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Honors & Awards

  • Anne T. and Robert M. Bass Fellowship, Stanford University (2014)
  • Outstanding Achievement Award for High Impact Paper, Hopkins Marine Station (2014-2015)

Professional Education

  • Doctor of Philosophy, Stanford University, BIO-PHD (2019)
  • Master of Arts, Stanford University, ED-MA (2005)
  • Bachelor of Arts, Brown University, Mathematics (2002)
  • Master of Science, Oregon State University, Ocean, Earth and Atmospheric Sciences (2014)

Stanford Advisors

All Publications

  • Minke whale feeding rate limitations suggest constraints on the minimum body size for engulfment filtration feeding. Nature ecology & evolution Cade, D. E., Kahane-Rapport, S. R., Gough, W. T., Bierlich, K. C., Linsky, J. M., Calambokidis, J., Johnston, D. W., Goldbogen, J. A., Friedlaender, A. S. 2023


    Bulk filter feeding has enabled gigantism throughout evolutionary history. The largest animals, extant rorqual whales, utilize intermittent engulfment filtration feeding (lunge feeding), which increases in efficiency with body size, enabling their gigantism. The smallest extant rorquals (7-10 m minke whales), however, still exhibit short-term foraging efficiencies several times greater than smaller non-filter-feeding cetaceans, raising the question of why smaller animals do not utilize this foraging modality. We collected 437 h of bio-logging data from 23 Antarctic minke whales (Balaenoptera bonaerensis) to test the relationship of feeding rates (λf) to body size. Here, we show that while ultra-high nighttime λf (mean ± s.d.: 165 ± 40 lunges h-1; max: 236 lunges h-1; mean depth: 28 ± 46 m) were indistinguishable from predictions from observations of larger species, daytime λf (mean depth: 72 ± 72 m) were only 25-40% of predicted rates. Both λf were near the maxima allowed by calculated biomechanical, physiological and environmental constraints, but these temporal constraints meant that maximum λf was below the expected λf for animals smaller than ~5 m-the length of weaned minke whales. Our findings suggest that minimum size for specific filter-feeding body plans may relate broadly to temporal restrictions on filtration rate and have implications for the evolution of filter feeding.

    View details for DOI 10.1038/s41559-023-01993-2

    View details for PubMedID 36914772

    View details for PubMedCentralID 3731297

  • Oceanic giants dance to atmospheric rhythms: Ephemeral wind-driven resource tracking by blue whales. Ecology letters Ryan, J. P., Benoit-Bird, K. J., Oestreich, W. K., Leary, P., Smith, K. B., Waluk, C. M., Cade, D. E., Fahlbusch, J. A., Southall, B. L., Joseph, J. E., Margolina, T., Calambokidis, J., DeVogelaere, A., Goldbogen, J. A. 2022


    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.

    View details for DOI 10.1111/ele.14116

    View details for PubMedID 36197736

  • Fast and Furious: Energetic Tradeoffs and Scaling of High-Speed Foraging in Rorqual Whales INTEGRATIVE ORGANISMAL BIOLOGY Gough, W. T., Cade, D. E., Czapanskiy, M. F., Potvin, J., Fish, F. E., Kahane-Rapport, S. R., Savoca, M. S., Bierlich, K. C., Johnston, D. W., Friedlaender, A. S., Szabo, A., Bejder, L., Goldbogen, J. A. 2022; 4 (1): obac038


    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.

    View details for DOI 10.1093/iob/obac038

    View details for Web of Science ID 000855481000001

    View details for PubMedID 36127894

    View details for PubMedCentralID PMC9475666

  • Blue whales increase feeding rates at fine-scale ocean features. Proceedings. Biological sciences Fahlbusch, J. A., Czapanskiy, M. F., Calambokidis, J., Cade, D. E., Abrahms, B., Hazen, E. L., Goldbogen, J. A. 2022; 289 (1981): 20221180


    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.

    View details for DOI 10.1098/rspb.2022.1180

    View details for PubMedID 35975432

  • Intra-seasonal variation in feeding rates and diel foraging behaviour in a seasonally fasting mammal, the humpback whale. Royal Society open science Nichols, R. C., Cade, D. E., Kahane-Rapport, S., Goldbogen, J., Stimpert, A., Nowacek, D., Read, A. J., Johnston, D. W., Friedlaender, A. 2022; 9 (7): 211674


    Antarctic humpback whales forage in summer, coincident with the seasonal abundance of their primary prey, the Antarctic krill. During the feeding season, humpback whales accumulate energy stores sufficient to fuel their fasting period lasting over six months. Previous animal movement modelling work (using area-restricted search as a proxy) suggests a hyperphagic period late in the feeding season, similar in timing to some terrestrial fasting mammals. However, no direct measures of seasonal foraging behaviour existed to corroborate this hypothesis. We attached high-resolution, motion-sensing biologging tags to 69 humpback whales along the Western Antarctic Peninsula throughout the feeding season from January to June to determine how foraging effort changes throughout the season. Our results did not support existing hypotheses: we found a significant reduction in foraging presence and feeding rates from the beginning to the end of the feeding season. During the early summer period, feeding occurred during all hours at high rates. As the season progressed, foraging occurred mostly at night and at lower rates. We provide novel information on seasonal changes in foraging of humpback whales and suggest that these animals, contrary to nearly all other animals that seasonally fast, exhibit high feeding rates soon after exiting the fasting period.

