Foraging behaviour and ecology of transient killer whales within a deep submarine canyon system.

Transient killer whales have been documented hunting marine mammals across a variety of habitats. However, relatively little has been reported about their predatory behaviours near deep submarine canyons and oceanic environments. We used a long-term database of sightings and encounters with these predators in and around the Monterey Submarine Canyon, California to describe foraging behaviour, diet, seasonal occurrence, and habitat use patterns. Transient killer whales belonging to the outer coast subpopulation were observed within the study area 261 times from 2006-2021. Occurrences, behaviours, and group sizes all varied seasonally, with more encounters occurring in the spring as grey whales migrated northward from their breeding and calving lagoons in Mexico (March-May). Groups of killer whales foraged exclusively in open water, with individuals within the groups following the contours of the submarine canyon as they searched for prey. Focal follows revealed that killer whales spent 51% of their time searching for prey (26% of their time along the shelf-break and upper slope of the canyon, and 25% in open water). The remainder of their time was spent pursuing prey (10%), feeding (23%), travelling (9%), socializing (6%), and resting (1%). Prey species during 87 observed predation events included California sea lions, grey whale calves, northern elephant seals, minke whales, common dolphins, Pacific white-sided dolphins, Dall's porpoise, harbour porpoise, harbour seals, and sea birds. The calculated kill rates (based on 270 hours of observing 50 predation events) were 0.26 California sea lions per killer whale over 24 hours, 0.11 grey whale calves, and 0.15 for all remaining prey species combined. These behavioural observations provide insights into predator-prey interactions among apex predators over submarine canyons and deep pelagic environments.

Otariid gammaherpesvirus 1 in South American fur seals (Arctocephalus australis) and a novel related herpesvirus in free-ranging South American sea lions (Otaria byronia): Prevalence and effects of age, sex, and sample type.

Otariid gammaherpesvirus 1 (OtGHV1) is associated with high rates of urogenital carcinoma in free-ranging California sea lions (Zalophus californianus; CSL), and until recently was reported only in the Northern Hemisphere. The objective of this study was to survey free-ranging South American sea lions (Otaria byronia; SASL) and South American fur seals (Arctocephalus australis: SAFS) in Punta San Juan, Peru for OtGHV1 and to determine prevalence characteristics. Twenty-one percent (14/67) of urogenital swabs collected over three years (2011, 2014, 2015) from live pinnipeds of both species tested positive with a pan-herpesvirus conventional PCR. Sequencing of SAFS amplicons revealed 100% homology to OtGHV1 at the DNA polymerase, glycoprotein B, and viral bcl2-like genes. Sequencing of SASL amplicons revealed a novel related virus, herein called Otariid gammaherpesvirus 8 (OtGHV8). For comparison of sample sites, urogenital, conjunctival, and oropharyngeal swabs collected from 136 live pinnipeds of both species at Punta San Juan between 2011-2018 were then assayed using quantitative PCR for a segment of the OtGHV1/8 DNA polymerase gene using a qPCR assay now determined to cross-react between the two viruses. In total, across both species, 38.6% (51/132) of urogenital swabs, 5.6% (4/71) of conjunctival swabs, and 1.1% (1/90) of oropharyngeal swabs were positive for OtGHV1/8, with SASL only positive on urogenital swabs. Results from SASL were complicated by the finding of OtGHV8, necessitating further study to determine prevalence of OtGHV1 versus OtGHV8 using an alternate assay. Results from SAFS suggest a potential relationship between OtGHV1 in SAFS and CSL. Though necropsy surveillance in SAFS is very limited, geographic patterns of OtGHV1-associated urogenital carcinoma in CSL and the tendency of herpesviruses to cause more detrimental disease in aberrant hosts suggests that it is possible that SAFS may be the definitive host of OtGHV1, which gives further insight into the diversity and phyogeography of this clade of related gammaherpesviruses.

Cardiorespiratory patterns of male South American sea lions (Otaria flavescens) resting on land.

