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.
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•Tags with both video and 3D accelerometry were deployed on feeding rorquals•Skull movement could be observed in concert with animal orientation and motion•Lunging whales displayed prey-dependent inter- and intra-species kinematic diversity•Humpback whales most likely sacrifice energy efficiency to increase foraging flexibility
Rorqual lunge feeding has been assumed to be kinematically similar across species. Cade et al. use new video and accelerometry tags to show that blue whales, krill specialists, and humpback whales, foraging generalists, have similarly stereotypical lunges on krill—but humpbacks feeding on fish demonstrate kinematic diversity in lunge timing.
Baleen whales influence their ecosystems through immense prey consumption and nutrient recycling
. 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
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
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
, and more than twice the global catch of marine fisheries today
. Larger whale populations may have supported higher productivity in large marine regions through enhanced nutrient recycling: our findings suggest mysticetes recycled 1.2 × 10
tonnes iron yr
in the Southern Ocean before whaling compared to 1.2 × 10
tonnes iron yr
recycled by whales today. The recovery of baleen whales and their nutrient recycling services
could augment productivity and restore ecosystem function lost during 20th century whaling
.
Ecological relationships of krill and whales have not been explored in the Western Antarctic Peninsula (WAP), and have only rarely been studied elsewhere in the Southern Ocean. In the austral autumn ...we observed an extremely high density (5.1 whales per km(2)) of humpback whales (Megaptera novaeangliae) feeding on a super-aggregation of Antarctic krill (Euphausia superba) in Wilhelmina Bay. The krill biomass was approximately 2 million tons, distributed over an area of 100 km(2) at densities of up to 2000 individuals m(-3); reports of such 'super-aggregations' of krill have been absent in the scientific literature for >20 years. Retentive circulation patterns in the Bay entrained phytoplankton and meso-zooplankton that were grazed by the krill. Tagged whales rested during daylight hours and fed intensively throughout the night as krill migrated toward the surface. We infer that the previously unstudied WAP embayments are important foraging areas for whales during autumn and, furthermore, that meso-scale variation in the distribution of whales and their prey are important features of this system. Recent decreases in the abundance of Antarctic krill around the WAP have been linked to reductions in sea ice, mediated by rapid climate change in this area. At the same time, baleen whale populations in the Southern Ocean, which feed primarily on krill, are recovering from past exploitation. Consideration of these features and the effects of climate change on krill dynamics are critical to managing both krill harvests and the recovery of baleen whales in the Southern Ocean.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
The seasonal movement of animals at broad spatial scales provides insight into life-history, ecology and conservation. By combining high-resolution satellite-tagged data with hierarchical Bayesian ...movement models, we can associate spatial patterns of movement with marine animal behavior. We used a multi-state mixture model to describe humpback whale traveling and area-restricted search states as they forage along the West Antarctic Peninsula. We estimated the change in the geography, composition and characteristics of these behavioral states through time. We show that whales later in the austral fall spent more time in movements associated with foraging, traveled at lower speeds between foraging areas, and shifted their distribution northward and inshore. Seasonal changes in movement are likely due to a combination of sea ice advance and regional shifts in the primary prey source. Our study is a step towards dynamic movement models in the marine environment at broad scales.
Many baleen whales undertake annual fasting and feeding cycles, resulting in substantial changes in their body condition, an important factor affecting fitness. As a measure of lipid-store body ...condition, tissue density of a few deep diving marine mammals has been estimated using a hydrodynamic glide model of drag and buoyancy forces. Here, we applied the method to shallow-diving humpback whales (Megaptera novaeangliae) in North Atlantic and Antarctic feeding aggregations. High-resolution 3-axis acceleration, depth and speed data were collected from 24 whales. Measured values of acceleration during 5 s glides were fitted to a hydrodynamic glide model to estimate unknown parameters (tissue density, drag term and diving gas volume) in a Bayesian framework. Estimated species-average tissue density (1031.6 ± 2.1 kg m-3, ±95% credible interval) indicates that humpback whale tissue is typically negatively buoyant although there was a large inter-individual variation ranging from 1025.2 to 1043.1 kg m-3. The precision of the individual estimates was substantially finer than the variation across different individual whales, demonstrating a progressive decrease in tissue density throughout the feeding season and comparably high lipid-store in pregnant females. The drag term (CDAm-1) was estimated to be relatively high, indicating a large effect of lift-related induced drag for humpback whales. Our results show that tissue density of shallow diving baleen whales can be estimated using the hydrodynamic gliding model, although cross-validation with other techniques is an essential next step. This method for estimating body condition is likely to be broadly applicable across a range of aquatic animals and environments.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
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
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.
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.
Celotno besedilo
Dostopno za:
BFBNIB, DOBA, IZUM, KILJ, NMLJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
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.
Animal behaviour can provide valuable information for wildlife management and conservation. Studying the detailed behaviour of marine mammals involves challenges not faced by most animal behaviour ...researchers due to the size, mobility and lack of continuous visibility of these animals. We describe several methods developed by marine mammal scientists to study behaviour, primarily of cetaceans, focusing on technological advances: unmanned aerial systems (UAS), satellite-linked telemetry, passive acoustics and multisensor high-resolution acoustic recording tags. We then go on to explain how the data collected by these methods have contributed to and informed conservation actions. We focus on examples including: satellite data informing the interactions between cetaceans and offshore oil and gas development; passive acoustics used to track distributions of several species of cetaceans, including their movements near shipping lanes; and high-resolution acoustic recording tags used to document responses of cetaceans to anthropogenic activities. Finally, we discuss recent efforts to link animal behaviour to individual fitness and, particularly for behavioural disturbances, to population-level consequences, which can be helpful for informing conservation efforts. The infusion of technological advancements into studies of cetacean behaviour combined with emerging analytical techniques brings us to the next 20+ years of studying these animals. These developments will improve our capabilities in areas such as testing whether their behaviour adheres to traditional behavioural theory, and will certainly assist the guiding of conservation efforts.
•Technological advances provide new insights into marine mammal behaviour.•We highlight four areas of technological advances.•Aerial systems, radiotelemetry, passive acoustics and acoustic recording tags.•These methods allow comparison between marine mammal behaviour and theory.•These crucial data are helping to inform conservation applications.
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.
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