Moriaki Yasuhara

Biotic and Human Vulnerability to Projected Changes in Ocean Biogeochemistry over the 21st Century

Mora and colleagues show that ongoing greenhouse gas emissions are likely to have a considerable effect on several biogeochemical properties of the worlds oceans, with potentially serious consequences for biodiversity and human welfare.

Abrupt climate change and collapse of deep-sea ecosystems

We investigated the deep-sea fossil record of benthic ostracodes during periods of rapid climate and oceanographic change over the past 20,000 years in a core from intermediate depth in the northwestern Atlantic. Results show that deep-sea benthic community “collapses” occur with faunal turnover of up to 50% during major climatically driven oceanographic changes. Species diversity as measured by the Shannon–Wiener index falls from 3 to as low as 1.6 during these events. Major disruptions in the benthic communities commenced with Heinrich Event 1, the Inter-Allerød Cold Period (IACP: 13.1 ka), the Younger Dryas (YD: 12.9–11.5 ka), and several Holocene Bond events when changes in deep-water circulation occurred. The largest collapse is associated with the YD/IACP and is characterized by an abrupt two-step decrease in both the upper North Atlantic Deep Water assemblage and species diversity at 13.1 ka and at 12.2 ka. The ostracode fauna at this site did not fully recover until ≈8 ka, with the establishment of Labrador Sea Water ventilation. Ecologically opportunistic slope species prospered during this community collapse. Other abrupt community collapses during the past 20 ka generally correspond to millennial climate events. These results indicate that deep-sea ecosystems are not immune to the effects of rapid climate changes occurring over centuries or less.

Temporal latitudinal-gradient dynamics and tropical instability of deep-sea species diversity

A benthic microfaunal record from the equatorial Atlantic Ocean over the past four glacial-interglacial cycles was investigated to understand temporal dynamics of deep-sea latitudinal species diversity gradients (LSDGs). The results demonstrate unexpected instability and high amplitude fluctuations of species diversity in the tropical deep ocean that are correlated with orbital-scale oscillations in global climate: Species diversity is low during glacial and high during interglacial periods. This implies that climate severely influences deep-sea diversity, even at tropical latitudes, and that deep-sea LSDGs, while generally present for the last 36 million years, were weakened or absent during glacial periods. Temporally dynamic LSDGs and unstable tropical diversity require reconsideration of current ecological hypotheses about the generation and maintenance of biodiversity as they apply to the deep sea, and underscore the potential vulnerability and conservation importance of tropical deep-sea ecosystems.