The goal of this study was to characterize the cardiorespiratory patterns of male South American sea lions (SASLs, Otaria flavescens) resting on land. We recorded respiratory and heart rate (n = 360 individuals studied) by observing the nostrils, chest movements and the impact of the heart on the thoracic wall. The sea lions breathe apneustically with a pause on inspiration, representing 74% of the respiratory cycle. The mean breathing frequency was 3.2 ± 1.0 breaths min-1, with a breathing cycle presenting periods of bradypneas, tachypneas, and long-term post-inspiratory pauses. The normal heart rate (nHR) was 73.4 ± 14.5 beats min-1 and no significant differences were observed between age classes. All animals showed variability in HR in relation to respiratory phases (Inspiration: 101.2 ± 18.4 beats min-1; post-inspiratory pause: 73.4 ± 14.5 beats min-1; expiration: 64.6 ± 17.7 beats min-1), consistent with respiratory sinus arrhythmia (RSA). The mean HR (measured during all respiratory phases) was 79.9 ± 22.7 beats min-1, and was significantly different between age classes. The total duration of respiratory cycle, and duration of both inspiration and expiration, decreased with an increment in ambient temperature, with no variation in the pause duration. Heart rate during pause and expiration was significantly higher during high temperatures. Similar changes in cardiorespiratory patterns have been reported in other pinnipeds. Our results showed ontogenetic differences in development and typical variations with environmental and behavioral variables.

Ultrasound images of the ascending aorta of anesthetized northern fur seals and Steller sea lions confirm that the aortic bulb maintains continuous blood flow.

The increased size and enhanced compliance of the aortic bulb-the enlargement of the ascending aorta-are believed to maintain blood flow in pinnipeds during extended periods of diastole induced by diving bradycardia. The aortic bulb has been described ex vivo in several species of pinnipeds, but in vivo measurements are needed to investigate the relationship between structure and function. We obtained ultrasound images using electrocardiogram-gated transesophageal echocardiography during anesthesia and after atropine administration to assess the relationship between aortic bulb anatomy and cardiac function (heart rate, stroke volume, cardiac output) in northern fur seals (Callorhinus ursinus) and Steller sea lions (Eumetopias jubatus). We observed that the aortic bulb in northern fur seals and Steller sea lions expands during systole and recoils over the entire diastolic period indicating that blood flow is maintained throughout the entire cardiac cycle as expected. The stroke volumes we measured in the fur seals and sea lions fit the values predicted based on body size in mammals and did not change with increased heart rates, suggesting that greater stroke volumes are not needed for aortic bulb function. Overall, our results suggest that peripheral vasoconstriction during diving is sufficient to modulate the volume of blood in the aortic bulb to ensure that flow lasts over the entire diastolic period. These results indicate that the shift of blood into the aortic bulb of pinnipeds is a fundamental mechanism caused by vasoconstriction while diving, highlighting the importance of this unique anatomical adaptation.

A multistate Langevin diffusion for inferring behavior-specific habitat selection and utilization distributions.

The identification of important habitat and the behavior(s) associated with it is critical to conservation and place-based management decisions. Behavior also links life-history requirements and habitat use, which are key to understanding why animals use certain habitats. Animal population studies often use tracking data to quantify space use and habitat selection, but they typically either ignore movement behavior (e.g., foraging, migrating, nesting) or adopt a two-stage approach that can induce bias and fail to propagate uncertainty. We develop a habitat-driven Langevin diffusion for animals that exhibit distinct movement behavior states, thereby providing a novel single-stage statistical method for inferring behavior-specific habitat selection and utilization distributions in continuous time. Practitioners can customize, fit, assess, and simulate our integrated model using the provided R package. Simulation experiments demonstrated that the model worked well under a range of sampling scenarios as long as observations were of sufficient temporal resolution. Our simulations also demonstrated the importance of accounting for different behaviors and the misleading inferences that can result when these are ignored. We provide case studies using plains zebra (Equus quagga) and Steller sea lion (Eumetopias jubatus) telemetry data. In the zebra example, our model identified distinct "encamped" and "exploratory" states, where the encamped state was characterized by strong selection for grassland and avoidance of other vegetation types, which may represent selection for foraging resources. In the sea lion example, our model identified distinct movement behavior modes typically associated with this marine central-place forager and, unlike previous analyses, found foraging-type movements to be associated with steeper offshore slopes characteristic of the continental shelf, submarine canyons, and seamounts that are believed to enhance prey concentrations. This is the first single-stage approach for inferring behavior-specific habitat selection and utilization distributions from tracking data that can be readily implemented with user-friendly software. As certain behaviors are often more relevant to specific conservation or management objectives, practitioners can use our model to help inform the identification and prioritization of important habitats. Moreover, by linking individual-level movement behaviors to population-level spatial processes, the multistate Langevin diffusion can advance inferences at the intersection of population, movement, and landscape ecology.

