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

Many studies have quantified the ecological impacts of individual foundation species (FS). However, emerging data suggest that FS often co-occur, potentially inhibiting or facilitating one another, thereby causing indirect, cascading effects on surrounding communities. Furthermore, global warming is accelerating, but little is known about how interactions between co-occurring FS vary with temperature. Shallow aquatic sedimentary systems are often dominated by three types of FS: slower-growing clonal angiosperms, faster-growing solitary seaweeds and shell-forming filter- and deposit-feeding bivalves. Here, we tested the impacts of one FS on another by analyzing manipulative interaction experiments from 148 papers with a global meta-analysis. We calculated 1942 (non-independent) Hedges' g effect sizes, from 11,652 extracted values over performance responses, such as abundances, growth or survival of FS and their associated standard deviations and replication levels. Standard aggregation procedures generated 511 independent Hedges' g that were classified into six types of reciprocal impacts between FS. We found that (i) seaweeds had consistent negative impacts on angiosperms across performance responses, organismal sizes, experimental approaches and ecosystem types; (ii) angiosperms and bivalves generally had positive impacts on each other (e.g. positive effects of angiosperms on bivalves were consistent across organismal sizes and experimental approaches, but angiosperm effect on bivalve growth and bivalve effect on angiosperm abundance were not significant); (iii) bivalves positively affected seaweeds (particularly on growth responses); (iv) there were generally no net effects of seaweeds on bivalves (except for the positive effect on growth) or angiosperms on seaweeds (except for the positive effect on ‘other processes’) and (v) bivalve interactions with other FS were typically more positive at higher temperatures, but angiosperm-seaweed interactions were not moderated by temperature. Synthesis. Despite variations in experimental and spatiotemporal conditions, the stronger positive interactions at higher temperatures suggest that facilitation, particularly involving bivalves, may become more important in a future warmer world. Importantly, addressing research gaps, such as the scarcity of FS interaction experiments from tropical and freshwater systems and for less studied species, as well as testing for density-dependent effects, could better inform aquatic ecosystem conservation and restoration efforts and broaden our knowledge of FS interactions in the Anthropocene.

Small animals (epifauna) are ubiquitous in marine systems. Epifauna have high secondary production and provide trophic linkages between primary producers and higher-order consumers, like fish. Despite their importance, little is known about how these animals respond to warming or how their communities vary across spatiotemporal gradients. Here we use mimics of turf seaweed and invasive kelp holdfast to test, in a 5-factorial field experiment, whether intertidal epifauna are facilitated by different habitat structures, temperature conditions, and along cooccurring spatiotemporal gradients. We found that facilitation of epifauna by intertidal turf seaweed peaked in summer, at low elevation, in older habitats and at a less wave-exposed site. However, epifauna were not affected by the presence of a secondary structure like kelp holdfast mimics or small temperature increases from passive solar heating of black and white mimics. There were many significant two-way, but few higher order interactions, showing stronger facilitation under specific environmental conditions, like at low elevation in summer, or low elevation in old habitats. These results highlight that turf-associated epifauna are controlled by vertical elevation, season, hydrodynamics, and habitat age, and appear to be resilient to small temperature increases. Findings are important to better understand linkages between primary producers and higher order consumers and system-wide productivity, and because fast growing turf, facilitated by global warming and eutrophication, are increasingly outcompeting slower growing large perennial canopy forming seaweeds, like kelp and rockweeds.

Aim: Ecologists traditionally study how contemporary local processes, such as biological interactions and physical stressors, affect the distribution and abundance of organisms. By comparison, biogeographers study the distribution of the same organisms, but focus on historic, larger-scale processes that can cause mass mortalities, such as earthquakes. Here we document cascading effects of rare biogeographical (seismic) and more common ecological (temperature-related) processes on the distribution and abundances of coastal foundation species. Location: Intertidal wave-exposed rocky reefs around Kaikōura, New Zealand, dominated by large, long-lived, and iconic southern bull kelps (Durvillaea antarctica and Durvillaea willana). Methods: In November 2016, a 7.8 Mw earthquake uplifted the coastline around Kaikōura by up to 2 m, and a year later the region experienced the hottest summer on record. Extensive sampling of intertidal communities over 15 km coastline were done shortly after the earthquake and heatwaves and 4 years after the earthquake. Results: Durvillaea lost 75% of its canopy to uplift and the heatwaves reduced canopies that had survived the uplift by an additional 35%. The survey done 4 years after the earthquake showed that Durvillaea had not recovered and that the intertidal zone in many places now was dominated by small turfs and foliose seaweed. Main conclusions: Cascading impacts from seismic uplift and heatwaves have destroyed populations of Durvillaea around Kaikōura. Surviving smaller and sparser Durvillaea patches will likely compromise capacity for self-replacement and lower resilience to future stressors. These results are discussed in a global biogeographical-ecological context of seismic activity and extreme heatwaves and highlight that these events, which are not particularly rare in a geological context, may have common long-lasting ecological legacies.

