Shallow Dives Research Articles

Acoustic wave propagation in depth-evolving sound-speed field using the lattice Boltzmann method

This study investigates the propagation of sound waves within deep-sea low-sound-speed channels using the lattice Boltzmann method, with a key focus on the influence of depth-dependent sound speed on wave propagation. The depth-variable sound speed condition is realized through the incorporation of an external force proportional to the density gradient. After the model verification, investigations into the two-dimensional spreading of sound sources reveal that the depth-dependent sound speed curves the wave propagation. When source depths differing from the low-sound-speed channel, wave paths deviate due to contrasting speeds above and below. When the sound source is situated within the low-sound-speed channel, waves exhibit converging patterns. The simulations also detail the total reflection behavior of sound waves. When the incident angle falls exceeds the critical angle, the waves remain intact within the low-sound-speed channel, thereby enabling the preservation of high amplitude acoustic signals even at remote locations. The subsequent simulations of sound wave propagation around obstacles demonstrate that the low-sound-speed channel also exhibits better signal transmission capabilities in the presence of obstacles. In a uniform sound speed environment, acoustic wave propagation around a submarine exhibits a symmetric pattern. By contrast, under depth-evolving speed conditions, submarines operating at various depths manifest distinct propagation characteristics, such as asymmetric wave propagation during shallow diving, as well as wave attenuation or even silencing when cruising within low-sound-speed channels. These findings underscore the profound implications of depth-evolving sound speed on underwater acoustic signal detection and transmission.

Bubble blasts! An adaptation for buoyancy regulation in shallow foraging gray whales.

Foraging efficiency is key to animal fitness. Consequently, animals evolved a variety of kinematic, morphological, physiological, and behavioral adaptations for efficient locomotion to reduce energy expenditure while moving to find, capture, and consume prey. Often suited to specific habitat and prey types, these adaptations correspond to the terrain or substrate the animal moves through. In aquatic systems, adaptations focus on overcoming drag, buoyancy, and hydrostatic forces. Buoyancy both benefits and hinders diving animals; in particular, shallow divers constantly contend with the costs of overcoming buoyancy to dive and maintain position. Pacific Coast Feeding Group (PCFG) gray whales forage in shallow habitats where they work against buoyancy to dive and feed using various foraging tactics. Bubble blasts (underwater exhalations) have been observed during several foraging tactics performed by PCFG whales. As exhalations aid buoyancy regulation in other diving animals, we hypothesize that bubble blasts are performed by longer, more buoyant whales in shallower water and that bubble blasts increase dive duration while accounting for size and tactic. We test our hypotheses using Bayesian linear mixed effects models and a 7-year dataset of drone footage containing concurrent individual morphological and behavioral data. We find that while headstanding - a stationary, head-down tactic - bubble blasts are performed by longer, more buoyant whales and extend the dive duration, whereas whales using forward-swimming tactics are less likely to bubble blast. Our results suggest that PCFG gray whales may use bubble blasts as a behavioral adaption to mitigate the cost of energetically expensive tactics in their shallow habitat foraging niche.

Breath-Hold Diving Injuries - A Primer for Medical Providers.

Breath-hold divers, also known as freedivers, are at risk of specific injuries that are unique from those of surface swimmers and compressed air divers. Using peer-reviewed scientific research and expert opinion, we created a guide for medical providers managing breath-hold diving injuries in the field. Hypoxia induced by prolonged apnea and increased oxygen uptake can result in an impaired mental state that can manifest as involuntary movements or full loss of consciousness. Negative pressure barotrauma secondary to airspace collapse can lead to edema and/or hemorrhage. Positive pressure barotrauma secondary to overexpansion of airspaces can result in gas embolism or air entry into tissues and organs. Inert gas loading into tissues from prolonged deep dives or repetitive shallow dives with short surface intervals can lead to decompression sickness. Inert gas narcosis at depth is commonly described as an altered state similar to that experienced by compressed air divers. Asymptomatic cardiac arrhythmias are common during apnea, normally reversing shortly after normal ventilation resumes. The methods of glossopharyngeal breathing (insufflation and exsufflation) can add to the risk of pulmonary overinflation barotrauma or loss of consciousness from decreased cardiac preload. This guide also includes information for medical providers who are tasked with providing medical support at an organized breath-hold diving event with a list of suggested equipment to facilitate diagnosis and treatment outside of the hospital setting.

