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

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

Effect of marine heatwaves and warming on kelp microbiota influence trophic interactions.

The range-expansion of tropical herbivores due to ocean warming can profoundly alter temperate reef communities by overgrazing the seaweed forests that underpin them. Such ecological interactions may be mediated by changes to seaweed-associated microbiota in response to warming, but empirical evidence demonstrating this is rare. We experimentally simulated ocean warming and marine heatwaves (MHWs) to quantify effects on two dominant temperate seaweed species and their microbiota, as well as grazing by a tropical herbivore. The kelp Ecklonia radiata's microbiota in sustained warming and MHW treatments was enriched with microorganisms associated with seaweed disease and tissue degradation. In contrast, the fucoid Sargassum linearifolium's microbiota was unaffected by temperature. Consumption by the tropical sea-urchin Tripneustes gratilla was greater on Ecklonia where the microbiota had been altered by higher temperatures, while Sargassum's consumption was unaffected. Elemental traits (carbon, nitrogen), chemical defences (phenolics) and tissue bleaching of both seaweeds were generally unaffected by temperature. Effects of warming and MHWs on seaweed holobionts (host plus its microbiota) are likely species-specific. The effect of increased temperature on Ecklonia's microbiota and subsequent increased consumption suggest that changes to kelp microbiota may underpin kelp-herbivore interactions, providing novel insights into potential mechanisms driving change in species' interactions in warming oceans.

Temperature sensitivity of detrital photosynthesis.

BACKGROUND AND AIMS: Kelp forests are increasingly considered blue carbon habitats for ocean-based biological carbon dioxide removal, but knowledge gaps remain in our understanding of their carbon cycle. Of particular interest is the remineralization of detritus, which can remain photosynthetically active. Here, we study a widespread, thermotolerant kelp (Ecklonia radiata) to explore detrital photosynthesis as a mechanism underlying temperature and light as two key drivers of remineralization. METHODS: We used meta-analysis to constrain the thermal optimum (Topt) of E. radiata. Temperature and light were subsequently controlled over a 119-day ex situ decomposition experiment. Flow-through experimental tanks were kept in darkness at 15 °C or under a subcompensating maximal irradiance of 8 µmol photons m-2 s-1 at 15, 20 or 25 °C. Photosynthesis of laterals (analogues to leaves) was estimated using closed-chamber oxygen evolution in darkness and under a saturating irradiance of 420 µmol photons m-2 s-1. KEY RESULTS: T opt of E. radiata is 18 °C across performance variables (photosynthesis, growth, abundance, size, mass and fertility), life stages (gametophyte and sporophyte) and populations. Our models predict that a temperature of >15 °C reduces the potential for E. radiata detritus to be photosynthetically viable, hence detrital Topt ≤ 15 °C. Detritus is viable under subcompensating irradiance, where it performs better than in darkness. Comparison of net and gross photosynthesis indicates that elevated temperature primarily decreases detrital photosynthesis, whereas darkness primarily increases detrital respiration compared with optimal experimental conditions, in which detrital photosynthesis can persist for ≥119 days. CONCLUSIONS: T opt of kelp detritus is ≥3 °C colder than that of the intact plant. Given that E. radiata is one of the most temperature-tolerant kelps, this suggests that photosynthesis is generally more thermosensitive in the detrital phase, which partly explains the enhancing effect of temperature on remineralization. In contrast to darkness, even subcompensating irradiance maintains detrital viability, elucidating the accelerating effect of depth and its concomitant light reduction on remineralization to some extent. Detrital photosynthesis is a meaningful mechanism underlying at least two drivers of remineralization, even below the photoenvironment inhabited by the attached alga.

Marine protected areas can be useful but are not a silver bullet for kelp conservation.