Disturbances often control community structure by removing large dominant species, allowing new species to colonize. Disturbances vary in intensity and extent, and their effects on resident communities can depend on local environmental conditions. We tested the effects of disturbance intensity and extent on different functional groups of understory species in kelp forests at 4 locations along an ocean climate gradient in Western Australia. We hypothesized that, compared to intact canopies, increasing disturbance intensities (50 and 100% of kelp removal) and extents (2, 4 and 8 m diameter) would promote light-dependent competitors (turf, foliose, articulated coralline and fucoid seaweeds) at the expense of less light-dependent functional groups (invertebrates and encrusting seaweeds). We also hypothesized that these effects would be most pronounced at warmer relative to cooler locations, where metabolic and ecological rates are faster. The first hypothesis was supported; light-dependent understory groups (turfs, in particular) increased, while less light-dependent groups (crusts in particular) decreased with increasing disturbance regimes. However, the second hypothesis was not supported; even though understory communities differed between locations and turf covers were highest at the warmest location, we found no significant interactions between locations and disturbance regimes. Importantly, our results revealed that even small-scale partial canopy loss can have significant effects on kelp-associated communities. The implied community-wide, density-dependent effects have implications for the management and conservation of kelp forests, because restoration of ecological functions must also consider the density of kelp forests, not simply their presence or absence.

During austral summers (DJF) 1934/35, 2017/18 and 2018/19, the New Zealand (NZ) region (approximately 4 million km2) experienced the most intense coupled ocean-atmosphere heatwaves on record. Average air temperature anomalies over land were + 1.7 to 2.1 °C while sea surface temperatures (SST) were 1.2 to 1.9 °C above average. All three heatwaves exhibited maximum SST anomalies west of the South Island of NZ. Atmospheric circulation anomalies showed a pattern of blocking centred over the Tasman Sea extending south-east of NZ, accompanied by strongly positive Southern Annular Mode conditions, and reduced trough activity over NZ. Rapid melt of seasonal snow occurred in all three cases. For the two most recent events, combined ice loss in the Southern Alps was estimated at 8.9 km3 (22% of the 2017 volume). Sauvignon blanc and Pinot noir wine grapes had above average berry number and bunch mass in 2018 but were below average in 2019. Summerfruit harvest (cherries and apricots) was 14 and 2 days ahead of normal in 2017/18 and 2018/19 respectively. Spring wheat simulations suggested earlier flowering and lower grain yields compared to average, and below-average yield and tuber quality in potatoes crops occurred. Major species disruption occurred in marine ecosystems. Hindcasts indicate that the heatwaves were either atmospherically driven or arose from combinations of atmospheric surface warming and oceanic heat advection.

We assess the trends and influences of non-indigenous and cryptogenic species (hereafter simply referred to as “NIS”) on Danish marine community compositions using three decades of quantitative monitoring data. Since the initiation of the Danish marine monitoring programmes in the 1980s, the number of marine NIS recorded in Denmark increased from 30 to 77 in 2014. Thus, of the total 77 marine NIS known from Denmark, 56 (73%) were captured in the standardized monitoring program, while the remaining 21 species (27%) were not detected (in particular gelatinous zooplankton, shallow water fish, parasitic invertebrates, and littoral angiosperms) because of limited spatial-temporal sampling efforts as well as methodological limitations. Significant exponential increases in records of non-native phytoplankton, benthic invertebrates and macroalgae (only in one region) relative to the total species records in the database were observed. Multivariate analyses of presence-absence data, indicated that the contribution of NIS to total community similarity increased over time, highlighting that NIS is becoming an increasingly important component of Danish marine communities. While the presence of NIS generally explained less than 10% of long-term changes in the community similarity across large regions, NIS presence showed local dominance. A correlation analysis indicated that changes in overall species composition within functional groups (phytoplankton, zooplankton, macroalgae and benthic fauna) were related to changes in NIS contribution but more strongly influenced by salinity, highlighting a well-described general positive relationship between salinity and species richness in Danish waters. The process described within this study could form the basis for the analysis of impact of NIS in marine water as required by the Marine Strategy Framework Directive (MSFD).

