Annual Review of Marine Science - Volume 10, 2018

My evolution from electrical engineering student to limnologist and then to oceanographer was a consequence of generous mentoring, which led to my use of the ¹⁵N tracer technique to measure nitrogen fixation in aquatic systems. The concept of new and regenerated production arose when I applied this method to measure nitrate and ammonium uptake in marine ecosystems. I then showed that enzyme kinetics could be applied to algal nitrogen uptake and used a silicate pump to explain silicate limitation of diatoms in coastal and equatorial upwelling systems. These concepts are now recognized as modern nutrient paradigms in biogeochemical oceanography. My interest in nutrients required field studies and led to my passion for the study of upwelling ecosystems and the establishment of two major international programs, with numerous advisors, collaborators, and students helping along the way.

Marine animals with complex life cycles may move passively or actively for fertilization, dispersal, predator avoidance, resource acquisition, and migration, and over scales from micrometers to thousands of kilometers. This diversity has catalyzed idiosyncratic and unfocused research, creating unsound paradigms regarding the role of movement in ecology and evolution. The emerging movement ecology paradigm offers a framework to consolidate movement research independent of taxon, life-history stage, scale, or discipline. This review applies the framework to movement among life-history stages in marine animals with complex life cycles to consolidate marine movement research and offer insights for scientists working in aquatic and terrestrial realms. Irrespective of data collection or simulation strategy, breaking each life-history stage down into the fundamental units of movement allows each unit to be studied independently or interactively with other units. Understanding these underlying mechanisms of movement within each life-history stage can then be used to construct lifetime movement paths. These paths can allow further investigation of the relative contributions and interdependencies of steps and phases across a lifetime and how these paths influence larger research topics, such as population-level movements.

Marine plankton elemental stoichiometric ratios can deviate from the Redfield ratio (106C:16N:1P); here, we examine physiological and biogeochemical mechanisms that lead to the observed variation across lineages, regions, and seasons. Many models of ecological stoichiometry blend together acclimative and adaptive responses to environmental conditions. These two pathways can have unique molecular mechanisms and stoichiometric outcomes, and we attempt to disentangle the two processes. We find that interactions between environmental conditions and cellular growth are key to understanding stoichiometric regulation, but the growth rates of most marine plankton populations are poorly constrained. We propose that specific physiological mechanisms have a strong impact on plankton and community stoichiometry in nutrient-rich environments, whereas biogeochemical interactions are important for the stoichiometry of the oligotrophic gyres. Finally, we outline key areas with missing information that is needed to advance understanding of the present and future ecological stoichiometry of ocean plankton.

Coccolithophores are major contributors to phytoplankton communities and ocean biogeochemistry and are strong modulators of the optical field in the sea. New discoveries are changing paradigms about these calcifiers. A new role for silicon in coccolithophore calcification is coupling carbonate and silicon cycles. Phosphorus and iron play key roles in regulating coccolithophore growth. Comparing molecular phylogenies with coccolith morphometrics is forcing the reconciliation of biological and geological observations. Mixotrophy may be a possible life strategy for deep-dwelling species, which has ramifications for biological pump and alkalinity pump paradigms. Climate, ocean temperatures, and pH appear to be affecting coccolithophores in unexpected ways. Global calcification is approximately 1–3% of primary productivity and affects CO2 budgets. New measurements of the backscattering cross section of coccolithophores have improved satellite-based algorithms and their application in case I and case II optical waters. Remote sensing has allowed the detection of basin-scale coccolithophore features in the Southern Ocean.

The well-lit upper layer of the open ocean is a dynamical environment that hosts approximately half of global primary production. In the remote parts of this environment, distant from the coast and from the seabed, there is no obvious spatially fixed reference frame for describing the dynamics of the microscopic drifting organisms responsible for this immense production of organic matter—the phytoplankton. Thus, a natural perspective for studying phytoplankton dynamics is to follow the trajectories of water parcels in which the organisms are embedded. With the advent of satellite oceanography, this Lagrangian perspective has provided valuable information on different aspects of phytoplankton dynamics, including bloom initiation and termination, spatial distribution patterns, biodiversity, export of carbon to the deep ocean, and, more recently, bottom-up mechanisms that affect the distribution and behavior of higher-trophic-level organisms. Upcoming submesoscale-resolving satellite observations and swarms of autonomous platforms open the way to the integration of vertical dynamics into the Lagrangian view of phytoplankton dynamics.