Mass Mortality of Sea Lions Caused by Highly Pathogenic Avian Influenza A(H5N1) Virus.

We report a massive mortality of 5,224 sea lions (Otaria flavescens) in Peru that seemed to be associated with highly pathogenic avian influenza A(H5N1) virus infection. The transmission pathway may have been through the close contact of sea lions with infected wild birds. We recommend evaluating potential virus transmission among sea lions.

Mercury and selenium concentrations in lanugo of free-ranging California sea lions in the southern Gulf of California, Mexico.

Total mercury ([THg]) and selenium ([TSe]) concentrations were determined in California sea lion (Zalophus californianus) lanugo from the Gulf of California in 2021 and 2022. Relationships with sex, morphometrics, and year were evaluated. Following toxicological thresholds of concern for piscivorous mammals, most pups had a [THg] < 10 ppm, one pup (2021) had a [THg] > 20 ppm, no pups had a [THg] > 30 ppm. Females had significantly higher [TSe] than males; sex did not influence [THg]. [THg] and [TSe] in 2022 were significantly higher in the general population and male cohorts compared to 2021. Significant negative correlations were observed between [THg], [TSe], and morphometrics (2021). These results indicate that, compared to other pinniped species, regional California sea lions may have a decreased likelihood of experiencing Hg-related adverse health effects. Year-related changes in element concentrations suggest continued monitoring of this population to assess pinniped, environmental, and potentially, human health.

Survival improvements of marine mammals in zoological institutions mirror historical advances in human longevity.

An intense public debate has fuelled governmental bans on marine mammals held in zoological institutions. The debate rests on the assumption that survival in zoological institutions has been and remains lower than in the wild, albeit the scientific evidence in support of this notion is equivocal. Here, we used statistical methods previously applied to assess historical improvements in human lifespan and data on 8864 individuals of four marine mammal species (harbour seal, Phoca vitulina; California sea lion, Zalophus californianus; polar bear, Ursus maritimus; common bottlenose dolphin, Tursiops truncatus) held in zoos from 1829 to 2020. We found that life expectancy increased up to 3.40 times, and first-year mortality declined up to 31%, during the last century in zoos. Moreover, the life expectancy of animals in zoos is currently 1.65-3.55 times longer than their wild counterparts. Like humans, these improvements have occurred concurrently with advances in management practices, crucial for population welfare. Science-based decisions will help effective legislative changes and ensure better implementation of animal care.

Time-calibrated phylogeny and full mitogenome sequence of the Galapagos sea lion (Zalophus wollebaeki) from scat DNA.

Background: The Galapagos sea lion, Zalophus wollebaeki, is an endemic and endangered otariid, which is considered as a sentinel species of ecosystem dynamics in the Galapagos archipelago. Mitochondrial DNA is an important tool in phylogenetic and population genetic inference. In this work we use Illumina sequencing to complement the mitogenomic resources for Zalophus genus-the other two species employed Sanger sequencing-by a complete mitochondrial genome and a molecular clock of this species, which is not present in any case. Materials and Methods: We used DNA obtained from a fresh scat sample of a Galapagos sea lion and shotgun-sequenced it on the Illumina NextSeq platform. The obtained raw reads were processed using the GetOrganelle software to filter the mitochondrial Zalophus DNA reads (∼16% survive the filtration), assemble them, and set up a molecular clock. Results: From the obtained 3,511,116 raw reads, we were able to assemble a full mitogenome of a length of 16,676 bp, consisting of 13 protein-coding genes (PCGs), 22 transfer RNAs (tRNA), and two ribosomal RNAs (rRNA). A time-calibrated phylogeny confirmed the phylogenetic position of Z. wollebaeki in a clade with Z. californianus, and Z. japonicus, and sister to Z. californianus; as well as establishing the divergence time for Z. wollebaeki 0.65 million years ago. Our study illustrates the possibility of seamlessly sequencing full mitochondrial genomes from fresh scat samples of marine mammals.

SCSLnO-SqueezeNet: Sine Cosine-Sea Lion Optimization enabled SqueezeNet for intrusion detection in IoT.