    View details for DOI 10.1098/rsos.211674

    View details for PubMedID 35814912

  • Acoustic signalling and behaviour of Antarctic minke whales (Balaenoptera bonaerensis). Royal Society open science Casey, C. B., Weindorf, S., Levy, E., Linsky, J. M., Cade, D. E., Goldbogen, J. A., Nowacek, D. P., Friedlaender, A. S. 2022; 9 (7): 211557


    Acoustic signalling is the predominant form of communication among cetaceans. Understanding the behavioural state of calling individuals can provide insights into the specific function of sound production; in turn, this information can aid the evaluation of passive monitoring datasets to estimate species presence, density, and behaviour. Antarctic minke whales are the most numerous baleen whale species in the Southern Ocean. However, our knowledge of their vocal behaviour is limited. Using, to our knowledge, the first animal-borne audio-video documentation of underwater behaviour in this species, we characterize Antarctic minke whale sound production and evaluate the association between acoustic behaviour, foraging behaviour, diel patterns and the presence of close conspecifics. In addition to the previously described downsweep call, we find evidence of three novel calls not previously described in their vocal repertoire. Overall, these signals displayed peak frequencies between 90 and 175 Hz and ranged from 0.2 to 0.8 s on average (90% duration). Additionally, each of the four call types was associated with measured behavioural and environmental parameters. Our results represent a significant advancement in understanding of the life history of this species and improve our capacity to acoustically monitor minke whales in a rapidly changing Antarctic region.

    View details for DOI 10.1098/rsos.211557

    View details for PubMedID 35911199

  • Feeding characteristics and prey profitability in five herring-feeding killer whales (Orcinus orca) in northern Norway MARINE MAMMAL SCIENCE Matika, A. F., Jourdain, E., Cade, D. E., Karoliussen, R., Hammond, P. S. 2022

    View details for DOI 10.1111/mms.12931

    View details for Web of Science ID 000777255100001

  • Scaling of maneuvering performance in baleen whales: larger whales outperform expectations. The Journal of experimental biology Segre, P. S., Gough, W. T., Roualdes, E. A., Cade, D. E., Czapanskiy, M. F., Fahlbusch, J., Kahane-Rapport, S. R., Oestreich, W. K., Bejder, L., Bierlich, K. C., Burrows, J. A., Calambokidis, J., Chenoweth, E. M., di Clemente, J., Durban, J. W., Fearnbach, H., Fish, F. E., Friedlaender, A. S., Hegelund, P., Johnston, D. W., Nowacek, D. P., Oudejans, M. G., Penry, G. S., Potvin, J., Simon, M., Stanworth, A., Straley, J. M., Szabo, A., Videsen, S. K., Visser, F., Weir, C. R., Wiley, D. N., Goldbogen, J. A. 2022; 225 (5)


    Despite their enormous size, whales make their living as voracious predators. To catch their much smaller, more maneuverable prey, they have developed several unique locomotor strategies that require high energetic input, high mechanical power output and a surprising degree of agility. To better understand how body size affects maneuverability at the largest scale, we used bio-logging data, aerial photogrammetry and a high-throughput approach to quantify the maneuvering performance of seven species of free-swimming baleen whale. We found that as body size increases, absolute maneuvering performance decreases: larger whales use lower accelerations and perform slower pitch-changes, rolls and turns than smaller species. We also found that baleen whales exhibit positive allometry of maneuvering performance: relative to their body size, larger whales use higher accelerations, and perform faster pitch-changes, rolls and certain types of turns than smaller species. However, not all maneuvers were impacted by body size in the same way, and we found that larger whales behaviorally adjust for their decreased agility by using turns that they can perform more effectively. The positive allometry of maneuvering performance suggests that large whales have compensated for their increased body size by evolving more effective control surfaces and by preferentially selecting maneuvers that play to their strengths.

