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Tasmania is an island state in south-eastern Australia that has a long and rich history of seaweed use, research, and development. It is a cool-temperate system with 750 macroalgal species currently described. Tasmanian Aboriginal peoples have lived on this land for at least 40,000 years utilising seaweed as food, shelter, water carriers and medicine, as well as for ceremonial reasons. Modern taxonomic investigations began with French naturalist Jacques-Julien Houtou de La Billardière in 1791, and there are 184 type specimens of seaweeds originating from Tasmania. Ecological and physiological studies of seaweed in Tasmania have focussed on the dominant large brown seaweeds (Laminariales and Fucales) and have contributed significantly to the global understanding of these systems, particularly related to community resilience, seaweed-urchin interactions, their habitat-forming role for other species, responses to global change, and restoration of lost habitat. Ocean warming and changing oceanography have caused a 95% decline in surface canopy cover of Macrocystis pyrifera in eastern Tasmania since the 1950s and led to a focus on restoring these lost forests. Tasmanian seaweed communities have a uniquely high proportion (up to ∼90%) of seaweeds that rely solely on CO2 for photosynthesis, which has implications for responses to ocean acidification. Tasmania has industries that use brown seaweeds for fucoidan extraction and beach-cast harvest for alginates, fertilisers, and feeds for agriculture. New aquaculture initiatives include integrated multi-trophic aquaculture, offshore kelp mariculture and Asparagopsis cultivation for bioactive products to reduce methane emissions in ruminants, as and the development of unexploited species including Caulerpa spp. for food.

In a future ocean, dissolved organic carbon (DOC) release by seaweed has been considered a pathway for organic carbon that is not incorporated into growth under carbon dioxide (CO2) enrichment/ocean acidification (OA). To understand the influence of OA on seaweed DOC release, a 21-day experiment compared the physiological responses of three seaweed species, two which operate CO2 concentrating mechanisms (CCMs), Ecklonia radiata (C. Agardh) J. Agardh and Lenormandia marginata (Hooker F. and Harvey) and one that only uses CO2 (non-CCM), Plocamium cirrhosum (Turner) M.J. Wynne. These two groups (CCM and non-CCM) are predicted to respond differently to OA dependent on their affinities for Ci (defined as CO2 + bicarbonate, HCO3−). Future ocean CO2 treatment did not drive changes to seaweed physiology—growth, Ci uptake, DOC production, photosynthesis, respiration, pigments, % tissue carbon, nitrogen, and C:N ratios—for any species, regardless of Ci uptake method. Our results further showed that Ci uptake method did not influence DOC release rates under OA. Our results show no benefit of elevated CO2 concentrations on the physiologies of the three species under OA and suggest that in a future ocean, photosynthetic CO2 fixation rates of these seaweeds will not increase with Ci concentration.

Kelp aquaculture is an emerging industry outside of Asia. To be successful, this industry requires a reliable production of seedstock, the optimisation of which greatly benefits from a detailed physiological understanding of the microscopic life-cycle stages of the cultured species. This study investigated the impact of six zoospore densities (10–278 mm−2) on the subsequent development of Ecklonia radiata gametophytes and sporophytes. The results showed that germination rates and sex ratio were unaffected by initial zoospore density, but there were significant effects on gametophyte size and sporophyte production. After two weeks, female gametophytes were largest at an initial zoospore density of 40 mm−2 while male gametophytes grew largest at densities below 40 mm−2, but after four weeks gametophyte size showed a negative relationship with initial zoospore density. Significantly more sporophytes developed at initial zoospore densities below 40 individuals mm−2 and no sporophytes were observed at the highest density (271 zoospores mm−2). These results clearly show the importance of initial zoospore density in optimising the nursery stage of kelp aquaculture.

The combined culture of fed species (bivalves, fish) and macroalgae, known as integrated multi-trophic aquaculture (IMTA), has been suggested as a method of mitigating localised nitrogen (N) increase from aquaculture, whilst simultaneously culturing macroalgae for commercial applications. The development of IMTA requires an understanding of the N ecophysiology of candidate macroalga species. We examined seasonal variations in ammonium (NH4+) uptake kinetics, carbon to nitrogen (C:N) ratio, pigment content and soluble tissue N of four macroalgae of the phylum Ochrophyta, Ecklonia radiata, Macrocystis pyrifera, Lessonia corrugata, and Phyllospora comosa, from Tasmania, Australia. This study aimed to determine, (1) if the N physiology of the four macroalgal species was suitable for IMTA applications and (2) whether the species had seasonal variations in N ecophysiology which would influence their suitability for IMTA. Macrocystis pyrifera, L. corrugata, and E. radiata exhibited saturable NH4+ uptake kinetics, with a maximum uptake rate (Vmax) during spring, summer and autumn of 200, 45.8 and 45 μmol gDW-1 h-1 and half-saturation constants (Ks) of 361.3, 104.2 and 121 μM, respectively. Phyllospora comosa exhibited biphasic uptake patterns for three out of four months sampled. There were no noticeable seasonal patterns in pigment content or soluble tissue N for any species. C:N ratios increased from spring (October) to autumn (March) in both E. radiata (28.34 – 47.83) and P. comosa (24.99 – 51.62), indicating progressive N limitation though summer and into autumn. Results suggest that M. pyrifera and P. comosa are most suitable for IMTA due to their high NH4+ uptake potential.

