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King penguins (Aptenodytes patagonicus) are an iconic Southern Ocean species, but the prey distributions that underpin their at-sea foraging tracks and diving behaviour remain unclear. We conducted simultaneous acoustic surveys off South Georgia and tracking of king penguins breeding ashore there in Austral summer 2017 to gain insight into habitat use and foraging behaviour. Acoustic surveys revealed ubiquitous deep scattering layers (DSLs; acoustically detected layers of fish and other micronekton that inhabit the mesopelagic zone) at c. 500 m and shallower ephemeral fish schools. Based on DNA extracted from penguin faecal samples, these schools were likely comprised of lanternfish (an important component of king penguin diets), icefish (Channichthyidae spp.) and painted noties (Lepidonotothen larseni). Penguins did not dive as deep as DSLs, but their prey-encounter depth-distributions, as revealed by biologging, overlapped at fine scale (10s of m) with depths of acoustically detected fish schools. We used neural networks to predict local scale (10 km) fish echo intensity and depth distribution at penguin dive locations based on environmental correlates, and developed models of habitat use. Habitat modelling revealed that king penguins preferentially foraged at locations predicted to have shallow and dense (high acoustic energy) fish schools associated with shallow and dense DSLs. These associations could be used to predict the distribution of king penguins from other colonies at South Georgia for which no tracking data are available, and to identify areas of potential ecological significance within the South Georgia and the South Sandwich Islands marine protected area.

King penguins (Aptenodytes patagonicus) are an iconic Southern Ocean species, but the prey distributions that underpin their at-sea foraging tracks and diving behaviour remain unclear. We conducted simultaneous acoustic surveys off South Georgia and tracking of king penguins breeding ashore there in Austral summer 2017 to gain insight into habitat use and foraging behaviour. Acoustic surveys revealed ubiquitous deep scattering layers (DSLs; acoustically detected layers of fish and other micronekton that inhabit the mesopelagic zone) at c. 500 m and shallower ephemeral fish schools. Based on DNA extracted from penguin faecal samples, these schools were likely comprised of lanternfish (an important component of king penguin diets), icefish (Channichthyidae spp.) and painted noties (Lepidonotothen larseni). Penguins did not dive as deep as DSLs, but their prey-encounter depth-distributions, as revealed by biologging, overlapped at fine scale (10s of m) with depths of acoustically detected fish schools. We used neural networks to predict local scale (10 km) fish echo intensity and depth distribution at penguin dive locations based on environmental correlates, and developed models of habitat use. Habitat modelling revealed that king penguins preferentially foraged at locations predicted to have shallow and dense (high acoustic energy) fish schools associated with shallow and dense DSLs. These associations could be used to predict the distribution of king penguins from other colonies at South Georgia for which no tracking data are available, and to identify areas of potential ecological significance within the South Georgia and the South Sandwich Islands marine protected area.

Animal telemetry is a powerful tool for observing marine animals and the physical environments that they inhabit, from coastal and continental shelf ecosystems to polar seas and open oceans. Satellite-linked biologgers and networks of acoustic receivers allow animals to be reliably monitored over scales of tens of meters to thousands of kilometers, giving insight into their habitat use, home range size, the phenology of migratory patterns and the biotic and abiotic factors that drive their distributions. Furthermore, physical environmental variables can be collected using animals as autonomous sampling platforms, increasing spatial and temporal coverage of global oceanographic observation systems. The use of animal telemetry, therefore, has the capacity to provide measures from a suite of essential ocean variables (EOVs) for improved monitoring of Earth's oceans. Here we outline the design features of animal telemetry systems, describe current applications and their benefits and challenges, and discuss future directions. We describe new analytical techniques that improve our ability to not only quantify animal movements but to also provide a powerful framework for comparative studies across taxa. We discuss the application of animal telemetry and its capacity to collect biotic and abiotic data, how the data collected can be incorporated into ocean observing systems, and the role these data can play in improved ocean management.

