Past and future decline of tropical pelagic biodiversity.

A major research question concerning global pelagic biodiversity remains unanswered: when did the apparent tropical biodiversity depression (i.e., bimodality of latitudinal diversity gradient [LDG]) begin? The bimodal LDG may be a consequence of recent ocean warming or of deep-time evolutionary speciation and extinction processes. Using rich fossil datasets of planktonic foraminifers, we show here that a unimodal (or only weakly bimodal) diversity gradient, with a plateau in the tropics, occurred during the last ice age and has since then developed into a bimodal gradient through species distribution shifts driven by postglacial ocean warming. The bimodal LDG likely emerged before the Anthropocene and industrialization, and perhaps ∼15,000 y ago, indicating a strong environmental control of tropical diversity even before the start of anthropogenic warming. However, our model projections suggest that future anthropogenic warming further diminishes tropical pelagic diversity to a level not seen in millions of years.

Climate change considerations are fundamental to management of deep-sea resource extraction.

Climate change manifestation in the ocean, through warming, oxygen loss, increasing acidification, and changing particulate organic carbon flux (one metric of altered food supply), is projected to affect most deep-ocean ecosystems concomitantly with increasing direct human disturbance. Climate drivers will alter deep-sea biodiversity and associated ecosystem services, and may interact with disturbance from resource extraction activities or even climate geoengineering. We suggest that to ensure the effective management of increasing use of the deep ocean (e.g., for bottom fishing, oil and gas extraction, and deep-seabed mining), environmental management and developing regulations must consider climate change. Strategic planning, impact assessment and monitoring, spatial management, application of the precautionary approach, and full-cost accounting of extraction activities should embrace climate consciousness. Coupled climate and biological modeling approaches applied in the water and on the seafloor can help accomplish this goal. For example, Earth-System Model projections of climate-change parameters at the seafloor reveal heterogeneity in projected climate hazard and time of emergence (beyond natural variability) in regions targeted for deep-seabed mining. Models that combine climate-induced changes in ocean circulation with particle tracking predict altered transport of early life stages (larvae) under climate change. Habitat suitability models can help assess the consequences of altered larval dispersal, predict climate refugia, and identify vulnerable regions for multiple species under climate change. Engaging the deep observing community can support the necessary data provisioning to mainstream climate into the development of environmental management plans. To illustrate this approach, we focus on deep-seabed mining and the International Seabed Authority, whose mandates include regulation of all mineral-related activities in international waters and protecting the marine environment from the harmful effects of mining. However, achieving deep-ocean sustainability under the UN Sustainable Development Goals will require integration of climate consideration across all policy sectors.

Climate-induced changes in the suitable habitat of cold-water corals and commercially important deep-sea fishes in the North Atlantic.

The deep sea plays a critical role in global climate regulation through uptake and storage of heat and carbon dioxide. However, this regulating service causes warming, acidification and deoxygenation of deep waters, leading to decreased food availability at the seafloor. These changes and their projections are likely to affect productivity, biodiversity and distributions of deep-sea fauna, thereby compromising key ecosystem services. Understanding how climate change can lead to shifts in deep-sea species distributions is critically important in developing management measures. We used environmental niche modelling along with the best available species occurrence data and environmental parameters to model habitat suitability for key cold-water coral and commercially important deep-sea fish species under present-day (1951-2000) environmental conditions and to project changes under severe, high emissions future (2081-2100) climate projections (RCP8.5 scenario) for the North Atlantic Ocean. Our models projected a decrease of 28%-100% in suitable habitat for cold-water corals and a shift in suitable habitat for deep-sea fishes of 2.0°-9.9° towards higher latitudes. The largest reductions in suitable habitat were projected for the scleractinian coral Lophelia pertusa and the octocoral Paragorgia arborea, with declines of at least 79% and 99% respectively. We projected the expansion of suitable habitat by 2100 only for the fishes Helicolenus dactylopterus and Sebastes mentella (20%-30%), mostly through northern latitudinal range expansion. Our results projected limited climate refugia locations in the North Atlantic by 2100 for scleractinian corals (30%-42% of present-day suitable habitat), even smaller refugia locations for the octocorals Acanella arbuscula and Acanthogorgia armata (6%-14%), and almost no refugia for P. arborea. Our results emphasize the need to understand how anticipated climate change will affect the distribution of deep-sea species including commercially important fishes and foundation species, and highlight the importance of identifying and preserving climate refugia for a range of area-based planning and management tools.