Our scientific understanding of climate change makes clear the necessity for both emission reduction and carbon dioxide removal CDR The ocean with its large surface area great depths and long coastlines is central to developing CDR approaches commensurate with the scale needed to limit warming to below 2 C Many proposed marine CDR approaches rely on spatial upscaling along with enhancement and or acceleration of the rates of naturally occurring processes One such approach is ocean afforestation which involves offshore transport and concurrent growth of nearshore macroalgae seaweed followed by their export into the deep ocean The purposeful occupation for months of open ocean waters by macroalgae which do not naturally occur there will probably affect offshore ecosystems through a range of biological threats including altered ocean chemistry and changed microbial physiology and ecology Here we present model simulations of ocean afforestation and link these to lessons from other examples of offshore dispersal including rafting plastic debris and discuss the ramifications for offshore ecosystems We explore what additional metrics are required to assess the ecological implications of this proposed CDR In our opinion these ecological metrics must have equal weight to CDR capacity in the development of initial trials pilot studies and potential licensing 2022 Springer Nature Limited

Lessonia (order Laminariales) is a kelp genus restricted to the temperate southern hemisphere, where species form dense forests from the low intertidal to 25 m depth at wave exposed sites. Lessonia spp. are among the most harvested kelps globally due to their importance in providing raw materials for food, cosmetics, bioactive and biomedical industries. Over-harvesting of natural beds can negatively affect Lessonia populations and the many species that depend on these habitats, including commercially important fish and molluscs, but good harvest management plans reduce these impacts on natural Lessonia stocks. However, the increasing demand for raw materials will likely only be met by aquaculture for which Lessonia shows high potential in pilot scale studies undertaken in Chile, New Zealand, and Australia. In this concise review, we highlight the current knowledge of Lessonia spp. taxonomy and distribution, life history, ecology and ecosystem services, wild harvest, aquaculture, and commercial applications. We discuss future research directions.

Most research investigating how ocean warming and acidification will impact marine species has focused on visually dominant species, such as kelps and corals, while ignoring visually cryptic species such as crustose coralline algae (CCA). CCA are important keystone species that provide settlement cues for invertebrate larvae and can be highly sensitive to global ocean change. However, few studies have assessed how CCA respond to low emission scenarios or conditions. In a laboratory experiment, we examined the responses of temperate CCA assemblages to combined warming and acidification projected under low, medium, and high emissions. Net calcification and net photosynthesis significantly declined in all emissions scenarios, while significant reductions in relative growth rates and increases in percentage bleaching were observed in the highest emission scenario. The negative responses of CCA to both low and medium emissions suggest that they may be adversely impacted by combined warming and acidification by 2030 if current emissions are sustained. This will have far reaching consequences for commercially important invertebrates that rely on them to induce settlement of larvae. These findings highlight the need to take rapid action to preserve these critical keystone species and the valuable services they provide.

Macrocystis pyrifera is a major habitat forming kelp in coastal ecosystems of temperate regions of the northern and southern hemispheres. We investigated the seasonal occurrence of adult sporophytes, morphological characteristics, and reproductive phenology at two sites within a wave-protected harbour and two wave-exposed sites in southern New Zealand every 3–4 months between 2012 and 2013. Seasonality in reproduction was assessed via the number of sporophylls, the occurrence of sori on sporophylls, and non-sporophyllous laminae (fertile pneumatocyst-bearing blades and fertile apical scimitars), meiospore release, and germination. We found that M. pyrifera was present and reproductive year-round in three of the four sites, and patterns were similar for the wave-exposure conditions. Sori were found on pneumatocyst-bearing blades and apical scimitars in addition to the sporophylls, and viable meiospores were released from all three types of laminae. Morphological variations between sites with different wave exposure indicate that sporophytes from wave-protected sites have bigger blades and holdfasts and are longer than those from wave-exposed sites. We discuss the implications of these biological variables for the ecology of M. pyrifera inhabiting different wave exposure environments in southern New Zealand.

Ensuring that global warming remains, Ocean afforestation is considered as an important method to remove gigatons of CO2 from the atmosphere. Here the authors use the Great Atlantic Sargassum Belt as a natural analogue to show that the efficacy of ocean afforestation is determined by complicated feedbacks with the Earth system.

The kelp Lessonia corrugata (Ochrophyta, Laminariales) is being developed for integrated multi-trophic aquaculture (IMTA) trials in the vicinity of salmon cages in Tasmania, Australia. Gametophytes are vegetally maintained before seeding on hatchery twine; however, the optimal temperature and light conditions for growth and sexual development are unknown. We measured vegetative size of female and male gametophytes and sexual development of females over a range of temperatures and irradiances using a temperature gradient table and neutral density light filters. Over a 4-week experiment, gametophytes were exposed to a combination of thermal (5.7–24.9 °C) and irradiance (10–100 μmol photons m−2 s−1) gradients, to assess biological performance. At the temperature extremes (hottest = 24.9 °C, coldest = 5.7 °C), we observed the critical thermal limits for this species and the results reveal a narrow optimal temperature range for growth and sexual development between 15.7 and 17.9 °C, with irradiances between 40 and 100 μmol photons m−2 s−1 resulting in fertile female gametophytes. Lessonia corrugata inhabits a small geographic range, found only around Tasmania, south of the Australian mainland, hence oceanic changes such as ongoing increases in sea surface temperatures (SSTs), and altered irradiance regimes may limit recruitment of the early microscopic life stages in the future. Our findings provide optimised culture conditions for aquaculture and information to predict the future geographic range of L. corrugata under ocean global change.

