Variability in kelp forest structure along a latitudinal gradient in ocean temperature

Subtidal forests comprised of kelps and other canopy-forming macroalgae represent critically important marine habitats. Kelp forests exhibit high rates of primary productivity, magnified secondary productivity, support high levels of biodiversity and provide various ecosystem services. Compared with many other regions, kelp forests around the UK have been largely understudied despite their recognised importance and the possible impacts of environmental change factors. We conducted surveys at 12 kelp-dominated open-coast sites within four regions in the UK, spanning ~ 9° in latitude and ~ 3 °C in mean sea temperature. We used a combination of quadrat-scale abundance and percent cover data as well as transect-scale canopy cover estimates to quantify ecological structure at multiple spatial scales. Kelp forest structure varied significantly between sites (nested within region) and also between regions. Regional-scale differences were principally driven by a higher abundance/cover of Alaria esculenta at the colder northern regions (i.e. north and west Scotland), and the presence of the Lusitanian kelp Laminaria ochroleuca at some sites in the southernmost region (i.e. southwest England) but nowhere else. The kelp Laminaria hyperborea dominated all sites and varied significantly between sites but not regions. All assemblage-level and population-level response variables were highly variable between sites within regions, suggesting that environmental factors varying at corresponding spatial scales (e.g. wave exposure, turbidity, sedimentation) are important drivers of pattern. The detection of regional-scale variability suggests that predicted changes in ocean climate, particularly increased sea temperature, may lead to changes in kelp forest structure in the future, with poleward range contractions (for A. esculenta) and expansions (for L. ochroleuca) likely. However, as the distribution-abundance patterns of the assemblage dominant L. hyperborea did not vary predictably with ocean temperature at this spatial scale, the fundamental structure of these habitats may be more influenced by localised factors, at least in the short-term. The relative importance of multiple, concurrent environmental change factors in structuring UK kelp forests remains largely unknown

Turning on the heat: ecological response to simulated warming in the sea

Significant warming has been observed in every ocean, yet our ability to predict the consequences of oceanic warming on marine biodiversity remains poor. Experiments have been severely limited because, until now, it has not been possible to manipulate seawater temperature in a consistent manner across a range of marine habitats. We constructed a "hot-plate'' system to directly examine ecological responses to elevated seawater temperature in a subtidal marine system. The substratum available for colonisation and overlying seawater boundary layer were warmed for 36 days, which resulted in greater biomass of marine organisms and a doubling of space coverage by a dominant colonial ascidian. The "hot-plate'' system will facilitate complex manipulations of temperature and multiple stressors in the field to provide valuable information on the response of individuals, populations and communities to environmental change in any aquatic habitat

Assemblage turnover and taxonomic sufficiency of subtidal macroalgae at multiplespatial scales

Spatial variability in the structure of subtidal macroalgal assemblages in southwest Australia was examined at multiple spatial scales using a three-factor hierarchal design. Spatial extents ranged from metres (between quadrats) to many hundreds of kilometres (between regions), and the study encompassed N2000 km of temperate coastline. In addition, the influence of taxonomic resolution, from species level data to class level, on spatial patterns was investigated to assess the potential evolutionary timescales of the pattern and for developing cost effective regionally applicable surrogates for biodiversity monitoring. Almost 300 species were identified from 14 sites, representing considerable biodiversity and a significant subset of the total benthic macroalgal diversity in the region (∼1000 species). Multivariate variability was significant at all spatial scales examined, but most prominent at smallest spatial scales, regardless of taxonomic resolution. Assemblage and species turnover was pronounced at scales of metres to hundreds of metres. Generally, small scale patchiness was a ubiquitous pattern for all individual taxa examined, regardless of taxonomic resolution, while variability at the scale of 10s of km was less important. Even so, differences in spatial variability between taxa were observed, and ecological and historical reasons for such differences are proposed. Taxonomic aggregation to family level had minimal effect on spatial patterns, but aggregation to order level led to changes in some aspects of patterns of assemblage structure. The unique and speciose macroalgal assemblages on subtidal reefs in southwest Australia are shaped by a complex array of historical and contemporary processes that act at multiple spatial (and temporal) scales. Understanding the relative importance of these processes requires that further manipulative and correlative work is conducted across a range of ecologically-important spatial scales

Multiple-scale interactions structure macroinvertebrate assemblages associated with kelp understory algae

Aim: Kelp forests provide habitat and food that supports a high diversity of flora and fauna. While numerous studies have described macroinvertebrates associated with kelp blades, stipes and holdfasts, a key kelp forest microhabitat, epilithic understory algae, remains poorly studied. Here, we used a macroecological approach and artificial seaweed units (ASUs) to explore the effects of ocean climate, wave exposure and habitat complexity on understory algal associated macroinvertebrate assemblages within Laminaria hyperborea forests in the United Kingdom. Location: 9° latitudinal gradient along the north and west coasts of the United Kingdom. Methods: Replicate ASUs comprising four different habitat complexities were deployed under mature L. hyperborea at 2 sites (along a wave exposure gradient, separated by km) within each of 4 locations (separated by 100s km) nested within two regions (warm and cold, spanning 9° of latitude). After 5 months in situ, the ASUs were collected and macroinvertebrates were identified to species level and enumerated. Results: Habitat complexity and wave exposure both influenced macroinvertebrate assemblage structure, but results also showed clear effects of ocean climate, with macroinvertebrate assemblages differing between warm and cool regions, primarily driven by higher diversity and evenness in the warmer region and greater abundance in the cooler region. Main conclusions: Predicted warming and a shift to less complex turf-forming algal assemblages are likely to alter the structure of macroinvertebrate assemblages associated with understory algae, with potential implications for kelp forest food web dynamics