Dongshaea marina gen. nov., sp. nov., a facultatively anaerobic marine bacterium that ferments glucose with gas production.

Two isolates of heterotrophic, facultatively anaerobic, marine bacteria, designated DM1 and DM2T, were recovered from a lagoon sediment sample of Dongsha Island, Taiwan. Cells were Gram-reaction-negative rods. Nearly all of the cells were non-motile and non-flagellated during the late exponential to early stationary phase of growth, while a few of the cells exhibited motility with monotrichous flagellation. The two isolates required NaCl for growth and grew optimally at about 30 °C, 2-3 % NaCl and pH 7-8. They grew aerobically and could achieve anaerobic growth by fermenting d-glucose or other carbohydrates with production of acids and the gases, including CO2 and H2. Ubiquinone Q-8 was the only respiratory quinone. Cellular fatty acids were predominated by C16 : 0, C18 : 1ω7c and C16 : 1ω7c. The major polar lipid was phosphatidylethanolamine. Strains DM1 and DM2T had DNA G+C contents of 52.0 and 51.8 mol%, respectively, as determined by HPLC analysis. Phylogenetic analyses based on 16S rRNA gene sequences clearly indicated that the two isolates formed a distinct genus-level lineage in the family Aeromonadaceae of the class Gammaproteobacteria and was an outgroup with respect to a stable supragenic clade comprising species of the genera Oceanimonas, Oceanisphaera and Zobellella. The phylogenetic data and those from chemotaxonomic, physiological and morphological characterizations support the establishment of a novel species and genus inside the family Aeromonadaceae, for which the name Dongshaea marina gen. nov., sp. nov. is proposed. The type strain is DM2T (=BCRC 81069T=JCM 32096T).

Biotic and human vulnerability to projected changes in ocean biogeochemistry over the 21st century.

Ongoing greenhouse gas emissions can modify climate processes and induce shifts in ocean temperature, pH, oxygen concentration, and productivity, which in turn could alter biological and social systems. Here, we provide a synoptic global assessment of the simultaneous changes in future ocean biogeochemical variables over marine biota and their broader implications for people. We analyzed modern Earth System Models forced by greenhouse gas concentration pathways until 2100 and showed that the entire world's ocean surface will be simultaneously impacted by varying intensities of ocean warming, acidification, oxygen depletion, or shortfalls in productivity. In contrast, only a small fraction of the world's ocean surface, mostly in polar regions, will experience increased oxygenation and productivity, while almost nowhere will there be ocean cooling or pH elevation. We compiled the global distribution of 32 marine habitats and biodiversity hotspots and found that they would all experience simultaneous exposure to changes in multiple biogeochemical variables. This superposition highlights the high risk for synergistic ecosystem responses, the suite of physiological adaptations needed to cope with future climate change, and the potential for reorganization of global biodiversity patterns. If co-occurring biogeochemical changes influence the delivery of ocean goods and services, then they could also have a considerable effect on human welfare. Approximately 470 to 870 million of the poorest people in the world rely heavily on the ocean for food, jobs, and revenues and live in countries that will be most affected by simultaneous changes in ocean biogeochemistry. These results highlight the high risk of degradation of marine ecosystems and associated human hardship expected in a future following current trends in anthropogenic greenhouse gas emissions.

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.

Drivers of coastal benthic communities in a complex environmental setting.

Analyzing the environmental factors affecting benthic communities in coastal areas is crucial for uncovering key factors that require conservation action. Here, we collected benthic and environmental (physical-chemical-historical and land-based) data for 433 transects in Taiwan. Using a k-means approach, five communities dominated by crustose coralline algae, turfs, stony corals, digitate, or bushy octocorals were first delineated. Conditional random forest models then identified physical, chemical, and land-based factors (e.g., light intensity, nitrite, and population density) relevant to community delineation and occurrence. Historical factors, including typhoons and temperature anomalies, had only little effect. The prevalent turf community correlated positively with chemical and land-based drivers, which suggests that anthropogenic impacts are causing a benthic homogenization. This mechanism may mask the effects of climate disturbances and regional differentiation of benthic assemblages. Consequently, management of nutrient enrichment and terrestrial runoff is urgently needed to improve community resilience in Taiwan amidst increasing challenges of climate change.

