we describe host-bacteria communities across multiple spatial scales (i.e., from 10s of m to 100s of km) in the understudied kelp, Eisenia cokeri, in Peru. We found that E. cokeri supports a distinct bacterial community compared to the surrounding seawater, but the structure of these communities varied markedly at the regional (~480 km), site (1-10 km), and individual (10s of m) scale. The marked regional-scale differences we observed may be driven by a range of processes, including temperature, upwelling intensity, or regional connectivity patterns. However, despite this variability, we observed consistency in the form of a persistent core community at the genus level. Here, the genera Arenicella, Blastopirellula, Granulosicoccus, and Litorimonas were found in >80% of samples and comprised ~53% of total sample abundance. These genera have been documented within bacterial communities associated with kelps and other seaweed species from around the world and may be important for host function and wider ecosystem health in general.

Population structure and reproductive states of the dogwhelk Nucella lapillus differ between artificial structures and natural rocky shores.

Artificial structures are an increasingly common feature of coastal marine environments. These structures are poor surrogates of natural rocky shores, and generally support less diverse communities and reduced population sizes. Little is known about sub-lethal effects of such structures in terms of demographic properties and reproductive potential, both of which may influence the dynamics and long-term viability of populations. This study examines the population structure, reproductive states and embryo production of Nucella lapillus populations on artificial structures and natural shores in Ireland and Wales. Population density was measured twice at six natural shores and six artificial structures: once in winter and once in spring. At each sampling, the shell height of 100 individuals from each site was measured. Monthly collections of adult specimens and egg capsules were made at each site from November-January and from March-May, in order to determine sex ratios, reproductive states, and embryo abundances. Artificial structures supported larger individuals and very few juveniles compared to natural shores. Between December and January, natural shores experienced a distinctive pulse in spawning activity followed by a decline in the proportion of females in a reproductive state, whereas on artificial structures the proportion of reproductive females remained relatively stable. Differences observed may be due to a lack of microhabitats on artificial structures, along with subtle variations in structure slope. Eco-engineering interventions, including the addition of refugia such as cracks and crevices, may allow N. lapillus populations on artificial structures to approximate those on natural shores.

Artificial rockpools: Seaweed colonisation and productivity vary between sites but are consistent across environmental contexts.

Artificial structures are widespread features of coastal environments, but are poor surrogates of natural rocky shores because they generally support depauperate assemblages with reduced population sizes. This has generated significant interest in eco-engineering solutions, including retrofitting seawalls with artificial rockpools to increase water retention and provide microhabitats. Although these have proven effective at individual sites, widespread uptake is contingent on evidence of consistent benefits across a range of contexts. In this study, Vertipools™ were retrofitted on eight seawalls in different environmental contexts (urban v rural and estuarine v marine) along the Irish Sea coastline and were monitored regularly for two years. Seaweed colonisation proceeded in a manner similar to patterns described for natural and artificial intertidal systems in general, consisting of early dominance by ephemeral species followed by the appearance and eventual establishment of perennial habitat-formers. After 24 months, species richness did not differ between contexts, but differed between sites. The units supported populations of large habitat-forming seaweeds at all sites. Productivity and community respiration of the colonising communities differed between sites by up to 0.5 mg O2 L-1 min-1, but not across environmental contexts. This study demonstrates that bolt-on rockpools attract similar levels of biotic colonisation and functioning in a variety of temperate environmental contexts, and could be considered for widespread implementation as an eco-engineering solution.

The value of ecosystem services in global marine kelp forests.

