Global patterns and predictions of seafloor biomass using random forests

A comprehensive seafloor biomass and abundance database has been constructed from 24 oceanographic institutions worldwide within the Census of Marine Life (CoML) field projects. The machine-learning algorithm, Random Forests, was employed to model and predict seafloor standing stocks from surface primary production, water-column integrated and export particulate organic matter (POM), seafloor relief, and bottom water properties. The predictive models explain 63% to 88% of stock variance among the major size groups. Individual and composite maps of predicted global seafloor biomass and abundance are generated for bacteria, meiofauna, macrofauna, and megafauna (invertebrates and fishes). Patterns of benthic standing stocks were positive functions of surface primary production and delivery of the particulate organic carbon (POC) flux to the seafloor. At a regional scale, the census maps illustrate that integrated biomass is highest at the poles, on continental margins associated with coastal upwelling and with broad zones associated with equatorial divergence. Lowest values are consistently encountered on the central abyssal plains of major ocean basins The shift of biomass dominance groups with depth is shown to be affected by the decrease in average body size rather than abundance, presumably due to decrease in quantity and quality of food supply. This biomass census and associated maps are vital components of mechanistic deep-sea food web models and global carbon cycling, and as such provide fundamental information that can be incorporated into evidence-based management.     

Spatial and body‐size dependent response of marine pelagic communities to projected global climate change

Temperature, oxygen, and food availability directly affect marine life. Climate models project a global warming of the ocean's surface (~+3 °C), a de‐oxygenation of the ocean's interior (~?3%) and a decrease in total marine net primary production (~?8%) under the ‘business as usual’ climate change scenario (RCP8.5). We estimated the effects of these changes on biological communities using a coupled biogeochemical (PISCES) – ecosystems (APECOSM) model forced by the physical outputs of the last generation of the IPSL‐CM Earth System Model. The APECOSM model is a size‐structured bio‐energetic model that simulates the 3D dynamical distributions of three interactive pelagic communities (epipelagic, mesopelagic, and migratory) under the effects of multiple environmental factors. The PISCES‐APECOSM model ran from 1850 to 2100 under historical forcing followed by RCP8.5. Our RCP8.5 simulation highlights significant changes in the spatial distribution, biomass, and maximum body‐size of the simulated pelagic communities. Biomass and maximum body‐size increase at high latitude over the course of the century, reflecting the capacity of marine organisms to respond to new suitable environment. At low‐ and midlatitude, biomass and maximum body‐size strongly decrease. In those regions, large organisms cannot maintain their high metabolic needs because of limited and declining food availability. This resource reduction enhances the competition and modifies the biomass distribution among and within the three communities: the proportion of small organisms increases in the three communities and the migrant community that initially comprised a higher proportion of small organisms is favored. The greater resilience of small body‐size organisms resides in their capacity to fulfill their metabolic needs under reduced energy supply and is further favored by the release of predation pressure due to the decline of large organisms. These results suggest that small body‐size organisms might be more resilient to climate change than large ones.     

Terrestrial subsidies to lake food webs: an experimental approach

Cross-ecosystem movements of material and energy are ubiquitous. Aquatic ecosystems typically receive material that also includes organic matter from the surrounding catchment. Terrestrial-derived (allochthonous) organic matter can enter aquatic ecosystems in dissolved or particulate form. Several studies have highlighted the importance of dissolved organic carbon to aquatic consumers, but less is known about allochthonous particulate organic carbon (POC). Similarly, most studies showing the effects of allochthonous organic carbon (OC) on aquatic consumers have investigated pelagic habitats; the effects of allochthonous OC on benthic communities are less well studied. Allochthonous inputs might further decrease primary production through light reduction, thereby potentially affecting autotrophic resource availability to consumers. Here, an enclosure experiment was carried out to test the importance of POC input and light availability on the resource use in a benthic food web of a clear-water lake. Corn starch (a C(4) plant) was used as a POC source due to its insoluble nature and its distinct carbon stable isotope value (δ(13)C). The starch carbon was closely dispersed over the bottom of the enclosures to study the fate of a POC source exclusively available to sediment biota. The addition of starch carbon resulted in a clear shift in the isotopic signature of surface-dwelling herbivorous and predatory invertebrates. Although the starch carbon was added solely to the sediment surface, the carbon originating from the starch reached zooplankton. We suggest that allochthonous POC can subsidize benthic food webs directly and can be further transferred to pelagic systems, thereby highlighting the importance of benthic pathways for pelagic habitats.     