Understanding the resilience and recovery processes of coastal marine ecosystems is of increasing importance in the face of increasing disturbances and stressors. Large-scale, catastrophic events can re-set the structure and functioning of ecosystems, and potentially lead to different stable states. Such an event occurred in south-eastern New Zealand when a Mw 7.8 earthquake lifted the coastline by up to 6 m. This caused widespread mortality of intertidal algal and invertebrate communities over 130 km of coast. This study involved structured and detailed sampling of three intertidal zones at 16 sites nested into four degree of uplift (none, 0.4–1, 1.5–2.5, and 4.5–6 m). Recovery of large brown algal assemblages, the canopy species of which were almost entirely fucoids, were devastated by the uplift, and recovery after 4 years was generally poor except at sites with < 1 m of uplift. The physical infrastructural changes to reefs were severe, with intertidal emersion temperatures frequently above 35°C and up to 50°C, which was lethal to remnant populations and recruiting algae. Erosion of the reefs composed of soft sedimentary rocks was severe. Shifting sand and gravel covered some lower reef areas during storms, and the nearshore light environment was frequently below compensation points for algal production, especially for the largest fucoid Durvillaea antarctica/poha. Low uplift sites recovered much of their pre-earthquake assemblages, but only in the low tidal zone. The mid and high tidal zones of all uplifted sites remained depauperate. Fucoids recruited well in the low zone of low uplift sites but then were affected by a severe heat wave a year after the earthquake that reduced their cover. This was followed by a great increase in fleshy red algae, which then precluded recruitment of large brown algae. The interactions of species’ life histories and the altered physical and ecological infrastructure on which they rely are instructive for attempts to lessen manageable stressors in coastal environments and help future-proof against the effects of compounded impacts.

Marine heatwaves (MHWs) can cause dramatic changes to ecologically, culturally, and economically important coastal ecosystems. To date, MHW studies have focused on geographically isolated regions or broad-scale global oceanic analyses, without considering coastal biogeographical regions and seasons. However, to understand impacts from MHWs on diverse coastal communities, a combined biogeographical-seasonal approach is necessary, because (1) bioregions reflect community-wide temperature tolerances and (2) summer or winter heatwaves likely affect communities differently. We therefore carried out season-specific Theil–Sen robust linear regressions and Pettitt change point analyses from 1982 to 2021 on the number of events, number of MHW days, mean intensity, maximum intensity, and cumulative intensity of MHWs, for each of the world’s 12 major coastal biogeographical realms. We found that 70% of 240 trend analyses increased significantly, 5% decreased and 25% were unaffected. There were clear differences between trends in metrics within biogeographical regions, and among seasons. For the significant increases, most change points occurred between 1998 and 2006. Regression slopes were generally positive across MHW metrics, seasons, and biogeographical realms as well as being highest after change point detection. Trends were highest for the Arctic, Northern Pacific, and Northern Atlantic realms in summer, and lowest for the Southern Ocean and several equatorial realms in other seasons. Our analysis highlights that future case studies should incorporate break point changes and seasonality in MHW analysis, to increase our understanding of how future, more frequent, and stronger MHWs will affect coastal ecosystems.

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

Description, Large protected areas, once successful in rescuing iconic wetlands from human activities, are undermined by plant invasions., The world has increasingly relied on protected areas (PAs) to rescue highly valued ecosystems from human activities, but whether PAs will fare well with bioinvasions remains unknown. By analyzing three decades of seven of the largest coastal PAs in China, including World Natural Heritage and/or Wetlands of International Importance sites, we show that, although PAs are achieving success in rescuing iconic wetlands and critical shorebird habitats from once widespread reclamation, this success is counteracted by escalating plant invasions. Plant invasions were not only more extensive in PAs than non-PA controls but also undermined PA performance by, without human intervention, irreversibly replacing expansive native wetlands (primarily mudflats) and precluding successional formation of new native marshes. Exotic species are invading PAs globally. This study across large spatiotemporal scales highlights that the consequences of bioinvasions for humanity’s major conservation tool may be more profound, far reaching, and critical for management than currently recognized.

Marine heatwaves (MHW) are becoming stronger and more frequent across the globe. MHWs affect the thermal physiology of all biological organisms, but wider ecosystem effects are particularly impactful when large habitat-forming foundation species such as kelps are affected. Many studies on impacts from MHWs on kelps have focused on temperature effects in isolation, except for a few studies that have integrated co-occurring stress from grazers, wave exposure and nutrient limitation. It is likely that many stressors act in concert with MHWs and exacerbate their effects. Here we analyzed satellite images over 60 months to assess temporal changes in abundance of surface canopies of the giant kelp Macrocystis pyrifera in the New Zealand coastal zone. The analysis encompassed the most extreme MHW on record (2017/18), across a 6° latitudinal gradient of four regions southward from the northern distributional limit of Macrocystis along mainland New Zealand. We tested the association of surface canopy cover of Macrocystis with sea surface temperature, temperature anomalies, chlorophyll-a (a proxy for nutrient availability) and water clarity (diffuse attenuation coefficient). We found a reduced cover of Macrocystis across all regions during and after the 2017/18 MHW, with least impact at the most southern region where the maximum temperatures did not exceed 18°C. There was also an important and significant interaction between temperature and water clarity, showing that temperature-induced kelp loss was greater when water clarity was poor. These results show that notable negative effects occurred across the coastal range of this foundation species and highlight the importance of studying MHW effects across latitudinal gradients and in concert with other co-occurring stressors.

