Academic Appointments

Honors & Awards

  • Young Investigator Award, Office of Naval Research (2016)
  • Hoagland Award for Teaching Innovation, Vice Provost for Undergraduate Education, Humanities and Sciences Dean's Office (2015)

Professional Education

  • BSc, University of Texas at Austin, Zoology (2002)
  • MSc, Scripps Institution of Oceanography, University of California - San Diego, Marine Biology (2005)
  • PhD, University of British Columbia, Zoology (2010)

2017-18 Courses

Stanford Advisees

All Publications

  • Independent evolution of baleen whale gigantism linked to Plio-Pleistocene ocean dynamics. Proceedings. Biological sciences Slater, G. J., Goldbogen, J. A., Pyenson, N. D. 2017; 284 (1855)


    Vertebrates have evolved to gigantic sizes repeatedly over the past 250 Myr, reaching their extreme in today's baleen whales (Mysticeti). Hypotheses for the evolution of exceptionally large size in mysticetes range from niche partitioning to predator avoidance, but there has been no quantitative examination of body size evolutionary dynamics in this clade and it remains unclear when, why or how gigantism evolved. By fitting phylogenetic macroevolutionary models to a dataset consisting of living and extinct species, we show that mysticetes underwent a clade-wide shift in their mode of body size evolution during the Plio-Pleistocene. This transition, from Brownian motion-like dynamics to a trended random walk towards larger size, is temporally linked to the onset of seasonally intensified upwelling along coastal ecosystems. High prey densities resulting from wind-driven upwelling, rather than abundant resources alone, are the primary determinant of efficient foraging in extant mysticetes and Late Pliocene changes in ocean dynamics may have provided an ecological pathway to gigantism in multiple independent lineages.

    View details for DOI 10.1098/rspb.2017.0546

    View details for PubMedID 28539520

  • 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

  • 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

  • Blue whales (Balaenoptera musculus) optimize foraging efficiency by balancing oxygen use and energy gain as a function of prey density. Science advances Hazen, E. L., Friedlaender, A. S., Goldbogen, J. A. 2015; 1 (9)


    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.

    View details for DOI 10.1126/sciadv.1500469

    View details for PubMedID 26601290

  • Stretchy nerves are an essential component of the extreme feeding mechanism of rorqual whales CURRENT BIOLOGY Vogl, A. W., Lillie, M. A., Piscitelli, M. A., Goldbogen, J. A., Pyenson, N. D., Shadwick, R. E. 2015; 25 (9): R360-R361


    Rorqual whales (Balaenopteridae) are among the largest vertebrates that have ever lived and include blue (Balaenoptera musculus) and fin (Balaenoptera physalus) whales. Rorquals differ from other baleen whales (Mysticeti) in possessing longitudinal furrows or grooves in the ventral skin that extend from the mouth to the umbilicus. This ventral grooved blubber directly relates to their intermittent lunge feeding strategy, which is unique among vertebrates and was potentially an evolutionary innovation that led to gigantism in this lineage [1]. This strategy involves the rorqual whale rapidly engulfing a huge volume of prey-laden water and then concentrating the prey by more slowly expelling the water through baleen plates (Figure 1A). The volume of water engulfed during a lunge can exceed the volume of the whale itself [2]. During engulfment, the whale accelerates, opens its jaw until it is almost perpendicular to the rostrum, and then the highly compliant floor of the oral cavity is inflated by the incoming water [3]. The floor of the oral cavity expands by inversion of the tongue and ballooning of the adjacent floor of the mouth into the cavum ventrale, an immense fascial pocket between the body wall and overlying blubber layer that reaches as far back as the umbilicus. The ventral grooved blubber in fin whales expands by an estimated 162% in the circumferential direction and 38% longitudinally [4]. In fin whales, multiple lunges can occur during a single dive, and the average time between lunges is just over forty seconds [3]. Here, we show that nerves in the floor of the oral cavity of fin whales are highly extensible.