Kelp forests are among the most valuable ecosystems on Earth, but they are increasingly being degraded and lost due to a range of human-related stressors, leading to recent calls for their improved management and conservation. One of the primary tools to conserve marine species and biodiversity is the establishment of marine protected areas (MPAs). International commitments to protect 30% of the world's ecosystems are gaining momentum, offering a promising avenue to secure kelp forests into the Anthropocene. However, a clear understanding of the efficacy of MPAs for conserving kelp forests in a changing ocean is lacking. In this perspective, we question whether strengthened global protection will create meaningful conservation outcomes for kelp forests. We explore the benefits of MPAs for kelp conservation under a suite of different stressors, focusing on empirical evidence from protected kelp forests. We show that MPAs can be effective against some drivers of kelp loss (e.g., overgrazing, kelp harvesting), particularly when they are maintained in the long-term and enforced as no-take areas. There is also some evidence that MPAs can reduce impacts of climate change through building resilience in multi-stressor situations. However, MPAs also often fail to provide protection against ocean warming, marine heatwaves, coastal darkening, and pollution, which have emerged as dominant drivers of kelp forest loss globally. Although well-enforced MPAs should remain an important tool to protect kelp forests, successful kelp conservation will require implementing an additional suite of management solutions that target these accelerating threats.

Host genotype and microbiome associations in co-occurring clonal and non-clonal kelp, Ecklonia radiata.

A fundamental question in holobiont biology is the extent to which microbiomes are determined by host characteristics regulated by their genotype. Studies on the interactions of host genotype and microbiomes are emerging but disentangling the role that host genotype has in shaping microbiomes remains challenging in natural settings. Host genotypes tend to be segregated in space and affected by different environments. Here we overcome this challenge by studying an unusual situation where host asexual (5 clonal lineages) and sexual genotypes (15 non-clonal lineages) of the same species co-occur under the same environment. This allowed us to partition the influence of morphological traits and genotype in shaping host-associated bacterial communities. Lamina-associated bacteria of co-occurring kelp sexual non-clonal (Ecklonia radiata) and asexual clonal (E. brevipes) morphs were compared to test whether host genotype influences microbiomes beyond morphology. Similarity of bacterial composition and predicted functions were evaluated among individuals within a single clonal genotype or among non-clonal genotypes of each morph. Higher similarity in bacterial composition and inferred functions were found among identical clones of E. brevipes compared to other clonal genotypes or unique non-clonal E. radiata genotypes. Additionally, bacterial diversity and composition differed significantly between the two morphs and were related with one morphological trait in E. brevipes (haptera). Thus, factors regulated by the host genotype (e.g. secondary metabolite production) likely drive differences in microbial communities between morphs. The strong association of genotype and microbiome found here highlights the importance of genetic relatedness of hosts in determining variability in their bacterial symbionts.

On the Challenges of Identifying Benthic Dominance on Anthropocene Coral Reefs.

The concept of dominance is frequently used to describe changes in rapidly reconfiguring ecosystems, but the definition of dominance can vary widely among studies. Using coral reefs as a model, we use extensive benthic composition data to explore how variability in applying dominance concepts can shape perceptions. We reveal that coral dominance is sensitive to the exclusion of key algal groups and the categorization of other benthic groups, with ramifications for detecting an ecosystem phase shift. For example, ignoring algal turf inflates the dominance of hard and soft corals in the benthic habitats underpinning reef ecosystems. We need a consensus on how dominance concepts are applied so that we can build a more comprehensive understanding of ecosystem shifts across a broad range of aquatic and terrestrial settings. For reefs, we highlight the benefits of comprehensive and inclusive surveys for evaluating and managing the altered ecosystem states that are emerging in the Anthropocene.

Research priorities for the sustainability of coral-rich western Pacific seascapes.