Security and privacy are regarded as the greatest priority in any real-world smart ecosystem built on the Internet of Things (IoT) paradigm. In this study, a SqueezeNet model for IoT threat detection is built using Sine Cosine Sea Lion Optimization (SCSLnO). The Base Station (BS) carries out intrusion detection. The Hausdorff distance is used to determine which features are important. Using the SqueezeNet model, attack detection is carried out, and the network classifier is trained using SCSLnO, which is developed by combining the Sine Cosine Algorithm (SCA) with Sea Lion Optimization (SLnO). BoT-IoT and NSL-KDD datasets are used for the analysis. In comparison to existing approaches, PSO-KNN/SVM, Voting Ensemble Classifier, Deep NN, and Deep learning, the accuracy value produced by devised method for the BoT-IoT dataset is 10.75%, 8.45%, 6.36%, and 3.51% higher when the training percentage is 90.

Assessing of the use of proteins A, G, and chimeric protein AG to detect marine mammal immunoglobulins.

In recent years, there has been an increase in infectious diseases in marine mammals, including brucellosis, infections of morbillivirus, herpesvirus, and poxvirus. Several serological diagnostic methods, including enzyme-linked immunosorbent assays, immunofluorescence assays (ELISA), and western blotting, have been used to detect antibodies against pathogens in marine mammals. However, options for commercial secondary antibodies used to detect antibodies in marine mammals are limited; therefore, the use of proteins A, G, or chimeric protein AG may provide a suitable alternative. This study aimed to assess the use of proteins A, G, and chimeric protein AG to detect marine mammal immunoglobulins. Currently, there are no comparative studies on the use of proteins A, G, and chimeric protein AG for the detection of immunoglobulins in marine mammals. In this study, we used ten pinnipeds' species (Baikal seal, California sea lion, harbor seal, northern fur seal, ringed seal, South American fur seal, South American sea lion, spotted seal, Steller sea lion, and walrus) and five cetacean species (beluga whale, bottlenose dolphin, harbor porpoise, killer whale, and Pacific white-sided dolphin) and compare binding ability to proteins A, G, or chimeric protein AG by ELISA. The results revealed that the immunoglobulins from pinniped and cetacean species reacted more strongly to protein A than protein G. In addition, the immunoglobulins of pinnipeds and cetaceans showed a strong binding ability to chimeric protein AG. These results suggest that proteins A, G, and chimeric protein AG would be used to help further develop serological assays.

Auditory masking in odobenid and otariid carnivoresa).

As the only living species within the odobenid lineage of carnivores, walruses (Odobenus rosmarus) have no close relatives from which auditory information can be extrapolated. Sea lions and fur seals in the otariid lineage are the nearest evolutionary outgroup. To advance understanding of odobenid and otariid hearing, we conducted behavioral testing with two walruses and one California sea lion (Zalophus californianus). Detection thresholds for airborne sounds were measured from 0.08 to at least 16 kHz in ambient noise conditions and then re-measured in the presence of octave-band white masking noise. Walruses were more sensitive than the sea lion at lower frequencies and less sensitive at higher frequencies. Critical ratios for the walruses ranged from 20 dB at 0.2 kHz to 32 dB at 10 kHz, while critical ratios for the sea lion ranged from 16 dB at 0.2 kHz to 35 dB at 32 kHz. The masking values for these species are comparable to one another and to those of terrestrial carnivores, increasing by about 3 dB per octave with increasing frequency. Despite apparent differences in hearing range and sensitivity, odobenids and otariids have a similar ability to hear signals in noisy conditions.

Mass mortality event in South American sea lions (Otaria flavescens) correlated to highly pathogenic avian influenza (HPAI) H5N1 outbreak in Chile.

In Chile, since January 2023, a sudden and pronounced increase in strandings and mortality has been observed among South American (SA) sea lions (Otaria flavescens), prompting significant concern. Simultaneously, an outbreak of highly pathogenic avian influenza H5N1 (HPAIV H5N1) in avian species has emerged since December 2022. To investigate the cause of this unexpected mortality, we conducted a comprehensive epidemiological and pathologic study. One hundred sixty-nine SA sea lions were sampled to ascertain their HPAIV H5N1 status, and long-term stranding trends from 2009 to 2023 were analyzed. In addition, two animals were necropsied. Remarkably, a significant surge in SA sea lion strandings was observed initiating in January 2023 and peaking in June 2023, with a count of 4,545 stranded and deceased animals. Notably, this surge in mortality correlates geographically with HPAIV outbreaks affecting wild birds. Among 168 sampled SA sea lions, 34 (20%) tested positive for Influenza A virus, and 21 confirmed for HPAIV H5N1 2.3.4.4b clade in tracheal/rectal swab pools. Clinical and pathological evaluations of the two necropsied stranded sea lions revealed prevalent neurological and respiratory signs, including disorientation, tremors, ataxia, and paralysis, as well as acute dyspnea, tachypnea, profuse nasal secretion, and abdominal breathing. The lesions identified in necropsied animals aligned with observed clinical signs. Detection of the virus via immunohistochemistry (IHC) and real-time PCR in the brain and lungs affirmed the findings. The findings provide evidence between the mass mortality occurrences in SA sea lions and HPAIV, strongly indicating a causal relationship. Further studies are needed to better understand the pathogenesis and transmission.