    View details for Web of Science ID 000353999000007

    View details for PubMedID 25942546

  • Mechanical challenges to freshwater residency in sharks and rays. journal of experimental biology Gleiss, A. C., Potvin, J., Keleher, J. J., Whitty, J. M., Morgan, D. L., Goldbogen, J. A. 2015; 218: 1099-1110


    Major transitions between marine and freshwater habitats are relatively infrequent, primarily as a result of major physiological and ecological challenges. Few species of cartilaginous fish have evolved to occupy freshwater habitats. Current thought suggests that the metabolic physiology of sharks has remained a barrier to the diversification of this taxon in freshwater ecosystems. Here, we demonstrate that the physical properties of water provide an additional constraint for this species-rich group to occupy freshwater systems. Using hydromechanical modeling, we show that occurrence in fresh water results in a two- to three-fold increase in negative buoyancy for sharks and rays. This carries the energetic cost of lift production and results in increased buoyancy-dependent mechanical power requirements for swimming and increased optimal swim speeds. The primary source of buoyancy, the lipid-rich liver, offers only limited compensation for increased negative buoyancy as a result of decreasing water density; maintaining the same submerged weight would involve increasing the liver volume by very large amounts: 3- to 4-fold in scenarios where liver density is also reduced to currently observed minimal levels and 8-fold without any changes in liver density. The first data on body density from two species of elasmobranch occurring in freshwater (the bull shark Carcharhinus leucas, Müller and Henle 1839, and the largetooth sawfish Pristis pristis, Linnaeus 1758) support this hypothesis, showing similar liver sizes as marine forms but lower liver densities, but the greatest negative buoyancies of any elasmobranch studied to date. Our data suggest that the mechanical challenges associated with buoyancy control may have hampered the invasion of freshwater habitats in elasmobranchs, highlighting an additional key factor that may govern the predisposition of marine organisms to successfully establish in freshwater habitats.

    View details for DOI 10.1242/jeb.114868

    View details for PubMedID 25573824

  • Feeding rates and under-ice foraging strategies of the smallest lunge filter feeder, the Antarctic minke whale (Balaenoptera bonaerensis). journal of experimental biology Friedlaender, A. S., Goldbogen, J. A., Nowacek, D. P., Read, A. J., Johnston, D., Gales, N. 2014; 217: 2851-2854


    Body size and feeding mode are two fundamental characteristics that determine foraging performance and ecological niche. As the smallest obligate lunge filter feeders, minke whales represent an ideal system for studying the physical and energetic limits of filter feeding in endotherms. We used multi-sensor suction cup tags to quantify the feeding performance of Antarctic minke whales. Foraging dives around and beneath sea ice contained up to 24 lunges per dive, the highest feeding rates for any lunge-feeding whale. Their small size allows minke whales access to krill in sea-ice environments not easily accessible to larger baleen whales. Furthermore, their ability to filter feed provides an advantage over other smaller sympatric krill predators such as penguins and seals that feed on individual prey. The unique combination of body size, feeding mechanism and sea-ice habitat of Antarctic minke whales defines a previously undocumented energetic niche that is unique among aquatic vertebrates.

    View details for DOI 10.1242/jeb.106682

    View details for PubMedID 25122916

  • Using accelerometers to determine the calling behavior of tagged baleen whales. journal of experimental biology Goldbogen, J. A., Stimpert, A. K., DeRuiter, S. L., Calambokidis, J., Friedlaender, A. S., Schorr, G. S., Moretti, D. J., Tyack, P. L., Southall, B. L. 2014; 217: 2449-2455


    Low-frequency acoustic signals generated by baleen whales can propagate over vast distances, making the assignment of calls to specific individuals problematic. Here, we report the novel use of acoustic recording tags equipped with high-resolution accelerometers to detect vibrations from the surface of two tagged fin whales that directly match the timing of recorded acoustic signals. A tag deployed on a buoy in the vicinity of calling fin whales and a recording from a tag that had just fallen off a whale were able to detect calls acoustically but did not record corresponding accelerometer signals that were measured on calling individuals. Across the hundreds of calls measured on two tagged fin whales, the accelerometer response was generally anisotropic across all three axes, appeared to depend on tag placement and increased with the level of received sound. These data demonstrate that high-sample rate accelerometry can provide important insights into the acoustic behavior of baleen whales that communicate at low frequencies. This method helps identify vocalizing whales, which in turn enables the quantification of call rates, a fundamental component of models used to estimate baleen whale abundance and distribution from passive acoustic monitoring.