The global ocean has warmed substantially over the past century, with far-reaching implications for marine ecosystems. Concurrent with long-term persistent warming, discrete periods of extreme regional ocean warming (marine heatwaves, MHWs) have increased in frequency. Here we quantify trends and attributes of MHWs across all ocean basins and examine their biological impacts from species to ecosystems. Multiple regions in the Pacific, Atlantic and Indian Oceans are particularly vulnerable to MHW intensification, due to the co-existence of high levels of biodiversity, a prevalence of species found at their warm range edges or concurrent non-climatic human impacts. The physical attributes of prominent MHWs varied considerably, but all had deleterious impacts across a range of biological processes and taxa, including critical foundation species (corals, seagrasses and kelps). MHWs, which will probably intensify with anthropogenic climate change, are rapidly emerging as forceful agents of disturbance with the capacity to restructure entire ecosystems and disrupt the provision of ecological goods and services in coming decades.

SummarySeagrasses are marine angiosperms that potentially provide habitat for crabs, shrimps and fish. However, these types of data are lacking for the seagrass species (Zostera muelleri/rimurēhia) that inhabit intertidal estuaries on the South Island of New Zealand.Abundances of crabs, shrimps and fish were therefore quantified from 361 non-destructive seine tows done in seagrass beds and bare mudflats in Duvauchelle bay and two sites in the Avon-Heathcote/Ihutai estuary between October 2019 and February 2020.A total of 2549 crabs, 5824 shrimps and 1149 fish (75% were juvenile flounders) were identified and counted in the seine-net and immediately released back in healthy condition to the exact location from where they were caught.Only few seagrass leaves were caught in the net and these leaves may have been previously uprooted drift fragments. The instant catch-and-release methodology therefore leaves, literally, nothing but a footprint.More fish taxa, including two species of pipefish, were found in seagrass beds in Duvauchelle bay than in the Avon-Heathcote estuary. Fish (minus juvenile flounders) were also more abundant in these seagrass beds. Furthermore, juvenile flounders and shrimps were more abundant in Duvauchelle bay compared to the Avon-Heathcote estuary, but were found in similar abundances in seagrass beds and on bare flats.It is possible that more fish were found in Duvauchelle seagrass beds because these beds have adjacent deeper areas, and may have high connectivity to seagrass beds in nearby bays. This hypothesis should be tested by sampling more seagrass beds in different types of estuaries and bays.By contrast, crabs were more abundant in the Avon-Heathcote estuary, where spider crabs were most abundant in the seagrass beds, but other crabs were found in similar abundances in seagrass beds and bare habitat. We hypothesize that crab abundances were higher in the Avon-Heathcote estuary because of lower fish predation pressure and/or larger populations of prey like mollusc and polychaetes.Our results suggests that (a) superficially similarZosterabeds in relatively close proximity can provide very different habitat values for fish and crustaceans, (b) seagrass beds with higher diversity and abundances of fish may be prioritized in conservation and management (assuming other important ecosystem functions are similar between beds), and (c) that pipefish may be useful indicator organisms, representing healthy, extensive, dense and connected seagrass beds.Abstract Figure

Coastal ecosystems are subjected to multiple co-occurring anthropogenic stressors which potentially interact to produce complex impacts on the structure and functioning of biological communities. Seagrass meadows are among the most rapidly declining coastal habitats on Earth. In particular, high nutrient loadings, enhanced sedimentation and competition from blooming seaweeds, like the invasive red alga Gracilaria vermiculophylla, are claimed to be associated with this decline. In this study, we tested for individual and potential interactive impacts on the intertidal seagrass Zostera noltei of these three stressors using a factorial field experiment. We measured seagrass shoots density and biomass (both above and below ground seagrass biomass) as proxies of seagrass physical condition. We also examined changes in fauna assemblages. The study suggested that sediment loading had the most detrimental impacts on health of the seagrass meadows. The effect of seaweed addition was negative and denso-dependent. Deleterious effects of nutrient enrichment were less evident. Non-additive interactions were also noticeable for some of the structural traits measured. Particularly, nutrient enrichment changed the effect of the other stressors for some of the responses measured. The effect of the treatments was also perceptible in the associated fauna assemblages. The ubiquitous grazer Peringia ulvae was more abundant in treatments with sediment loading, but those sediment effects in the density of the small grazer were modulated by nutrient and seaweed addition levels. Finally, we suggest that more studies should test for synergistic effects of co-occurring stressors to better understand the direction and intensity of changes triggered by human activities in natural ecosystems. project MarInfo Integrated Platform for Marine Data Acquisition and Analysis - North Portugal Regional Operational Programme (NORTE 2020), under the PORTUGAL 2020 Partnership Agreement, through the European Regional Development Fund (ERDF) [NORTE-01-0145-FEDER-000031] project SEEINGSHORE [NORTE-01-0145-FEDER-031893] NORTE 2020 through the ERDF Portugal 2020 through the ERDF European Union through the ERDF FCT Portuguese Foundation for Science and Technology European Commission info:eu-repo/semantics/publishedVersion