The meagre future of benthic fauna in a coastal sea—Benthic responses to recovery from eutrophication in a changing climate

Nutrient loading and climate change affect coastal ecosystems worldwide. Unravelling the combined effects of these pressures on benthic macrofauna is essential for understanding the future functioning of coastal ecosystems, as it is an important component linking the benthic and pelagic realms. In this study, we extended an existing model of benthic macrofauna coupled with a physical–biogeochemical model of the Baltic Sea to study the combined effects of changing nutrient loads and climate on biomass and metabolism of benthic macrofauna historically and in scenarios for the future. Based on a statistical comparison with a large validation dataset of measured biomasses, the model showed good or reasonable performance across the different basins and depth strata in the model area. In scenarios with decreasing nutrient loads according to the Baltic Sea Action Plan but also with continued recent loads (mean loads 2012–2014), overall macrofaunal biomass and carbon processing were projected to decrease significantly by the end of the century despite improved oxygen conditions at the seafloor. Climate change led to intensified pelagic recycling of primary production and reduced export of particulate organic carbon to the seafloor with negative effects on macrofaunal biomass. In the high nutrient load scenario, representing the highest recorded historical loads, climate change counteracted the effects of increased productivity leading to a hyperbolic response: biomass and carbon processing increased up to mid‐21st century but then decreased, giving almost no net change by the end of the 21st century compared to present. The study shows that benthic responses to environmental change are nonlinear and partly decoupled from pelagic responses and indicates that benthic–pelagic coupling might be weaker in a warmer and less eutrophic sea.     

Benthic-Pelagic Coupling: Effects on Nematode Communities along Southern European Continental Margins

Along a west-to-east axis spanning the Galicia Bank region (Iberian margin) and the Mediterranean basin, a reduction in surface primary productivity and in seafloor flux of particulate organic carbon was mirrored in the in situ organic matter quantity and quality within the underlying deep-sea sediments at different water depths (1200, 1900 and 3000 m). Nematode standing stock (abundance and biomass) and genus and trophic composition were investigated to evaluate downward benthic-pelagic coupling. The longitudinal decline in seafloor particulate organic carbon flux was reflected by a reduction in benthic phytopigment concentrations and nematode standing stock. An exception was the station sampled at the Galicia Bank seamount, where despite the maximal particulate organic carbon flux estimate, we observed reduced pigment levels and nematode standing stock. The strong hydrodynamic forcing at this station was believed to be the main cause of the local decoupling between pelagic and benthic processes. Besides a longitudinal cline in nematode standing stock, we noticed a west-to-east gradient in nematode genus and feeding type composition (owing to an increasing importance of predatory/scavenging nematodes with longitude) governed by potential proxies for food availability (percentage of nitrogen, organic carbon, and total organic matter). Within-station variability in generic composition was elevated in sediments with lower phytopigment concentrations. Standing stock appeared to be regulated by sedimentation rates and benthic environmental variables, whereas genus composition covaried only with benthic environmental variables. The coupling between deep-sea nematode assemblages and surface water processes evidenced in the present study suggests that it is likely that climate change will affect the composition and function of deep-sea nematodes.     

Protist diversity and function in the dark ocean – Challenging the paradigms of deep-sea ecology with special emphasis on foraminiferans and naked protists

The dark ocean and the underlying deep seafloor together represent the largest environment on this planet, comprising about 80% of the oceanic volume and covering more than two-thirds of the Earth's surface, as well as hosting a major part of the total biosphere. Emerging evidence suggests that these vast pelagic and benthic habitats play a major role in ocean biogeochemistry and represent an “untapped reservoir” of high genetic and metabolic microbial diversity. Due to its huge volume, the water column of the dark ocean is the largest reservoir of organic carbon in the biosphere and likely plays a major role in the global carbon budget. The dark ocean and the seafloor beneath it are also home to a largely enigmatic food web comprising little-known and sometimes spectacular organisms, mainly prokaryotes and protists. This review considers the globally important role of pelagic and benthic protists across all protistan size classes in the deep-sea realm, with a focus on their taxonomy, diversity, and physiological properties, including their role in deep microbial food webs. We argue that, given the important contribution that protists must make to deep-sea biodiversity and ecosystem processes, they should not be overlooked in biological studies of the deep ocean.     

