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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.

The effect of active sonars on marine mammal behaviour is a topic of considerable interest and scientific investigation. Some whales, including the largest species (blue whales, Balaenoptera musculus), can be impacted by mid-frequency (1-10 kHz) military sonars. Here we apply complementary experimental methods to provide the first experimentally controlled measurements of behavioural responses to military sonar and similar stimuli for a related endangered species, fin whales (Balaenoptera physalus). Analytical methods include: (i) principal component analysis paired with generalized additive mixed models; (ii) hidden Markov models; and (iii) structured expert elicitation using response severity metrics. These approaches provide complementary perspectives on the nature of potential changes within and across individuals. Behavioural changes were detected in five of 15 whales during controlled exposure experiments using mid-frequency active sonar or pseudorandom noise of similar frequency, duration and source and received level. No changes were detected during six control (no noise) sequences. Overall responses were more limited in occurrence, severity and duration than in blue whales and were less dependent upon contextual aspects of exposure and more contingent upon exposure received level. Quantifying the factors influencing marine mammal responses to sonar is critical in assessing and mitigating future impacts.

The giant rorqual whales are believed to have a massive food turnover driven by a high-intake lunge feeding style aptly described as the worlds largest biomechanical action. This high-drag feeding behavior is thought to limit dive times and constrain rorquals to target only the densest prey patches, making them vulnerable to disturbance and habitat change. Using biologging tags to estimate energy expenditure as a function of feeding rates on 23 humpback whales, we show that lunge feeding is energetically cheap. Such inexpensive foraging means that rorquals are flexible in the quality of prey patches they exploit and therefore more resilient to environmental fluctuations and disturbance. As a consequence, the food turnover and hence the ecological role of these marine giants have likely been overestimated.

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.

Empirical metabolic rate and oxygen consumption estimates for free-ranging whales have been limited to counting respiratory events at the surface. Because these observations were limited and generally viewed from afar, variability in respiratory properties was unknown and oxygen consumption estimates assumed constant breath-to-breath tidal volume and oxygen uptake. However, evidence suggests that cetaceans in human care vary tidal volume and breathing frequency to meet aerobic demand, which would significantly impact energetic estimates if the findings held in free-ranging species. In this study, we used suction cup-attached video tags positioned posterior to the nares of two humpback whales (Megaptera novaeangliae) and four Antarctic minke whales (Balaenoptera bonaerensis) to measure inhalation duration, relative nares expansion, and maximum nares expansion. Inhalation duration and nares expansion varied between and within initial, middle, and terminal breaths of surface sequences between dives. The initial and middle breaths exhibited the least variability and had the shortest durations and smallest nares expansions. In contrast, terminal breaths were highly variable, with the longest inhalation durations and the largest nares expansions. Our results demonstrate breath-to-breath variability in duration and nares expansion, suggesting differential oxygen exchange in each breath during the surface interval. With future validation, inhalation duration or nares area could be used alongside respiratory frequency to improve oxygen consumption estimates by accounting for breath-to-breath variation in wild whales.

Although gigantic body size and obligate filter feeding mechanisms have evolved in multiple vertebrate lineages (mammals and fishes), intermittent ram (lunge) filter feeding is unique to a specific family of baleen whales: rorquals. Lunge feeding is a high cost, high benefit feeding mechanism that requires the integration of unsteady locomotion (i.e., accelerations and maneuvers); the impact of scale on the biomechanics and energetics of this foraging mode continues to be the subject of intense study. The goal of our investigation was to use a combination of multi-sensor tags paired with UAS footage to determine the impact of morphometrics such as body size on kinematic lunging parameters such as fluking timing, maximum lunging speed, and deceleration during the engulfment period for a range of species from minke to blue whales. Our results show that, in the case of krill-feeding lunges and regardless of size, animals exhibit a skewed gradient between powered and fully unpowered engulfment, with fluking generally ending at the point of both the maximum lunging speed and mouth opening. In all cases, the small amounts of propulsive thrust generated by the tail were unable to overcome the high drag forces experienced during engulfment. Assuming this thrust to be minimal, we predicted the minimum speed of lunging across scale. To minimize the energetic cost of lunge feeding, hydrodynamic theory predicts slower lunge feeding speeds regardless of body size, with a lower boundary set by the ability of the prey to avoid capture. We used empirical data to test this theory and instead found that maximum foraging speeds remain constant and high (∼4 m s-1) across body size, even as higher speeds result in lower foraging efficiency. Regardless, we found an increasing relationship between body size and this foraging efficiency, estimated as the ratio of energetic gain from prey to energetic cost. This trend held across timescales ranging from a single lunge to a single day and suggests that larger whales are capturing more prey-and more energy-at a lower cost.