Associations between urogenital carcinoma and DECA-BDE (BDE-209) among wild California Sea lions (Zalophus californianus) and Steller Sea lions (Eumetopias jubatus).

Urogenital carcinoma (UGC) is prevalent among California sea lions (Zalophus californianus), while less is known concerning UGC among Steller sea lions (Eumetopias jubatus). Our objective was to investigate associations between UGC and polybrominated diphenyl ethers (PBDEs) among both sea lion species. Twenty-nine California sea lions and 20 Steller sea lions were lethally removed from the Columbia River Basin, Oregon, USA between 2020 and 2021, under Section 120 of the Marine Mammal Protection Act. UGC was diagnosed through gross necropsy and histopathology. Forty PBDE congeners were analyzed in blubber, including BDE-209, a potential carcinogen. Twenty (69 %) California sea lions and one (5 %) Steller sea lion were diagnosed with UGC. All cases were identified as early stage UGC, aside from one California sea lion with more advanced stage UGC. Among California sea lions, associations between PBDEs and UGC were analyzed using logistic regression. In the adjusted model, BDE-209 (log2-transformed) was associated with increased odds of UGC [Odds Ratio (OR): 4.68, 95 % confidence interval: 1.04, 21.0, OR p-value = 0.044). This is the first study to report BDE-209 concentrations in sea lion blubber. The percentages of California and Steller sea lions diagnosed with UGC were higher than expected for wild (non-stranded) sea lions. Our results suggested blubber BDE-209 was potentially associated with UGC in California sea lions in the Columbia River Basin.

Detection and quantification of microplastic pollution in the endangered Galapagos sea lion.

Marine debris pollution poses a significant global threat to biodiversity, with plastics being the primary debris type found in oceans due to their low-cost production and high demand worldwide. Microplastics (MPs, <5 mm in size) are highly bioavailable to a wide range of marine taxa, including marine mammals, through direct and indirect ingestion routes (i.e., trophic transfer). Recently, MP pollution has been detected on the Galapagos Marine Reserve, so in this study we developed a baseline framework for MP pollution in the Galapagos sea lion (GSL, Zalophus wollebaeki) through scat-based analysis. We collected 180 GSL scat samples from the southeast region following strict quality assurance/quality control protocols to detect, quantify and characterize physical-chemical properties of MPs through visual observations and µFT-IR spectroscopy. We recovered 81 MPs of varying sizes and colors in 37 % of samples (n = 66/180), consisting mostly of fibers (69 %, x¯ = 0.31 ± 0.57 particles scat-1). The number of particles per gram of sample wet weight ranged from 0.02 to 0.22 (x¯ = 0.04 ± 0.05 particles scat wet g-1). El Malecón and Punta Pitt rookeries at San Cristobal Island had the highest number of MPs (x¯ = 0.67 ± 0.51 and 0.43 ± 0.41 particles scat-1, respectively), and blue-colored particles were the most common in all samples. We identified eleven polymers in 46 particles, consisting mostly of polypropylene-polyethylene copolymer, polypropylene, cellulose, polyethylene, and polyvinyl chloride. The textile, fishing, and packaging industries are likely significant sources of microfibers into this insular ecosystem. Our results suggest that the GSL is exposed to MPs due to anthropogenic contamination that is subsequently transferred through trophic processes. These findings provide an important baseline framework and insights for future research on MP pollution in the region, as well as for management actions that will contribute to the long-term conservation of the GSL.

Pleural mesothelioma in a California sea lion (Zalophus californianus).

