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  1. Synopsis

    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.

     
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  2. Body condition is a crucial and indicative measure of an animal’s fitness, reflecting overall foraging success, habitat quality, and balance between energy intake and energetic investment toward growth, maintenance, and reproduction. Recently, drone-based photogrammetry has provided new opportunities to obtain body condition estimates of baleen whales in one, two or three dimensions (1D, 2D, and 3D, respectively) – a single width, a projected dorsal surface area, or a body volume measure, respectively. However, no study to date has yet compared variation among these methods and described how measurement uncertainty scales across these dimensions. This associated uncertainty may affect inference derived from these measurements, which can lead to misinterpretation of data, and lack of comparison across body condition measurements restricts comparison of results between studies. Here we develop a Bayesian statistical model using known-sized calibration objects to predict the length and width measurements of unknown-sized objects (e.g., a whale). We use the fitted model to predict and compare uncertainty associated with 1D, 2D, and 3D photogrammetry-based body condition measurements of blue, humpback, and Antarctic minke whales – three species of baleen whales with a range of body sizes. The model outputs a posterior predictive distribution of body condition measurements and allows for the construction of highest posterior density intervals to define measurement uncertainty. We find that uncertainty does not scale linearly across multi-dimensional measurements, with 2D and 3D uncertainty increasing by a factor of 1.45 and 1.76 compared to 1D, respectively. Each standardized body condition measurement is highly correlated with one another, yet 2D body area index (BAI) accounts for potential variation along the body for each species and was the most precise body condition metric. We hope this study will serve as a guide to help researchers select the most appropriate body condition measurement for their purposes and allow them to incorporate photogrammetric uncertainty associated with these measurements which, in turn, will facilitate comparison of results across studies. 
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  3. Abstract Bio-logging devices equipped with inertial measurement units—particularly accelerometers, magnetometers, and pressure sensors—have revolutionized our ability to study animals as necessary electronics have gotten smaller and more affordable over the last two decades. These animal-attached tags allow for fine scale determination of behavior in the absence of direct observation, particularly useful in the marine realm, where direct observation is often impossible, and recent devices can integrate more power hungry and sensitive instruments, such as hydrophones, cameras, and physiological sensors. To convert the raw voltages recorded by bio-logging sensors into biologically meaningful metrics of orientation (e.g., pitch, roll and heading), motion (e.g., speed, specific acceleration) and position (e.g., depth and spatial coordinates), we developed a series of MATLAB tools and online instructional tutorials. Our tools are adaptable for a variety of devices, though we focus specifically on the integration of video, audio, 3-axis accelerometers, 3-axis magnetometers, 3-axis gyroscopes, pressure, temperature, light and GPS data that are the standard outputs from Customized Animal Tracking Solutions (CATS) video tags. Our tools were developed and tested on cetacean data but are designed to be modular and adaptable for a variety of marine and terrestrial species. In this text, we describe how to use these tools, the theories and ideas behind their development, and ideas and additional tools for applying the outputs of the process to biological research. We additionally explore and address common errors that can occur during processing and discuss future applications. All code is provided open source and is designed to be useful to both novice and experienced programmers. 
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  4. Abstract Background Advances in biologging technology allow researchers access to previously unobservable behavioral states and movement patterns of marine animals. To relate behaviors with environmental variables, features must be evaluated at scales relevant to the animal or behavior. Remotely sensed environmental data, collected via satellites, often suffers from the effects of cloud cover and lacks the spatial or temporal resolution to adequately link with individual animal behaviors or behavioral bouts. This study establishes a new method for remotely and continuously quantifying surface ice concentration (SIC) at a scale relevant to individual whales using on-animal tag video data. Results Motion-sensing and video-recording suction cup tags were deployed on 7 Antarctic minke whales ( Balaenoptera bonaerensis ) around the Antarctic Peninsula in February and March of 2018. To compare the scale of camera-tag observations with satellite imagery, the area of view was simulated using camera-tag parameters. For expected conditions, we found the visible area maximum to be ~ 100m 2 which indicates that observations occur at an equivalent or finer scale than a single pixel of high-resolution visible spectrum satellite imagery. SIC was classified into one of six bins (0%, 1–20%, 21–40%, 41–60%, 61–80%, 81–100%) by two independent observers for the initial and final surfacing between dives. In the event of a disagreement, a third independent observer was introduced, and the median of the three observer’s values was used. Initial results ( n  = 6) show that Antarctic minke whales in the coastal bays of the Antarctic Peninsula spend 52% of their time in open water, and only 15% of their time in water with SIC greater than 20%. Over time, we find significant variation in observed SIC, indicating that Antarctic minke occupy an extremely dynamic environment. Sentinel-2 satellite-based approaches of sea ice assessment were not possible because of persistent cloud cover during the study period. Conclusion Tag-video offers a means to evaluate ice concentration at spatial and temporal scales relevant to the individual. Combined with information on underwater behavior, our ability to quantify SIC continuously at the scale of the animal will improve upon current remote sensing methods to understand the link between animal behavior and these dynamic environmental variables. 
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  5. null (Ed.)
    The largest animals are the rorquals, a group of whales which rapidly engulf large aggregations of small-bodied animals along with the water in which they are embedded, with the latter subsequently expulsed via filtration through baleen. Represented by species like the blue, fin, and humpback whales, rorquals can exist in a wide range of body lengths (8–30 m) and masses (4000–190,000 kg). When feeding on krill, kinematic data collected by whale-borne biologging sensors suggest that they first oscillate their flukes several times to accelerate towards their prey, followed by a coasting period with mouth agape as the prey-water mixture is engulfed in a process approximating a perfectly inelastic collision. These kinematic data, used along with momentum conservation and time-averages of a whale’s equation of motion, show the largest rorquals as generating significant body forces (10–40 kN) in order to set into forward motion enough engulfed water to at least double overall mass. Interestingly, a scaling analysis of these equations suggests significant reductions in the amount of body force generated per kilogram of body mass at the larger sizes. In other words, and in concert with the allometric growth of the buccal cavity, gigantism would involve smaller fractions of muscle mass to engulf greater volumes of water and prey, thereby imparting a greater efficiency to this unique feeding strategy. 
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  6. 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. 
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  7. 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.

     
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