While marine kelp forests have provided valuable ecosystem services for millennia, the global ecological and economic value of those services is largely unresolved. Kelp forests are diminishing in many regions worldwide, and efforts to manage these ecosystems are hindered without accurate estimates of the value of the services that kelp forests provide to human societies. Here, we present a global estimate of the ecological and economic potential of three key ecosystem services - fisheries production, nutrient cycling, and carbon removal provided by six major forest forming kelp genera (Ecklonia, Laminaria, Lessonia, Macrocystis, Nereocystis, and Saccharina). Each of these genera creates a potential value of between $64,400 and $147,100/hectare each year. Collectively, they generate between $465 and $562 billion/year worldwide, with an average of $500 billion. These values are primarily driven by fisheries production (mean $29,900, 904 Kg/Ha/year) and nitrogen removal ($73,800, 657 Kg N/Ha/year), though kelp forests are also estimated to sequester 4.91 megatons of carbon from the atmosphere/year highlighting their potential as blue carbon systems for climate change mitigation. These findings highlight the ecological and economic value of kelp forests to society and will facilitate better informed marine management and conservation decisions.

The population structure, sex ratio and reproductive potential of limpets (Patella spp.) on natural shores and artificial structures in the Irish Sea.

Artificial structures often support depauperate communities compared to natural rocky shores. Understanding variation in ecological success across shore types, particularly regarding habitat-forming species or those with structuring roles, is important to determine how artificial structure proliferation may influence ecosystem functioning and services. We investigated the population structure, sex ratio and reproductive potential of limpets on natural shores and artificial structures on Irish Sea coasts. Limpets were generally less abundant and Patella vulgata populations were often male dominated on artificial structures compared to natural shores, suggesting that shore type may influence these factors. P. vulgata length varied across sites within the Irish Sea (nested in coast and shore type) in autumn/winter, as well as temporally across sites along the Welsh coast. There was no difference in the proportion of P. vulgata in advanced stages of gonad development across shore types. The results suggest that rip-rap artificial structures may provide a habitat comparable to natural shores, however, the addition of ecological engineering interventions on artificial structures may allow limpet populations to better approximate those on natural shores.

Consistency and Variation in the Kelp Microbiota: Patterns of Bacterial Community Structure Across Spatial Scales.

Kelp species are distributed along ~ 25% of the world's coastlines and the forests they form represent some of the world's most productive and diverse ecosystems. Like other marine habitat-formers, the associated microbial community is fundamental for host and, in turn, wider ecosystem functioning. Given there are thousands of bacteria-host associations, determining which relationships are important remains a major challenge. We characterised the associated bacteria of two habitat-forming kelp species, Laminaria hyperborea and Saccharina latissima, from eight sites across a range of spatial scales (10 s of metres to 100 s of km) in the northeast Atlantic. We found no difference in diversity or community structure between the two kelps, but there was evidence of regional structuring (across 100 s km) and considerable variation between individuals (10 s of metres). Within sites, individuals shared few amplicon sequence variants (ASVs) and supported a very small proportion of diversity found across the wider study area. However, consistent characteristics between individuals were observed with individual host communities containing a small conserved "core" (8-11 ASVs comprising 25 and 32% of sample abundances for L. hyperborea and S. latissima, respectively). At a coarser taxonomic resolution, communities were dominated by four classes (Planctomycetes, Gammaproteobacteria, Alphaproteobacteria and Bacteroidia) that made up ~ 84% of sample abundances. Remaining taxa (47 classes) made up very little contribution to overall abundance but the majority of taxonomic diversity. Overall, our study demonstrates the consistent features of kelp bacterial communities across large spatial scales and environmental gradients and provides an ecologically meaningful baseline to track environmental change.

Biological Impacts of Marine Heatwaves.

Climatic extremes are becoming increasingly common against a background trend of global warming. In the oceans, marine heatwaves (MHWs)-discrete periods of anomalously warm water-have intensified and become more frequent over the past century, impacting the integrity of marine ecosystems globally. We review and synthesize current understanding of MHW impacts at the individual, population, and community levels. We then examine how these impacts affect broader ecosystem services and discuss the current state of research on biological impacts of MHWs. Finally, we explore current and emergent approaches to predicting the occurrence andimpacts of future events, along with adaptation and management approaches. With further increases in intensity and frequency projected for coming decades, MHWs are emerging as pervasive stressors to marine ecosystems globally. A deeper mechanistic understanding of their biological impacts is needed to better predict and adapt to increased MHW activity in the Anthropocene.