    View details for DOI 10.1242/jeb.243224

    View details for PubMedID 35234874

  • Evidence for Size-Selective Predation by Antarctic Humpback Whales FRONTIERS IN MARINE SCIENCE Cade, D. E., Kahane-Rapport, S. R., Wallis, B., Goldbogen, J. A., Friedlaender, A. S. 2022; 9
  • The effect of group size on individual behavior of bubble-net feeding humpback whales in the southern Gulf of Maine MARINE MAMMAL SCIENCE Mastick, N. C., Wiley, D., Cade, D. E., Ware, C., Parks, S. E., Friedlaender, A. S. 2022

    View details for DOI 10.1111/mms.12905

    View details for Web of Science ID 000740705000001

  • Baleen whale inhalation variability revealed using animal-borne video tags. PeerJ Nazario, E. C., Cade, D. E., Bierlich, K. C., Czapanskiy, M. F., Goldbogen, J. A., Kahane-Rapport, S. R., van der Hoop, J. M., San Luis, M. T., Friedlaender, A. S. 2022; 10: e13724


    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.

    View details for DOI 10.7717/peerj.13724

    View details for PubMedID 35880219

  • Tools for integrating inertial sensor data with video bio-loggers, including estimation of animal orientation, motion, and position ANIMAL BIOTELEMETRY Cade, D. E., Gough, W. T., Czapanskiy, M. F., Fahlbusch, J. A., Kahane-Rapport, S. R., Linsky, J. J., Nichols, R. C., Oestreich, W. K., Wisniewska, D. M., Friedlaender, A. S., Goldbogen, J. A. 2021; 9 (1)
  • Scaling of oscillatory kinematics and Froude efficiency in baleen whales. The Journal of experimental biology Gough, W. T., Smith, H. J., Savoca, M. S., Czapanskiy, M. F., Fish, F. E., Potvin, J., Bierlich, K. C., Cade, D. E., Clemente, J. D., Kennedy, J., Segre, P., Stanworth, A., Weir, C., Goldbogen, J. A. 2021


    High efficiency lunate-tail swimming with high-aspect-ratio lifting surfaces has evolved in many vertebrate lineages, from fish to cetaceans. Baleen whales (Mysticeti) are the largest swimming animals that exhibit this locomotor strategy and present an ideal study system to examine how morphology and the kinematics of swimming scale to the largest body sizes. We used data from whale-borne inertial sensors coupled with morphometric measurements from aerial drones to calculate the hydrodynamic performance of oscillatory swimming in six baleen whale species ranging in body length from 5-25m (fin whale, Balaenoptera physalus; Bryde's whale, Balaenoptera edeni; sei whale, Balaenoptera borealis; Antarctic minke whales, Balaenoptera bonaerensis; humpback whales, Megaptera novaeangliae; and blue whales, Balaenoptera musculus). We find that mass-specific thrust increases with both swimming speed and body size. Froude efficiency, defined as the ratio of useful power output to the rate of energy input (Sloop, 1978), generally increased with swimming speed but decreased on average with increasing body size. This finding is contrary to previous results in smaller animals where Froude efficiency increased with body size. Although our empirically-parameterized estimates for swimming baleen whale drag was higher than that of a simple gliding model, oscillatory locomotion at this scale exhibits generally high Froude efficiency as in other adept swimmers. Our results quantify the fine-scale kinematics and estimate the hydrodynamics of routine and energetically expensive swimming modes at the largest scale.

    View details for DOI 10.1242/jeb.237586

    View details for PubMedID 34109418

  • Modelling short-term energetic costs of sonar disturbance to cetaceans using high-resolution foraging data JOURNAL OF APPLIED ECOLOGY Czapanskiy, M. F., Savoca, M. S., Gough, W. T., Segre, P. S., Wisniewska, D. M., Cade, D. E., Goldbogen, J. A. 2021
  • Predator‐scale spatial analysis of intra‐patch prey distribution reveals the energetic drivers of rorqual whale super‐group formation Functional Ecology Cade, D. E., Seakamela , M., Findlay, K. P., Fukunaga , J., Kahane‐Rapport, S. R., Warren, J. D., Calambokidis, J., Fahlbusch, J. A., Friedlaender , A. S., Hazen, E. L., Kotze , D., McCue, S., Meÿer , M., Oestreich , W. K., Oudejans, M. G., Wilke, C., Goldbogen, J. A. 2021

    View details for DOI 10.1111/1365-2435.13763

  • Context-dependent variability in the predicted daily energetic costs of disturbance for blue whales. Conservation physiology Pirotta, E. n., Booth, C. G., Cade, D. E., Calambokidis, J. n., Costa, D. P., Fahlbusch, J. A., Friedlaender, A. S., Goldbogen, J. A., Harwood, J. n., Hazen, E. L., New, L. n., Southall, B. L. 2021; 9 (1): coaa137


    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.

