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Ocean circulation connects geographically distinct ecosystems across a wide range of spatial and temporal scales via exchanges of physical and biogeochemical properties. Remote oceanographic processes can be especially important for ecosystems in the Southern Ocean, where the Antarctic Circumpolar Current (ACC) transports properties across ocean basins through both advection and mixing. Recent tracking studies have indicated the existence of two large-scale, open ocean habitats in the Southern Ocean used by grey petrels (Procellaria cinerea) from two populations (i.e. Kerguelen and Antipodes islands) during their nonbreeding season for extended periods during austral summer (i.e. October to February). In this work, we use a novel combination of large-scale oceanographic observations, surface drifter data, satellite-derived primary productivity, numerical adjoint sensitivity experiments, and output from a biogeochemical state estimate to examine local and remote influences on these grey petrel habitats. Our aim is to understand the oceanographic features that control these isolated foraging areas and to evaluate their ecological value as oligotrophic open ocean habitats. We estimate the minimum local primary productivity required to support these populations to be much less than 1% of the estimated local primary productivity. The region in the southeast Indian Ocean used by the birds from Kerguelen is connected by circulation to the productive Kerguelen shelf. In contrast, the region in the south-central Pacific Ocean used by seabirds from the Antipodes is relatively isolated suggesting it is more influenced by local factors or the cumulative effects of many seasonal cycles. This work exemplifies the potential use of predator distributions and oceanographic data to highlight areas of the open ocean that may be more dynamic and productive than previously thought. Our results highlight the need to consider advective connections between ecosystems in the Southern Ocean

In 1999, the consortium on Estimating the Circulation and Climate of the Ocean (ECCO) set out to synthesize the hydrographic data collected by the World Ocean Circulation Experiment (WOCE) and the satellite sea surface height measurements into a complete and coherent description of the ocean, afforded by an ocean general circulation model. Twenty years later, the versatility of ECCO's estimation framework enables the production of global and regional ocean and sea-ice state estimates, that incorporate not only the initial suite of data and its successors, but nearly all data streams available today. New observations include measurements from Argo floats, marine mammal-based hydrography, satellite retrievals of ocean bottom pressure and sea surface salinity, as well as ice-tethered profiled data in polar regions. The framework also produces improved estimates of uncertain inputs, including initial conditions, surface atmospheric state variables, and mixing parameters. The freely available state estimates and related efforts are property-conserving, allowing closed budget calculations that are a requisite to detect, quantify, and understand the evolution of climate-relevant signals, as mandated by the Coupled Model Intercomparison Project Phase 6 (CMIP6) protocol. The solutions can be reproduced by users through provision of the underlying modeling and assimilation machinery. Regional efforts have spun off that offer increased spatial resolution to better resolve relevant processes. Emerging foci of ECCO are on a global sea level changes, in particular contributions from polar ice sheets, and the increased use of biogeochemical and ecosystem data to constrain global cycles of carbon, nitrogen and oxygen. Challenges in the coming decade include provision of uncertainties, informing observing system design, globally increased resolution, and moving toward a coupled Earth system estimation with consistent momentum, heat and freshwater fluxes between the ocean, atmosphere

© The Author(s), 2019. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Heimbach, P., Fukumori, I., Hills, C. N., Ponte, R. M., Stammer, D., Wunsch, C., Campin, J., Cornuelle, B., Fenty, I., Forget, G., Koehl, A., Mazloff, M., Menemenlis, D., Nguyen, A. T., Piecuch, C., Trossman, D., Verdy, A., Wang, O., & Zhang, H. Putting it all together: Adding value to the global ocean and climate observing systems with complete self-consistent ocean state and parameter estimates. Frontiers in Marine Science, 6 (2019):55, doi:10.3389/fmars.2019.00055., In 1999, the consortium on Estimating the Circulation and Climate of the Ocean (ECCO) set out to synthesize the hydrographic data collected by the World Ocean Circulation Experiment (WOCE) and the satellite sea surface height measurements into a complete and coherent description of the ocean, afforded by an ocean general circulation model. Twenty years later, the versatility of ECCO's estimation framework enables the production of global and regional ocean and sea-ice state estimates, that incorporate not only the initial suite of data and its successors, but nearly all data streams available today. New observations include measurements from Argo floats, marine mammal-based hydrography, satellite retrievals of ocean bottom pressure and sea surface salinity, as well as ice-tethered profiled data in polar regions. The framework also produces improved estimates of uncertain inputs, including initial conditions, surface atmospheric state variables, and mixing parameters. The freely available state estimates and related efforts are property-conserving, allowing closed budget calculations that are a requisite to detect, quantify, and understand the evolution of climate-relevant signals, as mandated by the Coupled Model Intercomparison Project Phase 6 (CMIP6) protocol. The solutions can be reproduced by users through provision of the underlying modeling and assimilation machinery. Regional efforts have spun off that offer increased spatial resolution to better resolve relevant processes. Emerging foci of ECCO are on a global sea level changes, in particular contributions from polar ice sheets, and the increased use of biogeochemical and ecosystem data to constrain global cycles of carbon, nitrogen and oxygen. Challenges in the coming decade include provision of uncertainties, informing observing system design, globally increased resolution, and moving toward a coupled Earth system estimation with consistent momentum, heat and freshwater fluxes between the ocean, atmosphere, Major support for ECCO is provided by NASA's Physical Oceanography program via a contract to JPL/Caltech, with additional support through NASA's Modeling, Analysis and Prediction program, the Cryosphere Science program, and the Computational Modeling and Cyberinfrastructure program. Supplemental funding was obtained throughout the years via standard grants to individual team members from NSF, NOAA, and ONR.