Global seaweed productivity.

The magnitude and distribution of net primary production (NPP) in the coastal ocean remains poorly constrained, particularly for shallow marine vegetation. Here, using a compilation of in situ annual NPP measurements across >400 sites in 72 geographic ecoregions, we provide global predictions of the productivity of seaweed habitats, which form the largest vegetated coastal biome on the planet. We find that seaweed NPP is strongly coupled to climatic variables, peaks at temperate latitudes, and is dominated by forests of large brown seaweeds. Seaweed forests exhibit exceptionally high per-area production rates (a global average of 656 and 1711 gC m-2 year-1 in the subtidal and intertidal, respectively), being up to 10 times higher than coastal phytoplankton in temperate and polar seas. Our results show that seaweed NPP is a strong driver of production in the coastal ocean and call for its integration in the oceanic carbon cycle, where it has traditionally been overlooked.

A global dataset of seaweed net primary productivity.

Net primary productivity (NPP) plays a pivotal role in the global carbon balance but estimating the NPP of underwater habitats remains a challenging task. Seaweeds (marine macroalgae) form the largest and most productive underwater vegetated habitat on Earth. Yet, little is known about the distribution of their NPP at large spatial scales, despite more than 70 years of local-scale studies being scattered throughout the literature. We present a global dataset containing NPP records for 246 seaweed taxa at 429 individual sites distributed on all continents from the intertidal to 55 m depth. All records are standardized to annual aerial carbon production (g C m-2 yr-1) and are accompanied by detailed taxonomic and methodological information. The dataset presented here provides a basis for local, regional and global comparative studies of the NPP of underwater vegetation and is pivotal for achieving a better understanding of the role seaweeds play in the global coastal carbon cycle.

Kelp carbon sink potential decreases with warming due to accelerating decomposition.

Cycling of organic carbon in the ocean has the potential to mitigate or exacerbate global climate change, but major questions remain about the environmental controls on organic carbon flux in the coastal zone. Here, we used a field experiment distributed across 28° of latitude, and the entire range of 2 dominant kelp species in the northern hemisphere, to measure decomposition rates of kelp detritus on the seafloor in relation to local environmental factors. Detritus decomposition in both species were strongly related to ocean temperature and initial carbon content, with higher rates of biomass loss at lower latitudes with warmer temperatures. Our experiment showed slow overall decomposition and turnover of kelp detritus and modeling of coastal residence times at our study sites revealed that a significant portion of this production can remain intact long enough to reach deep marine sinks. The results suggest that decomposition of these kelp species could accelerate with ocean warming and that low-latitude kelp forests could experience the greatest increase in remineralization with a 9% to 42% reduced potential for transport to long-term ocean sinks under short-term (RCP4.5) and long-term (RCP8.5) warming scenarios. However, slow decomposition at high latitudes, where kelp abundance is predicted to expand, indicates potential for increasing kelp-carbon sinks in cooler (northern) regions. Our findings reveal an important latitudinal gradient in coastal ecosystem function that provides an improved capacity to predict the implications of ocean warming on carbon cycling. Broad-scale patterns in organic carbon decomposition revealed here can be used to identify hotspots of carbon sequestration potential and resolve relationships between carbon cycling processes and ocean climate at a global scale.

Socioeconomic impacts of marine heatwaves: Global issues and opportunities.

Extreme climatic events, including marine heatwaves (MHWs), are altering ecosystems globally, often with profound socioeconomic impacts. We examine how MHWs have affected the provision of ecosystem services and evaluate the socioeconomic consequences for human society. Ecological impacts range from harmful algal blooms and mass mortality events to reconfigurations of entire ecosystems, affecting provisioning, habitat, regulating, and cultural ecosystem services globally. Reported economic costs of individual MHW events exceed US$800 million in direct losses or >US$3.1 billion in indirect losses of ecosystem services for multiple years. However, biological responses to MHWs can also increase human-ocean interactions, providing opportunities for coastal societies. Our study provides a global perspective on the far-reaching impacts of MHWs on human societies and highlights the urgent need to develop robust approaches to mitigation and adaptation.