    View details for DOI 10.1093/conphys/coaa137

    View details for PubMedID 33505702

    View details for PubMedCentralID PMC7816799

  • Remoras pick where they stick on blue whales. The Journal of experimental biology Flammang, B. E., Marras, S., Anderson, E. J., Lehmkuhl, O., Mukherjee, A., Cade, D. E., Beckert, M., Nadler, J. H., Houzeaux, G., Vazquez, M., Amplo, H. E., Calambokidis, J., Friedlaender, A. S., Goldbogen, J. A. 2020; 223 (Pt 20)


    Animal-borne video recordings from blue whales in the open ocean show that remoras preferentially adhere to specific regions on the surface of the whale. Using empirical and computational fluid dynamics analyses, we show that remora attachment was specific to regions of separating flow and wakes caused by surface features on the whale. Adhesion at these locations offers remoras drag reduction of up to 71-84% compared with the freestream. Remoras were observed to move freely along the surface of the whale using skimming and sliding behaviors. Skimming provided drag reduction as high as 50-72% at some locations for some remora sizes, but little to none was available in regions where few to no remoras were observed. Experimental work suggests that the Venturi effect may help remoras stay near the whale while skimming. Understanding the flow environment around a swimming blue whale will inform the placement of biosensor tags to increase attachment time for extended ecological monitoring.

    View details for DOI 10.1242/jeb.226654

    View details for PubMedID 33115921

  • The scale of the whale: using video-tag data to evaluate sea-surface ice concentration from the perspective of individual Antarctic minke whales ANIMAL BIOTELEMETRY Linsky, J. J., Wilson, N., Cade, D. E., Goldbogen, J. A., Johnston, D. W., Friedlaender, A. S. 2020; 8 (1)
  • A perfectly inelastic collision: Bulk prey engulfment by baleen whales and dynamical implications for the world's largest cetaceans AMERICAN JOURNAL OF PHYSICS Potvin, J., Cade, D. E., Werth, A. J., Shadwick, R. E., Goldbogen, J. A. 2020; 88 (10): 851–63

    View details for DOI 10.1119/10.0001771

    View details for Web of Science ID 000571786400010

  • Animal-Borne Metrics Enable Acoustic Detection of Blue Whale Migration. Current biology : CB Oestreich, W. K., Fahlbusch, J. A., Cade, D. E., Calambokidis, J., Margolina, T., Joseph, J., Friedlaender, A. S., McKenna, M. F., Stimpert, A. K., Southall, B. L., Goldbogen, J. A., Ryan, J. P. 2020


    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 (Figure1A) 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]).

    View details for DOI 10.1016/j.cub.2020.08.105

    View details for PubMedID 33007246

  • An Algorithmic Approach to Natural Behavior. Current biology : CB Hein, A. M., Altshuler, D. L., Cade, D. E., Liao, J. C., Martin, B. T., Taylor, G. K. 2020; 30 (11): R663–R675


    Uncovering the mechanisms and implications of natural behavior is a goal that unites many fields of biology. Yet, the diversity, flexibility, and multi-scale nature of these behaviors often make understanding elusive. Here, we review studies of animal pursuit and evasion - two special classes of behavior where theory-driven experiments and new modeling techniques are beginning to uncover the general control principles underlying natural behavior. A key finding of these studies is that intricate sequences of pursuit and evasion behavior can often be constructed through simple, repeatable rules that link sensory input to motor output: we refer to these rules as behavioral algorithms. Identifying and mathematically characterizing these algorithms has led to important insights, including the discovery of guidance rules that attacking predators use to intercept mobile prey, and coordinated neural and biomechanical mechanisms that animals use to avoid impending collisions. Here, we argue that algorithms provide a good starting point for studies of natural behavior more generally. Rather than beginning at the neural or ecological levels of organization, we advocate starting in the middle, where the algorithms that link sensory input to behavioral output can provide a solid foundation from which to explore both the implementation and the ecological outcomes of behavior. We review insights that have been gained through such an algorithmic approach to pursuit and evasion behaviors. From these, we synthesize theoretical principles and lay out key modeling tools needed to apply an algorithmic approach to the study of other complex natural behaviors.

    View details for DOI 10.1016/j.cub.2020.04.018

    View details for PubMedID 32516620

  • Whale sharks increase swimming effort while filter feeding, but appear to maintain high foraging efficiencies. The Journal of experimental biology Cade, D. E., Levenson, J. J., Cooper, R., de la Parra, R., Webb, D. H., Dove, A. D. 2020