    View details for DOI 10.1242/jeb.103259

    View details for PubMedID 24803468

  • Avoidance responses of minke whales to 1-4kHz naval sonar. Marine pollution bulletin Kvadsheim, P. H., Deruiter, S., Sivle, L. D., Goldbogen, J., Roland-Hansen, R., Miller, P. J., Lam, F. A., Calambokidis, J., Friedlaender, A., Visser, F., Tyack, P. L., Kleivane, L., Southall, B. 2017


    Minke whales are difficult to study and little information exists regarding their responses to anthropogenic sound. This study pools data from behavioural response studies off California and Norway. Data are derived from four tagged animals, of which one from each location was exposed to naval sonar signals. Statistical analyses were conducted using Mahalanobis distance to compare overall changes in parameters summarising dive behaviour, avoidance behaviour, and potential energetic costs of disturbance. Our quantitative analysis showed that both animals initiated avoidance behaviour, but responses were not associated with unusual dive behaviour. In one exposed animal the avoidance of the sonar source included a 5-fold increase in horizontal speed away from the source, implying a significant increase in metabolic rate. Despite the different environmental settings and exposure contexts, clear changes in behaviour were observed providing the first insights into the nature of responses to human noise for this wide-ranging species.

    View details for DOI 10.1016/j.marpolbul.2017.05.037

    View details for PubMedID 28552251

  • A MULTIVARIATE MIXED HIDDEN MARKOV MODEL FOR BLUE WHALE BEHAVIOUR AND RESPONSES TO SOUND EXPOSURE ANNALS OF APPLIED STATISTICS DeRuiter, S. L., Langrock, R., Skirbutas, T., Goldbogen, J. A., Calambokidis, J., Friedlaender, A. S., Southall, B. L. 2017; 11 (1): 362-392
  • How to tag a jellyfish? A methodological review and guidelines to successful jellyfish tagging JOURNAL OF PLANKTON RESEARCH Fossette, S., Katija, K., Goldbogen, J. A., Bograd, S., Patry, W., Howard, M. J., Knowles, T., Haddock, S. H., Bedell, L., Hazen, E. L., Robison, B. H., Mooney, T. A., Shorter, K. A., Bastian, T., Gleiss, A. C. 2016; 38 (6): 1347-1363
  • Studying cetacean behaviour: new technological approaches and conservation applications ANIMAL BEHAVIOUR Nowacek, D. P., Christiansen, F., Bejder, L., Goldbogen, J. A., Friedlaender, A. S. 2016; 120: 235-244
  • Development of an automated method of detecting stereotyped feeding events in multisensor data from tagged rorqual whales ECOLOGY AND EVOLUTION Allen, A. N., Goldbogen, J. A., Friedlaender, A. S., Calambokidis, J. 2016; 6 (20): 7522-7535

    View details for DOI 10.1002/ece3.2386

    View details for Web of Science ID 000386429200033

  • Discrimination of fast click-series produced by tagged Risso's dolphins (Grampus griseus) for echolocation or communication JOURNAL OF EXPERIMENTAL BIOLOGY Arranz, P., DeRuiter, S. L., Stimpert, A. K., Neves, S., Friedlaender, A. S., Goldbogen, J. A., Visser, F., Calambokidis, J., Southall, B. L., Tyack, P. L. 2016; 219 (18): 2898-2907

    View details for DOI 10.1242/jeb.144295

    View details for Web of Science ID 000384250600022

  • Prey-mediated behavioral responses of feeding blue whales in controlled sound exposure experiments ECOLOGICAL APPLICATIONS Friedlaender, A. S., Hazen, E. L., Goldbogen, J. A., Stimpert, A. K., Calambokidis, J., Southall, B. L. 2016; 26 (4): 1075-1085