Nearly a billion people depend on tropical seascapes. The need to ensure sustainable use of these vital areas is recognised, as one of 17 policy commitments made by world leaders, in Sustainable Development Goal (SDG) 14 ('Life below Water') of the United Nations. SDG 14 seeks to secure marine sustainability by 2030. In a time of increasing social-ecological unpredictability and risk, scientists and policymakers working towards SDG 14 in the Asia-Pacific region need to know: (1) How are seascapes changing? (2) What can global society do about these changes? and (3) How can science and society together achieve sustainable seascape futures? Through a horizon scan, we identified nine emerging research priorities that clarify potential research contributions to marine sustainability in locations with high coral reef abundance. They include research on seascape geological and biological evolution and adaptation; elucidating drivers and mechanisms of change; understanding how seascape functions and services are produced, and how people depend on them; costs, benefits, and trade-offs to people in changing seascapes; improving seascape technologies and practices; learning to govern and manage seascapes for all; sustainable use, justice, and human well-being; bridging communities and epistemologies for innovative, equitable, and scale-crossing solutions; and informing resilient seascape futures through modelling and synthesis. Researchers can contribute to the sustainability of tropical seascapes by co-developing transdisciplinary understandings of people and ecosystems, emphasising the importance of equity and justice, and improving knowledge of key cross-scale and cross-level processes, feedbacks, and thresholds.

Persistent thermally driven shift in the functional trait structure of herbivorous fishes: Evidence of top-down control on the rebound potential of temperate seaweed forests?

Extreme climatic events can reshape the functional structure of ecological communities, potentially altering ecological interactions and ecosystem functioning. While these shifts have been widely documented, evidence of their persistence and potential flow-on effects on ecosystem structure following relaxation of extreme events remains limited. Here, we investigate changes in the functional trait structure - encompassing dimensions of resource use, thermal affinity, and body size - of herbivorous fishes in a temperate reef system that experienced an extreme marine heatwave (MHW) and subsequent return to cool conditions. We quantify how changes in the trait structure modified the nature and intensity of herbivory-related functions (macroalgae, turf, and sediment removal), and explored the potential flow-on effects on the recovery dynamics of macroalgal foundation species. The trait structure of the herbivorous fish assemblage shifted as a result of the MHW, from dominance of cool-water browsing species to increased evenness in the distribution of abundance among temperate and tropical guilds supporting novel herbivory roles (i.e. scraping, cropping, and sediment sucking). Despite the abundance of tropical herbivorous fishes and intensity of herbivory-related functions declined following a period of cooling after the MHW, the underlying trait structure displayed limited recovery. Concomitantly, algal assemblages displayed a lack of recovery of the formerly dominant foundational species, the kelp Ecklonia radiata, transitioning to an alternative state dominated by turf and Sargassum spp. Our study demonstrates a legacy effect of an extreme MHW and exemplified the value of monitoring phenotypic (trait mediated) changes in the nature of core ecosystem processes to predict and adapt to the future configurations of changing reef ecosystems.

Maximization of fitness by phenological and phenotypic plasticity in range expanding rabbitfishes (Siganidae).

Global warming is modifying the phenology, life-history traits and biogeography of species around the world. Evidence of these effects have increased over recent decades; however, we still have a poor understanding of the possible outcomes of their interplay across global climatic gradients, hindering our ability to accurately predict the consequences of climate change in populations and ecosystems. We examined the effect that changes in biogeography can have on the life-history traits of two of the most successful range-extending fish species in the world: the tropical rabbitfishes Siganus fuscescens and Siganus rivulatus. Both species have established abundant populations at higher latitudes in the northern and southern hemispheres and have been identified as important ecological engineers with the potential to alter the community structure of seaweed forests (Laminariales and Fucales) in temperate regions. Life-history trait information from across their global distribution was compiled from the published literature and meta-analyses were conducted to assess changes in (i) the onset and duration of reproductive periods, (ii) size at maturity, (iii) fecundity, (iv) growth rates, (v) maximum body sizes and (vi) longevity in populations at the leading edge of range expansion in relation to sea surface temperature and primary productivity (a common proxy for nutritional resource levels). Populations at highest latitudes had shortened their reproductive periods and reduced growth rates, taking longer to reach sexual maturity and maximum sizes, but compensated this with higher fecundity per length class and longer lifespans than populations in warmer environments. Low primary productivity and temperature in the Mediterranean Sea resulted in lower growth rates and body sizes for S. rivulatus, but also lower length at maturity, increasing life-time reproductive output. The results suggest that plasticity in the phenology and life-history traits of range-expanding species would be important to enhance their fitness in high latitude environments, facilitating their persistence and possible further poleward expansions. Quantifying the magnitude and direction of these responses can improve our understanding and ability to forecast species redistributions and its repercussions in the functioning of temperate ecosystems.