In coastal ecosystems, seaweeds provide habitat and a food source for a variety of species including herbivores of commercial importance. In these systems seaweeds are the ultimate source of energy with any changes in the seaweeds invariably affecting species of higher trophic levels. Seaweeds are rich sources of nutritionally important compounds such as polyunsaturated fatty acids (PUFA) and are particularly rich in long-chain (≥ C20) PUFA (LC-PUFA). In southern Australia, the ‘Great Southern Reef’ has one of the most diverse assemblages of seaweeds in the world, which support highly productive fisheries and have been recognised as a promising resource of omega-3 LC-PUFA. Despite this, there is little information on the biochemical composition of most species and how it varies between sites and seasons. To address this knowledge gap, we undertook a survey to assess seasonal variability in the biochemical composition (fatty acids and nitrogen content) of abundant understory seaweeds across three sites in eastern Tasmania. The availability of nutritional compounds differed between sites and was primarily driven by differences in the biomass and the biochemical composition of the nutritious red seaweeds at each site. This variability may explain regional differences in the productivity of commercial fisheries. At the species level, seasonal changes in fatty acid composition were highly variable between species and sites, indicating that multiple environmental drivers influence fatty acid composition of seaweeds in this system. This finding suggests that commercial harvest of seaweeds from eastern Tasmania will need to consider species and site-specific variability in fatty acid composition.

Macroalgal growth in temperate coastal ecosystems is primarily regulated by light and inorganic nitrogen availability. The effect of light (photon irradiance) on NO3− and NH4+ uptake, and NO3− assi...

Marine heatwaves are extreme events that can have profound and lasting impacts on marine species. Field observations have shown seaweeds to be highly susceptible to marine heatwaves, but the physiological drivers of this susceptibility are poorly understood. Furthermore, the effects of marine heatwaves in conjunction with ocean warming and acidification are yet to be investigated. To address this knowledge gap, we conducted a laboratory culture experiment in which we tested the growth and physiological responses of Phyllospora comosa juveniles from the southern extent of its range (43–31°S) to marine heatwaves, ocean warming and acidification. We used a ‘collapsed factorial design’ in which marine heatwaves were superimposed on current (today's pH and temperature) and future (pH and temperature projected by 2100) ocean conditions. Responses were tested both during the heatwaves, and after a 7‐day recovery period. Heatwaves reduced net photosynthetic rates in both current and future conditions, while respiration rates were elevated under heatwaves in the current conditions only. Following the recovery period, there was little evidence of heatwaves having lasting negative effects on growth, photosynthesis or respiration. Exposure to heatwaves, future ocean conditions or both caused an increase in the degree of saturation of fatty acids. This adjustment may have counteracted negative effects of elevated temperatures by decreasing membrane fluidity, which increases at higher temperatures. Furthermore, P. comosa appeared to down‐regulate the energetically expensive carbon dioxide concentrating mechanism in the future conditions with a reduction in δ13C values detected in these treatments. Any saved energy arising from this down‐regulation was not invested in growth and was likely invested in the adjustment of fatty acid composition. This adjustment is a mechanism by which P. comosa and other seaweeds may tolerate the negative effects of ocean warming and marine heatwaves through benefits arising from ocean acidification.

Highlights: • First study to compare microplastic effects over a wide biogeographical range • Comparison between natural inorganic microparticles and plastic microparticles • Significant effects on byssus production, respiration and clearance rates, but small effect sizes • No ecologically relevant difference between impact of plastic and natural inorganic microparticles on Mytilidae Abstract: Microplastics are ubiquitous in the marine environment and studies on their effects on benthic filter feeders at least partly revealed a negative influence. However, it is still unclear whether the effects of microplastics differ from those of natural suspended microparticles, which constitute a common stressor in many coastal environments. We present a series of experiments that compared the effects of six-week exposures of marine mussels to two types of natural particles (red clay and diatom shells) to two types of plastic particles (Polymethyl Methacrylate and Polyvinyl Chloride). Mussels of the family Mytilidae from temperate regions (Japan, Chile, Tasmania) through subtropical (Israel) to tropical environments (Cabo Verde) were exposed to concentrations of 1.5 mg/L, 15 mg/L and 150 mg/L of the respective microparticles. At the end of this period, we found significant effects of suspended particles on respiration rate, byssus production and condition index of the animals. There was no significant effect on clearance rate and survival. Surprisingly, we observed only small differences between the effects of the different types of particles, which suggests that the mussels were generally equally robust towards exposure to variable concentrations of suspended solids regardless of whether they were natural or plastic. We conclude, that microplastics and suspended solids elicit similar effects on the tested response variables, and that both types of microparticles mainly cause acute responses rather than more persistent carry-over effects.

Marine coastal zones are highly productive, and dominated by engineer species (e.g. macrophytes, molluscs, corals) that modify the chemistry of their surrounding seawater via their metabolism, causing substantial fluctuations in oxygen, dissolved inorganic carbon, pH, and nutrients. The magnitude of these biologically driven chemical fluctuations is regulated by hydrodynamics, can exceed values predicted for the future open ocean, and creates chemical patchiness in subtidal areas at various spatial (µm to meters) and temporal (minutes to months) scales. Although the role of hydrodynamics is well explored for planktonic communities, its influence as a crucial driver of benthic organism and community functioning is poorly addressed, particularly in the context of ocean global change. Hydrodynamics can directly modulate organismal physiological activity or indirectly influence an organism's performance by modifying its habitat. This review addresses recent developments in (i) the influence of hydrodynamics on the biological activity of engineer species, (ii) the description of chemical habitats resulting from the interaction between hydrodynamics and biological activity, (iii) the role of these chemical habitat as refugia against ocean acidification and deoxygenation, and (iv) how species living in such chemical habitats may respond to ocean global change. Recommendations are provided to integrate the effect of hydrodynamics and environmental fluctuations in future research, to better predict the responses of coastal benthic ecosystems to ongoing ocean global change.

