Behavioral context of echolocation and prey-handling sounds produced by killer whales (Orcinus orca) during pursuit and capture of Pacific salmon (Oncorhynchus spp.).

Availability of preferred salmonid prey and a sufficiently quiet acoustic environment in which to forage are critical to the survival of resident killer whales (Orcinus orca) in the northeastern Pacific. Although piscivorous killer whales rely on echolocation to locate and track prey, the relationship between echolocation, movement, and prey capture during foraging by wild individuals is poorly understood. We used acoustic biologging tags to relate echolocation behavior to prey pursuit and capture during successful feeding dives by fish-eating killer whales in coastal British Columbia, Canada. The significantly higher incidence and rate of echolocation prior to fish captures compared to afterward confirms its importance in prey detection and tracking. Extremely rapid click sequences (buzzes) were produced before or concurrent with captures of salmon at depths typically exceeding 50 m, and were likely used by killer whales for close-range prey targeting, as in other odontocetes. Distinctive crunching and tearing sounds indicative of prey-handling behavior occurred at relatively shallow depths following fish captures, matching concurrent observations that whales surfaced with fish prior to consumption and often shared prey. Buzzes and prey-handling sounds are potentially useful acoustic signals for estimating foraging efficiency and determining if resident killer whales are meeting their energetic requirements.

CARDIAC EXAMINATIONS OF ANESTHETIZED STELLER SEA LIONS (EUMETOPIAS JUBATUS), NORTHERN FUR SEALS (CALLORHINUS URSINUS), AND A WALRUS (ODOBENUS ROSMARUS).

Pinniped hearts have been well described via dissection, but in vivo measurements of cardiac structure, function, and electrophysiology are lacking. Electrocardiograms (ECGs) were recorded under anesthesia from eight Steller sea lions (Eumetopias jubatus), five northern fur seals (Callorhinus ursinus), and one walrus (Odobenus rosmarus) to investigate cardiac electrophysiology in pinnipeds. In addition, echocardiograms were performed on all eight anesthetized Steller sea lions to evaluate in vivo cardiac structure and function. Measured and calculated ECG parameters included P-wave, PQ, QRS, and QT interval durations, P-, R-, and T-wave amplitudes, P- and T-wave polarities, and the mean electrical axis (MEA). Measured and calculated echocardiographic parameters included left ventricular internal diameter, interventricular septum thickness, and left ventricular posterior wall thickness in systole and diastole (using M-mode), left atrium and aortic root dimensions (using 2D), and maximum aortic and pulmonary flow velocities (using pulsed-wave spectral Doppler). ECG measurements were similar to those reported for other pinniped species, but there was considerable variation in the MEAs of Steller sea lions and northern fur seals. Echocardiographic measurements were similar to those reported for southern sea lions (Otaria flavenscens), including five out of eight Steller sea lions having a left atrial to aortic root ratio <1, which may indicate that they have an enlarged aortic root compared to awake terrestrial mammals. Isoflurane anesthesia likely affected some of the measurements as evidenced by the reduced fractional shortening found in Steller sea lions compared to awake terrestrial mammals. The values reported are useful reference points for assessing cardiac health in pinnipeds under human care.

REFERENCE RANGES AND AGE-RELATED AND DIVING EXERCISE EFFECTS ON HEMATOLOGY AND SERUM CHEMISTRY OF FEMALE STELLER SEA LIONS ( EUMETOPIAS JUBATUS).

Decreased health may have lowered the birth and survival rates of Steller sea lions ( Eumetopias jubatus) in the Gulf of Alaska and Aleutian Islands over the past 30 yr. Reference ranges for clinical hematology and serum chemistry parameters needed to assess the health of wild sea lion populations are limited. Here, blood parameters were serially measured in 12 captive female Steller sea lions ranging in age from 3 wk to 16 yr to establish baseline values and investigate age-related changes. Whether diving activity affects hematology parameters in animals swimming in the ocean compared with animals in a traditional aquarium setting was also examined. Almost all blood parameters measured exhibited significant changes with age. Many of the age-related changes reflected developmental life history changes, including a change in diet during weaning, an improvement of diving capacity, and the maturity of the immune system. Mean corpuscular hemoglobin and mean corpuscular volume were also higher in the ocean diving group compared with the aquarium group, likely reflecting responses to increased exercise regimes. These data provide ranges of hematology and serum chemistry values needed to evaluate and compare the health and nutritional status of captive and wild Steller sea lions.