Osteological profiling of femoral diaphysis and neck in aquatic, semiaquatic, and terrestrial carnivores and rodents: effects of body size and locomotor habits

The increased limb bone density documented previously for aquatic tetrapods has been proposed to be an adaptation to overcome buoyancy during swimming and diving. It can be achieved by increasing the amount of bone deposition or by reducing the amount of bone resorption, leading to cortical thickening, loss of medullary cavity, and compaction of trabecular bone. The present study examined the effects of locomotor habit, body size, and phylogeny on the densitometric, cross-sectional, and biomechanical traits of femoral diaphysis and neck in terrestrial, semiaquatic, and aquatic carnivores, and in terrestrial and semiaquatic rodents (12 species) by using peripheral quantitative computed tomography, three-point bending, and femoral neck loading tests. Groupwise differences were analyzed with the univariate generalized linear model and the multivariate linear discriminant analysis supplemented with hierarchical clustering. While none of the individual features could separate the lifestyles or species adequately, the combinations of multiple features produced very good or excellent classifications and clusterings. In the phocid seals, the aquatic niche allowed for lower femoral bone mineral densities than expected based on the body mass alone. The semiaquatic mammals mostly had high bone mineral densities compared to the terrestrial species, which could be considered an adaptation to overcome buoyancy during swimming and shallow diving. Generally, it seems that different osteological properties at the levels of mineral density and biomechanics could be compatible with the adaptation to aquatic, semiaquatic, or terrestrial niches.

High‐resolution diving data collected from foraging area reveal that leatherback turtles dive faster to forage longer

AbstractDespite multiple studies examining the diving behavior of leatherback sea turtles (Dermochelys coriacea) at coarse resolution over broad distances, there is still a paucity of high‐resolution diving data collected in areas where foraging has been confirmed. Short‐term (~1–3 h) deployments of suction cup tags with time–depth recorders (TDRs) on 10 free‐swimming leatherback turtles in a foraging area off Nova Scotia, Canada during August and September (2007–2014), captured a total of 161 dives. High‐resolution (1–5 s sampling rate) dive profile data indicated variability in diving behavior between and within individuals. On average, turtles spent 55.7% of their time diving and 44.3% at the surface. Turtles generally performed short (mean duration = 250.4 s [SD = 47.9 s]) and shallow dives (mean depth = 24.3 m [SD = 5.8 m]). We recorded a mean dive descent rate of 0.32 m/s, which is faster than values recorded for leatherbacks in tropical waters. This may reflect differences in environment, behavioral mode (e.g., foraging vs. inter‐nesting), and body condition. Linear mixed‐effects models suggest a significant positive correlation between descent rate and mean depth, maximum depth, and integrated vertical bottom movement (IVBM). Turtles with faster descent rates dove deeper and increased their predicted foraging behavior (IVBM, or the sum of absolute differences in depth changes while at the bottom portion of their dives). Models additionally showed that dive time, bottom time, and IVBM were all positively correlated to the post‐dive surfacing. This suggests that turtles required more time at the surface to recover and/or handle prey following longer dives characterized by increased vertical movement at the bottom portion of the dive. Dives were complex; the application of standard dive type/shape analysis may be over‐simplified and inappropriate for leatherbacks foraging in these habitats. These results portray a novel and detailed look at the foraging dynamics of a diving marine reptile.

