Submesoscale coupling of krill and whales revealed by aggregative Lagrangian coherent structures.

In the marine environment, dynamic physical processes shape biological productivity and predator-prey interactions across multiple scales. Identifying pathways of physical-biological coupling is fundamental to understand the functioning of marine ecosystems yet it is challenging because the interactions are difficult to measure. We examined submesoscale (less than 100 km) surface current features using remote sensing techniques alongside ship-based surveys of krill and baleen whale distributions in the California Current System. We found that aggregative surface current features, represented by Lagrangian coherent structures (LCS) integrated over temporal scales between 2 and 10 days, were associated with increased (a) krill density (up to 2.6 times more dense), (b) baleen whale presence (up to 8.3 times more likely) and (c) subsurface seawater density (at depths up to 10 m). The link between physical oceanography, krill density and krill-predator distributions suggests that LCS are important features that drive the flux of energy and nutrients across trophic levels. Our results may help inform dynamic management strategies aimed at reducing large whales ship strikes and help assess the potential impacts of environmental change on this critical ecosystem.

Oceanic giants dance to atmospheric rhythms: Ephemeral wind-driven resource tracking by blue whales.

Trophic transfer of energy through marine food webs is strongly influenced by prey aggregation and its exploitation by predators. Rapid aggregation of some marine fish and crustacean forage species during wind-driven coastal upwelling has recently been discovered, motivating the hypothesis that predators of these forage species track the upwelling circulation in which prey aggregation occurs. We examine this hypothesis in the central California Current Ecosystem using integrative observations of upwelling dynamics, forage species' aggregation, and blue whale movement. Directional origins of blue whale calls repeatedly tracked upwelling plume circulation when wind-driven upwelling intensified and aggregation of forage species was heightened. Our findings illustrate a resource tracking strategy by which blue whales may maximize energy gain amid ephemeral foraging opportunities. These findings have implications for the ecology and conservation of diverse predators that are sustained by forage populations whose behaviour is responsive to episodic environmental dynamics.

Blue whales increase feeding rates at fine-scale ocean features.

Marine predators face the challenge of reliably finding prey that is patchily distributed in space and time. Predators make movement decisions at multiple spatial and temporal scales, yet we have a limited understanding of how habitat selection at multiple scales translates into foraging performance. In the ocean, there is mounting evidence that submesoscale (i.e. less than 100 km) processes drive the formation of dense prey patches that should hypothetically provide feeding hot spots and increase predator foraging success. Here, we integrated environmental remote-sensing with high-resolution animal-borne biologging data to evaluate submesoscale surface current features in relation to the habitat selection and foraging performance of blue whales in the California Current System. Our study revealed a consistent functional relationship in which blue whales disproportionately foraged within dynamic aggregative submesoscale features at both the regional and feeding site scales across seasons, regions and years. Moreover, we found that blue whale feeding rates increased in areas with stronger aggregative features, suggesting that these features indicate areas of higher prey density. The use of fine-scale, dynamic features by foraging blue whales underscores the need to take these features into account when designating critical habitat and may help inform strategies to mitigate the impacts of human activities for the species.

An accelerometer-derived ballistocardiogram method for detecting heart rate in free-ranging marine mammals.

Physio-logging methods, which use animal-borne devices to record physiological variables, are entering a new era driven by advances in sensor development. However, existing datasets collected with traditional bio-loggers, such as accelerometers, still contain untapped eco-physiological information. Here, we present a computational method for extracting heart rate from high-resolution accelerometer data using a ballistocardiogram. We validated our method with simultaneous accelerometer-electrocardiogram tag deployments in a controlled setting on a killer whale (Orcinus orca) and demonstrate the predictions correspond with previously observed cardiovascular patterns in a blue whale (Balaenoptera musculus), including the magnitude of apneic bradycardia and increase in heart rate prior to and during ascent. Our ballistocardiogram method may be applied to mine heart rates from previously collected accelerometery data and expand our understanding of comparative cardiovascular physiology.

Baleen whale prey consumption based on high-resolution foraging measurements.

Baleen whales influence their ecosystems through immense prey consumption and nutrient recycling1-3. It is difficult to accurately gauge the magnitude of their current or historic ecosystem role without measuring feeding rates and prey consumed. To date, prey consumption of the largest species has been estimated using metabolic models3-9 based on extrapolations that lack empirical validation. Here, we used tags deployed on seven baleen whale (Mysticeti) species (n = 321 tag deployments) in conjunction with acoustic measurements of prey density to calculate prey consumption at daily to annual scales from the Atlantic, Pacific, and Southern Oceans. Our results suggest that previous studies3-9 have underestimated baleen whale prey consumption by threefold or more in some ecosystems. In the Southern Ocean alone, we calculate that pre-whaling populations of mysticetes annually consumed 430 million tonnes of Antarctic krill (Euphausia superba), twice the current estimated total biomass of E. superba10, and more than twice the global catch of marine fisheries today11. Larger whale populations may have supported higher productivity in large marine regions through enhanced nutrient recycling: our findings suggest mysticetes recycled 1.2 × 104 tonnes iron yr-1 in the Southern Ocean before whaling compared to 1.2 × 103 tonnes iron yr-1 recycled by whales today. The recovery of baleen whales and their nutrient recycling services2,3,7 could augment productivity and restore ecosystem function lost during 20th century whaling12,13.