Dive, food, and exercise effects on blood microparticles in Steller sea lions (Eumetopias jubatus): exploring a biomarker for decompression sickness.

Recent studies of stranded marine mammals indicate that exposure to underwater military sonar may induce pathophysiological responses consistent with decompression sickness (DCS). However, DCS has been difficult to diagnose in marine mammals. We investigated whether blood microparticles (MPs, measured as number/µl plasma), which increase in response to decompression stress in terrestrial mammals, are a suitable biomarker for DCS in marine mammals. We obtained blood samples from trained Steller sea lions (Eumetopias jubatus, 4 adult females) wearing time-depth recorders that dove to predetermined depths (either 5 or 50 meters). We hypothesized that MPs would be positively related to decompression stress (depth and duration underwater). We also tested the effect of feeding and exercise in isolation on MPs using the same blood sampling protocol. We found that feeding and exercise had no effect on blood MP levels, but that diving caused MPs to increase. However, blood MP levels did not correlate with diving depth, relative time underwater, and presumed decompression stress, possibly indicating acclimation following repeated exposure to depth.

Feeding kinematics and performance of basal otariid pinnipeds, Steller sea lions and northern fur seals: implications for the evolution of mammalian feeding.

Feeding performance studies can address questions relevant to feeding ecology and evolution. Our current understanding of feeding mechanisms for aquatic mammals is poor. Therefore, we characterized the feeding kinematics and performance of five Steller sea lions (Eumetopias jubatus) and six northern fur seals (Callorhinus ursinus). We tested the hypotheses that both species use suction as their primary feeding mode, and that rapid jaw opening was related to suction generation. Steller sea lions used suction as their primary feeding mode, but also used a biting feeding mode. In contrast, northern fur seals only used a biting feeding mode. Kinematic profiles of Steller sea lions were all indicative of suction feeding (i.e. a small gape, small gape angle, large depression of the hyolingual apparatus and lip pursing). However, jaw opening as measured by gape angle opening velocity (GAOV) was relatively slow in Steller sea lions. In contrast to Steller sea lions, the GAOV of northern fur seals was extremely fast, but their kinematic profiles indicated a biting feeding mode (i.e. northern fur seals exhibited a greater gape, a greater gape angle and minimal depression of the hyolingual apparatus compared with Steller sea lions). Steller sea lions produced both subambient and suprambient pressures at 45 kPa. In contrast, northern fur seals produced no detectable pressure measurements. Steller sea lions have a broader feeding repertoire than northern fur seals, which likely enables them to feed on a greater variety of prey, in more diverse habitats. Based on the basal phylogenetic position of northern fur seals, craniodental morphological data of the Callorhinus lineage, and the performance data provided in this study, we suggest that northern fur seals may be exhibiting their ancestral feeding mode.

A nutrigenomic approach to detect nutritional stress from gene expression in blood samples drawn from Steller sea lions.

Gene expression profiles are increasingly being used as biomarkers to detect the physiological responses of a number of species to disease, nutrition, and other stressors. However, little attention has been given to using gene expression to assess the stressors and physiological status of marine mammals. We sought to develop and validate a nutrigenomic approach to quantify nutritional stress in Steller sea lions (Eumetopias jubatus). We subjected 4 female Steller sea lions to 3 feeding regimes over 70-day trials (unrestricted food intake, acute nutritional stress, and chronic nutritional stress), and drew blood samples from each animal at the end of each feeding regime. We then extracted the RNA of white blood cells and measured the response of 8 genes known to react to diet restriction in terrestrial mammals. Overall, we found that the genomic response of Steller sea lions experiencing nutritional stress was consistent with how terrestrial mammals respond to dietary restrictions. Our nutritionally stressed sea lions down-regulated some cellular processes involved in immune response and oxidative stress, and up-regulated pro-inflammatory responses and metabolic processes. Nutrigenomics appears to be a promising means to monitor nutritional status and contribute to mitigation measures needed to assist in the recovery of Steller sea lions and other at-risk species of marine mammals.

Improving accuracy of DNA diet estimates using food tissue control materials and an evaluation of proxies for digestion bias.