Artificial shorelines lack natural structural complexity across scales.

From microbes to humans, habitat structural complexity plays a direct role in the provision of physical living space, and increased complexity supports higher biodiversity and ecosystem functioning across biomes. Coastal development and the construction of artificial shorelines are altering natural landscapes as humans seek socio-economic benefits and protection from coastal storms, flooding and erosion. In this study, we evaluate how much structural complexity is missing on artificial coastal structures compared to natural rocky shorelines, across a range of spatial scales from 1 mm to 10 s of m, using three remote sensing platforms (handheld camera, terrestrial laser scanner and uncrewed aerial vehicles). Natural shorelines were typically more structurally complex than artificial ones and offered greater variation between locations. However, our results varied depending on the type of artificial structure and the scale at which complexity was measured. Seawalls were deficient at all scales (approx. 20-40% less complex than natural shores), whereas rock armour was deficient at the smallest and largest scales (approx. 20-50%). Our findings reinforce concerns that hardening shorelines with artificial structures simplifies coastlines at organism-relevant scales. Furthermore, we offer much-needed insight into how structures might be modified to more closely capture the complexity of natural rocky shores that support biodiversity.

Fucus vesiculosus populations on artificial structures have potentially reduced fecundity and are dislodged at greater rates than on natural shores.

Artificial structures are widespread features of coastal marine environments. These structures, however, are poor surrogates of natural rocky shores, meaning they generally support depauperate assemblages with reduced population sizes. Little is known about sub-lethal effects of such structures, for example, in terms of demographic properties and reproductive potential that may affect the dynamics and long-term viability of populations. Such understanding is particularly important for ecosystem engineer species, such as the intertidal seaweed Fucus vesiculosus. In this study, F. vesiculosus was sampled on eight artificial structures and eight natural shores along the east coast of Ireland and the west coast of Wales. Algal percentage cover, biomass, density of individuals, and growth rate did not differ between artificial and natural shores. Growth and reproductive cycles were consistent with previous studies for this species. While there was considerable variation from site to site, on average, populations on natural shores produced a higher number of mature receptacles during the peak reproductive period in April, and lower rates of dislodgement than on artificial structures. As F. vesiculosus reach peak reproductive output after 24 months, this suggests that individuals may be removed from populations on artificial structures before reaching their full reproductive potential. In this case, this did not influence density, percentage cover, or biomass, which suggests that F. vesiculosus populations on artificial structures may function similarly to those on natural shores if supported by suitable source populations, but potentially may not persist otherwise.

Drivers and impacts of the most extreme marine heatwaves events.

Prolonged high-temperature extreme events in the ocean, marine heatwaves, can have severe and long-lasting impacts on marine ecosystems, fisheries and associated services. This study applies a marine heatwave framework to analyse a global sea surface temperature product and identify the most extreme events, based on their intensity, duration and spatial extent. Many of these events have yet to be described in terms of their physical attributes, generation mechanisms, or ecological impacts. Our synthesis identifies commonalities between marine heatwave characteristics and seasonality, links to the El Niño-Southern Oscillation, triggering processes and impacts on ocean productivity. The most intense events preferentially occur in summer, when climatological oceanic mixed layers are shallow and winds are weak, but at a time preceding climatological maximum sea surface temperatures. Most subtropical extreme marine heatwaves were triggered by persistent atmospheric high-pressure systems and anomalously weak wind speeds, associated with increased insolation, and reduced ocean heat losses. Furthermore, the most extreme events tended to coincide with reduced chlorophyll-a concentration at low and mid-latitudes. Understanding the importance of the oceanic background state, local and remote drivers and the ocean productivity response from past events are critical steps toward improving predictions of future marine heatwaves and their impacts.

