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.

Spatiotemporal variability in population demography and morphology of the habitat-forming macroalga Saccorhiza polyschides in the Western English Channel.

BACKGROUND AND AIMS: Large brown macroalgae serve as foundation organisms along temperate and polar coastlines, providing a range of ecosystem services. Saccorhiza polyschides is a warm-temperate kelp-like species found in the northeast Atlantic, which is suggested to have proliferated in recent decades across the southern UK, possibly in response to increasing temperatures, physical disturbance and reduced competition. However, little is known about S. polyschides with regard to ecological functioning and population dynamics across its geographical range. Here we examined the population demography of S. polyschides populations in southwest UK, located within the species' range centre, to address a regional knowledge gap and to provide a baseline against which to detect future changes. METHODS: Intertidal surveys were conducted during spring low tides at three sites along a gradient of wave exposure in Plymouth Sound (Western English Channel) over a period of 15 months. Density, cover, age, biomass and morphology of S. polyschides were quantified. Additionally, less frequent sampling of shallow subtidal reefs was conducted to compare intertidal and subtidal populations. KEY RESULTS: We recorded pronounced seasonality, with fairly consistent demographic patterns across sites and depths. By late summer, S. polyschides was a dominant habitat-former on both intertidal and subtidal reefs, with maximum standing stock exceeding 13 000 g wet weight m-2. CONCLUSIONS: Saccorhiza polyschides is a conspicuous and abundant member of rocky reef assemblages in the region, providing complex and abundant biogenic habitat for associated organisms and high rates of primary productivity. However, its short-lived pseudo-annual life strategy is in stark contrast to dominant long-lived perennial laminarian kelps. As such, any replacement or reconfiguration of habitat-forming macroalgae due to ocean warming will probably have implications for local biodiversity and community composition. More broadly, our study demonstrates the importance of high-resolution cross-habitat surveys to generate robust baselines of kelp population demography, against which the ecological impacts of climate change and other stressors can be reliably detected.

Marine heatwave duration and intensity interact to reduce physiological tipping points of kelp species with contrasting thermal affinities.

BACKGROUND AND AIMS: Marine heatwaves (MHWs) are widely recognized as pervasive drivers of ecosystem change, yet our understanding of how different MHW properties mediate ecological responses remains largely unexplored. Understanding MHW impacts on foundation species is particularly important, given their structural role in communities and ecosystems. METHODS: We simulated a series of realistic MHWs with different levels of intensity (Control: 14 °C, Moderate: 18 °C, Extreme: 22 °C) and duration (14 or 28 d) and examined responses of two habitat-forming kelp species in the southwest UK. Here, Laminaria digitata reaches its trailing edge and is undergoing a range contraction, whereas Laminaria ochroleuca reaches its leading edge and is undergoing a range expansion. KEY RESULTS: For both species, sub-lethal stress responses induced by moderate-intensity MHWs were exacerbated by longer duration. Extreme-intensity MHWs caused dramatic declines in growth and photosynthetic performance, and elevated bleaching, which were again exacerbated by longer MHW duration. Stress responses were most pronounced in L. ochroleuca, where almost complete tissue necrosis was observed by the end of the long-duration MHW. This was unexpected given the greater thermal safety margins assumed with leading edge populations. It is likely that prolonged exposure to sub-lethal thermal stress exceeded a physiological tipping point for L. ochroleuca, presumably due to depletion of internal reserves. CONCLUSIONS: Overall, our study showed that exposure to MHW profiles projected to occur in the region in the coming decades can have significant deleterious effects on foundation kelp species, regardless of their thermal affinities and location within respective latitudinal ranges, which would probably have consequences for entire communities and ecosystems.

Seasonal development of a tidal mixing front drives shifts in community structure and diversity of bacterioplankton.