A 25-year-old female California sea lion (Zalophus californianus) reared in an aquarium died following a history of anorexia, lethargy, abnormal protrusion of the skin, and oral respiration. At necropsy, multiple yellowish-white nodules with diameters of 0.1-0.5 cm were disseminated in the thoracic cavity and lungs. Histopathologically, the nodules were continuous with normal mesothelium and were characterized by the proliferation of spindle-shaped to polygonal neoplastic cells with prominent atypia. The neoplastic cells exhibited diffuse, strong staining for vimentin and partial, weak to moderate staining for cytokeratin AE1/AE3. Based on these findings, the lesions were diagnosed as pleural mesothelioma. This study reports the first case of pleural mesothelioma in California sea lion.

Input compensation of dolphin and sea lion auditory brainstem responses using frequency-modulated up-chirps.

Frequency-modulated "chirp" stimuli that offset cochlear dispersion (i.e., input compensation) have shown promise for increasing auditory brainstem response (ABR) amplitudes relative to traditional sound stimuli. To enhance ABR methods with marine mammal species known or suspected to have low ABR signal-to-noise ratios, the present study examined the effects of broadband chirp sweep rate and level on ABR amplitude in bottlenose dolphins and California sea lions. "Optimal" chirps were designed based on previous estimates of cochlear traveling wave speeds (using high-pass subtractive masking methods) in these species. Optimal chirps increased ABR peak amplitudes by compensating for cochlear dispersion; however, chirps with similar (or higher) frequency-modulation rates produced comparable results. The optimal chirps generally increased ABR amplitudes relative to noisebursts as threshold was approached, although this was more obvious when sound pressure level was used to equate stimulus levels (as opposed to total energy). Chirps provided progressively less ABR amplitude gain (relative to noisebursts) as stimulus level increased and produced smaller ABRs at the highest levels tested in dolphins. Although it was previously hypothesized that chirps would provide larger gains in sea lions than dolphins-due to the lower traveling wave speed in the former-no such pattern was observed.

Metal-specific biomagnification and trophic dilution in the coastal foodweb of the California sea lion (Zalophus californianus) off Bahía Magdalena, Mexico: The role of the benthic-pelagic foodweb in the trophic transfer of trace and toxic metals.

Trace metals concentrations along with stable isotopes ratios were measured in marine algae, sea grass, sponges, echinoderms, mollusks, crustaceans, fishes, and the California sea lion, to assess the bioaccumulation potential and detect potential risks for top predators off Bahia Magdalena, Mexico. We assessed the trophic magnification factor (TMF) to determine the potential for biomagnification of 11 trace metals. The concentrations of Fe and Zn were one order of magnitude higher than all other metals. Concentrations of As, Cu, Cd, Co, Cr, Fe, Mn and Ni correlated negatively with trophic level, supporting trophic dilution (TMF < 1, p > 0.05), while Zn and Hg had significant trophic magnification (TMF > 1, p < 0.05) when assessing only the benthic-pelagic foodweb. This research provides a baseline concentration of metals in multiple species, metal-specific foodweb bioaccumulation and biomagnification of mercury, underscoring the key role of the macrobenthic community as biovectors for trophic transfer of Hg through the foodweb to the California sea lion.

Deep dives and high tissue density increase mean dive costs in California sea lions (Zalophus californianus).

Diving is central to the foraging strategies of many marine mammals and seabirds. Still, the effect of dive depth on foraging cost remains elusive because energy expenditure is difficult to measure at fine temporal scales in wild animals. We used depth and acceleration data from eight lactating California sea lions (Zalophus californianus) to model body density and investigate the effect of dive depth and tissue density on rates of energy expenditure. We calculated body density in 5 s intervals from the rate of gliding descent. We modeled body density across depth in each dive, revealing high tissue densities and diving lung volumes (DLVs). DLV increased with dive depth in four individuals. We used the buoyancy calculated from dive-specific body-density models and drag calculated from swim speed to estimate metabolic power and cost of transport in 5 s intervals during descents and ascents. Deeper dives required greater mean power for round-trip vertical transit, especially in individuals with higher tissue density. These trends likely follow from increased mean swim speed and buoyant hinderance that increasingly outweighs buoyant aid in deeper dives. This suggests that deep diving is either a 'high-cost, high-reward' strategy or an energetically expensive option to access prey when prey in shallow waters are limited, and that poor body condition may increase the energetic costs of deep diving. These results add to our mechanistic understanding of how foraging strategy and body condition affect energy expenditure in wild breath-hold divers.