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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.

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.

Baleen whales influence their ecosystems through immense prey consumption and nutrient recycling 1 – 3 . It is difficult to accurately gauge the magnitude of their current or historic ecosystem role without measuring feeding rates and prey consumed. To date, prey consumption of the largest species has been estimated using metabolic models 3 – 9 based on extrapolations that lack empirical validation. Here, we used tags deployed on seven baleen whale (Mysticeti) species ( n  = 321 tag deployments) in conjunction with acoustic measurements of prey density to calculate prey consumption at daily to annual scales from the Atlantic, Pacific, and Southern Oceans. Our results suggest that previous studies 3 – 9 have underestimated baleen whale prey consumption by threefold or more in some ecosystems. In the Southern Ocean alone, we calculate that pre-whaling populations of mysticetes annually consumed 430 million tonnes of Antarctic krill ( Euphausia superba ), twice the current estimated total biomass of E. superba 10 , and more than twice the global catch of marine fisheries today 11 . Larger whale populations may have supported higher productivity in large marine regions through enhanced nutrient recycling: our findings suggest mysticetes recycled 1.2 × 10 4  tonnes iron yr −1 in the Southern Ocean before whaling compared to 1.2 × 10 3  tonnes iron yr −1 recycled by whales today. The recovery of baleen whales and their nutrient recycling services 2 , 3 , 7 could augment productivity and restore ecosystem function lost during 20th century whaling 12 , 13 . A combination of 3D whale locations and acoustic measurements of prey density is used here to show that whales’ consumption of krill is several times larger than often thought.

The unique engulfment filtration strategy of microphagous rorqual whales has evolved relatively recently (<5 Ma) and exploits extreme predator/prey size ratios to overcome the maneuverability advantages of swarms of small prey, such as krill. Forage fish, in contrast, have been engaged in evolutionary arms races with their predators for more than 100 million years and have performance capabilities that suggest they should easily evade whale-sized predators, yet they are regularly hunted by some species of rorqual whales. To explore this phenomenon, we determined, in a laboratory setting, when individual anchovies initiated escape from virtually approaching whales, then used these results along with in situ humpback whale attack data to model how predator speed and engulfment timing affected capture rates. Anchovies were found to respond to approaching visual looming stimuli at expansion rates that give ample chance to escape from a sea lion-sized predator, but humpback whales could capture as much as 30–60% of a school at once because the increase in their apparent (visual) size does not cross their prey’s response threshold until after rapid jaw expansion. Humpback whales are, thus, incentivized to delay engulfment until they are very close to a prey school, even if this results in higher hydrodynamic drag. This potential exaptation of a microphagous filter feeding strategy for fish foraging enables humpback whales to achieve 7× the energetic efficiency (per lunge) of krill foraging, allowing for flexible foraging strategies that may underlie their ecological success in fluctuating oceanic conditions.

Bulk-filter feeding is an energetically efficient strategy for resource acquisition and assimilation, and facilitates the maintenance of extreme body size as exemplified by baleen whales (Mysticeti) and multiple lineages of bony and cartilaginous fishes. Among mysticetes, rorqual whales (Balaenopteridae) exhibit an intermittent ram filter feeding mode, lunge feeding, which requires the abandonment of body-streamlining in favor of a high-drag, mouth-open configuration aimed at engulfing a very large amount of prey-laden water. Particularly while lunge feeding on krill (the most widespread prey preference among rorquals), the effort required during engulfment involve short bouts of high-intensity muscle activity that demand high metabolic output. We used computational modeling together with morphological and kinematic data on humpback (Megaptera noveaangliae), fin (Balaenoptera physalus), blue (Balaenoptera musculus) and minke (Balaenoptera acutorostrata) whales to estimate engulfment power output in comparison with standard metrics of metabolic rate. The simulations reveal that engulfment metabolism increases across the full body size of the larger rorqual species to nearly 50 times the basal metabolic rate of terrestrial mammals of the same body mass. Moreover, they suggest that the metabolism of the largest body sizes runs with significant oxygen deficits during mouth opening, namely, 20% over maximum VO2 at the size of the largest blue whales, thus requiring significant contributions from anaerobic catabolism during a lunge and significant recovery after a lunge. Our analyses show that engulfment metabolism is also significantly lower for smaller adults, typically one-tenth to one-half VO2|max. These results not only point to a physiological limit on maximum body size in this lineage, but also have major implications for the ontogeny of extant rorquals as well as the evolutionary pathways used by ancestral toothed whales to transition from hunting individual prey items to filter feeding on prey aggregations.