Contribution of benthic microalgae to the temporal variation in phytoplankton assemblages in a macrotidal system

Suspended marine benthic microalgae in the water column reflect the close relationship between the benthic and pelagic components of coastal ecosystems. In this study, a 12‐year phytoplankton time‐series was used to investigate the contribution of benthic microalgae to the pelagic system at a site along the French‐Atlantic coast. Furthermore, all taxa identified were allocated into different growth forms in order to study their seasonal patterns. The highest contribution of benthic microalgae was observed during the winter period, reaching up to 60% of the carbon biomass in the water column. The haptobenthic growth form showed the highest contribution in terms of biomass, dominant in the fall‐winter period when the turbidity and the river flow were high. The epipelic growth form did not follow any seasonal pattern. The epiphytic diatom Licmophora was most commonly found during summer. As benthic microalgae were found in the water column throughout the year, the temporal variation detected in the structure of pelagic assemblages in a macrotidal ecosystem was partly derived from the differentiated contribution of several benthic growth forms.     

Benthic–pelagic coupling: a comparison of the community structure of benthic and planktonic heterotrophic protists in shallow inlets of the southern Baltic

1. The taxonomic composition, abundance and biomass of heterotrophic protists (ciliates, heterotrophic flagellates (HF), rhizopods and actinopods) in the sediment and water column of shallow inlets of the Southern Baltic was studied under a variety of environmental conditions during 1996–1997. A shallow, highly eutrophic station and a deeper, less eutrophic station were compared.
2. Community biomass ranged from 0.12 to 0.34 μg C cm?3 in the water column and from 1.5 to 105 μg C cm?3 in the sediment. Heterotrophic protists dominated zooplankton biomass at both stations (73% and 84% mean contribution), while they were of minor importance within the zoobenthos. Expressed per unit area, benthic biomass contributed a significant part (44% and 49%) to the total heterotrophic protistan community at both stations.
3. Although the methodology for counting ciliates and HF was focussed on a high taxonomic resolution, the results reveal some general trends in the distribution of heterotrophic protists: protozooplankton biomass was dominated by flagellates (80% mean biomass contribution) at the shallow station and by ciliates (73% mean biomass contribution) at the deep station. In the benthos at both stations, ciliates were the dominant protozoans, followed by the hitherto little‐studied rhizopods (25% and 35% mean biomass contribution) and flagellates.
4. The degree of benthic–pelagic coupling differed between taxonomic groups. Benthic and pelagic communities of ciliates showed little taxonomic overlap. In contrast, many heterotrophic flagellate species were found both in the benthos and in the pelagic. These benthic–pelagic species contributed significantly to the biomass of HF in the water column. The planktonic rhizopod community consisted of a subset of those species found in the benthos.
5. The abundance of benthic and pelagic protists was positively correlated at the shallow station, but taxonomic data indicate that the direct exchange between benthic and pelagic communities was only partly responsible.     

Filter-feeding gelatinous macrozooplankton response to climate change and implications for benthic food supply and global carbon cycle

It is often suggested that gelatinous zooplankton may benefit from anthropogenic pressures of all kinds and in particular from climate change. Large pelagic tunicates, for example, are likely to be favored over other types of macrozooplankton due to their filter-feeding mode, which gives them access to small preys thought to be less affected by climate change than larger preys. In this study, we provide model-based estimate of potential community changes in macrozooplankton composition and estimate for the first time their effects on benthic food supply and on the ocean carbon cycle under two 21st-century climate-change scenarios. Forced with output from an Earth System Model climate projections, our ocean biogeochemical model simulates a large reduction in macrozooplankton biomass in response to anthropogenic climate change, but shows that gelatinous macrozooplankton are less affected than nongelatinous macrozooplankton, with global biomass declines estimated at −2.8% and −3.5%, respectively, for every 1°C of warming. The inclusion of gelatinous macrozooplankon in our ocean biogeochemical model has a limited effect on anthropogenic carbon uptake in the 21st century, but impacts the projected decline in particulate organic matter fluxes in the deep ocean. In subtropical oligotrophic gyres, where gelatinous zooplankton dominate macrozooplankton, the decline in the amount of organic matter reaching the seafloor is reduced by a factor of 2 when gelatinous macrozooplankton are considered (−17.5% vs. −29.7% when gelatinous macrozooplankton are not considered, all for 2100 under RCP8.5). The shift to gelatinous macrozooplankton in the future ocean therefore buffers the decline in deep carbon fluxes and should be taken into account when assessing potential changes in deep carbon storage and the risks that deep ecosystems may face when confronted with a decline in their food source.     