Bacterioplankton underpin biogeochemical cycles and an improved understanding of the patterns and drivers of variability in their distribution is needed to determine their wider functioning and importance. Sharp environmental gradients and dispersal barriers associated with ocean fronts are emerging as key determinants of bacterioplankton biodiversity patterns. We examined how the development of the Celtic Sea Front (CF), a tidal mixing front on the Northwest European Shelf affects bacterioplankton communities. We performed 16S-rRNA metabarcoding on 60 seawater samples collected from three depths (surface, 20 m and seafloor), across two research cruises (May and September 2018), encompassing the intra-annual range of the CF intensity. Communities above the thermocline of stratified frontal waters were clearly differentiated and less diverse than those below the thermocline and communities in the well-mixed waters of the Irish Sea. This effect was much more pronounced in September, when the CF was at its peak intensity. The stratified zone likely represents a stressful environment for bacterioplankton due to a combination of high temperatures and low nutrients, which fewer taxa can tolerate. Much of the observed variation was driven by Synechococcus spp. (cyanobacteria), which were more abundant within the stratified zone and are known to thrive in warm oligotrophic waters. Synechococcus spp. are key contributors to global primary productivity and carbon cycling and, as such, variability driven by the CF is likely to influence regional biogeochemical processes. However, further studies are required to explicitly link shifts in community structure to function and quantify their wider importance to pelagic ecosystems.

Identifying and protecting macroalgae detritus sinks toward climate change mitigation.

Harnessing natural solutions to mitigate climate change requires an understanding of carbon fixation, flux, and sequestration across ocean habitats. Recent studies have suggested that exported seaweed particulate organic carbon is stored within soft-sediment systems. However, very little is known about how seaweed detritus disperses from coastlines, or where it may enter seabed carbon stores, where it could become the target of conservation efforts. Here, focusing on regionally dominant seaweed species, we surveyed environmental DNA (eDNA) from natural coastal sediments, and studied their connectivity to seaweed habitats using a particle tracking model parameterized to reproduce seaweed detritus dispersal behavior based on laboratory observations of seaweed fragment degradation and sinking. Experiments showed that seaweed detritus density changed over time, differently across species. This, in turn, modified distances traveled by released fragments until they reached the seabed for the first time, during model simulations. Dispersal pathways connected detritus from the shore to the open ocean but, importantly, also to coastal sediments, and this was reflected by field eDNA evidence. Dispersion pathways were also affected by hydrodynamic conditions, varying in space and time. Both the properties and timing of released detritus, individual to each macroalgal population, and short-term near-seabed and medium-term water-column transport pathways, are thus seemingly important in determining the connectivity between seaweed habitats and potential sedimentary sinks. Studies such as this one, supported by further field verification of sedimentary carbon sequestration rates and source partitioning, are still needed to help quantify the role of seaweed in the ocean carbon cycle. Such studies will provide vital evidence to inform on the potential need to develop blue carbon conservation mechanisms, beyond wetlands.

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.

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.

Core Community Persistence Despite Dynamic Spatiotemporal Responses in the Associated Bacterial Communities of Farmed Pacific Oysters.

A breakdown in host-bacteria relationships has been associated with the progression of a number of marine diseases and subsequent mortality events. For the Pacific oyster, Crassostrea gigas, summer mortality syndrome (SMS) is one of the biggest constraints to the growth of the sector and is set to expand into temperate systems as ocean temperatures rise. Currently, a lack of understanding of natural spatiotemporal dynamics of the host-bacteria relationship limits our ability to develop microbially based monitoring approaches. Here, we characterised the associated bacterial community of C. gigas, at two Irish oyster farms, unaffected by SMS, over the course of a year. We found C. gigas harboured spatiotemporally variable bacterial communities that were distinct from bacterioplankton in surrounding seawater. Whilst the majority of bacteria-oyster associations were transient and highly variable, we observed clear patterns of stability in the form of a small core consisting of six persistent amplicon sequence variants (ASVs). This core made up a disproportionately large contribution to sample abundance (34 ± 0.14%), despite representing only 0.034% of species richness across the study, and has been associated with healthy oysters in other systems. Overall, our study demonstrates the consistent features of oyster bacterial communities across spatial and temporal scales and provides an ecologically meaningful baseline to track environmental change.