Ocean temperature drove changes in the mesopelagic fish community at the edge of the Pacific Warm Pool over the past 460,000 years.

Field observations and theoretical modeling suggest that ongoing anthropogenic ocean warming will lead to marine ecosystem degradation. Mesopelagic fish are a fundamental component of the pelagic ecosystem, and their role in linking the surface- and deep-ocean ecosystems is essential for the biological carbon pump. However, their response to a warmer ocean is unconstrained because of data scarcity. Using extraordinarily well-preserved fish otoliths, we reconstruct a continuous mesopelagic fish community record in the Pacific Warm Pool region over 460,000 years. Fish production and diversity followed hump-shaped temperature gradients, with lower tipping point temperatures for the diversity than the production by ~1.5° to 2.0°C. During warmer-than-present interglacial periods, both production and diversity declined drastically. Our findings imply that the temperature-sensitive mesopelagic fish community at the southwestern margin of the Pacific Warm Pool, and possibly other hydrographically similar regions, may be especially affected if ocean warming continues unabated in the future.

Three new species of Paratanaoidea (Crustacea, Tanaidacea) from the South China Sea.

Three new crustacean species of the order Tanaidacea are described from the South China Sea. Alloleptochelia falciformes sp. nov. (Family Leptocheliidae) can be distinguished from the most similar species, A. evansi (Edgar, 2012), by the much stouter body, and fewer articles on the antennule flagellum and relatively longer pereopods 46 dactylus of the male. Kalloleptochelia multiarticulata sp. nov. (Family Leptocheliidae) can be easily differentiated from its congeners by the male having significantly more antennule flagellum articles, a two-articled uropod exopod, one additional dorsal seta on the pereopods 23 merus, and one additional spiniform seta on the pereopod-6 carpus. Paranesotanais incisus sp. nov. (Family Nototanaidae) differs from its only congener, P. longicephalus Larsen & Shimomura, 2008, by the female having one robust inner seta on the palm of the cheliped propodus, and the male having one long and conspicuous notch extending from the outer surface to the ventrodistal corner on the cheliped carpus. Morphological keys and comparison tables of all known species of the genera Alloleptochelia and Kalloleptochelia, as well as a key of the genus Paranesotanais, are provided.

The BenBioDen database, a global database for meio-, macro- and megabenthic biomass and densities.

Benthic fauna refers to all fauna that live in or on the seafloor, which researchers typically divide into size classes meiobenthos (32/64 µm-0.5/1 mm), macrobenthos (250 µm-1 cm), and megabenthos (>1 cm). Benthic fauna play important roles in bioturbation activity, mineralization of organic matter, and in marine food webs. Evaluating their role in these ecosystem functions requires knowledge of their global distribution and biomass. We therefore established the BenBioDen database, the largest open-access database for marine benthic biomass and density data compiled so far. In total, it includes 11,792 georeferenced benthic biomass and 51,559 benthic density records from 384 and 600 studies, respectively. We selected all references following the procedure for systematic reviews and meta-analyses, and report biomass records as grams of wet mass, dry mass, or ash-free dry mass, or carbon per m2 and as abundance records as individuals per m2. This database provides a point of reference for future studies on the distribution and biomass of benthic fauna.

The SCOC database, a large, open, and global database with sediment community oxygen consumption rates.

Sediment community oxygen consumption (SCOC) rates provide important information about biogeochemical processes in marine sediments and the activity of benthic microorganisms and fauna. Therefore, several databases of SCOC data have been compiled since the mid-1990s. However, these earlier databases contained much less data records and were not freely available. Additionally, the databases were not transparent in their selection procedure, so that other researchers could not assess the quality of the data. Here, we present the largest, best documented, and freely available database of SCOC data compiled to date. The database is comprised of 3,540 georeferenced SCOC records from 230 studies that were selected following the procedure for systematic reviews and meta-analyses. Each data record states whether the oxygen consumption was measured ex situ or in situ, as total oxygen uptake, diffusive or advective oxygen uptake, and which measurement device was used. The database will be curated and updated annually to secure and maintain an up-to-date global database of SCOC data.