Effects of human footprint and biophysical factors on the body-size structure of fished marine species.

Marine fisheries in coastal ecosystems in many areas of the world have historically removed large-bodied individuals, potentially impairing ecosystem functioning and the long-term sustainability of fish populations. Reporting on size-based indicators that link to food-web structure can contribute to ecosystem-based management, but the application of these indicators over large (cross-ecosystem) geographical scales has been limited to either fisheries-dependent catch data or diver-based methods restricted to shallow waters (<20 m) that can misrepresent the abundance of large-bodied fished species. We obtained data on the body-size structure of 82 recreationally or commercially targeted marine demersal teleosts from 2904 deployments of baited remote underwater stereo-video (stereo-BRUV). Sampling was at up to 50 m depth and covered approximately 10,000 km of the continental shelf of Australia. Seascape relief, water depth, and human gravity (i.e., a proxy of human impacts) were the strongest predictors of the probability of occurrence of large fishes and the abundance of fishes above the minimum legal size of capture. No-take marine reserves had a positive effect on the abundance of fishes above legal size, although the effect varied across species groups. In contrast, sublegal fishes were best predicted by gradients in sea surface temperature (mean and variance). In areas of low human impact, large fishes were about three times more likely to be encountered and fishes of legal size were approximately five times more abundant. For conspicuous species groups with contrasting habitat, environmental, and biogeographic affinities, abundance of legal-size fishes typically declined as human impact increased. Our large-scale quantitative analyses highlight the combined importance of seascape complexity, regions with low human footprint, and no-take marine reserves in protecting large-bodied fishes across a broad range of species and ecosystem configurations.

Las pesquerías marinas de los ecosistemas costeros en muchas áreas del mundo históricamente han removido a individuos de gran tamaño, potencialmente perjudicando el funcionamiento ambiental y la sostenibilidad a largo plazo de las poblaciones de peces. Los reportes sobre los indicadores basados en el tamaño que se vinculan con la estructura de la red alimenticia pueden contribuir al manejo basado en el ecosistema, aunque la aplicación de estos indicadores a grandes (inter-ecosistemas) escalas geográficas ha estado limitada a datos de captura dependientes de las pesquerías o métodos basados en el buceo restringidos a aguas someras (<20 m), lo cual puede representar erróneamente la abundancia de peces de gran tamaño capturados para la pesca. Obtuvimos los datos de la estructura del tamaño corporal de 82 teleósteos marinos demersales focalizados por razones recreativas o comerciales tomados de 2,904 despliegues de video estéreo subacuático remoto con cebo (stereo-BRUV, en inglés). El muestreo se realizó hasta los 50 metros de profundidad y abarcó aproximadamente 10,000 km del talud continental de Australia. El relieve marino, la profundidad del agua y la gravedad humana (es decir, un indicador de los impactos humanos) fueron los pronosticadores más sólidos de la probabilidad de incidencia de los peces de gran tamaño y de la abundancia de peces por encima del tamaño legal mínimo de captura. Las reservas marinas de protección total tienen un efecto positivo sobre la abundancia de los peces que están por encima del tamaño legal, aunque el efecto varió según el grupo de especies. Como contraste, los peces de tamaño sublegal fueron pronosticados de mejor manera usando gradientes de la temperatura de la superficie marina (media y varianza). En las áreas con un impacto humano reducido, los peces de gran tamaño corporal tenían hasta tres veces mayor probabilidad de aparecer y los peces de tamaño legal eran aproximadamente cinco veces más abundantes. Para los grupos de especies conspicuas con afinidades contrastantes de hábitat, ambiente y biogeografía, la abundancia de peces de tamaño legal normalmente declinó conforme aumentó el impacto humano. Nuestros análisis cuantitativos a gran escala resaltan la importancia conjunta que tienen la complejidad marina, las regiones con una huella humana reducida y las reservas marinas de protección total para la protección de los peces de gran tamaño corporal en una extensa gama de especies y configuraciones ecosistémicas. Efectos de la Huella Humana y los Factores Biofísicos sobre la Estructura del Tamaño Corporal de Especies Marinas Capturadas para la Pesca.