Unlocking the depths: multiple factors contribute to risk for hypoxic blackout during deep freediving

PurposeTo examine the effect of freediving depth on risk for hypoxic blackout by recording arterial oxygen saturation (SpO2) and heart rate (HR) during deep and shallow dives in the sea.MethodsFourteen competitive freedivers conducted open-water training dives wearing a water-/pressure proof pulse oximeter continuously recording HR and SpO2. Dives were divided into deep (> 35 m) and shallow (10–25 m) post-hoc and data from one deep and one shallow dive from 10 divers were compared.ResultsMean ± SD depth was 53 ± 14 m for deep and 17 ± 4 m for shallow dives. Respective dive durations (120 ± 18 s and 116 ± 43 s) did not differ. Deep dives resulted in lower minimum SpO2 (58 ± 17%) compared with shallow dives (74 ± 17%; P = 0.029). Overall diving HR was 7 bpm higher in deep dives (P = 0.002) although minimum HR was similar in both types of dives (39 bpm). Three divers desaturated early at depth, of which two exhibited severe hypoxia (SpO2 ≤ 65%) upon resurfacing. Additionally, four divers developed severe hypoxia after dives.ConclusionsDespite similar dive durations, oxygen desaturation was greater during deep dives, confirming increased risk of hypoxic blackout with increased depth. In addition to the rapid drop in alveolar pressure and oxygen uptake during ascent, several other risk factors associated with deep freediving were identified, including higher swimming effort and oxygen consumption, a compromised diving response, an autonomic conflict possibly causing arrhythmias, and compromised oxygen uptake at depth by lung compression possibly leading to atelectasis or pulmonary edema in some individuals. Individuals with elevated risk could likely be identified using wearable technology.

Surface and diving metabolic rates, and dynamic aerobic dive limits (dADL) in near‐ and off‐shore bottlenose dolphins, Tursiops spp., indicate that deep diving is energetically cheap

AbstractHigh‐resolution dive depth and acceleration recordings from nearshore (Sarasota Bay, dive depth < 30 m), and offshore (Bermuda) bottlenose dolphins (Tursiops spp.) were used to estimate the diving metabolic rate (DMR) and the locomotor metabolic rate (LMR, L O2/min) during three phases of diving (descent, bottom, and ascent). For shallow dives (depth ≤ 30 m), we found no differences between the two ecotypes in the LMR during diving, nor during the postdive shallow interval between dives. For intermediate (30 m < depth ≤ 100 m) and deep dives (depth > 100 m), the LMR was significantly higher during ascent than during descent and the bottom phase by 59% and 9%, respectively. In addition, the rate of change in depth during descent and ascent (meters/second) increased with maximal dive depth. The dynamic aerobic dive limit (dADL) was calculated from the estimated DMR and the estimated predive O2 stores. For the Bermuda dolphins, the dADL decreased with dive depth, and was 18.7, 15.4, and 11.1 min for shallow, intermediate, and deep dives, respectively. These results provide a useful approach to understand the complex nature of physiological interactions between aerobic metabolism, energy use, and diving capacity.

Elevations of Extracellular Vesicles and Inflammatory Biomarkers in Closed Circuit SCUBA Divers.

Blood-borne extracellular vesicles and inflammatory mediators were evaluated in divers using a closed circuit rebreathing apparatus and custom-mixed gases to diminish some diving risks. "Deep" divers (n = 8) dove once to mean (±SD) 102.5 ± 1.2 m of sea water (msw) for 167.3 ± 11.5 min. "Shallow" divers (n = 6) dove 3 times on day 1, and then repetitively over 7 days to 16.4 ± 3.7 msw, for 49.9 ± 11.9 min. There were statistically significant elevations of microparticles (MPs) in deep divers (day 1) and shallow divers at day 7 that expressed proteins specific to microglia, neutrophils, platelets, and endothelial cells, as well as thrombospondin (TSP)-1 and filamentous (F-) actin. Intra-MP IL-1β increased by 7.5-fold (p < 0.001) after day 1 and 41-fold (p = 0.003) at day 7. Intra-MP nitric oxide synthase-2 (NOS2) increased 17-fold (p < 0.001) after day 1 and 19-fold (p = 0.002) at day 7. Plasma gelsolin (pGSN) levels decreased by 73% (p < 0.001) in deep divers (day 1) and 37% in shallow divers by day 7. Plasma samples containing exosomes and other lipophilic particles increased from 186% to 490% among the divers but contained no IL-1β or NOS2. We conclude that diving triggers inflammatory events, even when controlling for hyperoxia, and many are not proportional to the depth of diving.