Context-dependent variability in the predicted daily energetic costs of disturbance for blue whales.

Assessing the long-term consequences of sub-lethal anthropogenic disturbance on wildlife populations requires integrating data on fine-scale individual behavior and physiology into spatially and temporally broader, population-level inference. A typical behavioral response to disturbance is the cessation of foraging, which can be translated into a common metric of energetic cost. However, this necessitates detailed empirical information on baseline movements, activity budgets, feeding rates and energy intake, as well as the probability of an individual responding to the disturbance-inducing stressor within different exposure contexts. Here, we integrated data from blue whales (Balaenoptera musculus) experimentally exposed to military active sonar signals with fine-scale measurements of baseline behavior over multiple days or weeks obtained from accelerometry loggers, telemetry tracking and prey sampling. Specifically, we developed daily simulations of movement, feeding behavior and exposure to localized sonar events of increasing duration and intensity and predicted the effects of this disturbance source on the daily energy intake of an individual. Activity budgets and movements were highly variable in space and time and among individuals, resulting in large variability in predicted energetic intake and costs. In half of our simulations, an individual's energy intake was unaffected by the simulated source. However, some individuals lost their entire daily energy intake under brief or weak exposure scenarios. Given this large variation, population-level models will have to assess the consequences of the entire distribution of energetic costs, rather than only consider single summary statistics. The shape of the exposure-response functions also strongly influenced predictions, reinforcing the need for contextually explicit experiments and improved mechanistic understanding of the processes driving behavioral and physiological responses to disturbance. This study presents a robust approach for integrating different types of empirical information to assess the effects of disturbance at spatio-temporal and ecological scales that are relevant to management and conservation.

Animal-Borne Metrics Enable Acoustic Detection of Blue Whale Migration.

Linking individual and population scales is fundamental to many concepts in ecology [1], including migration [2, 3]. This behavior is a critical [4] yet increasingly threatened [5] part of the life history of diverse organisms. Research on migratory behavior is constrained by observational scale [2], limiting ecological understanding and precise management of migratory populations in expansive, inaccessible marine ecosystems [6]. This knowledge gap is magnified for dispersed oceanic predators such as endangered blue whales (Balaenoptera musculus). As capital breeders, blue whales migrate vast distances annually between foraging and breeding grounds, and their population fitness depends on synchrony of migration with phenology of prey populations [7, 8]. Despite previous studies of individual-level blue whale vocal behavior via bio-logging [9, 10] and population-level acoustic presence via passive acoustic monitoring [11], detection of the life history transition from foraging to migration remains challenging. Here, we integrate direct high-resolution measures of individual behavior and continuous broad-scale acoustic monitoring of regional song production (Figure 1A) to identify an acoustic signature of the transition from foraging to migration in the Northeast Pacific population. We find that foraging blue whales sing primarily at night, whereas migratory whales sing primarily during the day. The ability to acoustically detect population-level transitions in behavior provides a tool to more comprehensively study the life history, fitness, and plasticity of population behavior in a dispersed, capital breeding population. Real-time detection of this behavioral signal can also inform dynamic management efforts [12] to mitigate anthropogenic threats to this endangered population [13, 14]).

A low-cost, open-source inertial movement GPS logger for eco-physiology applications.

Open-source technology has been increasingly used for developing low-cost animal-borne bio-loggers; however, a gap remains for a bio-logger that records both inertial movement and GPS positions. We address this need with the Tapered Wings Logger (TWLogger), an archival bio-logger that records high-resolution (e.g. 50 Hz) tri-axial accelerometry and magnetometry, temperature and GPS. The TWLogger can be built for 90 USD, accepts user-defined sampling parameters, and with a 500 mA h battery weighs 25 g. We provide publicly available build instructions and custom analysis scripts. Bench tests recorded 50 Hz inertial movement and 2 min GPS for 31.8±2.2 h (mean±s.d., n=6) with GPS accuracy within 10.9±13.6 m. Field deployments on a medium-sized bird of prey in the wild achieved similar results (n=13). The customizable TWLogger has wide-ranging application across systems and thus offers a practical solution for eco-physiology applications.