Habitat configurations shape the trophic and energetic dynamics of reef fishes in a tropical-temperate transition zone: implications under a warming future.

Understanding the extent to which species' traits mediate patterns of community assembly is key to predict the effect of natural and anthropogenic disturbances on ecosystem functioning. Here, we apply a trait-based community assembly framework to understand how four different habitat configurations (kelp forests, Sargassum spp. beds, hard corals, and turfs) shape the trophic and energetic dynamics of reef fish assemblages in a tropical-temperate transition zone. Specifically, we tested (i) the degree of trait divergence and convergence in each habitat, (ii) which traits explained variation in species' abundances, and (iii) differences in standing biomass (kg ha-1), secondary productivity (kg ha-1 day-1) and turnover (% day-1). Fish assemblages in coral and kelp habitats displayed greater evidence of trait convergence, while turf and Sargassum spp. habitats displayed a higher degree of trait divergence, a pattern that was mostly driven by traits related to resource use and thermal affinity. This filtering effect had an imprint on the trophic and energetic dynamics of reef fishes, with turf habitats supporting higher fish biomass and productivity. However, these gains were strongly dependent on trophic guild, with herbivores/detritivores disproportionately contributing to among-habitat differences. Despite these perceived overall gains, turnover was decoupled for fishes that act as conduit of energy to higher trophic levels (i.e. microinvertivores), with coral habitats displaying higher rates of fish biomass replenishment than turf despite their lower productivity. This has important implications for biodiversity conservation and fisheries management, questioning the long-term sustainability of ecological processes and fisheries yields in increasingly altered marine habitats.

Persistence of seaweed forests in the anthropocene will depend on warming and marine heatwave profiles.

Marine heatwaves (MHWs), discrete periods of extreme warm water temperatures superimposed onto persistent ocean warming, have increased in frequency and significantly disrupted marine ecosystems. While field observations on the ecological consequences of MHWs are growing, a mechanistic understanding of their direct effects is rare. We conducted an outdoor tank experiment testing how different thermal stressor profiles impacted the ecophysiological performance of three dominant forest-forming seaweeds. Four thermal scenarios were tested: contemporary summer temperature (22°C), low persistent warming (24°C), a discrete MHW (22-27°C), and temperature variability followed by a MHW (22-24°C, 22-27°C). The physiological performance of seaweeds was strongly related to thermal profile and varied among species, with the highest temperature not always having the strongest effect. MHWs were highly detrimental for the fucoid Phyllospora comosa, whereas the laminarian kelp Ecklonia radiata showed sensitivity to extended thermal stress and demonstrated a cumulative temperature threshold. The fucoid Sargassum linearifolium showed resilience, albeit with signs of decline with bleached and degraded fronds, under all conditions, with stronger decline under stable control and warming conditions. The varying responses of these three co-occurring forest-forming seaweeds under different temperature scenarios suggests that the impact of ocean warming on near shore ecosystems may be complex and will depend on the specific thermal profile of rising water temperatures relative to the vulnerability of different species.

Leveraging the blue economy to transform marine forest restoration.

The UN Decade of Ecosystem Restoration is a response to the urgent need to substantially accelerate and upscale ecological restoration to secure Earth's sustainable future. Globally, restoration commitments have focused overwhelmingly on terrestrial forests. In contrast, despite a strong value proposition, efforts to restore seaweed forests lag far behind other major ecosystems and continue to be dominated by small-scale, short-term academic experiments. However, seaweed forest restoration can match the scale of damage and threat if moved from academia into the hands of community groups, industry, and restoration practitioners. Connecting two rapidly growing sectors in the Blue Economy-seaweed cultivation and the restoration industry-can transform marine forest restoration into a commercial-scale enterprise that can make a significant contribution to global restoration efforts.