The upper ocean dissolved inorganic carbon (DIC) concentration is regulated by advective and diffusive transport divergence, biological processes, freshwater, and air-sea CO2 fluxes. The relative importance of these mechanisms in the Southern Ocean is uncertain, as year-round observations in this area have been limited. We use a novel physical-biogeochemical state estimate of the Southern Ocean to construct a closed DIC budget of the top 650 m and investigate the spatial and temporal variability of the different components of the carbon system. The dominant mechanisms of variability in upper ocean DIC depend on location and time and space scales considered. Advective transport is the most influential mechanism and governs the local DIC budget across the 10 day-5 year timescales analyzed. Diffusive effects are nearly negligible. The large-scale transport structure is primarily set by upwelling and downwelling, though both the lateral ageostrophic and geostrophic transports are significant. In the Antarctic Circumpolar Current, the carbon budget components are also influenced by the presence of topography and biological hot spots. In the subtropics, evaporation and air-sea CO2 flux primarily balances the sink due to biological production and advective transport. Finally, in the subpolar region sea ice processes, which change the seawater volume and thus the DIC concentration, compensate the large impact of the advective transport and modulate the timing of biological activity and air-sea CO2 flux.

Marine ecosystems reflect the physical structure of their environment and of the biological processes they carry out. This leads to spatial heterogeneity and temporal variability, some of which is imposed externally, and some of which emerges from the biological processes themselves. The main focus of this thesis is on the formation of spatial patterns in the distribution of zooplankton arising from social interactions between individuals. In the Southern Ocean, krill often assemble in swarms and schools, the dynamics of which have important ecological consequences. I adopt a quantitative framework to describe the dynamics of predator and prey populations and to address the costs and benefits associated with social behavior. First, I formulate a model of resource utilization by a predator population with density-dependent reproduction. Second, I incorporate the predator- prey dynamics into a spatially-explicit model. Third, I derive a weakly nonlinear model for the spatial distribution of biomass and examine the formation of one-dimensional patterns driven by social tendencies. Fourth, I simulate the schooling behavior of zooplankton in a heterogeneous resource field. Finally, I consider two sources of temporal variability in ecosystem dynamics: transient amplification of small perturbations to stable equilibrium solutions, and climatic variability affecting the local biogeochemical environment., Prepared in collaboration with Woods Hole Oceanographic Institution, Woods Hole, MA. Sponsored in part by Grants OCE-0221369 and OCE-336839.