Steep redox gradient and biogeochemical cycling driven by deeply sourced fluids and gases in a terrestrial mud volcano.

Mud volcanoes provide an accessible channel through which deep subsurface environments can be observed. The manner in which deeply sourced materials shape biogeochemical processes and microbial communities in such geological features remains largely unknown. This study characterized redox transitions, biogeochemical fluxes and microbial communities for samples collected from a methane-rich mud volcano in southwestern Taiwan. Our results indicated that oxygen penetration was confined within the upper 4 mm of fluids/muds and counteracted by the oxidation of pyrite, dissolved sulfide, methane and organic matter at various degrees. Beneath the oxic zone, anaerobic sulfur oxidation, sulfate reduction, anaerobic methanotrophy and methanogenesis were compartmentalized into different depths in the pool periphery, forming a metabolic network that efficiently cycles methane and sulfur. Community members affiliated with various Proteobacteria capable of aerobic oxidation of sulfur, methane and methyl compounds were more abundant in the anoxic zone with diminished sulfate and high methane. These findings suggest either the requirement of alternative electron acceptors or a persistent population that once flourished in the oxic zone. Overall, this study demonstrates the distribution pattern for a suite of oxidative and reductive metabolic reactions along a steep redox gradient imposed by deep fluids in a mud volcano ecosystem.

Global Carbon Cycling on a Heterogeneous Seafloor.

Diverse biological communities mediate the transformation, transport, and storage of elements fundamental to life on Earth, including carbon, nitrogen, and oxygen. However, global biogeochemical model outcomes can vary by orders of magnitude, compromising capacity to project realistic ecosystem responses to planetary changes, including ocean productivity and climate. Here, we compare global carbon turnover rates estimated using models grounded in biological versus geochemical theory and argue that the turnover estimates based on each perspective yield divergent outcomes. Importantly, empirical studies that include sedimentary biological activity vary less than those that ignore it. Improving the relevance of model projections and reducing uncertainty associated with the anticipated consequences of global change requires reconciliation of these perspectives, enabling better societal decisions on mitigation and adaptation.

Biodiversity-ecosystem functioning relationships in long-term time series and palaeoecological records: deep sea as a test bed.

The link between biodiversity and ecosystem functioning (BEF) over long temporal scales is poorly understood. Here, we investigate biological monitoring and palaeoecological records on decadal, centennial and millennial time scales from a BEF framework by using deep sea, soft-sediment environments as a test bed. Results generally show positive BEF relationships, in agreement with BEF studies based on present-day spatial analyses and short-term manipulative experiments. However, the deep-sea BEF relationship is much noisier across longer time scales compared with modern observational studies. We also demonstrate with palaeoecological time-series data that a larger species pool does not enhance ecosystem stability through time, whereas higher abundance as an indicator of higher ecosystem functioning may enhance ecosystem stability. These results suggest that BEF relationships are potentially time scale-dependent. Environmental impacts on biodiversity and ecosystem functioning may be much stronger than biodiversity impacts on ecosystem functioning at long, decadal-millennial, time scales. Longer time scale perspectives, including palaeoecological and ecosystem monitoring data, are critical for predicting future BEF relationships on a rapidly changing planet.

Long-term observations of epibenthic fish zonation in the deep northern Gulf of Mexico.