The Southern Ocean ecosystem has undergone extensive changes in the past two centuries driven by industrial sealing and whaling, climate change and commercial fishing. However, following the end of commercial whaling, some populations of whales in this region are recovering. Baleen whales are reliant on Antarctic krill, which is also the largest Southern Ocean fishery. Since 1993, krill catch has increased fourfold, buoyed by nutritional supplement and aquaculture industries. In this Perspective, we approximate baleen whale consumption of Antarctic krill before and after whaling to examine if the ecosystem can support both humans and whales as krill predators. Our back-of-the-envelope calculations suggest that current krill biomass cannot support both an expanding krill fishery and the recovery of whale populations to pre-whaling sizes, highlighting an emerging human-wildlife conflict. We then provide recommendations for enhancing sustainability in this region by reducing encounters with whales and bolstering the krill population. The Southern Ocean ecosystem is recovering from 20th-century industrial whaling, while the krill fishery in this region has grown rapidly and may expand further, driven by demand for supplements and aquaculture feed. This Perspective discusses how current krill biomass is unlikely to support both a growing krill fishery and rebounding whale populations in the Southern Ocean.

For many marine organisms, especially large whales that cannot be studied in laboratory settings, our ability to obtain basic behavioral and physiological data is limited, because these organisms occupy offshore habitats and spend a majority of their time underwater. A class of multisensor, suction-cup-attached archival tags has revolutionized the study of large baleen whales, particularly with respect to the predatory strategies used by these gigantic bulk filter feeders to exploit abundant oceanic resources. By integrating these data with those from other disciplines, researchers have uncovered a diverse and extraordinary set of underwater behaviors, ranging from acrobatic diving maneuvers to extreme feeding events during which whales engulf volumes of prey-laden water that are much larger than their own body. This research framework not only improves our knowledge of the individual performance and behavior of these keystone predators but also informs our ability to understand the dynamics of complex marine ecosystems.

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.

Summary Despite their importance in determining the rate of both energy gain and expenditure, how the fine‐scale kinematics of foraging are modified in response to changes in prey abundance and distribution remain poorly understood in many animal ecosystems. In the marine environment, bulk‐filter feeders rely on dense aggregations of prey for energetically efficient foraging. Rorqual whales (Balaenopteridae) exhibit a unique form of filter feeding called lunge feeding, a process whereby discrete volumes of prey‐laden water are intermittently engulfed and filtered. In many large rorqual species the size of engulfed water mass is commensurate with the whale's body size, yet is engulfed in just a few seconds. This filter‐feeding mode thus requires precise coordination of the body and enlarged engulfment apparatus to maximize capture efficiency. Previous studies from whale‐borne tags revealed that many rorqual species perform rolling behaviours when foraging. It has been hypothesized that such acrobatic manoeuvres may be required for efficient prey capture when prey manifest in small discrete patches, but to date there has been no comprehensive analysis of prey patch characteristics during lunge feeding events. We developed a null hypothesis that blue whale kinematics are independent of prey patch characteristics. To test this hypothesis, we investigated the foraging performance of blue whales, the largest filter‐feeding predator and their functional response to variability in their sole prey source, krill using a generalized additive mixed model framework. We used a combination of animal‐borne movement sensors and hydroacoustic prey mapping to simultaneously quantify the three‐dimensional foraging kinematics of blue whales (Balaenoptera musculus) and the characteristics of targeted krill patches. Our analyses rejected our null hypothesis, showing that blue whales performed more acrobatic manoeuvres, including 180° and 360° rolling lunges, when foraging on low‐density krill patches. In contrast, whales targeting high‐density krill patches involved less manoeuvring during lunges and higher lunge feeding rates. These data demonstrate that blue whales exhibit a range of adaptive foraging strategies that maximize prey capture in different ecological contexts. Because first principles indicate that manoeuvres require more energy compared with straight trajectories, our data reveal a previously unrecognized level of complexity in predator–prey interactions that are not accounted for in optimal foraging and energetic efficiency models. Lay Summary

Among tremendous biodiversity within the California Current Ecosystem (CCE) are gigantic mysticetes (baleen whales) that produce structured sequences of sound described as song. From six years of passive acoustic monitoring within the central CCE we measured seasonal and interannual variations in the occurrence of blue ( Balaenoptera musculus ), fin ( Balaenoptera physalus ), and humpback ( Megaptera novaeangliae ) whale song. Song detection during 11 months of the year defines its prevalence in this foraging habitat and its potential use in behavioral ecology research. Large interannual changes in song occurrence within and between species motivates examination of causality. Humpback whales uniquely exhibited continuous interannual increases, rising from 34% to 76% of days over six years, and we examine multiple hypotheses to explain this exceptional trend. Potential influences of physical factors on detectability – including masking and acoustic propagation – were not supported by analysis of wind data or modeling of acoustic transmission loss. Potential influences of changes in local population abundance, site fidelity, or migration timing were supported for two of the interannual increases in song detection, based on extensive local photo ID data (17,356 IDs of 2,407 individuals). Potential influences of changes in foraging ecology and efficiency were supported across all years by analyses of the abundance and composition of forage species. Following detrimental food web impacts of a major marine heatwave that peaked during the first year of the study, foraging conditions consistently improved for humpback whales in the context of their exceptional prey-switching capacity. Stable isotope data from humpback and blue whale biopsy samples are consistent with observed interannual variations in the regional abundance and composition of forage species. This study thus indicates that major interannual changes in detection of baleen whale song may reflect underlying variations in forage species availability driven by energetic variations in ecosystem state.