    Whale sharks (Rhincodon typus Smith 1828) - the largest extant fish species - reside in tropical environments, making them an exception to the general rule that animal size increases with latitude. How this largest fish thrives in tropical environments that promote high metabolism but support less robust zooplankton communities has not been sufficiently explained. We used open-source inertial measurement units (IMU) to log 397 hours of whale shark behavior in Yucatan, Mexico, at a site of both active feeding and intense wildlife tourism. Here we show that the strategies employed by whale sharks to compensate for the increased drag of an open mouth are similar to ram-feeders five orders of magnitude smaller and one order of magnitude larger. Presumed feeding constituted 20% of the total time budget of four sharks, with individual feeding bouts lasting up to 11 consecutive hrs. Compared to normal, sub-surface swimming, three sharks increased their stroke rate and amplitude while surface feeding, while one shark that fed at depth did not demonstrate a greatly increased energetic cost. Additionally, based on time-depth budgets, we estimate that aerial surveys of shark populations should consider including a correction factor of 3 to account for the proportion of daylight hours that sharks are not visible at the surface. With foraging bouts generally lasting several hours, interruptions to foraging during critical feeding periods may represent substantial energetic costs to these endangered species, and this study presents baseline data from which management decisions affecting tourist interactions with whale sharks may be made.

    View details for DOI 10.1242/jeb.224402

    View details for PubMedID 32366692

  • Energetic and physical limitations on the breaching performance of large whales. eLife Segre, P. S., Potvin, J., Cade, D. E., Calambokidis, J., Di Clemente, J., Fish, F. E., Friedlaender, A. S., Gough, W. T., Kahane-Rapport, S. R., Oliveira, C., Parks, S. E., Penry, G. S., Simon, M., Stimpert, A. K., Wiley, D. N., Bierlich, K. C., Madsen, P. T., Goldbogen, J. A. 2020; 9


    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.

    View details for DOI 10.7554/eLife.51760

    View details for PubMedID 32159511

  • From a calf's perspective: humpback whale nursing behavior on two US feeding grounds PEERJ Tackaberry, J. E., Cade, D. E., Goldbogen, J. A., Wiley, D. N., Friedlaender, A. S., Stimpert, A. K. 2020; 8: e8538


    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 mother's 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 < 0.0001). In contrast, mothers had lower ODBA and reduced FSR during nursing events compared to non-nursing segments. These data provide the first characterization of accelerometer data of humpback whale nursing confirmed by animal-borne video tags and the first analysis of nursing events on feeding grounds. This is an important step in understanding the energetic consequences of lactation while foraging.

    View details for DOI 10.7717/peerj.8538

    View details for Web of Science ID 000517940900003

    View details for PubMedID 32181052

    View details for PubMedCentralID PMC7060748

  • Whale sharks increase swimming effort while filter feeding, but appear to maintain high foraging efficiencies. The Journal of experimental biology Cade, D. E., Levenson, J. J., Cooper, R., de la Parra, R., Webb, D. H., Dove, A. D. 2020


    Whale sharks (Rhincodon typus Smith 1828) - the largest extant fish species - reside in tropical environments, making them an exception to the general rule that animal size increases with latitude. How this largest fish thrives in tropical environments that promote high metabolism but support less robust zooplankton communities has not been sufficiently explained. We used open-source inertial measurement units (IMU) to log 397 hours of whale shark behavior in Yucatan, Mexico, at a site of both active feeding and intense wildlife tourism. Here we show that the strategies employed by whale sharks to compensate for the increased drag of an open mouth are similar to ram-feeders five orders of magnitude smaller and one order of magnitude larger. Presumed feeding constituted 20% of the total time budget of four sharks, with individual feeding bouts lasting up to 11 consecutive hrs. Compared to normal, sub-surface swimming, three sharks increased their stroke rate and amplitude while surface feeding, while one shark that fed at depth did not demonstrate a greatly increased energetic cost. Additionally, based on time-depth budgets, we estimate that aerial surveys of shark populations should consider including a correction factor of 3 to account for the proportion of daylight hours that sharks are not visible at the surface. With foraging bouts generally lasting several hours, interruptions to foraging during critical feeding periods may represent substantial energetic costs to these endangered species, and this study presents baseline data from which management decisions affecting tourist interactions with whale sharks may be made.

    View details for DOI 10.1242/jeb.224402

    View details for PubMedID 34005603

  • Behavioral responses of individual blue whales (Balaenoptera musculus) to mid-frequency military sonar. The Journal of experimental biology Southall, B. L., DeRuiter, S. L., Friedlaender, A., Stimpert, A. K., Goldbogen, J. A., Hazen, E., Casey, C., Fregosi, S., Cade, D. E., Allen, A. N., Harris, C. M., Schorr, G., Moretti, D., Guan, S., Calambokidis, J. 2019; 222 (Pt 5)