Maritime economy, ecosystem-based management and climate change adaptation and mitigation raise emerging needs on coastal ocean and biological observations. Integrated ocean observing aims at optimizing sampling strategies and cost-efficiency, sharing data and best practices, and maximizing the value of the observations for multiple purposes. Recently developed cost-effective, near real time technology such as gliders, radars, ferrybox, and shallow water Argo floats, should be used operationally to generate operational coastal sea observations and analysis. Furthermore, value of disparate coastal ocean observations can be unlocked with multi-dimensional integration on fitness-for-the-purpose, parameter and instrumental. Integration of operational monitoring with offline monitoring programs, such as those for research, ecosystem-based management and commercial purposes, is necessary to fill the gaps. Such integration should lead to a system of networks which can deliver data for all kinds of purposes. Detailed integration activities are identified which should enhance the coastal ocean and biological observing capacity. Ultimately a program is required which integrates physical, biogeochemical and biological observation of the ocean, from coastal to deep-sea environments, bringing together global, regional, and local observation efforts.

This publication is primarily aimed at stakeholders involved in ocean observing, spanning diverse roles from commissioning, managing, funding and coordinating, to developing, implementing, or advising on, ocean observation programmes. Such programmes will have strategic and policy drivers but their main purpose may vary from predominantly researchdriven scientific purposes to environmental monitoring for providing data and reporting to legally-binding regulations or directives. The main focus is on European capabilities but set in a global context with the various actors spanning a variety of geographical scales from national to regional and European. Key stakeholder organizations include environmental or other agencies; marine research institutions, their researchers and operators; international and regional ocean observing initiatives and programmes; national, regional and European policy makers and their advisors; national stations for observations; etc.). It will also be of interest to the wider marine and maritime research and policy community. The main aim of the publication is to increase the relevance of current (and future) European biological ocean observation capacity to strengthen global efforts towards our understanding of the ocean and enhance marine biodiversity conservation, for maintaining a healthy ocean for healthy societies. This document explains why biological ocean observations are needed to assess progress against national and international conservation targets, the Sustainable Development Goals (SDGs), the Blue Growth agenda and to contribute to key EU directives including the Marine Strategy Framework Directive (MSFD). To achieve this, the publication highlights the need of biological ocean observations to reflect clearly defined hypotheses about potential causes of change, including the combined impacts of local and global drivers, and to support the management of our impacts on the ocean. Additionally, it calls for flexible biological oce

Marine mammals help gather information on some of the harshest environments on the planet, through the use of miniaturized ocean sensors glued on their fur. Since 2004, hundreds of diving marine animals, mainly Antarctic and Arctic seals, have been fitted with a new generation of Argos tags developed by the Sea Mammal Research Unit of the University of St Andrews in Scotland, UK. These tags investigate the at-sea ecology of these animals while simultaneously collecting valuable oceanographic data. Some of the study species travel thousands of kilometres continuously diving to great depths (up to 2100 m). The resulting data are now freely available to the global scientific community at http://www.meop.net. Despite great progress in their reliability and data accuracy, the current generation of loggers while approaching standard ARGO quality specifications have yet to match them. Yet, improvements are underway; they involve updating the technology, implementing a more systematic phase of calibration and taking benefit of the recently acquired knowledge on the dynamical response of sensors. Together these efforts are rapidly transforming animal tagging into one of the most important sources of oceanographic data in polar regions and in many coastal areas

Since 2004, several hundreds of diving marine animals, mainly Antarctic and Arctic seals, were fitted with a new generation of Argos-CTD tags developed by the Sea Mammal Research Unit of the University of St. Andrews in Scotland. These tags can be used to investigate simultaneously the at-sea ecology of these animals while collecting valuable oceanographic data. Some of these species are able to travel thousands of kilometres, continuously diving to great depths (590 ± 200 m, with maxima around 2000m). Through the years, these animals have become an essential source of temperature and salinity profiles (MEOP-CTD database available at http://www.meop.net), especially for the polar oceans, complementing efficiently the Argo array. One region where the use of instrumented seals has been particularly successful is the Antarctic continental shelf. Recent contributions to the study of the Antarctic Bottom Water production area near Prydz Bay, the rapidly-thinning ice shelves in Amundsen Bay, or the stratification in the marginal ice zone, are demonstrating the rapidly growing value of these data for Polar Research.