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 individuals 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.

The considerable power needed for large whales to leap out of the water may represent the single most expensive burst maneuver found in nature. However, the mechanics and energetic costs associated with the breaching behaviors of large whales remain poorly understood. In this study we deployed whale-borne tags to measure the kinematics of breaching to test the hypothesis that these spectacular aerial displays are metabolically expensive. We found that breaching whales use variable underwater trajectories, and that high-emergence breaches are faster and require more energy than predatory lunges. The most expensive breaches approach the upper limits of vertebrate muscle performance, and the energetic cost of breaching is high enough that repeated breaching events may serve as honest signaling of body condition. Furthermore, the confluence of muscle contractile properties, hydrodynamics, and the high speeds required likely impose an upper limit to the body size and effectiveness of breaching whales.

Nursing influences growth rate and overall health of mammals; however, the behavior is difficult to study in wild cetaceans because it occurs below the surface and can thus be misidentified from surface observations. Nursing has been observed in humpback whales on the breeding and calving grounds, but the behavior remains unstudied on the feeding grounds. We instrumented three dependent calves (four total deployments) with combined video and 3D-accelerometer data loggers (CATS) on two United States feeding grounds to document nursing events. Two associated mothers were also tagged to determine if behavior diagnostic of nursing was evident in the mothers movement. Animal-borne video was manually analyzed and the average duration of successful nursing events was 23 s (±7 sd, n = 11). Nursing occurred at depths between 4.1-64.4 m (along the seafloor) and in close temporal proximity to foraging events by the mothers, but could not be predicted solely by relative positions of mother and calf. When combining all calf deployments, successful nursing was documented eleven times; totaling only 0.3% of 21.0 hours of video. During nursing events, calves had higher overall dynamic body acceleration (ODBA) and increased fluke-stroke rate (FSR) compared to non-nursing segments (Mixed effect models, ODBA: F1,107 = 13.57756, p = 0.0004, FSR: F1,107 = 32.31018, p

In terrestrial systems, the green wave hypothesis posits that migrating animals can enhance foraging opportunities by tracking phenological variation in high-quality forage across space (i.e., resource waves). To track resource waves, animals may rely on proximate cues and/or memory of long-term average phenologies. Although there is growing evidence of resource tracking in terrestrial migrants, such drivers remain unevaluated in migratory marine megafauna. Here we present a test of the green wave hypothesis in a marine system. We compare 10 years of blue whale movement data with the timing of the spring phytoplankton bloom resulting in increased prey availability in the California Current Ecosystem, allowing us to investigate resource tracking both contemporaneously (response to proximate cues) and based on climatological conditions (memory) during migrations. Blue whales closely tracked the long-term average phenology of the spring bloom, but did not track contemporaneous green-up. In addition, blue whale foraging locations were characterized by low long-term habitat variability and high long-term productivity compared with contemporaneous measurements. Results indicate that memory of long-term average conditions may have a previously underappreciated role in driving migratory movements of long-lived species in marine systems, and suggest that these animals may struggle to respond to rapid deviations from historical mean environmental conditions. Results further highlight that an ecological theory of migration is conserved across marine and terrestrial systems. Understanding the drivers of animal migration is critical for assessing how environmental changes will affect highly mobile fauna at a global scale.