    This study measured the degree of behavioral responses in blue whales (Balaenoptera musculus) to controlled noise exposure off the southern California coast. High-resolution movement and passive acoustic data were obtained from non-invasive archival tags (n=42) whereas surface positions were obtained with visual focal follows. Controlled exposure experiments (CEEs) were used to obtain direct behavioral measurements before, during and after simulated and operational military mid-frequency active sonar (MFAS), pseudorandom noise (PRN) and controls (no noise exposure). For a subset of deep-feeding animals (n=21), active acoustic measurements of prey were obtained and used as contextual covariates in response analyses. To investigate potential behavioral changes within individuals as a function of controlled noise exposure conditions, two parallel analyses of time-series data for selected behavioral parameters (e.g. diving, horizontal movement and feeding) were conducted. This included expert scoring of responses according to a specified behavioral severity rating paradigm and quantitative change-point analyses using Mahalanobis distance statistics. Both methods identified clear changes in some conditions. More than 50% of blue whales in deep-feeding states responded during CEEs, whereas no changes in behavior were identified in shallow-feeding blue whales. Overall, responses were generally brief, of low to moderate severity, and highly dependent on exposure context such as behavioral state, source-to-whale horizontal range and prey availability. Response probability did not follow a simple exposure-response model based on received exposure level. These results, in combination with additional analytical methods to investigate different aspects of potential responses within and among individuals, provide a comprehensive evaluation of how free-ranging blue whales responded to mid-frequency military sonar.

    View details for PubMedID 30833464

  • Behavioral responses of individual blue whales (Balaenoptera musculus) to mid-frequency military sonar JOURNAL OF EXPERIMENTAL BIOLOGY Southall, B. L., DeRuiter, S. L., Friedlaender, A., Stimpert, A. K., Goldbogen, J. A., Hazen, E., Casey, C., Fregosi, S., Cade, D. E., Allen, A. N., Harris, C. M., Schorr, G., Moretti, D., Guan, S., Calambokidis, J. 2019; 222 (5)

    View details for DOI 10.1242/jeb.190637

    View details for Web of Science ID 000461414600007

  • Scaling of swimming performance in baleen whales. The Journal of experimental biology Gough, W. T., Segre, P. S., Bierlich, K. C., Cade, D. E., Potvin, J. n., Fish, F. E., Dale, J. n., di Clemente, J. n., Friedlaender, A. S., Johnston, D. W., Kahane-Rapport, S. R., Kennedy, J. n., Long, J. H., Oudejans, M. n., Penry, G. n., Savoca, M. S., Simon, M. n., Videsen, S. K., Visser, F. n., Wiley, D. N., Goldbogen, J. A. 2019


    The scale-dependence of locomotor factors have long been studied in comparative biomechanics, but remain poorly understood for animals at the upper extremes of body size. Rorqual baleen whales include the largest animals, but we lack basic kinematic data about their movements and behavior below the ocean surface. Here we combined morphometrics from aerial drone photogrammetry, whale-borne inertial sensing tag data, and hydrodynamic modeling to study the locomotion of five rorqual species. We quantified changes in tail oscillatory frequency and cruising speed for individual whales spanning a threefold variation in body length, corresponding to an order of magnitude variation in estimated body mass. Our results showed that oscillatory frequency decreases with body length (∝ length-0.53) while cruising speed remains roughly invariant (∝ length0.08) at 2 m s-1 We compared these measured results for oscillatory frequency against simplified models of an oscillating cantilever beam (∝ length-1) and an optimized oscillating Strouhal vortex generator (∝ length-1). The difference between our length-scaling exponent and the simplified models suggests that animals are often swimming non-optimally in order to feed or perform other routine behaviors. Cruising speed aligned more closely with an estimate of the optimal speed required to minimize the energetic cost of swimming (∝ length0.07). Our results are among the first to elucidate the relationships between both oscillatory frequency and cruising speed and body size for free-swimming animals at the largest scale.

    View details for DOI 10.1242/jeb.204172

    View details for PubMedID 31558588

  • Filtration area scaling and evolution in mysticetes: trophic niche partitioning and the curious cases of sei and pygmy right whales BIOLOGICAL JOURNAL OF THE LINNEAN SOCIETY Werth, A. J., Potvin, J., Shadwick, R. E., Jensen, M. M., Cade, D. E., Goldbogen, J. A. 2018; 125 (2): 264–79
  • Determining forward speed from accelerometer jiggle in aquatic environments. The Journal of experimental biology Cade, D. E., Barr, K. R., Calambokidis, J. n., Friedlaender, A. S., Goldbogen, J. A. 2018; 221 (Pt 2)


    How fast animals move is critical to understanding their energetic requirements, locomotor capacity and foraging performance, yet current methods for measuring speed via animal-attached devices are not universally applicable. Here, we present and evaluate a new method that relates forward speed to the stochastic motion of biologging devices as tag jiggle, the amplitude of the tag vibrations as measured by high sample rate accelerometers, increases exponentially with increasing speed. We successfully tested this method in a flow tank using two types of biologging devices and in situ on wild cetaceans spanning ∼3 to >20 m in length using two types of suction cup-attached tag and two types of dart-attached tag. This technique provides some advantages over other approaches for determining speed as it is device-orientation independent and relies only on a pressure sensor and a high sample rate accelerometer, sensors that are nearly universal across biologging device types.