Marine heatwaves and decreased light availability interact to erode the ecophysiological performance of habitat-forming kelp species.

Coastal marine ecosystems are threatened by a range of anthropogenic stressors, operating at global, local, and temporal scales. We investigated the impact of marine heatwaves (MHWs) combined with decreased light availability over two seasons on the ecophysiological responses of three kelp species (Laminaria digitata, L. hyperborea, and L. ochroleuca). These species function as important habitat-forming foundation organisms in the northeast Atlantic and have distinct but overlapping latitudinal distributions and thermal niches. Under low-light conditions, summertime MHWs induced significant declines in biomass, blade surface area, and Fv/Fm values (a measure of photosynthetic efficiency) in the cool-water kelps L. digitata and L. hyperborea, albeit to varying degrees. Under high-light conditions, all species were largely resistant to simulated MHW activity. In springtime, MHWs had minimal impacts and in some cases promoted kelp performance, while reduced light availability resulted in lower growth rates. While some species were negatively affected by summer MHWs under low-light conditions (particularly L. digitata), they were generally resilient to MHWs under high-light conditions. As such, maintaining good environmental quality and water clarity may increase resilience of populations to summertime MHWs. Our study informs predictions of how habitat-forming foundation kelp species will be affected by interacting, concurrent stressors, typical of compound events that are intensifying under anthropogenic climate change.

Impacts of ocean warming on kelp forest ecosystems.

Kelp forests represent some of the most diverse and productive habitats on Earth, and provide a range of ecosystem goods and services on which human populations depend. As the distribution and ecophysiology of kelp species is strongly influenced by temperature, recent warming trends in many regions have been linked with concurrent changes in kelp populations, communities and ecosystems. Over the past decade, the number of reports of ocean warming impacts on kelp forests has risen sharply. Here, I synthesise recent studies to highlight general patterns and trends. While kelp responses to climate change vary greatly between ocean basins, regions and species, there is compelling evidence to show that ocean warming poses an unequivocal threat to the persistence and integrity of kelp forest ecosystems in coming decades.

Ground-based FTIR observation of hydrogen chloride (HCl) over Hefei, China, and comparisons with GEOS-Chem model data and other ground-based FTIR stations data.

In this study, the characterization of Hydrogen Chloride (HCl) seasonal variations and inter-annual linear trend are presented for the first time over the polluted region at Hefei (117°10'E, 31°54'N), China. The time series of HCl were retrieved by the mid-infrared (MIR) solar spectra recorded by the ground-based high-resolution Fourier transform infrared spectroscopy (FTIR) between July, 2015 and April, 2019. The magnitude of HCl reaches a peak in January (2.70 ± 0.16) × 1015 molecules*cm-2 and a minimum in September (2.27 ± 0.09) × 1015 molecules*cm-2. The four-year time series of HCl total column show a negative linear trend of (-1.83 ± 0.13) %. The FTIR data are compared with GEOS-Chem data in order to evaluate the performance of the GEOS-Chem model to simulate HCl. In general, total column FTIR data and GEOS-Chem model data are in a good agreement with a correlation coefficient of 0.82. GEOS-Chem model data present a good agreement with FTIR data in seasonal variation and inter-annul trend. The maximum differences occur in January and April with mean differences of 4%-6%. We also present HCl time series observed by 6 NDACC stations (Bremen, Toronto, Rikubetsu, Izana, Reunion.maido, Lauder) in low-middle-latitude sites of the northern and southern hemispheres and Hefei stations in order to investigate the seasonal and annual trends of HCl in low-middle-latitude sites. The HCl total column at the northern hemisphere stations reached the maximum in the late winter or early spring and the minimum in the early winter or late autumn. In general, the seasonal variations of HCl over Hefei is similar to that in other northern hemisphere mid-latitude FTIR stations.

Intra-Annual Variability in Responses of a Canopy Forming Kelp to Cumulative Low Tide Heat Stress: Implications for Populations at the Trailing Range Edge.