Carbon sequestration is defined as the secure storage of carbon-containing molecules for >100 years, and in the context of carbon dioxide removal for climate mitigation, the origin of this CO2 is from the atmosphere. On land, trees globally sequester substantial amounts of carbon in woody biomass, and an analogous role for seaweeds in ocean carbon sequestration has been suggested. The purposeful expansion of natural seaweed beds and aquaculture systems, including into the open ocean (ocean afforestation), has been proposed as a method of increasing carbon sequestration and use in carbon trading and offset schemes. However, to verify whether CO2 fixed by seaweeds through photosynthesis leads to carbon sequestration is extremely complex in the marine environment compared to terrestrial systems, because of the need to jointly consider: the comparatively rapid turnover of seaweed biomass, tracing the fate of carbon via particulate and dissolved organic carbon pathways in dynamic coastal waters, and the key role of atmosphere-ocean CO2 exchange. We propose a Forensic Carbon Accounting approach, in which a thorough analysis of carbon flows between the atmosphere and ocean, and into and out of seaweeds would be undertaken, for assessing the magnitude of CO2 removal and robust attribution of carbon sequestration to seaweeds.

Inorganic carbon, nitrogen and phosphorus are the main elements required by seaweeds for photosynthesis and growth. This review focusses mainly on nitrogen, but the roles of carbon and phosphorus, ...

Laboratory studies that test the responses of coastal organisms to ocean acidification (OA) typically use constant pH regimes which do not reflect coastal systems, such as seaweed beds, where pH fluctuates on diel cycles. Seaweeds that use CO2 as their sole inorganic carbon source (non-carbon dioxide concentrating mechanism species) are predicted to benefit from OA as concentrations of dissolved CO2 increase, yet this prediction has rarely been tested, and no studies have tested the effect of pH fluctuations on non-CCM seaweeds. We conducted a laboratory experiment in which two ecologically dominant non-CCM red seaweeds (Callophyllis lambertii and Plocamium dilatatum) were exposed to four pH treatments: two static, pHT 8.0 and 7.7 and two fluctuating, pHT 8.0 ± 0.3 and 7.7 ± 0.3. Fluctuating pH reduced growth and net photosynthesis in C. lambertii, while P. dilatatum was unaffected. OA did not benefit P. dilatatum, while C. lambertii displayed elevated net photosynthetic rates. We provide evidence that carbon uptake strategy alone cannot be used as a predictor of seaweed responses to OA and highlight the importance of species-specific sensitivity to [H+]. We also emphasize the importance of including realistic pH fluctuations in experimental studies on coastal organisms.

Our scientific understanding of climate change makes clear the necessity for both emission reduction and carbon dioxide removal (CDR). The ocean with its large surface area, great depths and long coastlines is central to developing CDR approaches commensurate with the scale needed to limit warming to below 2 °C. Many proposed marine CDR approaches rely on spatial upscaling along with enhancement and/or acceleration of the rates of naturally occurring processes. One such approach is 'ocean afforestation', which involves offshore transport and concurrent growth of nearshore macroalgae (seaweed), followed by their export into the deep ocean. The purposeful occupation for months of open ocean waters by macroalgae, which do not naturally occur there, will probably affect offshore ecosystems through a range of biological threats, including altered ocean chemistry and changed microbial physiology and ecology. Here, we present model simulations of ocean afforestation and link these to lessons from other examples of offshore dispersal, including rafting plastic debris, and discuss the ramifications for offshore ecosystems. We explore what additional metrics are required to assess the ecological implications of this proposed CDR. In our opinion, these ecological metrics must have equal weight to CDR capacity in the development of initial trials, pilot studies and potential licensing.

Macroalgae, with their various morphologies, are ubiquitous throughout the world’s oceans and provide ecosystem services to a multitude of organisms. Water motion is a fundamental physical parameter controlling the mass transfer of dissolved carbon and nutrients to and from the macroalgal surface, but measurements of flow speed and turbulence within and above macroalgal canopies are lacking. This information is becoming increasingly important as macroalgal canopies may act as refugia for calcifying organisms from ocean acidification (OA); and the extent to which they act as refugia is driven by water motion. Here we report on a field campaign to assess the flow speed and turbulence within and above natural macroalgal canopies at two depths (3 and 6 m) and two sites (Ninepin Point and Tinderbox) in Tasmania, Australia in relation to the canopy height and % cover of functional forms. Filamentous groups made up the greatest proportion (75%) at both sites and depth while foliose groups were more prevalent at 3 than at 6 m. Irrespective of background flows, depth or site, flow speeds within the canopies were –1 – a ∼90% reduction in flow speeds compared to above the canopy. Turbulent kinetic energy (TKE) within the canopies was up to two orders of magnitude lower (2 s–2) than above the canopies, with higher levels of TKE within the canopy at 3 compared to 6 m. The significant damping effect of flow and turbulence by macroalgae highlights the potential of these ecosystems to provide a refugia for vulnerable calcifying species to OA.

