Rorqual Lunge-Feeding Energetics Near and Away from the Kinematic Threshold of Optimal Efficiency

Humpback and blue whales are large baleen-bearing cetaceans which use a unique prey-acquisition strategy - lunge feeding - to engulf entire patches of large plankton or schools of forage fish and the water in which they are embedded. Dynamically, and while foraging on krill, lunge-feeding incurs metabolic expenditures estimated at up to 20.0 MJ. Because of prey abundance and its capture in bulk, lunge feeding is carried out at high acquired-to-expended energy ratios of up to 30 at the largest body sizes (∼27m). We use bio-logging tag data and the work-energy theorem to show that when krill-feeding at depth while using a wide range of prey approach swimming speeds (2 to 5m/s), rorquals generate significant and widely varying metabolic power output during engulfment, typically ranging from 10 to 50 times the basal metabolic rate of land mammals. At equal prey field density, such output variations lower their feeding efficiency 2- to 3-fold at high foraging speeds, thereby allowing slow and smaller rorquals to feed more efficiently than fast and larger rorquals. The analysis also shows how the slowest speeds of harvest so far measured may be connected to the biomechanics of the buccal cavity and the prey’s ability to collectively avoid engulfment. Such minimal speeds are important as they generate the most efficient lunges.

Insight into the kinematics of blue whale surface foraging through drone observations and prey data

To understand how predators optimize foraging strategies, extensive knowledge of predator behavior and prey distribution is needed. Blue whales employ an energetically demanding lunge feeding method that requires the whales to selectively feed where energetic gain exceeds energetic loss, while also balancing oxygen consumption, breath holding capacity, and surface recuperation time. Hence, blue whale foraging behavior is primarily driven by krill patch density and depth, but many studies have not fully considered surface feeding as a significant foraging strategy in energetic models. We collected predator and prey data on a blue whale (Balaenoptera musculus brevicauda) foraging ground in New Zealand in February 2017 to assess the distributional and behavioral response of blue whales to the distribution and density of krill prey aggregations. Krill density across the study region was greater toward the surface (upper 20 m), and blue whales were encountered where prey was relatively shallow and more dense. This relationship was particularly evident where foraging and surface lunge feeding were observed. Furthermore, New Zealand blue whales also had relatively short dive times (2.83 ± 0.27 SE min) as compared to other blue whale populations, which became even shorter at foraging sightings and where surface lunge feeding was observed. Using an unmanned aerial system (UAS; drone) we also captured unique video of a New Zealand blue whale’s surface feeding behavior on well-illuminated krill patches. Video analysis illustrates the whale’s potential use of vision to target prey, make foraging decisions, and orient body mechanics relative to prey patch characteristics. Kinematic analysis of a surface lunge feeding event revealed biomechanical coordination through speed, acceleration, head inclination, roll, and distance from krill patch to maximize prey engulfment. We compared these lunge kinematics to data previously reported from tagged blue whale lunges at depth to demonstrate strong similarities, and provide rare measurements of gape size, and krill response distance and time. These findings elucidate the predator-prey relationship between blue whales and krill, and provide support for the hypothesis that surface feeding by New Zealand blue whales is an important component to their foraging ecology used to optimize their energetic efficiency. Understanding how blue whales make foraging decisions presents logistical challenges, which may cause incomplete sampling and biased ecological knowledge if portions of their foraging behavior are undocumented. We conclude that surface foraging could be an important strategy for blue whales, and integration of UAS with tag-based studies may expand our understanding of their foraging ecology by examining surface feeding events in conjunction with behaviors at depth.

Extreme bradycardia and tachycardia in the world’s largest animal

The biology of the blue whale has long fascinated physiologists because of the animal’s extreme size. Despite high energetic demands from a large body, low mass-specific metabolic rates are likely powered by low heart rates. Diving bradycardia should slow blood oxygen depletion and enhance dive time available for foraging at depth. However, blue whales exhibit a high-cost feeding mechanism, lunge feeding, whereby large volumes of prey-laden water are intermittently engulfed and filtered during dives. This paradox of such a large, slowly beating heart and the high cost of lunge feeding represents a unique test of our understanding of cardiac function, hemodynamics, and physiological limits to body size. Here, we used an electrocardiogram (ECG)-depth recorder tag to measure blue whale heart rates during foraging dives as deep as 184 m and as long as 16.5 min. Heart rates during dives were typically 4 to 8 beats min−1 (bpm) and as low as 2 bpm, while after-dive surface heart rates were 25 to 37 bpm, near the estimated maximum heart rate possible. Despite extreme bradycardia, we recorded a 2.5-fold increase above diving heart rate minima during the powered ascent phase of feeding lunges followed by a gradual decrease of heart rate during the prolonged glide as engulfed water is filtered. These heart rate dynamics explain the unique hemodynamic design in rorqual whales consisting of a large-diameter, highly compliant, elastic aortic arch that allows the aorta to accommodate blood ejected by the heart and maintain blood flow during the long and variable pauses between heartbeats.