High dive efficiency in shallow water

Dive studies across mammals, birds, reptiles and fish often focus on deep dives, and shallow water diving has tended to be overlooked. For air-breathers, foraging in shallow water poses challenges since the lungs generate buoyancy, and shallow divers must trade off the extent of inhalation against the negative buoyancy needed to avoid floating to the surface. Using high-resolution depth loggers, we addressed this knowledge gap around the ecology of shallow water diving at a foraging site for hawksbill turtles (Eretmochelys imbricata) where depth was typically < 3 m. Contrary to predictions, dive durations were long, particularly at night (mean dive duration per turtle: 17–61 min, n = 12 turtles, n = 2576 nocturnal dives), despite warm water temperatures (24–37 °C). Dive efficiency (% time submerged) for hawksbills was 98%, the highest recorded for any air-breathing marine vertebrate including penguins (60–78%), seals (51–91%), cetaceans (68–87%), and other sea turtle species (68–95%). Hawksbills usually dive for much longer (42–286% increase) than green and loggerhead turtles when depth and temperature are accounted for. Hawksbill turtles likely forage in very shallow water to reduce predation risk from sharks: of 423 hawksbills captured by hand, none had any evidence of shark attack, although large sharks were present in nearby deeper water. Our results challenge the prediction that shallow water dives by air-breathers will usually be short and open the way for comparative studies of the ecology of shallow water diving in a range of other taxa. Our work emphasises the likely importance of predation risk in shaping patterns of habitat utilisation.

Stay the course: maintenance of consistent orientation by commuting penguins both underwater and at the water surface

Many marine vertebrates traverse more than hundreds of kilometres of the ocean. To efficiently achieve such long-distance movements, the ability to maintain orientation in a three-dimensional space is essential; however, it remains unevaluated in most species. In this study, we examined the bearing distributions of penguins undertaking long-distance foraging trips and compared their bearing consistency between underwater and at the water surface, as well as between night and day, to quantify their orientation ability. The subject species, king penguins, Aptenodytes patagonicus, from Possession Island, Crozet archipelago (46°25′S, 51°45′E; January to March 2011), showed high bearing consistency both during dives and at the water surface whilst commuting towards/from their main foraging area, the Antarctic polar front. Their bearing consistency was particularly high during and after shallow dives, irrespective of the time of day. Meanwhile, their bearings tended to vary during and after deep dives, particularly in the middle of the trip, probably owing to underwater foraging movements. However, the overall directions of deep dives during the commuting phases were similar to those of shallow dives and post-dive periods at the water surface. These findings indicate that king penguins employ compass mechanism(s) that are equivalently reliable both underwater and at the water surface, at any time of the day. This orientation ability appears to enable them to achieve long-distance trips under strong temporal constraints. Further studies on the fine-scale bearing distributions of other diving vertebrates are needed to better understand movement strategies in marine environments.

Case report: tracking data from foraging hawksbill turtles in the northern Red Sea

BackgroundHawksbill turtles (Eretmochelys imbricata) are Critically Endangered throughout their global range, and concerningly little is known about this species in the Red Sea. With large-scale coastal development projects underway in the northern Red Sea, it is critical to understand the movement and habitat use patterns of hawksbill turtles in this environmentally unique region, so that effective conservation strategies can be implemented. We satellite tagged three hawksbill turtles, one 63 cm curved carapace length adult male captured near Wahlei Island, one 55 cm turtle captured in the Gulf of Aqaba, and one 56 cm turtle suffering from a floating syndrome which was captured at Waqqadi Island, rehabilitated, and released at Waqqadi Island. Turtles were tracked for 156, 199, and 372 days between October 2020 and November 2021.ResultsWe calculated the home ranges and core use areas of hawksbill turtles using kernel-density estimations and found that each turtle showed high fidelity to their foraging sites. Home ranges calculated with GPS-derived locations ranged between 13.6 and 2.86 km2, whereas home ranges calculated with Argos-derived locations ranged from 38.98 to 286.45 km2. GPS-derived locations also revealed a higher proportion of time spent in coral and rock habitats compared to Argos, based on location overlap with the Allen Coral Reef Atlas. We also found that turtles were making shallow dives, usually remaining between 0 and 5 m.ConclusionsWhile the number of tracked turtles in this study was small, it represents an important contribution to the current understanding of spatial ecology among foraging hawksbill turtles globally, and provides the first-ever reported hawksbill turtle tracking data from the Red Sea. Our results suggest that protecting coral reef habitats and implementing boating speed limits near reefs could be effective conservation measures for foraging hawksbill turtles in the face of rapid coastal development.