Dissolved organic carbon (DOC) release by seaweeds (marine macroalgae) is a critical component of the coastal ocean biogeochemical carbon cycle but is an aspect of seaweed carbon physiology that we know relatively little about. Seaweed-derived DOC is found throughout coastal ecosystems and supports multiple food web linkages. Here, we discuss the mechanisms of DOC release by seaweeds and group them into passive (leakage, requires no energy) and active release (exudation, requires energy) with particular focus on the photosynthetic "overflow" hypothesis. The release of DOC from seaweeds was first studied in the 1960s, but subsequent studies use a range of units hindering evaluation: we convert published values to a common unit (μmol C · g DW−1 · h−1) allowing comparisons between seaweed phyla, functional groups, biogeographic region, and an assessment of the environmental regulation of DOC production. The range of DOC release rates by seaweeds from each phylum under ambient environmental conditions was 0–266.44 μmol C · g DW−1 · h−1 (Chlorophyta), 0–89.92 μmol C · g DW−1 · h−1 (Ochrophyta), and 0–41.28 μmol C · g DW−1· h−1 (Rhodophyta). DOC release rates increased under environmental factors such as desiccation, high irradiance, non-optimal temperatures, altered salinity, and elevated dissolved carbon dioxide (CO2) concentrations. Importantly, DOC release was highest by seaweeds that were desiccated (

Macrocystis pyrifera is a major habitat forming kelp in coastal ecosystems of temperate regions of the northern and southern hemispheres. We investigated the seasonal occurrence of adult sporophytes, morphological characteristics, and reproductive phenology at two sites within a wave-protected harbour and two wave-exposed sites in southern New Zealand every 3-4 months between 2012 and 2013. Seasonality in reproduction was assessed via the number of sporophylls, the occurrence of sori on sporophylls, and non-sporophyllous laminae (fertile pneumatocyst-bearing blades and fertile apical scimitars), meiospore release, and germination. We found that M. pyrifera was present and reproductive year-round in three of the four sites, and patterns were similar for the wave-exposure conditions. Sori were found on pneumatocyst-bearing blades and apical scimitars in addition to the sporophylls, and viable meiospores were released from all three types of laminae. Morphological variations between sites with different wave exposure indicate that sporophytes from wave-protected sites have bigger blades and holdfasts and are longer than those from wave-exposed sites. We discuss the implications of these biological variables for the ecology of M. pyrifera inhabiting different wave exposure environments in southern New Zealand.

Ocean warming, ocean acidification and overfishing are major threats to the structure and function of marine ecosystems. Driven by increasing anthropogenic emissions of CO2, ocean warming is leading to global redistribution of marine biota and altered ecosystem dynamics, while ocean acidification threatens the ability of calcifying marine organisms to form skeletons due to decline in saturation state of carbonate Ω and pH. In Tasmania, the interaction between overfishing of sea urchin predators and rapid ocean warming has caused a phase-shift from productive kelp beds to overgrazed sea urchin barren grounds, however potential impacts of ocean acidification on this system have not been considered despite this threat for marine ecosystems globally. Here we use automated loggers and point measures of pH, spanning kelp beds and barren grounds, to reveal that kelp beds have the capacity to locally ameliorate effects of ocean acidification, via photosynthetic drawdown of CO2, compared to unvegetated barren grounds. Based on meta-analysis of anticipated declines in physiological performance of grazing urchins to decreasing pH and assumptions of nil adaptation, future projection of OA across kelp-barrens transition zones reveals that kelp beds could act as important pH refugia, with urchins potentially becoming increasingly challenged at distances >40 m from kelp beds. Using spatially explicit simulation of physicochemical feedbacks between grazing urchins and their kelp prey, we show a stable mosaicked expression of kelp patches to emerge on barren grounds. Depending on the adaptative capacity of sea urchins, future declines in pH appear poised to further alter phase-shift dynamics for reef communities; thus, assessing change in spatial-patterning of reef-scapes may indicate cascading ecological impacts of ocean acidification.

Highlights: • Mussel BCI was 14% lower after exposure to PVC particles in comparison to red clay. • Byssus production, respiration and survival did not differ between particle types. • Mussels suffered little from the five-week particle exposure up to 150 mg l−1. Abstract: Effects of microplastics on marine taxa have become a focal point in marine experimental biology. Almost all studies so far, however, assessed the influence of microplastics on animals only in relation to a zero-particle group. Documented microplastic impacts may thus be overestimated, since many marine species also experience natural suspended solids as a stressor. Here, we compared the effects of polyvinyl chloride (PVC) and red clay (mean for both particles: ~12–14 μm) on the Mediterranean mussel Mytilus galloprovincialis across three particle concentrations (1.5, 15, 150 mg l−1). Exposure to PVC for 35 days lowered mussel body condition index by 14% in relation to clay, but no difference in byssus production, respiration and survival rates emerged between the two particle types. This suggests that the effects of synthetic particles on filter feeders may emulate those of natural suspended solids, and highlights the importance of including natural particles in microplastic exposure studies.

Kelp forests around the world are under increasing pressure from anthropogenic stressors. A widespread consequence is that in many places, complex and highly productive kelp habitats have been replaced by structurally simple and less productive turf algae habitats. Turf algae habitats resist re‐establishment of kelp via recruitment inhibition; however, little is known about the specific mechanisms involved. One potential factor is the chemical environment within the turf algae and into which kelp propagules settle and develop. Using laboratory trials, we illustrate that the chemical microenvironment (O2 concentration and pH) 0.0–50 mm above the substratum within four multispecies macroalgal assemblages (including a turf‐sediment assemblage and an Ecklonia radiata kelp‐dominated assemblage) are characterized by elevated O2 and pH relative to the surrounding seawater. Notably however, O2 and pH were significantly higher within turf‐sediment assemblages than in kelp‐dominated assemblages, and at levels that have previously been demonstrated to impair the photosynthetic or physiological capacity of kelp propagules. Field observations of the experimental assemblages confirmed that recruitment of kelp was significantly lower into treatments with dense turf algae than in the kelp‐dominated assemblages. We demonstrate differences between the chemical microenvironments of kelp and turf algae assemblages that correlate with differences in kelp recruitment, highlighting how degradation of kelp habitats might result in the persistence of turf algae habitats and the localized absence of kelp.