Ecologists are increasingly interested in quantifying consumer diets based on food DNA in dietary samples and high-throughput sequencing of marker genes. It is tempting to assume that food DNA sequence proportions recovered from diet samples are representative of consumer's diet proportions, despite the fact that captive feeding studies do not support that assumption. Here, we examine the idea of sequencing control materials of known composition along with dietary samples in order to correct for technical biases introduced during amplicon sequencing and biological biases such as variable gene copy number. Using the Ion Torrent PGM(©) , we sequenced prey DNA amplified from scats of captive harbour seals (Phoca vitulina) fed a constant diet including three fish species in known proportions. Alongside, we sequenced a prey tissue mix matching the seals' diet to generate tissue correction factors (TCFs). TCFs improved the diet estimates (based on sequence proportions) for all species and reduced the average estimate error from 28 ± 15% (uncorrected) to 14 ± 9% (TCF-corrected). The experimental design also allowed us to infer the magnitude of prey-specific digestion biases and calculate digestion correction factors (DCFs). The DCFs were compared with possible proxies for differential digestion (e.g. fish protein%, fish lipid%) revealing a strong relationship between the DCFs and percent lipid of the fish prey, suggesting prey-specific corrections based on lipid content would produce accurate diet estimates in this study system. These findings demonstrate the value of parallel sequencing of food tissue mixtures in diet studies and offer new directions for future research in quantitative DNA diet analysis.

Steller sea lions (Eumetopias jubatus) have greater blood volumes, higher diving metabolic rates and a longer aerobic dive limit when nutritionally stressed.

Marine mammal foraging behaviour inherently depends on diving ability. Declining populations of Steller sea lions may be facing nutritional stress that could affect their diving ability through changes in body composition or metabolism. Our objective was to determine whether nutritional stress (restricted food intake resulting in a 10% decrease in body mass) altered the calculated aerobic dive limit (cADL) of four captive sea lions diving in the open ocean, and how this related to changes in observed dive behaviour. We measured diving metabolic rate (DMR), blood O2 stores, body composition and dive behaviour prior to and while under nutritional restriction. We found that nutritionally stressed sea lions increased the duration of their single long dives, and the proportion of time they spent at the surface during a cycle of four dives. Nutritionally stressed sea lions lost both lipid and lean mass, resulting in potentially lower muscle O2 stores. However, total body O2 stores increased due to rises in blood O2 stores associated with having higher blood volumes. Nutritionally stressed sea lions also had higher mass-specific metabolic rates. The greater rise in O2 stores relative to the increase in mass-specific DMR resulted in the sea lions having a longer cADL when nutritionally stressed. We conclude that there was no negative effect of nutritional stress on the diving ability of sea lions. However, nutritional stress did lower foraging efficiency and require more foraging time to meet energy requirements due to increases in diving metabolic rates and surface recovery times.

Prevalence of Chinook salmon is higher for southern than for northern resident killer whales in summer hot-spot feeding areas.

Differences in the availability of prey may explain the low numbers of southern resident killer whales and the increase in northern resident killer whales in British Columbia and Washington State. However, in-situ data on the availability of their preferred prey (Chinook salmon, Oncorhynchus tshawytscha) in the core feeding areas used by these two populations of fish-eating killer whales have been lacking to test this hypothesis. We used multi-frequency echosounders (38, 70, 120, and 200 kHz) to estimate densities of adult Chinook (age-4+, > 81 cm) within 16 hot-spot feeding areas used by resident killer whales during summer 2020 in the Salish Sea and North Island Waters. We found Chinook were generally concentrated within 50 m from the bottom in the deep waters, and tended to be absent near the surface in the shallow waters (< 50 m). In general, the densities of Chinook we encountered were highest as the fish entered the Salish Sea (from Swiftsure Bank in the south) and Johnstone Strait (from Queen Charlotte Strait to the north)-and declined as fish migrated eastward along the shoreline of Vancouver Island. Median densities of Chinook for all sampled areas combined were 0.4 ind.·1000 m-2 in northern resident foraging areas, and 0.9 ind.·1000 m-2 in southern resident killer whale areas (p < 0.05, Mann-Whitney U test). Thus, Chinook salmon were twice as prevalent within the hot-spot feeding areas of southern versus northern resident killer whales. This implies that southern resident killer whales have greater access to Chinook salmon compared to northern residents during summer-and that any food shortage southern residents may be encountering is occurring at other times of year, or elsewhere in their range.

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.

Chinook salmon depth distributions on the continental shelf are shaped by interactions between location, season, and individual condition.