Declining oxygen in the global ocean and coastal waters

Beneath the waves, oxygen disappears As plastic waste pollutes the oceans and fish stocks decline, unseen below the surface another problem grows: deoxygenation. Breitburg et al. review the evidence for the downward trajectory of oxygen levels in increasing areas of the open ocean and coastal waters. Rising nutrient loads coupled with climate change—each resulting from human activities—are changing ocean biogeochemistry and increasing oxygen consumption. This results in destabilization of sediments and fundamental shifts in the availability of key nutrients. In the short term, some compensatory effects may result in improvements in local fisheries, such as in cases where stocks are squeezed between the surface and elevated oxygen minimum zones. In the longer term, these conditions are unsustainable and may result in ecosystem collapses, which ultimately will cause societal and economic harm. Science, this issue p. eaam7240 BACKGROUND Oxygen concentrations in both the open ocean and coastal waters have been declining since at least the middle of the 20th century. This oxygen loss, or deoxygenation, is one of the most important changes occurring in an ocean increasingly modified by human activities that have raised temperatures, CO2 levels, and nutrient inputs and have altered the abundances and distributions of marine species. Oxygen is fundamental to biological and biogeochemical processes in the ocean. Its decline can cause major changes in ocean productivity, biodiversity, and biogeochemical cycles. Analyses of direct measurements at sites around the world indicate that oxygen-minimum zones in the open ocean have expanded by several million square kilometers and that hundreds of coastal sites now have oxygen concentrations low enough to limit the distribution and abundance of animal populations and alter the cycling of important nutrients. ADVANCES In the open ocean, global warming, which is primarily caused by increased greenhouse gas emissions, is considered the primary cause of ongoing deoxygenation. Numerical models project further oxygen declines during the 21st century, even with ambitious emission reductions. Rising global temperatures decrease oxygen solubility in water, increase the rate of oxygen consumption via respiration, and are predicted to reduce the introduction of oxygen from the atmosphere and surface waters into the ocean interior by increasing stratification and weakening ocean overturning circulation. In estuaries and other coastal systems strongly influenced by their watershed, oxygen declines have been caused by increased loadings of nutrients (nitrogen and phosphorus) and organic matter, primarily from agriculture; sewage; and the combustion of fossil fuels. In many regions, further increases in nitrogen discharges to coastal waters are projected as human populations and agricultural production rise. Climate change exacerbates oxygen decline in coastal systems through similar mechanisms as those in the open ocean, as well as by increasing nutrient delivery from watersheds that will experience increased precipitation. Expansion of low-oxygen zones can increase production of N2O, a potent greenhouse gas; reduce eukaryote biodiversity; alter the structure of food webs; and negatively affect food security and livelihoods. Both acidification and increasing temperature are mechanistically linked with the process of deoxygenation and combine with low-oxygen conditions to affect biogeochemical, physiological, and ecological processes. However, an important paradox to consider in predicting large-scale effects of future deoxygenation is that high levels of productivity in nutrient-enriched coastal systems and upwelling areas associated with oxygen-minimum zones also support some of the world’s most prolific fisheries. OUTLOOK Major advances have been made toward understanding patterns, drivers, and consequences of ocean deoxygenation, but there is a need to improve predictions at large spatial and temporal scales important to ecosystem services provided by the ocean. Improved numerical models of oceanographic processes that control oxygen depletion and the large-scale influence of altered biogeochemical cycles are needed to better predict the magnitude and spatial patterns of deoxygenation in the open ocean, as well as feedbacks to climate. Developing and verifying the next generation of these models will require increased in situ observations and improved mechanistic understanding on a variety of scales. Models useful for managing nutrient loads can simulate oxygen loss in coastal waters with some skill, but their ability to project future oxygen loss is often hampered by insufficient data and climate model projections on drivers at appropriate temporal and spatial scales. Predicting deoxygenation-induced changes in ecosystem services and human welfare requires scaling effects that are measured on individual organisms to populations, food webs, and fisheries stocks; considering combined effects of deoxygenation and other ocean stressors; and placing an increased research emphasis on developing nations. Reducing the impacts of other stressors may provide some protection to species negatively affected by low-oxygen conditions. Ultimately, though, limiting deoxygenation and its negative effects will necessitate a substantial global decrease in greenhouse gas emissions, as well as reductions in nutrient discharges to coastal waters. Low and declining oxygen levels in the open ocean and coastal waters affect processes ranging from biogeochemistry to food security. The global map indicates coastal sites where anthropogenic nutrients have exacerbated or caused O2 declines to <2 mg liter−1 (<63 μmol liter−1) (red dots), as well as ocean oxygen-minimum zones at 300 m of depth (blue shaded regions). [Map created from data provided by R. Diaz, updated by members of the GO2NE network, and downloaded from the World Ocean Atlas 2009]. Oxygen is fundamental to life. Not only is it essential for the survival of individual animals, but it regulates global cycles of major nutrients and carbon. The oxygen content of the open ocean and coastal waters has been declining for at least the past half-century, largely because of human activities that have increased global temperatures and nutrients discharged to coastal waters. These changes have accelerated consumption of oxygen by microbial respiration, reduced solubility of oxygen in water, and reduced the rate of oxygen resupply from the atmosphere to the ocean interior, with a wide range of biological and ecological consequences. Further research is needed to understand and predict long-term, global- and regional-scale oxygen changes and their effects on marine and estuarine fisheries and ecosystems.

CLIMATIC INFLUENCES ON DEEP-SEA OSTRACODE (CRUSTACEA) DIVERSITY FOR THE LAST THREE MILLION YEARS

Ostracodes are small, bivalved crustaceans with the finest-scale fossil resolution of any metazoan, rivaled only by the fossil record of the protistan Foraminifera. This article presents a synthesis of the patterns and possible causes of alpha species diversity variation in benthic deep-sea ostracodes at drilling sites in the North Atlantic and Arctic Oceans. Taken together, these sites represent a period of great climatic variability covering the past three million years. Sediment cores taken from the Mid-Atlantic Ridge show a positive correlation between warm temperatures and high species diversity. These Mid-Atlantic Ridge cores, at the same latitude as northern Spain, show the same positive correlation during the last two glacial-interglacial cycles (200-0 ka [thousands of years ago]) as they do during the pre-glacial Pliocene 2.85-2.4 Ma (millions of years ago). This positive correlation is also found in Pliocene cores from the Rockall Plateau, at the same latitude as Ireland. During the last 200 thousand years, however, this correlation is reversed in cores taken from both the Rockall and Iceland Plateaus. The discovery of high diversity during colder periods in recent high-latitude Rockall and Iceland cores seems to be explained by spikes in diversity caused by ice-rafting events, which would not affect the lower-latitude Mid-Atlantic Ridge. The Heinrich ice-rafting events reduce North Atlantic surface temperatures and salinity every approximately 6-12 ka, dramatically decreasing surface productivity. This increase in diversity during Heinrich events may be explained either by a negative correlation between surface productivity and benthic diversity or by increase in diversity caused by moderate disturbance when ice rafted debris fall to the bottom of the ocean.