Satellite passive ocean color instruments have provided an unbroken ∼20-year record of global ocean plankton properties, but this measurement approach has inherent limitations in terms of spatial-temporal sampling and ability to resolve vertical structure within the water column. These limitations can be addressed by coupling ocean color data with measurements from a spaceborne lidar. Airborne lidars have been used for decades to study ocean subsurface properties, but recent breakthroughs have now demonstrated that plankton properties can be measured with a satellite lidar. The satellite lidar era in oceanography has arrived. Here, we present a review of the lidar technique, its applications in marine systems, a perspective on what can be accomplished in the near future with an ocean- and atmosphere-optimized satellite lidar, and a vision for a multiplatform virtual constellation of observational assets that would enable a three-dimensional reconstruction of global ocean ecosystems.

Carbonate precipitation has been a common life strategy for marine organisms for 3.7 billion years, as, therefore, has their construction of reefs. As favored by modern corals, reef-forming organisms have typically adopted a niche in warm, shallow, well-lit, tropical marine waters, where they are capable of building vast carbonate edifices. Because fossil reefs form water aquifers and hydrocarbon reservoirs, considerable effort has been dedicated to understanding their anatomy and morphology. Remote sensing has a particular role to play here. Interpretation of satellite images has done much to reveal the grand spatial and temporal tapestry of tropical reefs. Comparative sedimentology, whereby modern environments are contrasted with the rock record to improve interpretation, has been particularly transformed by observations made from orbit. Satellite mapping has also become a keystone technology to quantify the coral reef crisis—it can be deployed not only directly to quantify the distribution of coral communities, but also indirectly to establish a climatology for their physical environment. This article reviews the application of remote sensing to tropical coralgal reefs in order to communicate how this fast-growing technology might be central to addressing the coral reef crisis and to look ahead at future developments in the science.

In this review, we show how climate affects species, communities, and ecosystems, and why many responses from the species to the biome level originate from the interaction between the species’ ecological niche and changes in the environmental regime in both space and time. We describe a theory that allows us to understand and predict how marine species react to climate-induced changes in ecological conditions, how communities form and are reconfigured, and so how biodiversity is arranged and may respond to climate change. Our study shows that the responses of species to climate change are therefore intelligible—that is, they have a strong deterministic component and can be predicted.

Empirical data on food web dynamics and predator-prey interactions underpin ecosystem models, which are increasingly used to support strategic management of marine resources. These data have traditionally derived from stomach content analysis, but new and complementary forms of ecological data are increasingly available from biochemical tracer techniques. Extensive opportunities exist to improve the empirical robustness of ecosystem models through the incorporation of biochemical tracer data and derived indices, an area that is rapidly expanding because of advances in analytical developments and sophisticated statistical techniques. Here, we explore the trophic information required by ecosystem model frameworks (species, individual, and size based) and match them to the most commonly used biochemical tracers (bulk tissue and compound-specific stable isotopes, fatty acids, and trace elements). Key quantitative parameters derived from biochemical tracers include estimates of diet composition, niche width, and trophic position. Biochemical tracers also provide powerful insight into the spatial and temporal variability of food web structure and the characterization of dominant basal and microbial food web groups. A major challenge in incorporating biochemical tracer data into ecosystem models is scale and data type mismatches, which can be overcome with greater knowledge exchange and numerical approaches that transform, integrate, and visualize data.