Prolonged high-temperature extreme events in the ocean, marine heatwaves, can have severe and long-lasting impacts on marine ecosystems, fisheries and associated services. This study applies a marine heatwave framework to analyse a global sea surface temperature product and identify the most extreme events, based on their intensity, duration and spatial extent. Many of these events have yet to be described in terms of their physical attributes, generation mechanisms, or ecological impacts. Our synthesis identifies commonalities between marine heatwave characteristics and seasonality, links to the El Niño-Southern Oscillation, triggering processes and impacts on ocean productivity. The most intense events preferentially occur in summer, when climatological oceanic mixed layers are shallow and winds are weak, but at a time preceding climatological maximum sea surface temperatures. Most subtropical extreme marine heatwaves were triggered by persistent atmospheric high-pressure systems and anomalously weak wind speeds, associated with increased insolation, and reduced ocean heat losses. Furthermore, the most extreme events tended to coincide with reduced chlorophyll-a concentration at low and mid-latitudes. Understanding the importance of the oceanic background state, local and remote drivers and the ocean productivity response from past events are critical steps toward improving predictions of future marine heatwaves and their impacts. Concepts and analyses were developed during three workshops organized by an international working group on marine heatwaves (https://www.marineheatwaves.org) funded by a University of Western Australia Research Collaboration Award and a Natural Environment Research Council (UK) International Opportunity Fund (NE/N00678X/1). D.A.S. is supported by a UKRI Future Leaders Fellowship (MR/S032827/1). The Australian Research Council supported T.W. (FT110100174 and DP170100023) and A.S.T. (FT160100495). N.J.H. and L.V.A. are supported by the ARC Centre of Excellence for Climate Extremes (CE170100023). M.S.T was supported by the Brian Mason Trust. P.J.M. is supported by a Marie Curie Career Integration Grant (PCIG10-GA-2011–303685) and a Natural Environment Research Council (UK) Grant (NE/J024082/1). E.C.J.O. was supported by National Sciences and Engineering Research Council of Canada (NSERC) Discovery Grant RGPIN-2018-05255 and Marine Environmental Observation, Prediction and Response Network (MEOPAR) project 1-02-02-059.1. C.C.U. acknowledges financial support through the Early Career Scientist Endowed Fund, George E. Thibault Early Career Scientist Fund, and The Joint Initiative Awards Fund from the Andrew W. Mellon Foundation at WHOI. M.G.D. received funding by the Spanish Ministry for the Economy, Industry and Competitiveness Ramón y Cajal 2017 grant reference RYC-2017-22964. NOAA High Resolution SST data provided by the NOAA/OAR/ESRL PSD, Boulder, Colorado, USA, from their Web site at https://www.esrl.noaa.gov/psd/. All datasets used in this work are freely available. NOAA high resolution SST were provided by NOAA/OAR/ESRL PSD (Boulder, CO, USA) from their website (www.esrl.noaa.gov/psd/). Processed MHW severity data are stored at the National Computing Infrastructure, Australia and is available on request. Python and MATLAB codes for processing raw SST data, analysing and plotting datasets and most extremes MHW characteristics are available at https://github.com/alexsengupta/ExtremeExtremes.git.

AIM: Understanding the relative importance of climatic and non‐climatic distribution drivers for co‐occurring, functionally similar species is required to assess potential consequences of climate change. This understanding is, however, lacking for most ecosystems. We address this knowledge gap and forecast changes in distribution for habitat‐forming seaweeds in one of the world's most species‐rich temperate reef ecosystems. LOCATION: The Great Southern Reef. The full extent of Australia's temperate coastline. METHODS: We assessed relationships between climatic and non‐climatic environmental data known to influence seaweed, and the presence of 15 habitat‐forming seaweeds. Distributional data (herbarium records) were analysed with MAXENT and generalized linear and additive models, to construct species distribution models at 0.2° spatial resolution, and project possible distribution shifts under the RCP 6.0 (medium) and 2.6 (conservative) emissions scenarios of ocean warming for 2100. RESULTS: Summer temperatures, and to a lesser extent winter temperatures, were the strongest distribution predictors for temperate habitat‐forming seaweeds in Australia. Projections for 2100 predicted major poleward shifts for 13 of the 15 species, on average losing 78% (range: 36%–100%) of their current distributions under RCP 6.0 and 62% (range: 27%–100%) under RCP 2.6. The giant kelp (Macrocystis pyrifera) and three prominent fucoids (Durvillaea potatorum, Xiphophora chondrophylla and Phyllospora comosa) were predicted to become extinct from Australia under RCP 6.0. Many species currently distributed up the west and east coasts, including the dominant kelp Ecklonia radiata (71% and 49% estimated loss for RPC 6.0 and 2.6, respectively), were predicted to become restricted to the south coast. MAIN CONCLUSIONS: In close accordance with emerging observations in Australia and globally, our study predicted major range contractions of temperate seaweeds in coming decades. These changes will likely have significant impacts on marine biodiversity and ecosystem functioning because large seaweeds are foundation species for 100s of habitat‐associated plants and animals, many of which are socio‐economically important and endemic to southern Australia.