Human-induced marine ecological degradation: micropaleontological perspectives

We analyzed published downcore microfossil records from 150 studies and reinterpreted them from an ecological degradation perspective to address the following critical but still imperfectly answered questions: (1) How is the timing of human-induced degradation of marine ecosystems different among regions? (2) What are the dominant causes of human-induced marine ecological degradation? (3) How can we better document natural variability and thereby avoid the problem of shifting baselines of comparison as degradation progresses over time? The results indicated that: (1) ecological degradation in marine systems began significantly earlier in Europe and North America (∼1800s) compared with Asia (post-1900) due to earlier industrialization in European and North American countries, (2) ecological degradation accelerated globally in the late 20th century due to post-World War II economic growth, (3) recovery from the degraded state in late 20th century following various restoration efforts and environmental regulations occurred only in limited localities. Although complex in detail, typical signs of ecological degradation were diversity decline, dramatic changes in total abundance, decrease in benthic and/or sensitive species, and increase in planktic, resistant, toxic, and/or introduced species. The predominant cause of degradation detected in these microfossil records was nutrient enrichment and the resulting symptoms of eutrophication, including hypoxia. Other causes also played considerable roles in some areas, including severe metal pollution around mining sites, water acidification by acidic wastewater, and salinity changes from construction of causeways, dikes, and channels, deforestation, and land clearance. Microfossils enable reconstruction of the ecological history of the past 102–103 years or even more, and, in conjunction with statistical modeling approaches using independent proxy records of climate and human-induced environmental changes, future research will enable workers to better address Shifting Baseline Syndrome and separate anthropogenic impacts from background natural variability.

Temperature impacts on deep-sea biodiversity

Temperature is considered to be a fundamental factor controlling biodiversity in marine ecosystems, but precisely what role temperature plays in modulating diversity is still not clear. The deep ocean, lacking light and in situ photosynthetic primary production, is an ideal model system to test the effects of temperature changes on biodiversity. Here we synthesize current knowledge on temperature–diversity relationships in the deep sea. Our results from both present and past deep‐sea assemblages suggest that, when a wide range of deep‐sea bottom‐water temperatures is considered, a unimodal relationship exists between temperature and diversity (that may be right skewed). It is possible that temperature is important only when at relatively high and low levels but does not play a major role in the intermediate temperature range. Possible mechanisms explaining the temperature–biodiversity relationship include the physiological‐tolerance hypothesis, the metabolic hypothesis, island biogeography theory, or some combination of these. The possible unimodal relationship discussed here may allow us to identify tipping points at which on‐going global change and deep‐water warming may increase or decrease deep‐sea biodiversity. Predicted changes in deep‐sea temperatures due to human‐induced climate change may have more adverse consequences than expected considering the sensitivity of deep‐sea ecosystems to temperature changes.

Combining marine macroecology and palaeoecology in understanding biodiversity: microfossils as a model

There is growing interest in the integration of macroecology and palaeoecology towards a better understanding of past, present, and anticipated future biodiversity dynamics. However, the empirical basis for this integration has thus far been limited. Here we review prospects for a macroecology–palaeoecology integration in biodiversity analyses with a focus on marine microfossils [i.e. small (or small parts of) organisms with high fossilization potential, such as foraminifera, ostracodes, diatoms, radiolaria, coccolithophores, dinoflagellates, and ichthyoliths]. Marine microfossils represent a useful model system for such integrative research because of their high abundance, large spatiotemporal coverage, and good taxonomic and temporal resolution. The microfossil record allows for quantitative cross‐scale research designs, which help in answering fundamental questions about marine biodiversity, including the causes behind similarities in patterns of latitudinal and longitudinal variation across taxa, the degree of constancy of observed gradients over time, and the relative importance of hypothesized drivers that may explain past or present biodiversity patterns. The inclusion of a deep‐time perspective based on high‐resolution microfossil records may be an important step for the further maturation of macroecology. An improved integration of macroecology and palaeoecology would aid in our understanding of the balance of ecological and evolutionary mechanisms that have shaped the biosphere we inhabit today and affect how it may change in the future.

An Arctic and Subarctic ostracode database: biogeographic and paleoceanographic applications

A new Arctic Ostracode Database-2015 (AOD-2015) provides census data for 96 species of benthic marine Ostracoda from 1340 modern surface sediments from the Arctic Ocean and subarctic seas. Ostracoda is a meiofaunal, Crustacea group that secretes a bivalved calcareous (CaCO3) shell commonly preserved in sediments. Arctic and subarctic ostracode species have ecological limits controlled by temperature, salinity, oxygen, sea ice, food, and other habitat-related factors. Unique species ecology, shell chemistry (Mg/Ca ratios, stable isotopes), and limited stratigraphic ranges make them a useful tool for paleoceanographic reconstructions and biostratigraphy. The database, described here, will facilitate the investigation of modern ostracode biogeography, regional community structure, and ecology. These data, when compared to downcore faunal data from sediment cores, will provide a better understanding of how the Arctic has been affected by climatic and oceanographic change during the Quaternary. Images of all species and biogeographic distribution maps for selected species are presented, with brief discussion of representative species’ biogeographic and ecological significance. Publication of AOD-2015 is open-sourced and will be available online at several public websites with latitude, longitude, water depth, and bottom water temperature for most samples. It includes material from Arctic abyssal plains and submarine ridges, continental slopes, and shelves of the Kara, Laptev, East Siberian, Chukchi, Beaufort Seas, and several subarctic regions.

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.