Acoustic sampling of Antarctic krill with simulated underwater buoyancy gliders: Does the sawtooth dive pattern work?

Autonomous underwater gliders may be viable adjuncts to or in some cases replacements for ship-based oceanographic sampling. Gliders and ships acoustically sample the water column differently, with ships sampling all depths simultaneously in a single vertical pulse and gliders sampling shorter vertical segments of the water column in an up and down, sawtooth pattern. We simulated gliders following this flight pattern to sample the densities at depth of Antarctic krill (Euphausia superba), a patchily-distributed crustacean that is targeted by an international fishery. Krill densities from ship-based surveys conducted between 2001 to 2011 were treated as the “true” population densities sampled by the simulated gliders. Depth-integrated densities estimated from the glider sampling were compared to the population densities for each year. Coverage probabilities (the proportion of population means within a standard deviation of the glider sample means) for gliders diving to 150 m were near 100% in most years, better than the nominal 68%. Gliders diving to a maximum depth of 150 m estimated the annual population means better than gliders diving deeper because shallow dives provided more samples for a given length of trackline. Modeling the zero and non-zero data as separate distributions (the delta approach), an alternative to the lognormal CV approach used in this study, resulted in less accurate estimates of krill population densities. These results suggest that the sawtooth flight pattern of gliders can produce density estimates of krill comparable to the annual time series of density estimates from ship-based surveys. Gliders may also be useful to survey other patchily-distributed pelagic organisms.

Remarkable consistency of spinal cord microvasculature in highly adapted diving odontocetes.

Odontocetes are breath-hold divers with a suite of physiological, anatomical, and behavioral adaptations that are highly derived and vastly different from those of their terrestrial counterparts. Because of these adaptations for diving, odontocetes were originally thought to be exempt from the harms of nitrogen gas embolism while diving. However, recent studies have shown that these mammals may alter their dive behavior in response to anthropogenic sound, leading to the potential for nitrogen supersaturation and bubble formation which may cause decompression sickness in the central nervous system (CNS). We examined the degree of interface between blood, gases, and neural tissues in the spinal cord by quantifying its microvascular characteristics in five species of odontocetes (Tursiops truncatus, Delphinus delphis, Grampus griseus, Kogia breviceps, and Mesoplodon europaeus) and a model terrestrial species (the pig-Sus scrofa domesticus) for comparison. This approach allowed us to compare microvascular characteristics (microvascular density, branching, and diameter) at several positions (cervical, thoracic, and lumbar) along the spinal cord from odontocetes that are known to be either deep or shallow divers. We found no significant differences (p < 0.05 for all comparisons) in microvessel density (9.30-11.18%), microvessel branching (1.60-2.12 branches/vessel), or microvessel diameter (11.83-16.079µm) between odontocetes and the pig, or between deep and shallow diving odontocete species. This similarity of spinal cord microvasculature anatomy in several species of odontocetes as compared to the terrestrial mammal is in contrast to the wide array of remarkable physio-anatomical adaptations marine mammals have evolved within their circulatory system to cope with the physiological demands of diving. These results, and other studies on CNS lipids, indicate that the spinal cords of odontocetes do not have specialized features that might serve to protect them from Type II DCS.