Sighting rates and prey of Minke Whales (Balaenoptera acutorostrata) and other cetaceans off Cormorant Island, British Columbia

From June to August 2012, we conducted over 500 h of visual surveys from Cormorant Island, British Columbia, to determine behaviour and habitat use patterns of nearby cetaceans. Seven species were documented, but Minke Whales (Balaenoptera acutorostrata) were by far the most common and were observed lunge feeding at the surface on 15 occasions. In addition, this species was documented surface lunge feeding on Pacific Herring (Clupea pallasi) and Pacific Sand Lance (Ammodytes personatus) on 32 occasions during vessel-based cetacean surveys around Cormorant Island between 2010 and 2014. Although Minke Whales are relatively uncommon in British Columbia, these results indicate that they can regularly be found in specific feeding areas during the summer.

Lunge Feeding in Rorqual Whales

The largest animals are baleen filter feeders that exploit large aggregations of small-bodied plankton. Although this feeding mechanism has evolved multiple times in marine vertebrates, rorqual whales exhibit a distinct lunge filter feeding mode that requires extreme physiological adaptations—most of which remain poorly understood. Here, we review the biomechanics of the lunge feeding mechanism in rorqual whales that underlies their extraordinary foraging performance and gigantic body size.

A new genus and species of eomysticetid (Cetacea: Mysticeti) and a reinterpretation of ‘Mauicetus’ lophocephalus Marples, 1956: transitional baleen whales from the upper Oligocene of New Zealand

The early evolution of toothless baleen whales (Chaeomysticeti) remains elusive despite a robust record of Eocene-Oligocene archaeocetes and toothed mysticetes. Eomysticetids, a group of archaic longirostrine and putatively toothless baleen whales fill in a crucial morphological gap between well-known toothed mysticetes and more crownward Neogene Mysticeti. A historically important but perplexing cetacean is “Mauicetus” lophocephalus (upper Oligocene South Island, New Zealand). The discovery of new skulls and skeletons of eomysticetids from the Oligocene Kokoamu Greensand and Otekaike Limestone permit a redescription and modern reinterpretation of “Mauicetus” lophocephalus, and indicating that this species may have retained adult teeth. A new genus and species, Tokarahia kauaeroa, is erected on the basis of a well-preserved subadult to adult skull with mandibles, tympanoperiotics, and cervical and thoracic vertebrae, ribs, sternum, and forelimbs from the Otekaike Limestone (>25.2 Ma). “Mauicetus” lophocephalus is relatively similar and recombined as Tokarahia lophocephalus. Phylogenetic analysis supports inclusion of Tokarahia within the Eomysticetidae alongside Eomysticetus, Micromysticetus, Yamatocetus, and Tohoraata, and strongly supports monophyly of Eomysticetidae. Tokarahia lacked extreme rostral kinesis of extant Mysticeti and primitively retained a delicate archaeocete-like posterior mandible and synovial temporomandibular joint, suggesting that Tokarahia was capable of at most, limited lunge feeding in contrast to extant Balaenopteridae, and utilized an alternative as-yet unspecified feeding strategy.

A new Miocene baleen whale from the Peruvian desert

The Pisco-Ica and Sacaco basins of southern Peru are renowned for their abundance of exceptionally preserved fossil cetaceans, several of which retain traces of soft tissue and occasionally even stomach contents. Previous work has mostly focused on odontocetes, with baleen whales currently being restricted to just three described taxa. Here, we report a new Late Miocene rorqual (family Balaenopteridae), Incakujira anillodefuego gen. et sp. nov., based on two exceptionally preserved specimens from the Pisco Formation exposed at Aguada de Lomas, Sacaco Basin, southern Peru. Incakujira overall closely resembles modern balaenopterids, but stands out for its unusually gracile ascending process of the maxilla, as well as a markedly twisted postglenoid process of the squamosal. The latter likely impeded lateral (omega) rotation of the mandible, in stark contrast with the highly flexible craniomandibular joint of extant lunge-feeding rorquals. Overall, Incakujira expands the still meagre Miocene record of balaenopterids and reveals a previously underappreciated degree of complexity in the evolution of their iconic lunge-feeding strategy.