Oxygen loss in the ocean, termed deoxygenation, is a major consequence of climate change and is exacerbated by other aspects of global change. An average global loss of 2% or more has been recorded in the open ocean over the past 50–100 years, but with greater oxygen declines in intermediate waters (100–600 m) of the North Pacific, the East Pacific, tropical waters, and the Southern Ocean. Although ocean warming contributions to oxygen declines through a reduction in oxygen solubility and stratification effects on ventilation are reasonably well understood, it has been a major challenge to identify drivers and modifying factors that explain different regional patterns, especially in the tropical oceans. Changes in respiration, circulation (including upwelling), nutrient inputs, and possibly methane release contribute to oxygen loss, often indirectly through stimulation of biological production and biological consumption. Microbes mediate many feedbacks in oxygen minimum zones that can either exacerbate or ameliorate deoxygenation via interacting nitrogen, sulfur, and carbon cycles. The paleo-record reflects drivers of and feedbacks to deoxygenation that have played out through the Phanerozoic on centennial, millennial, and hundred-million-year timescales. Natural oxygen variability has made it difficult to detect the emergence of a climate-forced signal of oxygen loss, but new modeling efforts now project emergence to occur in many areas in 15–25 years. Continued global deoxygenation is projected for the next 100 or more years under most emissions scenarios, but with regional heterogeneity. Notably, even small changes in oxygenation can have significant biological effects. New efforts to systematically observe oxygen changes throughout the open ocean are needed to help address gaps in understanding of ocean deoxygenation patterns and drivers.

Climate sensitivity represents the global mean temperature change caused by changes in the radiative balance of climate; it is studied for both present/future (actuo) and past (paleo) climate variations, with the former based on instrumental records and/or various types of model simulations. Paleo-estimates are often considered informative for assessments of actuo-climate change caused by anthropogenic greenhouse forcing, but this utility remains debated because of concerns about the impacts of uncertainties, assumptions, and incomplete knowledge about controlling mechanisms in the dynamic climate system, with its multiple interacting feedbacks and their potential dependence on the climate background state. This is exacerbated by the need to assess actuo- and paleoclimate sensitivity over different timescales, with different drivers, and with different (data and/or model) limitations. Here, we visualize these impacts with idealized representations that graphically illustrate the nature of time-dependent actuo- and paleoclimate sensitivity estimates, evaluating the strengths, weaknesses, agreements, and differences of the two approaches. We also highlight priorities for future research to improve the use of paleo-estimates in evaluations of current climate change.

The role of marine bioaerosols in cloud formation and climate is currently so uncertain that even the sign of the climate forcing is unclear. Marine aerosols form through direct emissions and through the conversion of gas-phase emissions to aerosols in the atmosphere. The composition and size of aerosols determine how effective they are in catalyzing the formation of water droplets and ice crystals in clouds by acting as cloud condensation nuclei and ice nucleating particles, respectively. Marine organic aerosols may be sourced both from recent regional phytoplankton blooms that add labile organic matter to the surface ocean and from long-term global processes, such as the upwelling of old refractory dissolved organic matter from the deep ocean. Understanding the formation of marine aerosols and their propensity to catalyze cloud formation processes are challenges that must be addressed given the major uncertainties associated with aerosols in climate models.

Over the last four decades, more than 500 sites of seafloor hydrothermal venting have been identified in a range of tectonic environments. These vents represent the seafloor manifestation of hydrothermal convection of seawater through the permeable oceanic basement that is driven by a subsurface heat source. Hydrothermal circulation has fundamental effects on the transfer of heat and mass from the lithosphere to the hydrosphere, the composition of seawater, the physical and chemical properties of the oceanic basement, and vent ecosystems at and below the seafloor. In this review, we compare and contrast the vent fluid chemistry from hydrothermal fields in a range of tectonic settings to assess the relative roles of fluid-mineral equilibria, phase separation, magmatic input, seawater entrainment, and sediment cover in producing the observed range of fluid compositions. We focus particularly on hydrothermal activity in those tectonic environments (e.g., mid-ocean ridge detachment faults, back-arc basins, and island arc volcanoes) where significant progress has been made in the last decade in documenting the variations in vent fluid composition.

Plankton are transported onshore, providing subsidies of food and new recruits to surf-zone and intertidal communities. The transport of plankton to the surf zone is influenced by wind, wave, and tidal forcing, and whether they enter the surf zone depends on alongshore variation in surf-zone hydrodynamics caused by the interaction of breaking waves with coastal morphology. Areas with gently sloping shores and wide surf zones typically have orders-of-magnitude-higher concentrations of plankton in the surf zone and dense larval settlement in intertidal communities because of the presence of bathymetric rip currents, which are absent in areas with steep shores and narrow surf zones. These striking differences in subsidies have profound consequences; areas with greater subsidies support more productive surf-zone communities and possibly more productive rocky intertidal communities. Recognition of the importance of spatial subsidies for rocky community dynamics has recently advanced ecological theory, and incorporating surf-zone hydrodynamics would be an especially fruitful line of investigation.