Increased plant biomass is observed in terrestrial systems due to rising levels of atmospheric CO2, but responses of marine macroalgae to CO2 enrichment are unclear. The 200% increase in CO2 by 2100 is predicted to enhance the productivity of fleshy macroalgae that acquire inorganic carbon solely as CO2 (non‐carbon dioxide‐concentrating mechanism [CCM] species—i.e., species without a carbon dioxide‐concentrating mechanism), whereas those that additionally uptake bicarbonate (CCM species) are predicted to respond neutrally or positively depending on their affinity for bicarbonate. Previous studies, however, show that fleshy macroalgae exhibit a broad variety of responses to CO2 enrichment and the underlying mechanisms are largely unknown. This physiological study compared the responses of a CCM species (Lomentaria australis) with a non‐CCM species (Craspedocarpus ramentaceus) to CO2 enrichment with regards to growth, net photosynthesis, and biochemistry. Contrary to expectations, there was no enrichment effect for the non‐CCM species, whereas the CCM species had a twofold greater growth rate, likely driven by a downregulation of the energetically costly CCM(s). This saved energy was invested into new growth rather than storage lipids and fatty acids. In addition, we conducted a comprehensive literature synthesis to examine the extent to which the growth and photosynthetic responses of fleshy macroalgae to elevated CO2 are related to their carbon acquisition strategies. Findings highlight that the responses of macroalgae to CO2 enrichment cannot be inferred solely from their carbon uptake strategy, and targeted physiological experiments on a wider range of species are needed to better predict responses of macroalgae to future oceanic change.

To assess the suitability of southern-Australian macroalgae as potential marine resources for fatty acids (FA), and in particular polyunsaturated fatty acids (PUFA), analysis of 61 species, comprising of 11 Chlorophyta, 17 Phaeophyceae (Ochrophyta) and 33 Rhodophyta, was conducted. Total fatty acid (TFA) concentrations varied considerably (between 0.6 and 7.8 in % of dry weight (DW)) between species, with on average the highest concentrations being in the Phaeophyceae, then the Chlorophyta, and with the Rhodophyta recording the lowest average concentrations. Results revealed significant differences in the fatty acid profiles of the three algal groups. Most species exhibit high proportions of PUFA in their fatty acid profile and a low ratio of n-6/n-3 PUFA. These properties highlight the potential for southern-Australian macroalgae to be used for these FA in food, animal feed and nutraceutical applications.

Ocean acidification is a global phenomenon, but it is overlaid by pronounced regional variability modulated by local physics, chemistry and biology. Recognition of its multifaceted nature and the interplay of acidification with other ocean drivers has led to international and regional initiatives to establish observation networks and develop unifying principles for biological responses. There is growing awareness of the threat presented by ocean acidification to ecosystem services and the socio-economic consequences are becoming increasingly apparent and quantifiable. In this higher-CO2 world, future challenges involve better design and rigorous testing of adaptation, mitigation and intervention options to offset the effects of ocean acidification at scales ranging from local to regional.

International audience; Marine life is controlled by multiple physical and chemical drivers and by diverse ecological processes. Many of these oceanic properties are being altered by climate change and other anthropogenic pressures. Hence, identifying the influences of multifaceted ocean change, from local to global scales, is a complex task. To guide policy‐making and make projections of the future of the marine biosphere, it is essential to understand biological responses at physiological, evolutionary and ecological levels. Here, we contrast and compare different approaches to multiple driver experiments that aim to elucidate biological responses to a complex matrix of ocean global change. We present the benefits and the challenges of each approach with a focus on marine research, and guidelines to navigate through these different categories to help identify strategies that might best address research questions in fundamental physiology, experimental evolutionary biology and community ecology. Our review reveals that the field of multiple driver research is being pulled in complementary directions: the need for reductionist approaches to obtain process‐oriented, mechanistic understanding and a requirement to quantify responses to projected future scenarios of ocean change. We conclude the review with recommendations on how best to align different experimental approaches to contribute fundamental information needed for science‐based policy formulation.

Seaweeds are able to modify the chemical environment at their surface, in a micro‐zone called the diffusive boundary layer (DBL), via their metabolic processes controlled by light intensity. Depending on the thickness of the DBL, sessile invertebrates such as calcifying bryozoans or tube‐forming polychaetes living on the surface of the blades can be affected by the chemical variations occurring in this microlayer. Especially in the context of ocean acidification (OA), these microhabitats might be considered as a refuge from lower pH, because during the day photosynthesis temporarily raises the pH to values higher than in the mainstream seawater. We assessed the thickness and the characteristics of the DBL at two pH levels (today's average surface ocean pH 8.1 and a reduced pH predicted for the end of the century, pH 7.7) and seawater flows (slow, 0.5 and fast, >8 cm/s) on Ecklonia radiata (kelp) blades. Oxygen and pH profiles from the blade surface to the mainstream seawater were measured with O2 and pH microsensors for both bare blades and blades colonized by the bryozoan Membranipora membranacea. The DBL was thicker in slow flow compared with fast flow and the presence of bryozoans increased the DBL thickness and shaped the DBL gradient in dark conditions. Net production was increased in the low pH condition, increasing the amount of oxygen in the DBL in both bare and epiphytized blades. This increase drove the daily pH fluctuations at the blade surface, shifting them towards higher values compared with today's pH. The presence of bryozoans led to lower oxygen concentrations in the DBL and more complex pH fluctuations at the blade surface, particularly at pH 7.7. Overall, this study, based on microprofiles, shows that, in slow flow, DBL microenvironments at the surface of the kelps may constitute a refuge from OA with pH values higher than those of the mainstream seawater. For calcifying organisms, it could also represent training ground for harsh conditions, with broad daily pH and oxygen fluctuations. These chemical microenvironments, biologically shaped by the macrophytes, are of great interest for the resilience of coastal ecosystems in the context of global change. -- Keywords : Bryozoan ; Ecomechanics ; Epiphytism ; Hydrodynamic ; Microhabitat ; pH ; Seaweed.