BACKGROUND: Ecological and physical conditions vary with depth in aquatic ecosystems, resulting in gradients of habitat suitability. Although variation in vertical distributions among individuals provides evidence of habitat selection, it has been challenging to disentangle how processes at multiple spatio-temporal scales shape behaviour. METHODS: We collected thousands of observations of depth from > 300 acoustically tagged adult Chinook salmon Oncorhynchus tshawytscha, spanning multiple seasons and years. We used these data to parameterize a machine-learning model to disentangle the influence of spatial, temporal, and dynamic oceanographic variables while accounting for differences in individual condition and maturation stage. RESULTS: The top performing machine learning model used bathymetric depth ratio (i.e., individual depth relative to seafloor depth) as a response. We found that bathymetry, season, maturation stage, and spatial location most strongly influenced Chinook salmon depth. Chinook salmon bathymetric depth ratios were deepest in shallow water, during winter, and for immature individuals. We also identified non-linear interactions among covariates, resulting in spatially-varying effects of zooplankton concentration, lunar cycle, temperature and oxygen concentration. CONCLUSIONS: Our results suggest Chinook salmon vertical habitat use is a function of ecological interactions, not physiological constraints. Temporal and spatial variation in depth distributions could be used to guide management decisions intended to reduce fishery impacts on Chinook salmon. More generally, our findings demonstrate how complex interactions among bathymetry, seasonality, location, and life history stage regulate vertical habitat selection.

Aquaculture related humpback whale entanglements in coastal waters of British Columbia from 2008-2021.

Over the past two decades, increasing numbers of humpback whales have been returning to feed in the inshore waters of British Columbia (BC) where marine aquaculture farms are situated. This has led to growing concerns that the presence of aquaculture farms may pose an entanglement threat to humpback whales. However, it is not known whether aquaculture facilities attract humpback whales, or whether there are factors that increase the likelihood of humpback whale, becoming entangled and dying. We examined eight reports of humpback whales interacting with Atlantic salmon farms in BC from 2008 to 2021 to evaluate the conditions that may have contributed to their entanglements. Of the eight entangled humpbacks, three individuals died and five were successfully disentangled and released. All were young animals (1 calf, 7 subadults). Multiple factors were associated with two or more of the reported incidents. These included facility design, environmental features, seasonality, humpback whale age, and feeding behaviour. We found that humpback whales were most commonly entrapped in the predator nets of the aquaculture facilities (6/8 incidents), and were less often entangled in anchor support lines (2/8). The presence of salmon smolts did not appear to be an attractant for humpback whales given that half of the reported entanglements (4/8) occurred at fallowed salmon farms. Almost all of the entanglements (7/8) occurred in late winter (prior to the seasonal return of humpbacks) and during late fall (after most humpbacks have migrated south). Overall, the number of humpback whales impacted by fish farms was small compared to the numbers that return to BC (> 7,000) and accounted for <6% of all types of reported entanglements in BC. Human intervention was required to release humpback whales at fish farms, which points to the need to have well-established protocols to minimize entanglements and maximize successful releases.

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.

Killer whale respiration rates.

Measuring breathing rates is a means by which oxygen intake and metabolic rates can be estimated to determine food requirements and energy expenditure of killer whales (Orcinus orca) and other cetaceans. This relatively simple measure also allows the energetic consequences of environmental stressors to cetaceans to be understood but requires knowing respiration rates while they are engaged in different behaviours such as resting, travelling and foraging. We calculated respiration rates for different behavioural states of southern and northern resident killer whales using video from UAV drones and concurrent biologging data from animal-borne tags. Behavioural states of dive tracks were predicted using hierarchical hidden Markov models (HHMM) parameterized with time-depth data and with labeled tracks of drone-identified behavioural states (from drone footage that overlapped with the time-depth data). Dive tracks were sequences of dives and surface intervals lasting ≥ 10 minutes cumulative duration. We calculated respiration rates and estimated oxygen consumption rates for the predicted behavioural states of the tracks. We found that juvenile killer whales breathed at a higher rate when travelling (1.6 breaths min-1) compared to resting (1.2) and foraging (1.5)-and that adult males breathed at a higher rate when travelling (1.8) compared to both foraging (1.7) and resting (1.3). The juveniles in our study were estimated to consume 2.5-18.3 L O2 min-1 compared with 14.3-59.8 L O2 min-1 for adult males across all behaviours based on estimates of mass-specific tidal volume and oxygen extraction. Our findings confirm that killer whales take single breaths between dives and indicate that energy expenditure derived from respirations requires using sex, age, and behavioural-specific respiration rates. These findings can be applied to bioenergetics models on a behavioural-specific basis, and contribute towards obtaining better predictions of dive behaviours, energy expenditure and the food requirements of apex predators.