Summer and Winter Marine Heatwaves Favor an Invasive Over Native Seaweeds.

Marine heatwaves (MHWs) are emerging as forceful agents of ecosystem change and are increasing in frequency, duration, and intensity with climate change. During MHWs, physiological thresholds of native species may be exceeded while the performance of invasive species with warm affinities may be enhanced. As a consequence, MHWs could significantly alter an ecosystem's invasive dynamics, but such interactions are poorly understood. Following a 10-d acclimation period, we investigated the physiological resistance and resilience of an intertidal rock pool assemblage invaded by the seaweed Sargassum muticum to realistic 14-d marine heatwave scenarios (+1.5°C, +2.0°C, +3.5°C) followed by a 14-d recovery period. We conducted mesocosm experiments in both summer and winter to investigate temporal variability of MHWs. MHW treatments had clear negative impacts on native seaweeds (Fucus serratus and Chondrus crispus) while enhancing the performance of S. muticum. This pattern was consistent across season indicating that acclimation to cooler ambient temperatures results in winter MHWs having significant impacts on native species. As climate warming advances, this may ultimately lead to changes in competitive interactions and potentially exclusion of native species, while invasive species may proliferate and become more conspicuous within temperate rocky shore environments.

Environmental factors influencing primary productivity of the forest-forming kelp Laminaria hyperborea in the northeast Atlantic.

Rates and drivers of primary productivity are well understood for many terrestrial ecosystems, but remain poorly resolved for many marine ecosystems, particularly those within in coastal benthic environments. We quantified net primary productivity (NPP) using two methods as well as carbon standing stock within kelp forests (Laminaria hyperborea) at multiple subtidal habitats in the United Kingdom (UK). Study sites spanned 9° in latitude and encompassed a gradient in average temperature of ~ 2.5 °C. In addition to temperature, we measured other factors (e.g. light intensity, water motion, nutrients, sea urchin density) that may influence productivity. Although estimates of NPP were highly variable between sites, ranging from 166 to 738 g C m-2 yr-1, our study-wide average of 340 g C m-2 yr-1 indicated that L. hyperborea forests are highly productive. We observed clear differences between NPP and carbon standing stock between our cold northernmost sites and our warm southernmost sites, with NPP and standing stock being around 1.5 and 2.5 times greater in the northern sites, respectively. Ocean temperature was identified as a likely driver of productivity, with reduced NPP and standing stock observed in warmer waters. Light availability was also strongly linked with carbon accumulation and storage, with increased light levels positively correlated with NPP and standing stock. Across its geographical range, total NPP from L. hyperborea is estimated at ~ 7.61 Tg C yr-1. This biomass production is likely to be important for local food webs, as a trophic subsidy to distant habitats and for inshore carbon cycling and (potentially) carbon sequestration. However, given the strong links with temperature, continued ocean warming in the northeast Atlantic may reduce primary productivity of this foundation species, as optimal temperatures for growth and performance are surpassed.

A global assessment of marine heatwaves and their drivers.

Marine heatwaves (MHWs) can cause devastating impacts to marine life. Despite the serious consequences of MHWs, our understanding of their drivers is largely based on isolated case studies rather than any systematic unifying assessment. Here we provide the first global assessment under a consistent framework by combining a confidence assessment of the historical refereed literature from 1950 to February 2016, together with the analysis of MHWs determined from daily satellite sea surface temperatures from 1982-2016, to identify the important local processes, large-scale climate modes and teleconnections that are associated with MHWs regionally. Clear patterns emerge, including coherent relationships between enhanced or suppressed MHW occurrences with the dominant climate modes across most regions of the globe - an important exception being western boundary current regions where reports of MHW events are few and ocean-climate relationships are complex. These results provide a global baseline for future MHW process and prediction studies.