Genotype-Environment mismatch of kelp forests under climate change.

Climate change is increasingly impacting ecosystems globally. Understanding adaptive genetic diversity and whether it will keep pace with projected climatic change is necessary to assess species' vulnerability and design efficient mitigation strategies such as assisted adaptation. Kelp forests are the foundations of temperate reefs globally but are declining in many regions due to climate stress. A lack of knowledge of kelp's adaptive genetic diversity hinders assessment of vulnerability under extant and future climates. Using 4245 single nucleotide polymorphisms (SNPs), we characterized patterns of neutral and putative adaptive genetic diversity for the dominant kelp in the southern hemisphere (Ecklonia radiata) from ~1000 km of coastline off Western Australia. Strong population structure and isolation-by-distance was underpinned by significant signatures of selection related to temperature and light. Gradient forest analysis of temperature-linked SNPs under selection revealed a strong association with mean annual temperature range, suggesting adaptation to local thermal environments. Critically, modelling revealed that predicted climate-mediated temperature changes will probably result in high genomic vulnerability via a mismatch between current and future predicted genotype-environment relationships such that kelp forests off Western Australia will need to significantly adapt to keep pace with projected climate change. Proactive management techniques such as assisted adaptation to boost resilience may be required to secure the future of these kelp forests and the immense ecological and economic values they support.

Homogenization and miniaturization of habitat structure in temperate marine forests.

Humans are rapidly transforming the structural configuration of the planet's ecosystems, but these changes and their ecological consequences remain poorly quantified in underwater habitats. Here, we show that the loss of forest-forming seaweeds and the rise of ground-covering 'turfs' across four continents consistently resulted in the miniaturization of underwater habitat structure, with seascapes converging towards flattened habitats with smaller habitable spaces. Globally, turf seascapes occupied a smaller architectural trait space and were structurally more similar across regions than marine forests, evidencing habitat homogenization. Surprisingly, such habitat convergence occurred despite turf seascapes consisting of vastly different species richness and with different taxa providing habitat architecture, as well as across disparate drivers of marine forest decline. Turf seascapes contained high sediment loads, with the miniaturization of habitat across 100s of km in mid-Western Australia resulting in reefs retaining an additional ~242 million tons of sediment (four orders of magnitude more than the sediments delivered fluvially annually). Together, this work demonstrates that the replacement of marine forests by turfs is a generalizable phenomenon that has profound consequences for the ecology of temperate reefs.

Local flexibility in feeding behaviour and contrasting microhabitat use of an omnivore across latitudes.

As the environment is getting warmer and species are redistributed, consumers can be forced to adjust their interactions with available prey, and this could have cascading effects within food webs. To better understand the capacity for foraging flexibility, our study aimed to determine the diet variability of an ectotherm omnivore inhabiting kelp forests, the sea urchin Echinus esculentus, along its entire latitudinal distribution in the northeast Atlantic. Using a combination of gut content and stable isotope analyses, we determined the diet and trophic position of sea urchins at sites in Portugal (42° N), France (49° N), southern Norway (63° N), and northern Norway (70° N), and related these results to the local abundance and distribution of putative food items. With mean estimated trophic levels ranging from 2.4 to 4.6, omnivory and diet varied substantially within and between sites but not across latitudes. Diet composition generally reflected prey availability within epiphyte or understorey assemblages, with local affinities demonstrating that the sea urchin adjusts its foraging to match the small-scale distribution of food items. A net "preference" for epiphytic food sources was found in northern Norway, where understorey food was limited compared to other regions. We conclude that diet change may occur in response to food source redistribution at multiple spatial scales (microhabitats, sites, regions). Across these scales, the way that key consumers alter their foraging in response to food availability can have important implication for food web dynamics and ecosystem functions along current and future environmental gradients.