An integrated system to assess marine extinctions.

More than 20 global marine extinctions and over 700 local extinctions have reportedly occurred during the past 500 years. However, available methods to determine how many of these species can be confidently declared true disappearances tend to be data-demanding, time-consuming, and not applicable to all taxonomic groups or scales of marine extinctions (global [G] and local [L]). We developed an integrated system to assess marine extinctions (ISAME) that can be applied to any taxonomic group at any geographic scale. We applied the ISAME method to 10 case studies to illustrate the possible ways in which the extinction status of marine species can be categorized as unverified, possibly extinct, or extinct. Of the 10 case studies we assessed, the ISAME method concludes that 6 should be categorized as unverified extinctions due to problems with species' identity and lack of reliable evidence supporting their disappearance (periwinkle-Littoraria flammea [G], houting-Coregonus oxyrinchus [G], long-spined urchin-Diadema antillarum [L], smalltooth sawfish-Pristis pectinata [L], and largetooth sawfish-P. pristis [L]). In contrast, ISAME classified the Guadalupe storm-petrel (Oceanodroma macrodactyla [G]) and the lost shark (Carcharhinus obsolerus [G]) as possibly extinct because the available evidence indicates that their extinction is plausible-while the largetooth sawfish [L] and Steller's sea cow (Hydrodamalis gigas [G]) were confirmed to be extinct. Determining whether a marine population or species is actually extinct or still extant is needed to guide conservation efforts and prevent further biodiversity losses.

Data on the diets of Salish Sea harbour seals from DNA metabarcoding.

Marine trophic ecology data are in high demand as natural resource agencies increasingly adopt ecosystem-based management strategies that account for complex species interactions. Harbour seal (Phoca vitulina) diet data are of particular interest because the species is an abundant predator in the northeast Pacific Ocean and Salish Sea ecosystem that consumes Pacific salmon (Oncorhynchus spp.). A multi-agency effort was therefore undertaken to produce harbour seal diet data on an ecosystem scale using, 1) a standardized set of scat collection and analysis methods, and 2) a newly developed DNA metabarcoding diet analysis technique designed to identify prey species and quantify their relative proportions in seal diets. The DNA-based dataset described herein contains records from 4,625 harbour seal scats representing 52 haulout sites, 7 years, 12 calendar months, and a total of 11,641 prey identifications. Prey morphological hard parts analyses were conducted alongside, resulting in corresponding hard parts data for 92% of the scat DNA samples. A custom-built prey DNA sequence database containing 201 species (192 fishes, 9 cephalopods) is also provided.

Dive behaviour impacts the ability of heart rate to predict oxygen consumption in Steller sea lions (Eumetopias jubatus) foraging at depth.

The predictive relationship between heart rate (f(H)) and oxygen consumption (VO2) has been derived for several species of marine mammals swimming horizontally or diving in tanks to shallow depths. However, it is unclear how dive activity affects the f(H):VO2 relationship and whether the existing equations apply to animals diving to deeper depths. We investigated these questions by simultaneously measuring the f(H) and VO2 of Steller sea lions (Eumetopias jubatus) under different activity states (surface resting or diving), types of dives (single dives or dive bouts), and depths (10 or 40 m). We examined the relationship over dives only and also over dive cycles (dive + surface interval). We found that f(H) could only predict VO2 over a complete single dive cycle or dive bout cycle (i.e. surface intervals had to be included). The predictive equation derived for sea lions resting on the surface did not differ from that for single dive cycles. However, the equation derived over dive bout cycles (multiple dives + surface intervals) differed from those for single dive cycles or surface resting, with similar f(H) for multiple dive bout equations yielding higher predicted VO2 than that for single dive bout cycles (or resting). The f(H):VO2 relationships were not significantly affected by dive duration, dive depth, water temperature or cumulative food consumed under the conditions tested. Ultimately, our results demonstrate that f(H) can be used to predict activity-specific metabolic rates of diving Steller sea lions, but only over complete dive cycles that include a post-dive surface recovery period.