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

© 2020, The Author(s). 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

Resistance, Extinction and Everything in Between - The Diverse Responses of Seaweeds to Marine Heatwaves

Globally, anomalously warm temperature events have increased by 34% in frequency and 17% in duration from 1925 to 2016 with potentially major impacts on coastal ecosystems. These “marine heatwaves” (MHWs) have been linked to changes in primary productivity, community composition and biogeography of seaweeds, which often control ecosystem function and services. Here we journalarticle the literature on seaweed responses to MHWs, including 58 observations related to resistance, bleaching, changes in abundance, species invasions and local to regional extinctions. More records existed for canopy-forming kelps and bladed and filamentous turf-forming seaweeds than for canopy-forming fucoids, geniculate coralline turf and crustose coralline algae. Turf-forming seaweeds, especially invasive seaweeds, generally increased in abundance after a MHW, whereas native canopy-forming kelps and fucoids typically declined in abundance. We also found four examples of regional extinctions of kelp and fucoids following specific MHWs, events that likely have long term consequences for ecological structure and functioning. Although a relatively small number of studies have described impacts of MHWs on seaweed, the broad range of documented responses highlights the necessity of better baseline information regarding seaweed distributions and performance, and the need to study specific characteristics of MHWs that affect the vulnerability and resilience of seaweeds to these increasingly important climatic perturbations. A major challenge will be to disentangle impacts caused by the extreme temperature increases of MHWs itself from co-occurring potential stressors including altered current patterns, increasing herbivory, changes in water clarity and nutrient content, solar radiation and desiccation stress in the intertidal zone. With future increases anticipated in the intensity, duration and frequencies of MHWs, we expect to see more replacements of large long-lived habitat forming seaweeds with smaller ephemeral seaweeds, reducing the habitat structure and effective services seaweed-dominated reefs can provide

Ecological performance differs between range centre and trailing edge populations of a cold-water kelp:implications for estimating net primary productivity

Kelp forests are extensive, widely distributed and highly productive. However, despite their importance, reliable estimates of net primary productivity (NPP) are currently unknown for most species and regions. In particular, how performance and subsequent NPP change throughout a species range is lacking. Here, we attempted to resolve this by examining growth and performance of the boreal kelp, Laminaria digitata, from range centre and trailing edge regions in the United Kingdom. During the peak growth season (March/April), range-centre individuals were up to three times heavier and accumulated biomass twice as fast as their trailing-edge counterparts. This was not apparent during the reduced growth season (August/September), when populations within both regions had similar biomass profiles. In total, annual NPP estimates were considerably lower for trailing-edge (181±34 g C m−2 year−1) compared to range-centre (344±33 g C m−2 year−1) populations. Our first-order UK estimates of total standing stock and NPP for L. digitata suggest this species makes a significant contribution to coastal carbon cycling. Further work determining the ultimate fate of this organic matter is needed to understand the overall contribution of kelp populations to regional and global carbon cycles. Nevertheless, we highlight the need for large-scale sampling across multiple populations and latitudes to accurately evaluate kelp species’ contributions to coastal carbon cycling

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

Linking environmental variables with regional-scale variability in ecological structure and standing stock of carbon within UK kelp forests

Kelp forests represent some of the most productive and diverse habitats on Earth. Understanding drivers of ecological patterns at large spatial scales is critical for effective management and conservation of marine habitats. We surveyed kelp forests dominated by Laminaria hyperborea (Gunnerus) Foslie 1884 across 9° latitude and \u3e1000 km of coastline and measured a number of physical parameters at multiple scales to link ecological structure and standing stock of carbon with environmental variables. Kelp density, biomass, morphology and age were generally greater in exposed sites within regions, highlighting the importance of wave exposure in structuring L. hyperborea populations. At the regional scale, wave-exposed kelp canopies in the cooler regions (the north and west of Scotland) were greater in biomass, height and age than in warmer regions (southwest Wales and England). The range and maximal values of estimated standing stock of carbon contained within kelp forests was greater than in historical studies, suggesting that this ecosystem property may have been previously undervalued. Kelp canopy density was positively correlated with large-scale wave fetch and fine-scale water motion, whereas kelp canopy biomass and the standing stock of carbon were positively correlated with large-scale wave fetch and light levels and negatively correlated with temperature. As light availability and summer temperature were important drivers of kelp forest biomass, effective management of human activities that may affect coastal water quality is necessary to maintain ecosystem functioning, while increased temperatures related to anthropogenic climate change may impact the structure of kelp forests and the ecosystem services they provide