Sarcothalia lanceata is a broad-bladed, New Zealand red alga. The tetrasporic life stage contains a lambda-carrageenan that has a strong potential for commercial utilisation. The male and female gametophytic life stages contain kappa-II carrageenan that also has commercial potential. However, fundamental information on the growth and variation in carrageenan content of this species in the wild is necessary to underpin possible aquaculture of this species. Therefore, growth in blade length and width was assessed for male and female gametophytes, and tetrasporophytes in two S. lanceata populations in New Zealand’s South Island, and monthly seawater nutrient and tissue nutrients, and carrageenan constituent sugar levels were analysed. Blade length did not vary significantly over time or between life history phases; however, blade width varied significantly for both factors with most temporal variation occurring in male blades. Analysis of seawater and tissue nutrient levels suggested that these populations are nutrient limited year-round. Constituent sugars varied temporally and between life history phases in most cases. Relative 3,6-anhydrogalactitol peracetate (for the males and females) and galactitol peracetate levels (all life stages) were higher in winter than in summer, while glucitol peracetate and xylitol peracetate levels were generally lower in winter than summer. The distinctive morphologies of the male and female plants of this species mean there is potential to cultivate and then separately harvest male and female plants in addition to tetrasporophytic plants. Male blades contain more 3,6-anhydrogalactitol peracetate and galactitol peracetate than female blades, and their morphology may make them less prone to breakage in an aquaculture situation. Experimentation on spore release and growth and the potential effects of nutrient manipulation on early stages are suggested as the direction of future research.

A series of semi-continuous incubation experiments were conducted with the coccolithophore Emiliania huxleyi strain NIWA1108 (Southern Ocean isolate) to examine the effects of five environmental drivers (nitrate and phosphate concentrations, irradiance, temperature, and partial pressure of CO2 (pCO2)) on both the physiological rates and elemental composition of the coccolithophore. Here, we report the alteration of the elemental composition of E. huxleyi in response to the changes in these environmental drivers. A series of dose–response curves for the cellular elemental composition of E. huxleyi were fitted for each of the five drivers across an environmentally representative gradient. The importance of each driver in regulating the elemental composition of E. huxleyi was ranked using a semi-quantitative approach. The percentage variations in elemental composition arising from the change in each driver between present-day and model-projected conditions for the year 2100 were calculated. Temperature was the most important driver controlling both cellular particulate organic and inorganic carbon content, whereas nutrient concentrations were the most important regulator of cellular particulate nitrogen and phosphorus of E. huxleyi. In contrast, elevated pCO2 had the greatest influence on cellular particulate inorganic carbon to organic carbon ratio, resulting in a decrease in the ratio. Our results indicate that the different environmental drivers play specific roles in regulating the elemental composition of E. huxleyi with wide-reaching implications for coccolithophore-related marine biogeochemical cycles, as a consequence of the regulation of E. huxleyi physiological processes.

Invasive seaweeds have the potential to disrupt ecosystem functioning if they are unsuitable hosts for the small mobile invertebrates (epifauna) that are an important trophic link between benthic primary producers and higher trophic levels. The Asian kelp Undaria pinnatifida has successfully invaded many coastal regions worldwide. We compared the epifaunal assemblage on U. pinnatifida with epifauna of 7 co-occurring, canopy-forming native brown seaweed species in southern New Zealand to help understand the effect of the invasive species on shallow subtidal ecosystems. The density, diversity and composition of epifauna across the 8 seaweeds were much more strongly related to host morphology than to the geographic origin of the host. U. pinnatifida and several native seaweeds with similarly simple morphologies supported relatively depauperate epifaunal assemblages dominated by copepods. More structurally complex native seaweeds supported more abundant and diverse epifaunal assemblages containing lower proportions of copepods and higher proportions of amphipods and other epifaunal groups. Our results indicate that abundances of epifauna at the ecosystem level will be reduced if U. pinnatifida displaces more structurally complex native seaweed species that host more diverse and dense epifaunal assemblages. The findings suggest that morphological complexity may be key to predicting the impacts of invasive seaweeds on epifaunal assemblages, and potentially on food webs, in other geographic regions.

The threat posed by ocean acidification (OA) to the diversity and productivity of New Zealand marine ecosystems is assessed in a synthesis of published trends and impacts. A 20-year time series in Subantarctic water, and a national coastal monitoring programme, provide insight into pH variability, and context for experimental design, modelling and projections. A review of the potential impact of changes in the carbonate system on the major phyla in New Zealand waters confirms international observations that calcifying organisms, and particularly their early life-history stages, are vulnerable. The synthesis considers ecosystem and socio-economic impacts, and identifies current knowledge gaps and future research directions, including mechanistic studies of OA sensitivity. Advanced ecosystem models of OA, that incorporate the indirect effects of OA and interactions with other climate stressors, are required for robust projection of the future status of New Zealand marine ecosystems.