Anthropogenic climate change is driving the redistribution of species at a global scale. For marine species, populations at trailing edges often live very close to their upper thermal limits and, as such, poleward range contractions are one of the most pervasive effects of ongoing and predicted warming. However, the mechanics of processes driving such contractions are poorly understood. Here, we examined the response of the habitat forming kelp, Laminaria digitata, to realistic terrestrial heatwave simulations akin to those experienced by intertidal populations persisting at the trailing range edge in the northeast Atlantic (SW England). We conducted experiments in both spring and autumn to determine temporal variability in the effects of heatwaves. In spring, heatwave scenarios caused minimal stress to L. digitata but in autumn all scenarios tested resulted in tissue being nonviable by the end of each assay. The effects of heatwave scenarios were only apparent after consecutive exposures, indicating erosion of resilience over time. Monthly field surveys corroborated experimental evidence as the prevalence of bleaching (an indication of physiological stress and tissue damage) in natural populations was greatest in autumn and early winter. Overall, our data showed that L. digitata populations in SW England persist close to their upper physiological limits for emersion stress in autumn. As the intensity of extreme warming events is likely to increase with anthropogenic climate change, thermal conditions experienced during periods of emersion will soon exceed physiological thresholds and will likely induce widespread mortality and consequent changes at the population level.

Carbon assimilation and transfer through kelp forests in the NE Atlantic is diminished under a warmer ocean climate.

Global climate change is affecting carbon cycling by driving changes in primary productivity and rates of carbon fixation, release and storage within Earth's vegetated systems. There is, however, limited understanding of how carbon flow between donor and recipient habitats will respond to climatic changes. Macroalgal-dominated habitats, such as kelp forests, are gaining recognition as important carbon donors within coastal carbon cycles, yet rates of carbon assimilation and transfer through these habitats are poorly resolved. Here, we investigated the likely impacts of ocean warming on coastal carbon cycling by quantifying rates of carbon assimilation and transfer in Laminaria hyperborea kelp forests-one of the most extensive coastal vegetated habitat types in the NE Atlantic-along a latitudinal temperature gradient. Kelp forests within warm climatic regimes assimilated, on average, more than three times less carbon and donated less than half the amount of particulate carbon compared to those from cold regimes. These patterns were not related to variability in other environmental parameters. Across their wider geographical distribution, plants exhibited reduced sizes toward their warm-water equatorward range edge, further suggesting that carbon flow is reduced under warmer climates. Overall, we estimated that Laminaria hyperborea forests stored ~11.49 Tg C in living biomass and released particulate carbon at a rate of ~5.71 Tg C year-1 . This estimated flow of carbon was markedly higher than reported values for most other marine and terrestrial vegetated habitat types in Europe. Together, our observations suggest that continued warming will diminish the amount of carbon that is assimilated and transported through temperate kelp forests in NE Atlantic, with potential consequences for the coastal carbon cycle. Our findings underline the need to consider climate-driven changes in the capacity of ecosystems to fix and donate carbon when assessing the impacts of climate change on carbon cycling.

Python-based dynamic scheduling assistant for atmospheric measurements by Bruker instruments using OPUS.

Atmospheric remote sensing by instruments such as spectrometers and interferometers often requires scheduling that is dependent on external factors, for example; time and solar (or lunar) zenith angle. Such instruments manufactured by Bruker often use the software package OPUS, which, while useful, is not flexible enough for automatic, repeated, atmospheric measurements of this nature. In this brief paper, we describe ASAP, a Python tool developed to run our network of Fourier transform interferometers in New Zealand and Antarctica. It allows the automated scheduling of measurements by time, lunar, or solar zenith angle while accounting for weather or other external parameters. There is a wide range of useful functions, all packaged in a simple graphical user interface; it is available on request.

Community responses to seawater warming are conserved across diverse biological groupings and taxonomic resolutions.