Global patterns of bacterial beta-diversity in seafloor and seawater ecosystems

Background

Marine microbial communities have been essential contributors to global biomass, nutrient cycling, and biodiversity since the early history of Earth, but so far their community distribution patterns remain unknown in most marine ecosystems.

Methodology/Principal Findings

The synthesis of 9.6 million bacterial V6-rRNA amplicons for 509 samples that span the global ocean''s surface to the deep-sea floor shows that pelagic and benthic communities greatly differ, at all taxonomic levels, and share <10% bacterial types defined at 3% sequence similarity level. Surface and deep water, coastal and open ocean, and anoxic and oxic ecosystems host distinct communities that reflect productivity, land influences and other environmental constraints such as oxygen availability. The high variability of bacterial community composition specific to vent and coastal ecosystems reflects the heterogeneity and dynamic nature of these habitats. Both pelagic and benthic bacterial community distributions correlate with surface water productivity, reflecting the coupling between both realms by particle export. Also, differences in physical mixing may play a fundamental role in the distribution patterns of marine bacteria, as benthic communities showed a higher dissimilarity with increasing distance than pelagic communities.

Conclusions/Significance

This first synthesis of global bacterial distribution across different ecosystems of the World''s oceans shows remarkable horizontal and vertical large-scale patterns in bacterial communities. This opens interesting perspectives for the definition of biogeographical biomes for bacteria of ocean waters and the seabed.     

Zebra mussels mediate benthic–pelagic coupling by biodeposition and changing detrital stoichiometry

1. The zebra mussel ( Dreissena polymorpha ) is one of the most successful invasive species; it has colonised many aquatic systems in Europe and North America with strong impacts on various ecosystem processes. The effect of D. polymorpha filtration on pelagic seston concentrations has been quantified in several studies, but the magnitude and stoichiometry of the transfer of sestonic biomass into benthic detritus by D. polymorpha and the accompanying enrichment of the benthic habitat is still under-investigated.
2. We studied biodeposition by zebra mussels in two series of laboratory experiments with the food algae Cryptomonas erosa and Scenedesmus obliquus . We also measured the year-round biodeposition rate under natural conditions in the oligotrophic Lake Constance.
3. In all experiments, zebra mussel biodeposition was linearly related to seston concentration. In the field, the relationship changed with a seasonal shift in algal composition and lower biodeposition rates during the spring algal bloom.
4. For both algal species in laboratory experiments, biodeposited material was depleted in phosphorous at an algal concentration ≤0.6 mg ash-free dry mass L⁻¹, but not at higher concentrations. This effect was not observed in the field, probably because of high variation in C : N : P stoichiometry.
5. By mediating the transfer of pelagic resources into the benthos zebra mussels provide a sufficient amount of detritus for benthic invertebrates, especially during summer. Thus, material biodeposited by the mussels might increase benthic secondary production from pelagic resources, and zebra mussels are important mediators of this flux of organic matter from the pelagic zone into the benthos.     

Benthic carbon is inefficiently transferred in the food webs of two eutrophic shallow lakes

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Bottom‐up and top‐down effects of browning and warming on shallow lake food webs