Plasticity in the morphometrics and movements of an Antarctic apex predator, the leopard seal

Animals that display plasticity in behavioral, ecological, and morphological traits are better poised to cope with environmental disturbances. Here, we examined individual plasticity and intraspecific variation in the morphometrics, movement patterns, and dive behavior of an enigmatic apex predator, the leopard seal (Hydrurga leptonyx). Satellite/GPS tags and time-depth recorders were deployed on 22 leopard seals off the Western Antarctic Peninsula. Adult female leopard seals were significantly larger (454±59 kg) and longer (302±11 cm) than adult males (302±22 kg, 276±11 cm). As females were 50% larger than their male counterparts, leopard seals are therefore one of the most extreme examples of female-biased sexual size dimorphism in marine mammals. Female leopard seals also spent more time hauled-out on land and ice than males. In the austral spring/summer, three adult female leopard seals hauled-out on ice for 10+ days, which likely represent the first satellite tracks of parturition and lactation for the species. While we found sex-based differences in morphometrics and haul-out durations, other variables, including maximum distance traveled and dive parameters, did not vary by sex. Regardless of sex, some leopard seals remained in near-shore habitats, traveling less than 50 kilometers, while other leopard seals traveled up to 1,700 kilometers away from the tagging location. Overall, leopard seals were short (3.0±0.7 min) and shallow (29±8 m) divers. However, within this general pattern, some individual leopard seals primarily used short, shallow dives, while others switched between short, shallow dives and long, deep dives. We also recorded the single deepest and longest dive made by any leopard seal—1, 256 meters for 25 minutes. Together, our results showcased high plasticity among leopard seals tagged in a single location. These flexible behaviors and traits may offer leopard seals, an ice-associated apex predator, resilience to the rapidly changing Southern Ocean.

Foraging patterns vary with the degree of sociality among Common Loon (&lt;i&gt;Gavia immer&lt;/i&gt;) overwintering on a freshwater lake

Little is known about the behaviour of Common Loon (Gavia immer) during the critical overwintering period, let alone the behaviour of the small, but increasing number of loons that overwinter on freshwater lakes in North America. We examined the diurnal time-activity budgets of Common Loon overwintering on a large reservoir in northwest South Carolina between 2018 and 2020. Similar to previous studies of breeding individuals and individuals that overwinter in marine waters, loons (n = 132) overwintering on this reservoir spent most of their time (52%) foraging. However, we found distinct differences in the activity budgets of individuals associated with their degree of sociality. Solitary birds (individuals spending 0–20% of time within 20 m of conspecifics) spent significantly more time foraging than did those that were either loosely-social (&gt;20–&lt;70% of time within 20 m of conspecifics) or strongly-social (70–100% of time). Although solitary loons made as many foraging dives as social birds, their dives were much longer, likely reflecting dives for larger predatory fish. In contrast, social individuals made much shorter, shallower dives, often foraging on shallower baitfish that they appear to pursue to the water surface and consume collectively. Such findings add to our understanding of loon winter behaviour and raise interesting questions regarding social behaviour and the short- and long-term trade-offs associated with social foraging in this species.

Pulmonary Barotrauma in a BUD/S Candidate Following Shallow Dives Using the MK 25 Rebreather.

Pulmonary barotrauma of ascent is a well-recognized risk of compressed air diving in the civilian and military diving communities. Chest imaging is a beneficial adjunct to clinical evaluation in screening select individuals for fitness to dive, evaluating dive-related injuries, and safely returning individuals to diving duty. We present a case of a 26-year-old male U.S. Navy Ensign and Basic Underwater Demolition/SEAL (BUD/S) candidate who experienced pulmonary barotrauma following two shallow dives to a maximum depth of 18 feet of seawater using the MK-25 100% oxygen rebreather. A chest radiograph showed a left upper lobe peripheral wedge-shaped opacity abutting the pleural surface. A subsequent computerized tomography (CT) scan of the chest showed multifocal areas of peripheral pulmonary hemorrhage associated with small pneumatoceles. Two months after the diving injury, chest CT showed resolution of the pulmonary hemorrhage and pneumatoceles. Diving-related pulmonary barotrauma occurs most often secondary to breath-holding on ascent by inexperienced divers and is one of the most common diving injuries seen in BUD/S candidates. The risk of pulmonary barotrauma may be decreased through adequate training and thorough medical screening to include assessing for symptoms of infection before every dive. In cases of acute pulmonary barotrauma, chest radiographs may be used to screen for thoracic injury. Chest CT with inspiratory and expiratory sequences should be used to screen dive candidates on a case-by-case basis and to evaluate lung injury and predisposing pulmonary conditions following pulmonary barotrauma.