Marine invasive macroalgae can have severe local-scale impacts on ecological communities. The kelp Undaria pinnatifida is one of the most successful marine invasive species worldwide, and is widely regarded as one of the worst. Here, we review research on Undaria in Australasia, where the kelp is established throughout much of New Zealand and south-eastern Australia. The presence of Undaria for at least three decades in these locations makes Australasia one of the longest-invaded bioregions globally, and a valuable case study for considering Undaria's invasion success and associated impacts. In Australasia, Undaria has primarily invaded open spaces, turf communities, and gaps in native canopies within a relatively narrow elevation band on rocky shores. Despite its high biomass, Undaria has relatively few direct impacts on native species, and can increase community-wide attributes such as primary productivity and the provision of biogenic habitat. Therefore, Australasian Undaria research provides an example of a decoupling between the success and impact of an invasive species. Undaria will most likely continue to spread along thousands of kilometres of rocky coastline in temperate Australasia, due to its tolerance to large variations in temperature, ability to exploit disturbances to local communities, and the continued transfer among regions via vessel movements and aquaculture activities. However, the spread of Undaria remains difficult to manage as eradication is challenging and seldom successful. Therefore, understanding potential invasion pathways, maintaining native canopy-forming species that limit Undaria success, and effectively managing anthropogenic vectors of Undaria spread, should be key management priorities.

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.

Facilitation cascades occur when a habitat-forming species facilitates another habitat-forming species with synergistic effects on biodiversity. This review summarizes the current state of knowledge of facilitation cascades in marine environments, describing (1) the geographic distribution of studies documenting facilitation cascades; (2) the range of habitats and organisms involved; (3) the mechanisms underpinning their establishment; and (4) the sources of their spatial and temporal patterns and variability. We synthesize this knowledge to identify ways in which facilitation cascades could be integrated with other ecological theories regarding community assembly and function, and also to explore how they may be integrated into conservation and management strategies. Finally, we identify critical knowledge gaps and future research directions for improving our understanding of the mechanisms regulating the establishment and persistence of facilitation cascades in marine environments.

Detailed research has documented gradual changes to biological communities attributed to increases in global average temperatures. However, localized and abrupt temperature anomalies associated with heatwaves may cause more rapid biological changes. We analysed temperature data from the South Island of New Zealand and investigated whether the hot summer of 2017/18 affected species of bull kelp, Durvillaea antarctica, D. poha and D. willana. Durvillaea spp. are large iconic seaweed that inhabit the low intertidal zone of exposed coastlines, where they underpin biodiversity and ecosystem functioning. Sea surface temperatures during the summer of 2017/18 included the strongest marine heatwaves recorded in 38 years of existing oceanic satellite data for this region. Air temperatures were also high, and, coupled with small wave heights, resulted in strong desiccation stress during daytime low tides. Before-After analysis of drone images of four reef platforms (42, 42, 44 and 45 ˚S) was used to evaluate changes to bull kelp over the hot summer. Bull kelp loss varied among species and reefs, with the greatest (100%) loss of D. poha at Pile Bay in Lyttelton Harbour (44 ˚S). In Pile Bay, sea surface temperature exceeded 23 ˚C and air temperatures exceeded 30 ˚C, while Durvillaea was exposed for up to 3 hours per day during low tide. Follow-up surveys showed that all bull kelps were eliminated from Pile Bay, and from all reefs within and immediately outside of Lyttelton Harbour. Following the localized extinction of bull kelp in Pile Bay, the invasive kelp Undaria pinnatifida recruited in high densities (average of 120 m-2). We conclude that bull kelps are likely to experience additional mortalities in the future because heatwaves are predicted to increase in magnitude and durations. Losses of the endemic D. poha are particularly concerning due to its narrow distributional range.

Marine heatwaves (MHWs) are extreme climatic events in oceanic systems that can have devastating impacts on ecosystems, causing abrupt ecological changes and socioeconomic consequences. Several prominent MHWs have attracted scientific and public interest, and recent assessments have documented global and regional increases in their frequency. However, for proactive marine management, it is critical to understand how patterns might change in the future. Here, we estimate future changes in MHWs to the end of the 21st century, as simulated by the CMIP5 global climate model projections. Significant increases in MHW intensity and count of annual MHW days are projected to accelerate, with many parts of the ocean reaching a near-permanent MHW state by the late 21st century. The two greenhouse gas (GHG) emission scenarios considered (Representative Concentration Pathway 4.5 and 8.5) strongly affect the projected intensity of MHW events, the proportion of the globe exposed to permanent MHW states, and the occurrence of the most extreme MHW events. Comparison with simulations of a natural world, without anthropogenic forcing, indicate that these trends have emerged from the expected range of natural variability within the first half of the 21st century. This discrepancy implies a degree of “anthropogenic emergence,” with a departure from the natural MHW conditions that have previously shaped marine ecosystems for centuries or even millennia. Based on these projections we expect impacts on marine ecosystems to be widespread, significant and persistent through the 21st century. This research was supported by the Australian Research Council grants CE170100023 and FT170100106, Natural Environment Research Council International Opportunity Fund NE/N00678X/1, National Sciences and Engineering Research Council of Canada Discovery Grant RGPIN-2018-05255, and Brian Mason (Impacts of an unprecedented marine heatwave). This project was partially supported through funding from the Earth Systems and Climate Change Hub of the Australian Government’s National Environmental Science Program.