Thesis (Ph. D.)--Joint Program in Physical Oceanography (Massachusetts Institute of Technology, Dept. of Earth, Atmospheric, and Planetary Sciences; and the Woods Hole Oceanographic Institution), 2008., Includes bibliographical references (p. [221]-232)., Marine ecosystems reflect the physical structure of their environment and the biological processes they carry out. This leads to spatial heterogeneity and temporal variability, some of which is imposed externally and some of which emerges from the ecological mechanisms themselves. The main focus of this thesis is on the formation of spatial patterns in the distribution of zooplankton arising from social interactions between individuals. In the Southern Ocean, krill often assemble in swarms and schools, the dynamics of which have important ecological consequences. Mathematical and numerical models are employed to study the interplay of biological and physical processes that contribute to the observed patchiness. The evolution of social behavior is simulated in a theoretical framework that includes zooplankton population dynamics, swimming behavior, and some aspects of the variability inherent to fluid environments. First, I formulate a model of resource utilization by a stage-structured predator population with density-dependent reproduction. Second, I incorporate the predator-prey dynamics into a spatially-explicit model, in which aggregations develop spontaneously as a result of linear instability of the uniform distribution. In this idealized ecosystem, benefits related to the local abundance of mates are offset by the cost of having to share resources with other group members. Third, I derive a weakly nonlinear approximation for the steady-state distributions of predator and prey biomass that captures the spatial patterns driven by social tendencies. Fourth, I simulate the schooling behavior of zooplankton in a variable environment; when turbulent flows generate patchiness in the resource field, schools can forage more efficiently than individuals., (cont.) Taken together, these chapters demonstrate that aggregation/ schooling can indeed be the favored behavior when (i) reproduction (or other survival measures) increases with density in part of the range and (ii) mixing of prey into patches is rapid enough to offset the depletion. In the final two chapters, I consider sources of temporal variability in marine ecosystems. External perturbations amplified by nonlinear ecological interactions induce transient ex-cursions away from equilibrium; in predator-prey dynamics the amplitude and duration of these transients are controlled by biological processes such as growth and mortality. In the Southern Ocean, large-scale winds associated with ENSO and the Southern Annular Mode cause convective mixing, which in turn drives air-sea fluxes of carbon dioxide and oxygen. Whether driven by stochastic fluctuations or by climatic phenomena, variability of the biogeochemical/physical environment has implications for ecosystem dynamics., by Ariane Verdy., Ph.D.

Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy at the Massachusetts Institute of Technology and the Woods Hole Oceanographic Institution February 2008, Marine ecosystems reflect the physical structure of their environment and the biological processes they carry out. This leads to spatial heterogeneity and temporal variability, some of which is imposed externally and some of which emerges from the ecological mechanisms themselves. The main focus of this thesis is on the formation of spatial patterns in the distribution of zooplankton arising from social interactions between individuals. In the Southern Ocean, krill often assemble in swarms and schools, the dynamics of which have important ecological consequences. Mathematical and numerical models are employed to study the interplay of biological and physical processes that contribute to the observed patchiness. The evolution of social behavior is simulated in a theoretical framework that includes zooplankton population dynamics, swimming behavior, and some aspects of the variability inherent to fluid environments. First, I formulate a model of resource utilization by a stage-structured predator population with density-dependent reproduction. Second, I incorporate the predator-prey dynamics into a spatially-explicit model, in which aggregations develop spontaneously as a result of linear instability of the uniform distribution. In this idealized ecosystem, benefits related to the local abundance of mates are offset by the cost of having to share resources with other group members. Third, I derive a weakly nonlinear approximation for the steady-state distributions of predator and prey biomass that captures the spatial patterns driven by social tendencies. Fourth, I simulate the schooling behavior of zooplankton in a variable environment; when turbulent flows generate patchiness in the resource field, schools can forage more efficiently than individuals. Taken together, these chapters demonstrate that aggregation/ schooling can indeed be the favored behavior when (i) reproduction (or other survival measures) increases with density in part of the range and (ii) mixing o, Funding was provided by the Academic Programs Office of the MIT-WHOI Joint Program, an Ocean Ventures Fund Award, an Anonymous Ys Endowed Science Fellowship, and by NSF grants OCE-0221369 and OCE-336839.

Author Posting. © Springer, 2008. This is the author's version of the work. It is posted here by permission of Springer for personal use, not for redistribution. The definitive version was published in Bulletin of Mathematical Biology 70 (2008): 1634-1659, doi:10.1007/s11538-008-9312-7., Ecological systems with asymptotically stable equilibria may exhibit significant transient dynamics following perturbations. In some cases, these transient dynamics include the possibility of excursions away from the equilibrium before the eventual return; systems that exhibit such amplification of perturbations are called reactive. Reactivity is a common property of ecological systems, and the amplification can be large and long-lasting. The transient response of a reactive ecosystem depends on the parameters of the underlying model. To investigate this dependence, we develop sensitivity analyses for indices of transient dynamics (reactivity, the amplification envelope, and the optimal perturbation) in both continuous- and discrete-time models written in matrix form. The sensitivity calculations require expressions, some of them new, for the derivatives of equilibria, eigenvalues, singular values, and singular vectors, obtained using matrix calculus. Sensitivity analysis provides a quantitative framework for investigating the mechanisms leading to transient growth. We apply the methodology to a predator-prey model and a size-structured food web model. The results suggest predator-driven and prey-driven mechanisms for transient amplification resulting from multispecies interactions., Financial support provided by NSF grant DEB-0343820, NOAA grant NA03-NMF4720491, the Ocean Life Institute of the Woods Hole Oceanographic Institution, and the Academic Programs Office of the MIT-WHOI Joint Program in Oceanography.