    Behavioral response studies provide significant insights into the nature, magnitude, and consequences of changes in animal behavior in response to some external stimulus. Controlled exposure experiments (CEEs) to study behavioral response have faced challenges in quantifying the importance of and interaction among individual variability, exposure conditions, and environmental covariates. To investigate these complex parameters relative to blue whale behavior and how it may change as a function of certain sounds, we deployed multi-sensor acoustic tags and conducted CEEs using simulated mid-frequency active sonar (MFAS) and pseudo-random noise (PRN) stimuli, while collecting synoptic, quantitative prey measures. In contrast to previous approaches that lacked such prey data, our integrated approach explained substantially more variance in blue whale dive behavioral responses to mid-frequency sounds (r2 = 0.725 vs. 0.14 previously). Results demonstrate that deep-feeding whales respond more clearly and strongly to CEEs than those in other behavioral states, but this was only evident with the increased explanatory power provided by incorporating prey density and distribution as contextual covariates. Including contextual variables increases the ability to characterize behavioral variability and empirically strengthens previous findings that deep-feeding blue whales respond significantly to mid-frequency sound exposure. However, our results are only based on a single behavioral state with a limited sample size, and this analytical framework should be applied broadly across behavioral states. The increased capability to describe and account for individual response variability by including environmental variables, such as prey, that drive foraging behavior underscores the importance of integrating these and other relevant contextual parameters in experimental designs. Our results suggest the need to measure and account for the ecological dynamics of predator-prey interactions when studying the effects of anthropogenic disturbance in feeding animals.

    View details for DOI 10.1002/15-0783

    View details for Web of Science ID 000378092900009

    View details for PubMedID 27509749

  • Multiple-stage decisions in a marine central-place forager ROYAL SOCIETY OPEN SCIENCE Friedlaender, A. S., Johnston, D. W., Tyson, R. B., Kaltenberg, A., Goldbogen, J. A., Stimpert, A. K., Curtice, C., Hazen, E. L., Halpin, P. N., Read, A. J., Nowacek, D. P. 2016; 3 (5)


    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.

    View details for DOI 10.1098/rsos.160043

    View details for Web of Science ID 000377969800009

    View details for PubMedID 27293784

  • Using morphology to infer physiology: case studies on rorqual whales (Balaenopteridae) CANADIAN JOURNAL OF ZOOLOGY Goldbogen, J. A., Shadwick, R. E., Lillie, M. A., Piscitelli, M. A., Potvin, J., Pyenson, N. D., Vogl, A. W. 2015; 93 (9): 687-700
  • Prey density and distribution drive the three-dimensional foraging strategies of the largest filter feeder FUNCTIONAL ECOLOGY Goldbogen, J. A., Hazen, E. L., Friedlaender, A. S., Calambokidis, J., DeRuiter, S. L., Stimpert, A. K., Southall, B. L. 2015; 29 (7): 951-961
  • Feeding performance by sympatric blue and fin whales exploiting a common prey resource MARINE MAMMAL SCIENCE Friedlaender, A. S., Goldbogen, J. A., Hazen, E. L., Calambokidis, J., Southall, B. L. 2015; 31 (1): 345-354

    View details for DOI 10.1111/mms.12134

    View details for Web of Science ID 000346769200018

  • Simultaneous tracking of blue whales and large ships demonstrates limited behavioral responses for avoiding collision ENDANGERED SPECIES RESEARCH McKenna, M. F., Calambokidis, J., Oleson, E. M., Laist, D. W., Goldbogen, J. A. 2015; 27 (3): 219-232

    View details for DOI 10.3354/esr00666

    View details for Web of Science ID 000353916900003

  • Acoustic and foraging behavior of a Baird's beaked whale, Berardius bairdii, exposed to simulated sonar SCIENTIFIC REPORTS Stimpert, A. K., DeRuiter, S. L., Southall, B. L., Moretti, D. J., Falcone, E. A., Goldbogen, J. A., Friedlaender, A., Schorr, G. S., Calambokidis, J. 2014; 4

    View details for DOI 10.1038/srep07031

    View details for Web of Science ID 000344762200006

  • The device that revolutionized marine organismal biology. journal of experimental biology Goldbogen, J. A., Meir, J. U. 2014; 217: 167-168

    View details for DOI 10.1242/jeb.092189

    View details for PubMedID 24431140

  • Tracking fitness in marine vertebrates: current knowledge and opportunities for future research MARINE ECOLOGY PROGRESS SERIES Crossin, G. T., Cooke, S. J., Goldbogen, J. A., Phillips, R. A. 2014; 496: 1-17

    View details for DOI 10.3354/meps10691

    View details for Web of Science ID 000330356500001