abstract
High-resolution tri-axial accelerometry biologging tags have quantitatively described behaviors in baleen whale species that forage using lunges and continuous ram filtration. However, detailed quantitative descriptions of foraging behaviors do not exist for gray whales, a unique baleen whale species that primarily uses benthic suction feeding with a rolling component. We deployed suction cup biologging tags on Pacific Coast Feeding Group (PCFG) gray whales to quantify foraging behavior at the broad state (dive) and foraging tactic (roll event) scales. Hidden Markov Models were used to describe three distinct states using turn angle, dive duration, pseudotrack tortuosity, and presence of roll events that can be interpreted as forage, search, and transit behavior. Classification and Regression Tree models best described foraging tactics (headstands, benthic digs, and side swims) using median pitch, depth to total length ratio, and absolute value of the median roll. On average, PCFG gray whales spent more time searching and performed more left-rolled foraging tactics at shallower depths at night compared to during the day, potentially to track prey above them in the water column. Describing foraging behavior in PCFG gray whales enables examination of links between behavioral budgets, energetics, and the physiological impact of threats facing this group.

abstract
Hidden Markov models (HMMs) are popular models to identify a finite number of latent states from sequential data. However, fitting them to large data sets can be computationally demanding because most likelihood maximization techniques require iterating through the entire underlying data set for every parameter update. We propose a novel optimization algorithm that updates the parameters of an HMM without iterating through the entire data set. Namely, we combine a partial E step with variance-reduced stochastic optimization within the M step. We prove the algorithm converges under certain regularity conditions. Using simulations and a case study of kinematic data from eight killer whales (Orcinus orca) off the western coast of Canada, we show that our algorithm converges in fewer epochs and to regions of higher likelihood compared to standard numerical optimization techniques. Our algorithm allows practitioners to fit complicated HMMs to large time-series data sets more efficiently than existing baselines.

abstract
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. 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. 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. 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.  

abstract
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.

abstract
In the northeastern Pacific, sightings of small numbers of killer whales of unknown ecotype have been sporadically reported during open ocean marine mammal surveys, pelagic birding expeditions, and high seas fishing operations. However, it is unknown whether these oceanic killer whales belong to a mammal-eating ecotype of killer whale, an offshore fish-eating ecotype, or an offshore generalist type. We attempted to determine the ecotype of 49 unknown individuals observed during 9 encounters from 1997-2021 in the deep oceanic waters far from the coastlines of California and Oregon (>65 km) based on their foraging behaviors, prey species consumed, morphologies, and the prevalence of cookiecutter shark bite scars. We hypothesize that these killer whales may represent a distinct oceanic subpopulation of transient killer whales or an undescribed oceanic population that feeds on marine mammals and sea turtles in the oceanic open ocean waters beyond the continental shelf break.

abstract
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 estimated respiration rates and calculated 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 consumed 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.

abstract
Coho Salmon Oncorhynchus kisutch provide an important resource for recreational, commercial, and Indigenous fisheries in the Pacific Northwest. The goal of this study was to improve our understanding of how marine mam-mal predation may be impacting the survival and productivity of Coho Salmon in the Strait of Georgia, British Columbia. Specifically, we quantified the impact of harbor seal Phoca vitulina predation on juvenile Coho Salmon during their first several months at sea. Early marine survival is believed to be the limiting factor for the recovery of Coho Salmon populations in this region. To estimate the number of juvenile Coho Salmon consumed by harbor seals, we developed a mathematical model that integrates predator diet data and salmon population and mortality dynamics. Our analysis estimated that harbor seals consumed an annual average of 46−59% of juvenile Coho Salmon between 2004–2016, providing the first quantitative estimate of seal predation in the Strait of Georgia. Marine mammal predation on juvenile Coho Salmon is potentially a very important factor limiting survival and recovery of Coho Salmon in the Strait of Georgia.

abstract
The age at which sea lions wean can vary significantly between years and among populations. It is an important life-history parameter that is influenced by environmental conditions and can drive changes in sea lion numbers. However, knowing when weaning begins and ends is difficult to determine. We developed a method (using Fourier analysis) to identify the lactation period from changes in the δ15N profiles of vibrissae from juvenile California sea lions—born in three colonies in Mexico. We sectioned vibrissae from 15 juvenile California sea lions (aged approximately 12 months) into 33–74 segments of similar weight. We measured δ15N and δ13C for each vibrissa segment and assigned dates to each one using site-specific vibrissa growth rates. We also compared the δ15N profiles that corresponded to the pup stage on the juvenile vibrissae with δ15N values of adult female vibrissae from the same colonies to validate the dietary transition from milk to fish identified by the Fourier analysis. We found that pups began supplementing their milk diet with fish at different times between colonies—ranging from 3 to 5 months old (San Esteban Island), 5–7 months old (Santa Margarita Island), and > 12 months old (Los Islotes Island). All pups were > 1 year old when weaned. The longer lactation period in Mexico contrasts with the shorter 10–11 months age at weaning recorded at northern colonies of California sea lions along the US Pacific coast (San Miguel Island). The difference in lactation duration among regions likely reflects latitudinal differences in marine productivity, and a lower nutrition quality of prey available to California sea lions in Mexico. Our study augments the limited knowledge of weaning in California sea lions and provides a means to determine weaning in other species.

abstract
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 Washing-ton 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.

abstract
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 hump-back 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 hump-backs, 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

abstract
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 electrocardiogramgated 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.

abstract
The enlarged size of the aortic bulb is thought to enhance the ability of marine mammals to remain underwater for extended periods. However, a convincing link between aortic bulb size and diving capacity has not been established. Using new and existing data, we examined the relationships between body size, maximum and routine dive duration, and aortic bulb size of pinnipeds. Comparisons among seven species of pinnipeds showed that the diameter of the aortic bulb increases allometrically with body mass (aortic bulb diameter = 0.58 x body mass0.41). We also found a linear relationship between routine dive duration and relative aortic bulb diameter (routine dive duration = 0.20 x relative aortic bulb diameter – 3.30), but no apparent relationship with maximum dive duration. Our results indicate that relative aortic bulb diameter influences diving capacity, providing further evidence that the aortic bulb is an adaptation to diving. Specifically, the relative diameter of the aortic bulb partially determines how long pinnipeds can routinely remain underwater. This has implications for the ability of different species of marine mammals to adapt to projected environmental changes and effectively forage or evade threats in altered habitats.

abstract
Abducting and sexually coercing pups are hypothesized to be a means for immature male otariids to learn how to secure, control, and mate with females—and increase their future reproductive success. However, the impact of unintentional pup deaths and the disruption of mother-pup bonds due to the behavior of juvenile and subadult males on population growth is unclear, especially for species such as Guadalupe fur seals (Arctocephalus townsendi), which are recovering from the catastrophic effects of sealing that occurred over a century ago. We observed juvenile and subadult male Guadalupe fur seals at the male-dominated San Benito Archipelago engaging in aggressive behaviors such as harassment, abduction, and attempted copulation with pups. These actions are presumed to benefit the fitness of individual subadult males, but can have severe consequences for the pups, including physical trauma, separation from their mothers, interrupted nursing, and even death. Such detrimental effects can hinder the recovery of the Guadalupe fur seal population, particularly at the San Benito Archipelago, where such behaviors are more prevalent compared to the breeding population on Guadalupe Island.