Addressing the potential changes in species distribution in response to global change requires evaluating current species-wide patterns of habitat utilization and how these are determined by the local environment. These efforts, however, can be limited by a lack of data on distribution of the species, which are particularly difficult to collect in the case of air-breathing marine predators in high latitude systems. We present here a retrospective analysis on the distributional patterns of both Weddell (Leptonychotes weddellii) and crabeater seals (Lobodon carcinophaga) throughout their circumpolar ranges, using satellite telemetry data obtained over a time span of three decades (1990 to 2010s). Raw Argos tracks (nWeddell = 132; ncrabeater = 115), were filtered using a Switching State Space Model, and the tracking-derived behaviors (transit, foraging) were used to construct localized habitat models based on sea ice and bathymetry derived variables using binary Generalized Additive Mixed Models. We found differences in the effects and significance that environmental covariates had on the distribution at the local scale. The development of local habitat models for the entire range of these species will allow us to understand the ability of each species to cope with the anticipated environmental changes in their habitat.

McIntyre et al. (2011; Mar Ecol Prog Ser 441:257−272) concluded that climate change-related ocean warming may lead to deeper foraging dives by southern elephant seals as their prey is forced into deeper depths. They further assert that fitness for the seals will be reduced because of greater physiological costs for deep dives and the assumption that deep foraging is less successful. Their conclusions are based on an observed correlation between a temperature index and elephant seal diving depth but do not include any observations of prey. We recently observed pronounced differences in the vertical distribution of pelagic biota—biota that may well include elephant seal prey—across the same frontal zone considered by McIntyre et al. (2011) and believe that their suggested link between temperature and diving depth is actually a link between predators and distinct prey fields—a reflection of adaptive foraging behaviour in a complex and dynamic pelagic system. As such, the analysis of McIntyre et al. (2011) is uninformative about likely impacts of ocean warming.

The increasing need for continuous monitoring of the world oceans has stimulated the development of a range of autonomous sampling platforms. One novel addition to these approaches is a small, relatively inexpensive data-relaying device that can be deployed on marine mammals to provide vertical oceanographic profiles throughout the upper 2000 m of the water column. When an animal dives, the CTD-Satellite Relay Data Logger (CTD-SRDL) records vertical profiles of temperature, conductivity and pressure. Data are compressed once the animal returns to the surface where it is located by, and relays data to, the Argos satellite system. The technical challenges met in the design of the CTD-SRDL are the maximising of energy efficiency and minimising size, whilst simultaneously maintaining the reliability of an instrument that cannot be recovered and is required to survive its lifetime attached to a marine mammal. The CTD-SRDLs record temperature and salinity with an accuracy of better than 0.005 degrees C and 0.02 respectively. However, due to the limited availability of reference data, real-time data from remote places are often associated with slightly higher errors. The potential to collect large numbers of profiles cost-effectively makes data collection using CTD-SRDL technology particularly beneficial in regions where traditional oceanographic measurements are scarce or even absent. Depending on the CTD-SRDL configuration, it is possible to sample and transmit hydrographic profiles on a daily basis, providing valuable and often unique information for a real-time ocean observing system.

Polar regions are particularly sensitive to climate change, with the potential for significant feedbacks between ocean circulation, sea ice, and the ocean carbon cycle. However, the difficulty in obtaining in situ data means that our ability to detect and interpret change is very limited, especially in the Southern Ocean, where the ocean beneath the sea ice remains almost entirely unobserved and the rate of sea-ice formation is poorly known. Here, we show that southern elephant seals (Mirounga leonina) equipped with oceanographic sensors can measure ocean structure and water mass changes in regions and seasons rarely observed with traditional oceanographic platforms. In particular, seals provided a 30-fold increase in hydrographic profiles from the sea-ice zone, allowing the major fronts to be mapped south of 60 degrees S and sea-ice formation rates to be inferred from changes in upper ocean salinity. Sea-ice production rates peaked in early winter (April-May) during the rapid northward expansion of the pack ice and declined by a factor of 2 to 3 between May and August, in agreement with a three-dimensional coupled ocean-sea-ice model. By measuring the high-latitude ocean during winter, elephant seals fill a "blind spot" in our sampling coverage, enabling the establishment of a truly global ocean-observing system.