An increasing number of studies report impacts from invasive species on community metrics or ecosystem functions. We draw attention to an issue arising whenever impact is measured on a community where the invader is an integrated part: should or shouldn’t the attributes of the invader itself be included in the data-analysis? We identify many examples from the published literature showing inconsistency in whether or not data for the invader is included or excluded, and discuss potential implications for ecological interpretations. We also provide a case study to show that the invasive seaweed Undaria pinnatifida can be interpreted to have strong or no impact on seaweed communities, depending on its inclusion or exclusion in the data analysis. We conclude that it is critical for studies to (1) clearly state in the methods section, if the invaders are included or excluded from the data-analysis, (2) acknowledge potential differences in outcomes when comparing results based on different methods, and (3) analyze, if possible, impacts both with and without the invader. Finally, we note that this ‘inclusion versus exclusion’ conundrum is not only relevant to invasion biology, but to any field where the test-object of interest can be an integrated part of the response, such as when impact of seaweed blooms are analysed on community productivity or community effects are quantified over time from ecological pulse-perturbation experiments.

Large scale disturbances associated with anthropogenic activities or natural disasters can destroy primary habitat-forming species like corals, seagrasses and seaweeds. However, little research has documented if and on how large-scale disturbances affect secondary habitat formers, such as epiphytes and small animals that depend on biogenic habitats. Here we quantified changes in the abundance of both primary and secondary habitat-forming seaweeds as well as seaweed-associated invertebrates before and after a 7.8 Mw earthquake that uplifted four intertidal reef platforms by 0.5–0.8 m on the Kaikōura coastline in New Zealand. We found that the dominant primary (Hormosirabanksii and three Cystophora species) and secondary (obligate and facultative epiphytes) habitat-forming seaweeds were all decimated and that mobile seaweed-associated animals were significantly less abundant (per gram of seaweed biomass) after the earthquake. Importantly, epiphytes became functionally extinct after the earthquake, as less than 0.1 % of the populations survived, whereas primary habitat formers survived in suitable microhabitats, like water covered tide-pools and tidal channels. Based on these results we also discuss possible cascading ecosystem effects and future scenarios for natural recovery vs. active restoration that could speed up the recovery of habitat-forming species on degraded reefs.

Genetic diversity confers adaptive capacity to populations under changing conditions but its role in mediating impacts of climate change remains unresolved for most ecosystems. This lack of knowledge is particularly acute for foundation species, where impacts may cascade throughout entire ecosystems. We combined population genetics with eco-physiological and ecological field experiments to explore relationships among latitudinal patterns in genetic diversity, physiology and resilience of a kelp ecosystem to climate stress. A subsequent ‘natural experiment’ illustrated the possible influence of latitudinal patterns of genetic diversity on ecosystem vulnerability to an extreme climatic perturbation (marine heatwave). There were strong relationships between physiological versatility, ecological resilience and genetic diversity of kelp forests across latitudes, and genetic diversity consistently outperformed other explanatory variables in contributing to the response of kelp forests to the marine heatwave. Population performance and vulnerability to a severe climatic event were thus strongly related to latitudinal patterns in genetic diversity, with the heatwave extirpating forests with low genetic diversity. Where foundation species control ecological structure and function, impacts of climatic stress can cascade through the ecosystem and, consequently, genetic diversity could contribute to ecosystem vulnerability to climate change.

Carbon flow is essential to the function of all ecosystems, yet there is little mechanistic understanding of how non-indigenous macroalgae alter rates of carbon fixation in marine ecosystems. The spread of fast-growing non-indigenous species, such as the annual kelp Undaria pinnatifida (Harvey) Suringer, can potentially change trophic links through variable photosynthetic parameters relative to indigenous species. Here we use in situ photorespirometry to compare rates of net primary productivity (NPP) of assemblages dominated by U. pinnatifida and two native canopy-forming species, Cystophora torulosa and Durvillaea antarctica . The three assemblages had different light-use dynamics across a full light range, with the indigenous macroalgae showing no sign of saturated NPP at high irradiance, but with U. pinnatifida showing saturated NPP beyond 1000 μmol m − 2 s − 1 . Using incident irradiance collected over a full year, we show small differences in modeled average daily NPP during spring between U. pinnatifida assemblages (8 g C m − 2 day − 1 ) and those dominated by C. torulosa (7 g C m − 2 day − 1 ), whereas D. antarctica assemblages were the most productive (10 g C m − 2 day − 1 ). The proportion of NPP provided by the sub-canopy component of assemblages varied between the canopy-forming species, where D. antarctica had a low sub-canopy contribution compared to stands dominated by U. pinnatifida or C. torulosa . High biomass turnover associated with the annual life history of U. pinnatifida has the potential to increase carbon export to surrounding ecosystems compared to perennial fucoid species. Therefore, U. pinnatifida may have a positive effect on carbon flow, fixing similar quantities of carbon in a 6-month period as the native C. torulosa in a year. It appears that U. pinnatifida has the potential to contribute a great deal of carbon and alter the biomass export regime as it spreads across shallow coastal habitats.