    View details for PubMedID 29191861

  • Determining forward speed from accelerometer jiggle in aquatic environments JOURNAL OF EXPERIMENTAL BIOLOGY Cade, D. E., Barr, K. R., Calambokidis, J., Friedlaender, A. S., Goldbogen, J. A. 2018; 221 (2)

    View details for DOI 10.1242/jeb.170449

    View details for Web of Science ID 000424076300018

  • Context-dependent lateralized feeding strategies in blue whales CURRENT BIOLOGY Friedlaender, A. S., Herbert-Read, J. E., Hazen, E. L., Cade, D. E., Calambokidis, J., Southall, B. L., Stimpert, A. K., Goldbogen, J. A. 2017; 27 (22): R1206–R1208


    Lateralized behaviors benefit individuals by increasing task efficiency in foraging and anti-predator behaviors [1-4]. The conventional lateralization paradigm suggests individuals are left or right lateralized, although the direction of this laterality can vary for different tasks (e.g. foraging or predator inspection/avoidance). By fitting tri-axial movement sensors to blue whales (Balaenoptera musculus), and by recording the direction and size of their rolls during lunge feeding events, we show how these animals differ from such a paradigm. The strength and direction of individuals' lateralization were related to where and how the whales were feeding in the water column. Smaller rolls (≤180°) predominantly occurred at depth (>70 m), with whales being more likely to rotate clockwise around their longest axis (right lateralized). Larger rolls (>180°), conversely, occurred more often at shallower depths (<70 m) and were more likely to be performed anti-clockwise (left lateralized). More acrobatic rolls are typically used to target small, less dense krill patches near the water's surface [5,6], and we posit that the specialization of lateralized feeding strategies may enhance foraging efficiency in environments with heterogeneous prey distributions.

    View details for PubMedID 29161554

  • How Baleen Whales Feed: The Biomechanics of Engulfment and Filtration. Annual review of marine science Goldbogen, J. A., Cade, D. E., Calambokidis, J., Friedlaender, A. S., Potvin, J., Segre, P. S., Werth, A. J. 2017; 9: 367-386


    Baleen whales are gigantic obligate filter feeders that exploit aggregations of small-bodied prey in littoral, epipelagic, and mesopelagic ecosystems. At the extreme of maximum body size observed among mammals, baleen whales exhibit a unique combination of high overall energetic demands and low mass-specific metabolic rates. As a result, most baleen whale species have evolved filter-feeding mechanisms and foraging strategies that take advantage of seasonally abundant yet patchily and ephemerally distributed prey resources. New methodologies consisting of multi-sensor tags, active acoustic prey mapping, and hydrodynamic modeling have revolutionized our ability to study the physiology and ecology of baleen whale feeding mechanisms. Here, we review the current state of the field by exploring several hypotheses that aim to explain how baleen whales feed. Despite significant advances, major questions remain about the processes that underlie these extreme feeding mechanisms, which enabled the evolution of the largest animals of all time.

    View details for DOI 10.1146/annurev-marine-122414-033905

    View details for PubMedID 27620830

  • Humpback whale "super-groups" - A novel low-latitude feeding behaviour of Southern Hemisphere humpback whales (Megaptera novaeangliae) in the Benguela Upwelling System. PloS one Findlay, K. P., Seakamela, S. M., Meÿer, M. A., Kirkman, S. P., Barendse, J., Cade, D. E., Hurwitz, D., Kennedy, A. S., Kotze, P. G., McCue, S. A., Thornton, M., Vargas-Fonseca, O. A., Wilke, C. G. 2017; 12 (3)


    Southern Hemisphere humpback whales (Megaptera novaeangliae) generally undertake annual migrations from polar summer feeding grounds to winter calving and nursery grounds in subtropical and tropical coastal waters. Evidence for such migrations arises from seasonality of historic whaling catches by latitude, Discovery and natural mark returns, and results of satellite tagging studies. Feeding is generally believed to be limited to the southern polar region, where Antarctic krill (Euphausia superba) has been identified as the primary prey item. Non-migrations and / or suspended migrations to the polar feeding grounds have previously been reported from a summer presence of whales in the Benguela System, where feeding on euphausiids (E. lucens), hyperiid amphipods (Themisto gaudichaudii), mantis shrimp (Pterygosquilla armata capensis) and clupeid fish has been described. Three recent research cruises (in October/November 2011, October/November 2014 and October/November 2015) identified large tightly-spaced groups (20 to 200 individuals) of feeding humpback whales aggregated over at least a one-month period across a 220 nautical mile region of the southern Benguela System. Feeding behaviour was identified by lunges, strong milling and repetitive and consecutive diving behaviours, associated bird and seal feeding, defecations and the pungent "fishy" smell of whale blows. Although no dedicated prey sampling could be carried out within the tightly spaced feeding aggregations, observations of E. lucens in the region of groups and the full stomach contents of mantis shrimp from both a co-occurring predatory fish species (Thyrsites atun) and one entangled humpback whale mortality suggest these may be the primary prey items of at least some of the feeding aggregations. Reasons for this recent novel behaviour pattern remain speculative, but may relate to increasing summer humpback whale abundance in the region. These novel, predictable, inter-annual, low latitude feeding events provide considerable potential for further investigation of Southern Hemisphere humpback feeding behaviours in these relatively accessible low-latitude waters.