Polar regions are particularly sensitive to climate change, with the potential for significant feedbacks between ocean circulation, sea ice, and the ocean carbon cycle. However, the difficulty in obtaining in situ data means that our ability to detect and interpret change is very limited, especially in the Southern Ocean, where the ocean beneath the sea ice remains almost entirely unobserved and the rate of sea-ice formation is poorly known. Here, we show that southern elephant seals (Mirounga leonina) equipped with oceanographic sensors can measure ocean structure and water mass changes in regions and seasons rarely observed with traditional oceanographic platforms. In particular, seals provided a 30-fold increase in hydrographic profiles from the sea-ice zone, allowing the major fronts to be mapped south of 60°S and sea-ice formation rates to be inferred from changes in upper ocean salinity. Sea-ice production rates peaked in early winter (April–May) during the rapid northward expansion of the pack ice and declined by a factor of 2 to 3 between May and August, in agreement with a three-dimensional coupled ocean–sea-ice model. By measuring the high-latitude ocean during winter, elephant seals fill a “blind spot” in our sampling coverage, enabling the establishment of a truly global ocean-observing system.

Due to the low accessibility of the region, most of the seasonally ice-covered Southern Ocean remains unobserved during winter. Here we show that southern elephant seals (Mirounga leonina), equipped with oceanographic sensors, can measure winter hydrography with unprecedented temporal and spatial resolution. Seals provided a 30-fold increase in hydrographic profiles from the sea ice zone, allowing the major fronts to be mapped south of 60°S. Sea-ice formation rates in the East Antarctic sector were estimated from salinity increase in the upper water column and compared to simulations with a circumpolar finite-element sea-ice ocean model. From the observations, peak freezing rates of 2.5-3 cm/d were estimated for the period from late April to early May during the rapid northward expansion of the ice cover. While the pack ice becomes more compact, sea ice formation declines to 1 cm/d in June/July, and virtually stops by the end of August, when the maximum ice extent is reached. Modeled and observed freezing rates agree remarkably well, so that the model can be used to complete the seasonal cycle. By sampling the ocean during winter, elephant seals fill a blind spot in our sampling coverage, bringing us closer to a truly global ocean observing system.

Responses by marine top predators to environmental variability have previously been almost impossible to observe directly. By using animal-mounted instruments simultaneously recording movements, diving behavior, and in situ oceanographic properties, we studied the behavioral and physiological responses of southern elephant seals to spatial environmental variability throughout their circumpolar range. Improved body condition of seals in the Atlantic sector was associated with Circumpolar Deep Water upwelling regions within the Antarctic Circumpolar Current, whereas High-Salinity Shelf Waters or temperature/salinity gradients under winter pack ice were important in the Indian and Pacific sectors. Energetic consequences of these variations could help explain recently observed population trends, showing the usefulness of this approach in examining the sensitivity of top predators to global and regional-scale climate variability.

Responses by marine top predators to environmental variability have previously been almost impossible to observe directly. By using animal-mounted instruments simultaneously recording movements, diving behavior, and in situ oceanographic properties, we studied the behavioral and physiological responses of southern elephant seals to spatial environmental variability throughout their circumpolar range. Improved body condition of seals in the Atlantic sector was associated with Circumpolar Deep Water upwelling regions within the Antarctic Circumpolar Current, whereas High-Salinity Shelf Waters or temperature/salinity gradients under winter pack ice were important in the Indian and Pacific sectors. Energetic consequences of these variations could help explain recently observed population trends, showing the usefulness of this approach in examining the sensitivity of top predators to global and regional-scale climate variability.