A total of 172 bottom trawl/skimmer samples (183 to 3655-m depth) from three deep-sea studies, R/V Alaminos cruises (1964-1973), Northern Gulf of Mexico Continental Slope (NGoMCS) study (1983-1985) and Deep Gulf of Mexico Benthos (DGoMB) program (2000 to 2002), were compiled to examine temporal and large-scale changes in epibenthic fish species composition. Based on percent species shared among samples, faunal groups (≥10% species shared) consistently reoccurred over time on the shelf-break (ca. 200 m), upper-slope (ca. 300 to 500 m) and upper-to-mid slope (ca. 500 to 1500 m) depths. These similar depth groups also merged when the three studies were pooled together, suggesting that there has been no large-scale temporal change in depth zonation on the upper section of the continental margin. Permutational multivariate analysis of variance (PERMANOVA) also detected no significant species changes on the limited sites and areas that have been revisited across the studies (P>0.05). Based on the ordination of the species shared among samples, species replacement was a continuum along a depth or macrobenthos biomass gradient. Despite the well-known, close, negative relationship between water depth and macrofaunal biomass, the fish species changed more rapidly at depth shallower than 1,000 m, but the rate of change was surprisingly slow at the highest macrofaunal biomass (>100 mg C m(-2)), suggesting that the composition of epibenthic fishes was not altered in response to the extremely high macrofaunal biomass in the upper Mississippi and De Soto Submarine Canyons. An alternative is that the pattern of fish species turnover is related to the decline in macrofaunal biomass, the presumptive prey of the fish, along the depth gradient.

Anthropogenic "Litter" and macrophyte detritus in the deep Northern Gulf of Mexico.

A deep-sea trawl survey of the Northern Gulf of Mexico has documented the abundance and diversity of human-generated litter and natural detrital plant material, from the outer margin of the continental shelf out to the Sigsbee abyssal plain. Plastics were the most frequently encountered type of material. Litter and debris were encountered more frequently in the eastern than in the western GoM. Land-derived plant material was located primarily within the head of the Mississippi Canyon, whereas ocean-derived plant material was spread evenly throughout the NE GoM. Human discards were principally from ships offshore. Some of the material was contained in metal cans that sank to the sea floor, probably in order to conform to international agreements that prohibit disposal of toxic material and plastics. The Mississippi Canyon was a focal point for litter, perhaps due to topography, currents or proximity to shipping lanes.

Exploring the role of environmental variables in shaping patterns of seabed biodiversity composition in regional-scale ecosystems.

1. Environmental variables are often used as indirect surrogates for mapping biodiversity because species survey data are scant at regional scales, especially in the marine realm. However, environmental variables are measured on arbitrary scales unlikely to have simple, direct relationships with biological patterns. Instead, biodiversity may respond nonlinearly and to interactions between environmental variables.2. To investigate the role of the environment in driving patterns of biodiversity composition in large marine regions, we collated multiple biological survey and environmental data sets from tropical NE Australia, the deep Gulf of Mexico and the temperate Gulf of Maine. We then quantified the shape and magnitude of multispecies responses along >30 environmental gradients and the extent to which these variables predicted regional distributions. To do this, we applied a new statistical approach, Gradient Forest, an extension of Random Forest, capable of modelling nonlinear and threshold responses.3. The regional-scale environmental variables predicted an average of 13-35% (up to 50-85% for individual species) of the variation in species abundance distributions. Important predictors differed among regions and biota and included depth, salinity, temperature, sediment composition and current stress. The shapes of responses along gradients also differed and were nonlinear, often with thresholds indicative of step changes in composition. These differing regional responses were partly due to differing environmental indicators of bioregional boundaries and, given the results to date, may indicate limited scope for extrapolating bio-physical relationships beyond the region of source data sets.4.Synthesis and applications. Gradient Forest offers a new capability for exploring relationships between biodiversity and environmental gradients, generating new information on multispecies responses at a detail not available previously. Importantly, given the scarcity of data, Gradient Forest enables the combined use of information from disparate data sets. The gradient response curves provide biologically informed transformations of environmental layers to predict and map expected patterns of biodiversity composition that represent sampled composition better than uninformed variables. The approach can be applied to support marine spatial planning and management and has similar applicability in terrestrial realms.Gradient Forest offers a new capability for exploring relationships between biodiversity and environmental gradients, generating new information on multispecies responses at a detail not available previously. Importantly, given the scarcity of data, Gradient Forest enables the combined use of information from disparate data sets. The gradient response curves provide biologically informed transformations of environmental layers to predict and map expected patterns of biodiversity composition that represent sampled composition better than uninformed variables. The approach can be applied to support marine spatial planning and management and has similar applicability in terrestrial realms.

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.