Electrocardiographic Scaling Reveals Differences in Electrocardiogram Interval Durations Between Marine and Terrestrial Mammals.

Although the ability of marine mammals to lower heart rates for extended periods when diving is well documented, it is unclear whether marine mammals have electrophysiological adaptations that extend beyond overall bradycardia. We analyzed electrocardiographic data from 50 species of terrestrial mammals and 19 species of marine mammals to determine whether the electrical activity of the heart differs between these two groups of mammals. We also tested whether physiological state (i.e., anesthetized or conscious) affects electrocardiogram (ECG) parameters. Analyses of ECG waveform morphology (heart rate, P-wave duration, and PQ, PR, QRS, and QT intervals) revealed allometric relationships between body mass and all ECG intervals (as well as heart rate) for both groups of mammals and specific differences in ECG parameters between marine mammals and their terrestrial counterparts. Model outputs indicated that marine mammals had 19% longer P-waves, 24% longer QRS intervals, and 21% shorter QT intervals. In other words, marine mammals had slower atrial and ventricular depolarization, and faster ventricular repolarization than terrestrial mammals. Heart rates and PR intervals were not significantly different between marine and terrestrial mammals, and physiological state did not significantly affect any ECG parameter. On average, ECG interval durations of marine and terrestrial mammals scaled with body mass to the power of 0.21 (range: 0.19-0.23) rather than the expected 0.25-while heart rate scaled with body mass to the power of -0.22 and was greater than the widely accepted -0.25 derived from fractal geometry. Our findings show clear differences between the hearts of terrestrial and marine mammals in terms of cardiac timing that extend beyond diving bradycardia. They also highlight the importance of considering special adaptations (such as breath-hold diving) when analyzing allometric relationships.

Growth and development of North Pacific gray whales (Eschrichtius robustus).

Understanding variability in growth patterns of marine mammals provides insights into the health of individuals and status of populations. Body growth of gray whales (Eschrichtius robustus) has been described for particular life stages, but has not been quantified across all ages. We derived a comprehensive growth equation for gray whales by fitting a two-phased growth model to age-specific length data of eastern North Pacific gray whales that were captured, stranded, or harvested between 1926 and 1997. To predict mass-at-age, we used the allometric relationship between mass and length. We found that on average (± SD), calves were 4.6 ± 0.043 m and 972 ± 26 kg at birth, and reached 8.5 ± 0.095 m and 6,019 ± 196 kg by the end of their first year of life (n = 118). Thus, calves almost double (2×) in length and octuple (8×) in mass while nursing, and are effectively about two-thirds of their asymptotic adult length and one-third of their maximum mass when weaned. The large sample of aged individuals (n = 730) indicates that gray whales live up to ~48 years and have a life expectancy of < 30 years. Adult females attain a mean (± SD) asymptotic size of 13.1 ± 0.048 m and 20,758 ± 222 kg, while the smaller males average 12.6 ± 0.048 m and 19,938 ± 222 kg at ~40 years of age. Females are thereby ~4% longer and heavier than males. These age-specific estimates of body size can be used to estimate food requirements and assess nutritional status of individuals.

Bowhead whales use two foraging strategies in response to fine-scale differences in zooplankton vertical distribution.

As zooplanktivorous predators, bowhead whales (Balaena mysticetus) must routinely locate patches of prey that are energy-rich enough to meet their metabolic needs. However, little is known about how the quality and quantity of prey might influence their feeding behaviours. We addressed this question using a new approach that included: (1) multi-scale biologging and unmanned aerial system observations of bowhead whales in Cumberland Sound, Nunavut (Canada), and (2) an optical plankton counter (OPC) and net collections to identify and enumerate copepod prey species through the water column. The OPC data revealed two prey layers comprised almost exclusively of lipid-rich calanoid copepods. The deep layer contained fewer, but larger, particles (10% greater overall biomass) than the shallow prey layer. Dive data indicated that the whales conducted long deep Square-shaped dives (80% of dives; averaging depth of 260.4 m) and short shallow Square-shaped dives (16%; averaging depth of 22.5 m) to feed. The whales tended to dive proportionally more to the greater biomass of zooplankton that occurred at depth. Combining behavioural recordings with prey sampling showed a more complex feeding ecology than previously understood, and provides a means to evaluate the energetic balance of individuals under current environmental conditions.