Sowerby's beaked whale biosonar and movement strategy indicate deep-sea foraging niche differentiation in mesoplodont whales.

ABSTRACTClosely related species are expected to diverge in foraging strategy, reflecting the evolutionary drive to optimize foraging performance. The most speciose cetacean genus, Mesoplodon, comprises beaked whales with little diversity in external morphology or diet, and overlapping distributions. Moreover, the few studied species of beaked whales (Ziphiidae) show very similar foraging styles with slow, energy-conserving movement during long, deep foraging dives. This raises the question of what factors drive their speciation. Using data from animal-attached tags and aerial imagery, we tested the hypothesis that two similar-sized mesoplodonts, Sowerby's (Mesoplodon bidens) and Blainville's (Mesoplodon densirostris) beaked whales, exploit a similar low-energy niche. We show that, compared with the low-energy strategist Blainville's beaked whale, Sowerby's beaked whale lives in the fast lane. While targeting a similar mesopelagic/bathypelagic foraging zone, they consistently swim and hunt faster, perform shorter deep dives, and echolocate at a faster rate with higher frequency clicks. Further, extensive near-surface travel between deep dives challenges the interpretation of beaked whale shallow inter-foraging dives as a management strategy for decompression sickness. The distinctively higher frequency echolocation clicks do not hold apparent foraging benefits. Instead, we argue that a high-speed foraging style influences dive duration and echolocation behaviour, enabling access to a distinct prey population. Our results demonstrate that beaked whales exploit a broader diversity of deep-sea foraging and energetic niches than hitherto suspected. The marked deviation of Sowerby's beaked whales from the typical ziphiid foraging strategy has potential implications for their response to anthropogenic sounds, which appears to be strongly behaviourally driven in other ziphiids.

Community composition and spatial energetics of mesopelagic fishes and squids in the eastern Bering Sea as influenced by habitat variables

Micronekton of the mesopelagic zone (200 m–1000 m depth) play a central role in most aspects of global marine ecology linking epipelagic and deep scattering layers as planktivores, carnivores and principal prey to both shallow diving and deep living apex predators. Yet, despite this critical role and conspicuous presence, little is known regarding most aspects of mesopelagic community ecology, including habitat associations at sub-basin spatial scales. Here we address several aspects of these data deficiencies in the eastern Bering Sea mesopelagic – a highly productive subarctic marine ecosystem home to multiple protected predator species and large commercial fisheries. Through 41 midwater trawls conducted at 250 m, 500 m and 1000 m depths during May of 1999 and 2000, and a series of generalized linear models and cluster analyses, we describe the influence of remotely sensed oceanographic habitat variables on the diel distribution, relative abundance and community composition of 30 species of mesopelagic fish and squid in 3 contrasting study areas. We then project the total energy available to predators at depth and across the eastern Bering Sea Basin based on the identified habitat constraints. Fishes from the families Bathylagidae and Myctophidae, and squids from the Gonatidae dominated the 72 species catch biomass (n = 225,000, 2,100 kg). The six most abundant species were equally divided between each of the three dominant families and are also the most highly represented mesopelagic species in eastern Bering Sea pelagic predator diets. Their association with surface habitat variables describe a complex and multifaceted relationship between physical/biological predictors and the total mesopelagic energy field during the spring bloom period in this ecosystem. Leuroglossus schmidti, Bathylagus pacificus, Stenobrachius leucopsaurus, S. nannochir, Gonatopsis borealis and Eogonatus tinro contrast in the suite of physical and biological variables that they respond to, and also in the character of their response. Although L. schmidti is of moderate caloric value, it expressed the greatest overall energy signature at all depths due to its overwhelming biomass, and response to the presence of specific, variable habitat conditions across extensive portions of the study area. Interannual variability in habitat variables forecasted a pronounced redistribution of dominant species and thereby total mesopelagic energy content – providing a potential mechanism for shifts in central-place predator habitat use, energy expenditure, and diet. The known relevance of mesopelagic species as prey and the energetic potential for apex predators foraging in the Basin illustrates a cautionary narrative regarding unregulated harvest of biota from this poorly understood and critically important oceanic zone.