Water motion is the least studied of the primary drivers of seaweed productivity. In 50% of field studies, seaweed growth and productivity is reduced in slow compared to fast flows and this is partly explained by the reduced transport of dissolved nutrients (e.g. nitrogen) across thicker diffusion boundary layers (DBLs) that form in slow flows. Alternative/additional explanations include increased light heterogeneity stimulating productivity at wave-exposed sites, and a drag-induced enhancement of inorganic carbon uptake and allocation to structural material. Higher growth rates at wave-sheltered sites may be explained by the accumulation of nitrogen regenerated by associated fauna within DBLs, whereas reduced growth in slow flows may be a result of OH- or O2 accumulation rather than reduced nutrient supply.

Ongoing ocean global change due to anthropogenic activities is causing multiple chemical and physical seawater properties to change simultaneously, which may affect the physiology of marine phytoplankton. The coccolithophore Emiliania huxleyi is a model species often employed in the study of the marine carbon cycle. The effect of ocean acidification (OA) on coccolithophore calcification has been extensively studied; however, physiological responses to multiple environmental drivers are still largely unknown. Here we examined two‐way and multiple driver effects of OA and other key environmental drivers—nitrate, phosphate, irradiance, and temperature—on the growth, photosynthetic, and calcification rates, and the elemental composition of E. huxleyi. In addition, changes in functional gene expression were examined to understand the molecular mechanisms underpinning the physiological responses. The single driver manipulation experiments suggest decreased nitrate supply being the most important driver regulating E. huxleyi physiology, by significantly reducing the growth, photosynthetic, and calcification rates. In addition, the interaction of OA and decreased nitrate supply (projected for year 2100) had more negative synergistic effects on E. huxleyi physiology than all other two‐way factorial manipulations, suggesting a linkage between the single dominant driver (nitrate) effects and interactive effects with other drivers. Simultaneous manipulation of all five environmental drivers to the conditions of the projected year 2100 had the largest negative effects on most of the physiological metrics. Furthermore, functional genes associated with inorganic carbon acquisition (RubisCO, AEL1, and δCA) and calcification (CAX3, AEL1, PATP, and NhaA2) were most downregulated by the multiple driver manipulation, revealing linkages between responses of functional gene expression and associated physiological metrics. These findings together indicate that for more holistic projections of coccolithophore responses to future ocean global change, it is necessary to understand the relative importance of environmental drivers both individually (i.e., mechanistic understanding) and interactively (i.e., cumulative effect) on coccolithophore physiology.

Local and global changes associated with anthropogenic activities are impacting marine and terrestrial ecosystems. Macroalgae, especially habitat-forming species like kelp, play critical roles in temperate coastal ecosystems. However, their abundance and distribution patterns have been negatively affected by warming in many regions around the globe. Along with global change, coastal ecosystems are also impacted by local drivers such as eutrophication. The interaction between global and local drivers might modulate kelp responses to environmental change. This study examines the regulatory effect of NO3− on the thermal plasticity of the giant kelp Macrocystis pyrifera. To do this, thermal performance curves (TPCs) of key temperature-dependant traits–growth, photosynthesis, NO3− assimilation and chlorophyll a fluorescence–were examined under nitrate replete and deplete conditions in a short-term incubation. We found that thermal plasticity was modulated by NO3− but different thermal responses were observed among traits. Our study reveals that nitrogen, a local driver, modulates kelp responses to high seawater temperatures, ameliorating the negative impacts on physiological performance (i.e. growth and photosynthesis). However, this effect might be species-specific and vary among biogeographic regions – thus, further work is needed to determine the generality of our findings to other key temperate macroalgae that are experiencing temperatures close to their thermal tolerance due to climate change.

In New Zealand, the kelp Undaria pinnatifida has colonized nearly all major coastal ports and has spread to colonize diverse habitats over a range of hydrodynamic environments. We report one of the earliest surveys of the distribution of this species in Macrocystis pyrifera dominated kelp forest habitats in New Zealand alongside experiments comparing the photosynthetic physiology of U. pinnatifida to nine native co-occurring macroalgal species. Undaria pinnatifida was identified at four locations that had previously been reported to be uninvaded and was found throughout the depth range of the kelp forest. Highest densities were observed at shallow depths, reflecting the commonly accepted mode of invasion via sheltered rock pools. Photosynthesis versus Irradiance (P vs. E) curves were constructed for 10 dominant brown macroalgal species. Undaria pinnatifida exhibited greater photosynthetic ability than native macroalgae with maximal rates of photosynthesis at least 1.6-times higher than the habitat forming native kelp species Macrocystis pyrifera and Ecklonia radiata and more than 2.5 times higher than eight other native species. The photosynthetic efficiency of U. pinnatifida under low light was at least as high as macroalgae common on deeper sections of surveyed reefs (e.g. M. pyrifera and E. radiata). Collectively, this information shows that U. pinnatifida has superior photosynthetic performance across all ecologically relevant light intensities. Our data show that the photosynthetic physiology of U. pinnatifida is not likely to limit invasion into deeper sections of reef. This trend could continue, with the potential to fundamentally alter the structure of these wave-exposed kelp communities.

Ecklonia radiata is a globally widespread kelp, dominating temperate reefs throughout the southern hemisphere. Throughout much of its range it is the only laminarian kelp and hence plays a key role in facilitating biodiversity and driving food webs, and underpins immense ecological and socioeconomic values. This review synthesises the growing literature on E. radiata from its phylogeny and distribution through to its biology, ecology and recent changes. It provides an assessment of the state of knowledge and identifies gaps in our understanding of this important species. Despite being tolerant to a wide range of abiotic conditions, recent environmental change has caused direct and indirect loss of E. radiata forests, with extensive areas transitioning to turf and urchin barrens. Ongoing climate change may require application of multi-faceted and novel strategies to increase resistance and resilience to future conditions. By integrating variation across space, time and environmental change, this review provides a synthesis of the current status and possible future trajectories of E. radiata forests.