Temperature variability is a major driver of ecological pattern, with recent changes in average and extreme temperatures having significant impacts on populations, communities and ecosystems. In the marine realm, very few experiments have manipulated temperature in situ, and current understanding of temperature effects on community dynamics is limited. We developed new technology for precise seawater temperature control to examine warming effects on communities of bacteria, microbial eukaryotes (protists) and metazoans. Despite highly contrasting phylogenies, size spectra and diversity levels, the three community types responded similarly to seawater warming treatments of +3°C and +5°C, highlighting the critical and overarching importance of temperature in structuring communities. Temperature effects were detectable at coarse taxonomic resolutions and many taxa responded positively to warming, leading to increased abundances at the community-level. Novel field-based experimental approaches are essential to improve mechanistic understanding of how ocean warming will alter the structure and functioning of diverse marine communities.

The effects of warming on the ecophysiology of two co-existing kelp species with contrasting distributions.

The northeast Atlantic has warmed significantly since the early 1980s, leading to shifts in species distributions and changes in the structure and functioning of communities and ecosystems. This study investigated the effects of increased temperature on two co-existing habitat-forming kelps: Laminaria digitata, a northern boreal species, and Laminaria ochroleuca, a southern Lusitanian species, to shed light on mechanisms underpinning responses of trailing and leading edge populations to warming. Kelp sporophytes collected from southwest United Kingdom were maintained under 3 treatments: ambient temperature (12 °C), +3 °C (15 °C) and +6 °C (18 °C) for 16 days. At higher temperatures, L. digitata showed a decline in growth rates and Fv/Fm, an increase in chemical defence production and a decrease in palatability. In contrast, L. ochroleuca demonstrated superior growth and photosynthesis at temperatures higher than current ambient levels, and was more heavily grazed. Whilst the observed decreased palatability of L. digitata held at higher temperatures could reduce top-down pressure on marginal populations, field observations of grazer densities suggest that this may be unimportant within the study system. Overall, our study suggests that shifts in trailing edge populations will be primarily driven by ecophysiological responses to high temperatures experienced during current and predicted thermal maxima, and although compensatory mechanisms may reduce top-down pressure on marginal populations, this is unlikely to be important within the current biogeographical context. Better understanding of the mechanisms underpinning climate-driven range shifts is important for habitat-forming species like kelps, which provide organic matter, create biogenic structure and alter environmental conditions for associated communities.

Species traits and climate velocity explain geographic range shifts in an ocean-warming hotspot.

Species' ranges are shifting globally in response to climate warming, with substantial variability among taxa, even within regions. Relationships between range dynamics and intrinsic species traits may be particularly apparent in the ocean, where temperature more directly shapes species' distributions. Here, we test for a role of species traits and climate velocity in driving range extensions in the ocean-warming hotspot of southeast Australia. Climate velocity explained some variation in range shifts, however, including species traits more than doubled the variation explained. Swimming ability, omnivory and latitudinal range size all had positive relationships with range extension rate, supporting hypotheses that increased dispersal capacity and ecological generalism promote extensions. We find independent support for the hypothesis that species with narrow latitudinal ranges are limited by factors other than climate. Our findings suggest that small-ranging species are in double jeopardy, with limited ability to escape warming and greater intrinsic vulnerability to stochastic disturbances.

The tropicalization of temperate marine ecosystems: climate-mediated changes in herbivory and community phase shifts.

Climate-driven changes in biotic interactions can profoundly alter ecological communities, particularly when they impact foundation species. In marine systems, changes in herbivory and the consequent loss of dominant habitat forming species can result in dramatic community phase shifts, such as from coral to macroalgal dominance when tropical fish herbivory decreases, and from algal forests to 'barrens' when temperate urchin grazing increases. Here, we propose a novel phase-shift away from macroalgal dominance caused by tropical herbivores extending their range into temperate regions. We argue that this phase shift is facilitated by poleward-flowing boundary currents that are creating ocean warming hotspots around the globe, enabling the range expansion of tropical species and increasing their grazing rates in temperate areas. Overgrazing of temperate macroalgae by tropical herbivorous fishes has already occurred in Japan and the Mediterranean. Emerging evidence suggests similar phenomena are occurring in other temperate regions, with increasing occurrence of tropical fishes on temperate reefs.