Carbon export is facilitated by sea urchins transforming kelp detritus.

With the increasing imperative for societies to act to curb climate change by increasing carbon stores and sinks, it has become critical to understand how organic carbon is produced, released, transformed, transported, and sequestered within and across ecosystems. In freshwater and open-ocean systems, shredders play a significant and well-known role in transforming and mobilizing carbon, but their role in the carbon cycle of coastal ecosystems is largely unknown. Marine plants such as kelps produce vast amounts of detritus, which can be captured and consumed by shedders as it traverses the seafloor. We measured capture and consumption rates of kelp detritus by sea urchins across four sampling periods and over a range of kelp detritus production rates and sea urchin densities, in northern Norway. When sea urchin densities exceeded 4 m-2, the sea urchins captured and consumed a high percentage (ca. 80%) of kelp detritus on shallow reefs. We calculated that between 1.3 and 10.8 kg of kelp m-2 are shredded annually from these reefs. We used a hydrodynamic dispersal model to show that transformation of kelp blades to sea urchin feces increased its export distance fourfold. Our findings show that sea urchins can accelerate and extend the export of carbon to neighboring areas. This collector-shredder pathway could represent a significant flow of small particulate carbon from kelp forests to deep-sea areas, where it can subsidize benthic communities or contribute to the global carbon sink.

Detrital carbon production and export in high latitude kelp forests.

The production and fate of seaweed detritus is a major unknown in the global C-budget. Knowing the quantity of detritus produced, the form it takes (size) and its timing of delivery are key to understanding its role as a resource subsidy to secondary production and/or its potential contribution to C-sequestration. We quantified the production and release of detritus from 10 Laminaria hyperborea sites in northern Norway (69.6° N). Kelp biomass averaged 770 ± 100 g C m-2 while net production reached 499 ± 50 g C m-2 year-1, with most taking place in spring when new blades were formed. Production of biomass was balanced by a similar formation of detritus (478 ± 41 g C m-2 year-1), and both were unrelated to wave exposure when compared across sites. Distal blade erosion accounted for 23% of the total detritus production and was highest during autumn and winter, while dislodgment of whole individuals and/or whole blades corresponded to 24% of the detritus production. Detachment of old blades constituted the largest source of kelp detritus, accounting for > 50% of the total detrital production. Almost 80% of the detritus from L. hyperborea was thus in the form of whole plants or blades and > 60% of that was delivered as a large pulse within 1-2 months in spring. The discrete nature of the delivery suggests that the detritus cannot be retained and consumed locally and that some is exported to adjacent deep areas where it may subsidize secondary production or become buried into deep marine sediments as blue carbon.

Overwintering tropical herbivores accelerate detritus production on temperate reefs.

The tropicalization of temperate marine ecosystems can lead to increased herbivory rates, reducing the standing stock of seaweeds and potentially causing increases in detritus production. However, long-term studies analysing these processes associated with the persistence of tropical herbivores in temperate reefs are lacking. We assessed the seasonal variation in abundances, macrophyte consumption, feeding modes and defecation rates of the range-extending tropical rabbitfish Siganus fuscescens and the temperate silver drummer Kyphosus sydneyanus and herring cale Olisthops cyanomelas on tropicalized reefs of Western Australia. Rabbitfish overwintered in temperate reefs, consumed more kelp and other macrophytes in all feeding modes, and defecated more during both summer and winter than the temperate herbivores. Herbivory and defecation increased with rabbitfish abundance, but this was dependent on temperature, with higher rates attained by big schools during summer and lower rates in winter. Still, rabbitfish surpassed temperate herbivores, leading to a fivefold acceleration in the transformation of macrophyte standing stock to detritus, a function usually attributed to sea urchins in kelp forests. Our results suggest that further warming and tropicalization will not only increase primary consumption and affect the habitat structure of temperate reefs but also increase detritus production, with the potential to modify energy pathways.