Estuarine turbidity maxima (ETMs) are generated by a large suite of hydrodynamic and sediment dynamic processes, leading to longitudinal convergence of cross-sectionally integrated and tidally averaged transport of cohesive and noncohesive suspended particulate matter (SPM). The relative importance of these processes for SPM trapping varies substantially among estuaries depending on topography, fluvial and tidal forcing, and SPM composition. The high-frequency dynamics of ETMs are constrained by interactions with the low-frequency dynamics of the bottom pool of easily erodible sediments. Here, we use a transport decomposition to present processes that lead to convergent SPM transport, and review trapping mechanisms that lead to ETMs at the landward limit of the salt intrusion, in the freshwater zone, at topographic transitions, and by lateral processes within the cross section. We use model simulations of example estuaries to demonstrate the complex concurrence of ETM formation mechanisms. We also discuss how changes in SPM trapping mechanisms, often caused by direct human interference, can lead to the generation of hyperturbid estuaries.

The oceanic bottom boundary layer extracts energy and momentum from the overlying flow, mediates the fate of near-bottom substances, and generates bedforms that retard the flow and affect benthic processes. The bottom boundary layer is forced by winds, waves, tides, and buoyancy and is influenced by surface waves, internal waves, and stratification by heat, salt, and suspended sediments. This review focuses on the coastal ocean. The main points are that (a) classical turbulence concepts and modern turbulence parameterizations provide accurate representations of the structure and turbulent fluxes under conditions in which the underlying assumptions hold, (b) modern sensors and analyses enable high-quality direct or near-direct measurements of the turbulent fluxes and dissipation rates, and (c) the remaining challenges include the interaction of waves and currents with the erodible seabed, the impact of layer-scale two- and three-dimensional instabilities, and the role of the bottom boundary layer in shelf-slope exchange.

Internal waves are widespread features of global oceans that play critical roles in mixing and thermohaline circulation. Similarly to surface waves, internal waves can travel long distances, ultimately breaking along continental margins. These breaking waves can transport deep ocean water and associated constituents (nutrients, larvae, and acidic low-oxygen waters) onto the shelf and locally enhance turbulence and mixing, with important effects on nearshore ecosystems. We are only beginning to understand the role internal waves play in shaping nearshore ecosystems. Here, I review the physics of internal waves in shallow waters and identify two commonalities among internal waves in the nearshore: exposure to deep offshore waters and enhanced turbulence and mixing. I relate these phenomena to important ecosystem processes ranging from extreme events to fertilization success to draw general conclusions about the influence of internal waves on ecosystems and the effects of internal waves in a changing climate.

Mixing efficiency is the ratio of the net change in potential energy to the energy expended in producing the mixing. Parameterizations of efficiency and of related mixing coefficients are needed to estimate diapycnal diffusivity from measurements of the turbulent dissipation rate. Comparing diffusivities from microstructure profiling with those inferred from the thickening rate of four simultaneous tracer releases has verified, within observational accuracy, 0.2 as the mixing coefficient over a 30-fold range of diapycnal diffusivities. Although some mixing coefficients can be estimated from pycnocline measurements, at present mixing efficiency must be obtained from channel flows, laboratory experiments, and numerical simulations. Reviewing the different approaches demonstrates that estimates and parameterizations for mixing efficiency and coefficients are not converging beyond the at-sea comparisons with tracer releases, leading to recommendations for a community approach to address this important issue.

Cold ocean temperature anomalies have been observed in the mid- to high-latitude North Atlantic on interannual to centennial timescales. Most notably, a large region of persistently low surface temperatures accompanied by a sharp reduction in ocean heat content was evident in the subpolar gyre from the winter of 2013–2014 to 2016, and the presence of this feature at a time of pervasive warming elsewhere has stimulated considerable debate. Here, we review the role of air-sea interaction and ocean processes in generating this cold anomaly and place it in a longer-term context. We also discuss the potential impacts of surface temperature anomalies for the atmosphere, including the North Atlantic Oscillation and European heat waves; contrast the behavior of the Atlantic with the extreme warm surface event that occurred in the North Pacific over a similar timescale; and consider the possibility that these events represent a response to a change in atmospheric planetary wave forcing.