Productivity and trophic structure of aquatic ecosystems result from a complex interplay of bottom‐up and top‐down forces that operate across benthic and pelagic food web compartments. Projected global changes urge the question how this interplay will be affected by browning (increasing input of terrestrial dissolved organic matter), nutrient enrichment and warming. We explored this with a process‐based model of a shallow lake food web consisting of benthic and pelagic components (abiotic resources, primary producers, grazers, carnivores), and compared model expectations with the results of a browning and warming experiment in nutrient‐poor ponds harboring a boreal lake community. Under low nutrient conditions, the model makes three major predictions. (a) Browning reduces light and increases nutrient supply; this decreases benthic and increases pelagic production, gradually shifting productivity from the benthic to the pelagic habitat. (b) Because of active habitat choice, fish exert top‐down control on grazers and benefit primary producers primarily in the more productive of the two habitats. (c) Warming relaxes top‐down control of grazers by fish and decreases primary producer biomass, but effects of warming are generally small compared to effects of browning and nutrient supply. Experimental results were consistent with most model predictions for browning: light penetration, benthic algal production, and zoobenthos biomass decreased, and pelagic nutrients and pelagic algal production increased with browning. Also consistent with expectations, warming had negative effects on benthic and pelagic algal biomass and weak effects on algal production and zoobenthos and zooplankton biomass. Inconsistent with expectations, browning had no effect on zooplankton and warming effects on fish depended on browning. The model is applicable also to nutrient‐rich systems, and we propose that it is a useful tool for the exploration of the consequences of different climate change scenarios for productivity and food web dynamics in shallow lakes, the worldwide most common lake type.     

Food sources and lipid retention of zooplankton in subarctic ponds

1. Subarctic ponds are seasonal aquatic habitats subject to short summers but often have surprisingly numerous planktonic consumers relative to phytoplankton productivity. Because subarctic ponds have low pelagic productivity but a high biomass of benthic algae, we hypothesised that benthic mats provide a complementary and important food source for the zooplankton. To test this, we used a combination of fatty acid and stable isotope analyses to evaluate the nutritional content of benthic and pelagic food and their contributions to the diets of crustacean zooplankton in 10 Finnish subarctic ponds. 2. Benthic mats and seston differed significantly in total lipids, with seston (62.5 μg mg^?1) having approximately eight times higher total lipid concentrations than benthic mats (7.0 μg mg^?1). Moreover, the two potential food sources differed in their lipid quality, with benthic organic matter completely lacking some nutritionally important polyunsaturated fatty acids (PUFA), most notably docosahexaenoic acid and arachidonic acid. 3. Zooplankton had higher PUFA concentrations (27–67 μg mg^?1) than either of the food sources (mean benthic mats: 1.2 μg mg^?1; mean seston: 9.9 μg mg^?1), indicating that zooplankton metabolically regulate their accumulation of PUFA. In addition, when each pond was evaluated independently, the zooplankton was consistently more ¹³C‐depleted (δ¹³C ?20 to ?33‰) than seston (?23 to ?29‰) or benthic (?15 to ?27‰) food sources. In three ponds, a subset of the zooplankton (Eudiaptomus graciloides, Bosmina sp., Daphnia sp. and Branchinecta paludosa) showed evidence of feeding on both benthic and planktonic resources, whereas in most (seven out of 10) ponds the zooplankton appeared to feed primarily on plankton. 4. Our results indicate that pelagic primary production was consistently the principal food resource of most metazoans. While benthic mats were highly productive, they did not appear to be a major food source for zooplankton. The pond zooplankton, faced by strong seasonal food limitation, acquires particular dietary elements selectively.     

Benthic food web structure in the southeastern Chukchi Sea: an assessment using ��13C and ��15N analyses

The benthos of the southeastern Chukchi Sea shelf is typified by high faunal abundance and biomass resulting from settlement of a large proportion of seasonal phytoplankton under highly nutritious offshore Bering Shelf Anadyr Water (BSAW). In contrast, inshore Alaska Coastal Water (ACW) is much less productive. Yet the Chukchi Bight and Kotzebue Sound, located under ACW in the southeastern Chukchi Sea, contain a substantial faunal abundance and biomass of invertebrates, fishes and marine mammals. We examined food web structure to gain an understanding of how a relatively rich benthic fauna with a high biomass can be supported under ACW with a supposedly low flux of carbon to the benthos. We measured stable isotope (δ¹³C and δ¹⁵N) values of selected organisms (from zooplankton to fishes) as markers of food sources and trophic position to compare fauna on the shelf under BSAW with that in the Chukchi Bight and Kotzebue Sound under ACW. Relative isotope position of organisms in all three regions was similar, even though some pelagic species within the Sound were depleted in δ¹³C compared to the other regions. We attribute the depletion to the influence of terrestrially derived carbon. We suggest that the hydrodynamics along an oceanic front between the Chukchi Shelf and the Chukchi Bight support the advection of nutrient-rich POC into the Bight and Sound as additional food sources to local production. We conclude that local conditions and multiple POC sources in the Bight and Sound support the substantial population of benthic invertebrates and the fishes, seabirds, and marine mammals that feed on them.     