    View details for DOI 10.1371/journal.pone.0172002

    View details for PubMedID 28249036

    View details for PubMedCentralID PMC5332018

  • Hydrodynamic properties of fin whale flippers predict maximum rolling performance. journal of experimental biology Segre, P. S., Cade, D. E., Fish, F. E., Potvin, J., Allen, A. N., Calambokidis, J., Friedlaender, A. S., Goldbogen, J. A. 2016; 219: 3315-3320


    Maneuverability is one of the most important and least understood aspects of animal locomotion. The hydrofoil-like flippers of cetaceans are thought to function as control surfaces that effect maneuvers, but quantitative tests of this hypothesis have been lacking. Here, we constructed a simple hydrodynamic model to predict the longitudinal-axis roll performance of fin whales, and we tested its predictions against kinematic data recorded by on-board movement sensors from 27 free-swimming fin whales. We found that for a given swimming speed and roll excursion, the roll velocity of fin whales calculated from our field data agrees well with that predicted by our hydrodynamic model. Although fluke and body torsion may further influence performance, our results indicate that lift generated by the flippers is sufficient to drive most of the longitudinal-axis rolls used by fin whales for feeding and maneuvering.

    View details for PubMedID 27591304

  • Kinematic Diversity in Rorqual Whale Feeding Mechanisms CURRENT BIOLOGY Cade, D. E., Friedlaender, A. S., Calambokidis, J., Goldbogen, J. A. 2016; 26 (19): 2617-2624


    Rorqual whales exhibit an extreme lunge filter-feeding strategy characterized by acceleration to high speed and engulfment of a large volume of prey-laden water [1-4]. Although tagging studies have quantified the kinematics of lunge feeding, the timing of engulfment relative to body acceleration has been modeled conflictingly because it could never be directly measured [5-7]. The temporal coordination of these processes has a major impact on the hydrodynamics and energetics of this high-cost feeding strategy [5-9]. If engulfment and body acceleration are temporally distinct, the overall cost of this dynamic feeding event would be minimized. However, greater temporal overlap of these two phases would theoretically result in higher drag and greater energetic costs. To address this discrepancy, we used animal-borne synchronized video and 3D movement sensors to quantify the kinematics of both the skull and body during feeding events. Krill-feeding blue and humpback whales exhibited temporally distinct acceleration and engulfment phases, with humpback whales reaching maximum gape earlier than blue whales. In these whales, engulfment coincided largely with body deceleration; however, humpback whales pursuing more agile fish demonstrated highly variable coordination of skull and body kinematics in the context of complex prey-herding techniques. These data suggest that rorquals modulate the coordination of acceleration and engulfment to optimize foraging efficiency by minimizing locomotor costs and maximizing prey capture. Moreover, this newfound kinematic diversity observed among rorquals indicates that the energetic efficiency of foraging is driven both by the whale's engulfment capacity and the comparative locomotor capabilities of predator and prey. VIDEO ABSTRACT.

    View details for DOI 10.1016/j.cub.2016.07.037

    View details for Web of Science ID 000385690800022

    View details for PubMedID 27666966

  • Insights into the Underwater Behavior, Species Interactions, and Biomechanics of Baleen Whales using Suction-Cup Attached Video and Inertial Sensors Goldbogen, J., Cade, D., Calambokidis, J., Stimpert, A., Friedlaender, A. FEDERATION AMER SOC EXP BIOL. 2016
  • Depths, migration rates and environmental associations of acoustic scattering layers in the Gulf of California DEEP-SEA RESEARCH PART I-OCEANOGRAPHIC RESEARCH PAPERS Cade, D. E., Benoit-Bird, K. J. 2015; 102: 78-89
  • An automatic and quantitative approach to the detection and tracking of acoustic scattering layers LIMNOLOGY AND OCEANOGRAPHY-METHODS Cade, D. E., Benoit-Bird, K. J. 2014; 12: 742-756