Vertebral Trabecular Bone Mechanical Properties Vary Among Functional Groups of Cetaceans.

SynopsisSince their appearance in the fossil record 34 million years ago, modern cetaceans (dolphins, whales, and porpoises) have radiated into diverse habitats circumglobally, developing vast phenotypic variations among species. Traits such as skeletal morphology and ecologically linked behaviors denote swimming activity; trade-offs in flexibility and rigidity along the vertebral column determine patterns of caudal oscillation. Here, we categorized 10 species of cetaceans (families Delphinidae and Kogiidae; N = 21 animals) into functional groups based on vertebral centra morphology, swimming speeds, diving behavior, and inferred swimming patterns. We quantified trabecular bone mechanical properties (yield strength, apparent stiffness, and resilience) among functional groups and regions of the vertebral column (thoracic, lumbar, and caudal). We extracted 6 mm3 samples from vertebral bodies and tested them in compression in 3 orientations (rostrocaudal, dorsoventral, and mediolateral) at 2 mm min−1. Overall, bone from the pre-fluke/fluke boundary had the greatest yield strength and resilience, indicating that the greatest forces are translated to the tail during caudal oscillatory swimming. Group 1, composed of 5 shallow-diving delphinid species, had the greatest vertebral trabecular bone yield strength, apparent stiffness, and resilience of all functional groups. Conversely, Group 3, composed of 2 deep-diving kogiid species, had the least strong, stiff, and resilient bone, while Group 2 (3 deep-diving delphinid species) exhibited intermediate values. These data suggest that species that incorporate prolonged glides during deep descents in the water column actively swim less, and place relatively smaller loads on their vertebral columns, compared with species that execute shallower dives. We found that cetacean vertebral trabecular bone properties differed from the properties of terrestrial mammals; for every given bone strength, cetacean bone was less stiff by comparison. This relative lack of material rigidity within vertebral bone may be attributed to the non-weight-bearing locomotor modes of fully aquatic mammals.

Cuvier’s beaked whale foraging dives identified via machine learning using depth and triaxial acceleration

Knowledge of Cuvier’s beaked whale Ziphius cavirostris behavior has expanded through the utilization of animal-borne tags. However, many tag types do not record sound—thus preventing echolocation click detections to identify foraging—or have short deployments that sample a limited range of behaviors. As the quantity of such non-acoustic tag data grows, so too does the need for robust methods of detecting foraging from non-acoustic data. We used 692 dives from 5 sound-recording tags on Cuvier’s beaked whales in southern California, USA, to develop extreme gradient boosting tree models to detect foraging based on 1 Hz depth and 16 Hz triaxial acceleration data. We performed repeated 10-fold cross validation using classification accuracy to tune 500 models with randomly partitioned training and testing datasets. An average of 99.9 and 99.2% of training and testing dataset dives, respectively, were correctly classified across the 500 models. Dives without associated sound recordings (n = 2069 from 7 whales including 4 non-acoustic tags) were classified via a model that maximized training information using dive depth and duration, ascent and descent rates, bottom-phase average vertical speed, and roll circular variance during dive descents and bottom phases. Of all long, deep dives (conventionally assumed to include foraging), 2.4% were classified as non-foraging dives, while 0.3% of short, shallow dives were classified as foraging dives. Results confirm that conventional depth and/or duration classifiers provide reasonable estimates of longer-term foraging patterns. However, additional variables previously listed enhance foraging detections for unusual dives (notably non-foraging deep dives) for southern California Cuvier’s beaked whales.