Resource partitioning is an important process driving habitat use and foraging strategies in sympatric species that potentially compete. Differences in foraging behavior are hypothesized to contribute to species coexistence by facilitating resource partitioning, but little is known on the multiple mechanisms for partitioning that may occur simultaneously. Studies are further limited in the marine environment, where the spatial and temporal distribution of resources is highly dynamic and subsequently difficult to quantify. We investigated potential pathways by which foraging behavior may facilitate resource partitioning in two of the largest co-occurring and closely related species on Earth, blue (Balaenoptera musculus) and humpback (Megaptera novaeangliae) whales. We integrated multiple long-term datasets (line-transect surveys, whale-watching records, net sampling, stable isotope analysis, and remote-sensing of oceanographic parameters) to compare the diet, phenology, and distribution of the two species during their foraging periods in the highly productive waters of Monterey Bay, California, USA within the California Current Ecosystem. Our long-term study reveals that blue and humpback whales likely facilitate sympatry by partitioning their foraging along three axes: trophic, temporal, and spatial. Blue whales were specialists foraging on krill, predictably targeting a seasonal peak in krill abundance, were present in the bay for an average of 4.7 months, and were spatially restricted at the continental shelf break. In contrast, humpback whales were generalists apparently feeding on a mixed diet of krill and fishes depending on relative abundances, were present in the bay for a more extended period (average of 6.6 months), and had a broader spatial distribution at the shelf break and inshore. Ultimately, competition for common resources can lead to behavioral, morphological, and physiological character displacement between sympatric species. Understanding the mechanisms for species coexistence is both fundamental to maintaining biodiverse ecosystems, and provides insight into the evolutionary drivers of morphological differences in closely related species.

Air-breathing marine animals face a complex set of physical challenges associated with diving that affect the decisions of how to optimize feeding. Baleen whales (Mysticeti) have evolved bulk-filter feeding mechanisms to efficiently feed on dense prey patches. Baleen whales are central place foragers where oxygen at the surface represents the central place and depth acts as the distance to prey. Although hypothesized that baleen whales will target the densest prey patches anywhere in the water column, how depth and density interact to influence foraging behaviour is poorly understood. We used multi-sensor archival tags and active acoustics to quantify Antarctic humpback whale foraging behaviour relative to prey. Our analyses reveal multi-stage foraging decisions driven by both krill depth and density. During daylight hours when whales did not feed, krill were found in deep high-density patches. As krill migrated vertically into larger and less dense patches near the surface, whales began to forage. During foraging bouts, we found that feeding rates (number of feeding lunges per hour) were greatest when prey was shallowest, and feeding rates decreased with increasing dive depth. This strategy is consistent with previous models of how air-breathing diving animals optimize foraging efficiency. Thus, humpback whales forage mainly when prey is more broadly distributed and shallower, presumably to minimize diving and searching costs and to increase feeding rates overall and thus foraging efficiency. Using direct measurements of feeding behaviour from animal-borne tags and prey availability from echosounders, our study demonstrates a multi-stage foraging process in a central place forager that we suggest acts to optimize overall efficiency by maximizing net energy gain over time. These data reveal a previously unrecognized level of complexity in predator-prey interactions and underscores the need to simultaneously measure prey distribution in marine central place forager studies.

Terrestrial predators can modulate the energy used for prey capture to maximize efficiency, but diving animals face the conflicting metabolic demands of energy intake and the minimization of oxygen depletion during a breath hold. It is thought that diving predators optimize their foraging success when oxygen use and energy gain act as competing currencies, but this hypothesis has not been rigorously tested because it has been difficult to measure the quality of prey that is targeted by free-ranging animals. We used high-resolution multisensor digital tags attached to foraging blue whales (Balaenoptera musculus) with concurrent acoustic prey measurements to quantify foraging performance across depth and prey density gradients. We parameterized two competing physiological models to estimate energy gain and expenditure based on foraging decisions. Our analyses show that at low prey densities, blue whale feeding rates and energy intake were low to minimize oxygen use, but at higher prey densities feeding frequency increased to maximize energy intake. Contrary to previous paradigms, we demonstrate that blue whales are not indiscriminate grazers but instead switch foraging strategies in response to variation in prey density and depth to maximize energetic efficiency.

Mid-frequency military (1-10 kHz) sonars have been associated with lethal mass strandings of deep-diving toothed whales, but the effects on endangered baleen whale species are virtually unknown. Here, we used controlled exposure experiments with simulated military sonar and other mid-frequency sounds to measure behavioural responses of tagged blue whales (Balaenoptera musculus) in feeding areas within the Southern California Bight. Despite using source levels orders of magnitude below some operational military systems, our results demonstrate that mid-frequency sound can significantly affect blue whale behaviour, especially during deep feeding modes. When a response occurred, behavioural changes varied widely from cessation of deep feeding to increased swimming speed and directed travel away from the sound source. The variability of these behavioural responses was largely influenced by a complex interaction of behavioural state, the type of mid-frequency sound and received sound level. Sonar-induced disruption of feeding and displacement from high-quality prey patches could have significant and previously undocumented impacts on baleen whale foraging ecology, individual fitness and population health.