While invasive species often threaten biodiversity and human well-being, their potential to enhance functioning by offsetting the loss of native habitat has rarely been considered. We manipulated the abundance of the nonnative, habitat-forming seaweed Gracilaria vermiculophylla in large plots (25 m2) on southeastern US intertidal landscapes to assess impacts on multiple ecosystem functions underlying coastal ecosystem services. We document that in the absence of native habitat formers, this invasion has an overall positive, density-dependent impact across a diverse set of ecosystem processes (e.g., abundance and richness of nursery taxa, flow attenuation). Manipulation of invader abundance revealed both thresholds and saturations in the provisioning of ecosystem functions. Taken together, these findings call into question the focus of traditional invasion research and management that assumes negative effects of nonnatives, and emphasize the need to consider context-dependence and integrative measurements when assessing the impact of an invader, including density dependence, multifunctionality, and the status of native habitat formers. This work supports discussion of the idea that where native foundation species have been lost, invasive habitat formers may be considered as sources of valuable ecosystem functions.

One of the greatest impediments to detecting changes in species distributions in response to ocean warming is the lack of baseline data. In a recent article, we compared old (1940–1959) and new (1990–2009) herbarium records of Australian seaweeds and found a net southward shift in the latitude of northernmost collections of temperate species, implying a flora-wide poleward retreat over the past five decades. Huisman & Millar (2013) criticised our methods, contending that a comparison of herbarium records from different time periods cannot be used to infer changes in species distributions without field-based validation. However, our analysis compared the median position of extreme records of random species from random locations rather than focusing on particular species and their possible loss from specific sites. Hence, ground-truthing ‘extinctions' are of limited value to the interpretation of our analysis. Moreover, subtidal ground-truthing over biogeographic scales is not logistically possible and even...

Impacts of marine invaders on local biodiversity have not been analyzed across inva- sive species and invaded habitats. We conducted a meta-analysis of 56 field experiments pub- lished in 29 papers that examined the effects of marine invaders on local species richness, diver- sity, and/or evenness. We show that invaders, across studies, typically have negative effects on biodiversity within a trophic level but positive effects on biodiversity of higher trophic levels. For example, both plants and sessile filter-feeders had positive effects on richness and diversity of mobile consumers. The contrasting negative and positive effects on similar versus higher trophic levels are potentially manifested through community-wide antagonism (competition and con- sumption) versus facilitation (habitat and food provisioning) interactions, respectively. These relation ships extended to functional interactions, as sessile invaders had negative effects on the biodiversity of sessile communities (intra-functional interactions) but positive effects on the biodi- versity of mobile communities (inter-functional interactions). Our analyses highlight the impor- tance of pairing attributes of the invader and the impacted organisms to obtain simple predictions of how the diversity of entire communities may respond to species invasions on local scales. We also note that our analysis did not require information on co-evolutionary history but that such data, coupled with long-term large-scale mensurative data, are needed to gain a comprehensive predictive insight into invasion impact.

Recent research has shown that co-occurring primary and secondary habitat-forming species typically support higher biodiversity than do monocultures of the primary habitat-former alone. However, these ‘habitat cascades’ may not be universal and it is important to know whether, when and where positive effects on biodiversity from secondary habitat-forming species change to negative effects. Here, we tested how anthropogenic stressors (fertilisation and sedimentation) and unattached secondary habitat-forming Ulva seaweeds affected the primary habitat-forming seagrass, Zostera muelleri, and its associated invertebrates in the Avon–Heathcote Estuary, New Zealand. We experimentally stressed Zostera by adding different fertilisation and sediment levels. Fertilisation had little impact, whereas even low sedimentation levels had strong negative effects on Zostera and its associated fauna. In a second experiment, sediments and Ulva were added to seagrass beds and unvegetated mudflats to test whether sediment stress modifies habitat cascades. We found again strong negative effects of sediments on Zostera, irrespective of spatio-temporal conditions, and that negative effects of sediments on invertebrates were enhanced in the presence of the secondary habitat former. These results highlighted that anthropogenic stressors can destabilise habitat cascades; processes that may be of particular importance in estuaries that are characterised by low biodiversity and stressful environmental conditions.