Nitrogen cycling in coastal marine ecosystems

It is generally considered that nitrogen availability is one of the major factors regulating primary production in temperate coastal marine environments. Coastal regions often receive large anthropogenic inputs of nitrogen that cause eutrophication. The impact of these nitrogen additions has a profound effect in estuaries and coastal lagoons where water exchange is limited. Such increased nutrient loading promotes the growth of phytoplankton and fast growing pelagic macroalgae while rooted plants (sea-grasses) and benthic are suppressed due to reduced light availability. This shift from benthic to pelagic primary production introduces large diurnal variations in oxygen concentrations in the water column. In addition oxygen consumption in the surface sediments increases due to the deposition of readily degradable biomass. In this review the physico-chemical and biological factors regulating nitrogen cycling in coastal marine ecosystems are considered in relation to developing effective management programmes to rehabilitate seagrass communities in lagoons currently dominated by pelagic macroalgae and/or cyanobacteria.     

Coral reefs modify their seawater carbon chemistry – case study from a barrier reef (Moorea,French Polynesia)

Changes in the carbonate chemistry of coral reef waters are driven by carbon fluxes from two sources: concentrations of CO₂ in the atmospheric and source water, and the primary production/respiration and calcification/dissolution of the benthic community. Recent model analyses have shown that, depending on the composition of the reef community, the air‐sea flux of CO₂ driven by benthic community processes can exceed that due to increases in atmospheric CO₂ (ocean acidification). We field test this model and examine the role of three key members of benthic reef communities in modifying the chemistry of the ocean source water: corals, macroalgae, and sand. Building on data from previous carbon flux studies along a reef‐flat transect in Moorea (French Polynesia), we illustrate that the drawdown of total dissolved inorganic carbon (C_T) due to photosynthesis and calcification of reef communities can exceed the draw down of total alkalinity (A_T) due to calcification of corals and calcifying algae, leading to a net increase in aragonite saturation state (Ω_a). We use the model to test how changes in atmospheric CO₂ forcing and benthic community structure affect the overall calcification rates on the reef flat. Results show that between the preindustrial period and 1992, ocean acidification caused reef flat calcification rates to decline by an estimated 15%, but loss of coral cover caused calcification rates to decline by at least three times that amount. The results also show that the upstream–downstream patterns of carbonate chemistry were affected by the spatial patterns of benthic community structure. Changes in the ratio of photosynthesis to calcification can thus partially compensate for ocean acidification, at least on shallow reef flats. With no change in benthic community structure, however, ocean acidification depressed net calcification of the reef flat consistent with findings of previous studies.     

Climate and productivity shape fish and invertebrate community structure in subarctic lakes

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Global reductions in seafloor biomass in response to climate change

Seafloor organisms are vital for healthy marine ecosystems, contributing to elemental cycling, benthic remineralization, and ultimately sequestration of carbon. Deep‐sea life is primarily reliant on the export flux of particulate organic carbon from the surface ocean for food, but most ocean biogeochemistry models predict global decreases in export flux resulting from 21st century anthropogenically induced warming. Here we show that decadal‐to‐century scale changes in carbon export associated with climate change lead to an estimated 5.2% decrease in future (2091–2100) global open ocean benthic biomass under RCP8.5 (reduction of 5.2 Mt C) compared with contemporary conditions (2006–2015). Our projections use multi‐model mean export flux estimates from eight fully coupled earth system models, which contributed to the Coupled Model Intercomparison Project Phase 5, that have been forced by high and low representative concentration pathways (RCP8.5 and 4.5, respectively). These export flux estimates are used in conjunction with published empirical relationships to predict changes in benthic biomass. The polar oceans and some upwelling areas may experience increases in benthic biomass, but most other regions show decreases, with up to 38% reductions in parts of the northeast Atlantic. Our analysis projects a future ocean with smaller sized infaunal benthos, potentially reducing energy transfer rates though benthic multicellular food webs. More than 80% of potential deep‐water biodiversity hotspots known around the world, including canyons, seamounts, and cold‐water coral reefs, are projected to experience negative changes in biomass. These major reductions in biomass may lead to widespread change in benthic ecosystems and the functions and services they provide.     

Variable littoral‐pelagic coupling as a food‐web response to seasonal changes in pelagic primary production

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