Walruses live in the rapidly changing Arctic. David Rosen is trying to figure out how climate change will impact the health of young walruses. 

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A Window into the Lives of Resident Killer Whales

This summer, a team of researchers from the University of British Columbia, together with the Hakai Institute, set out to determine how fish-eating killer whales find their food, and whether there is a shortage of Chinook salmon available to killer whales in the Salish Sea.  Here is a peek into what the researchers saw. <see full story>

Struggle in the Strait

Is seal predation driving salmon declines in the Strait of Georgia?

The decline of Chinook salmon is affecting predators like killer whales, as well as human fisheries. Are harbor seals to blame? New research examines decades of population data to gain insights into whether there is a relationship between harbor seal numbers and salmon stocks. The results show a surprising trend. <see full story>

Do Foraging Strategies Matter?

Are female northern fur seals finding enough food to help their pups survive?

How efficiently do female northern fur seals forage for themselves and their pups? Do their decisions help or hinder their pups’ chances of survival at sea? New Consortium research examines the foraging strategies of female northern fur seals and their implications for declining populations in the North Pacific Ocean.  <see full story>


Reference ranges and age-related and diving exercise effects on hematology and serum chemistry of female Steller sea lions (Eumetopias jubatus).
Gerlinsky, C. D., M. Haulena, A. W. Trites and D. A. S. Rosen. 2018.
Journal of Zoo and Wildlife Medicine 49(1):18-29.
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.

keywords     Diving, Eumetopias jubatus, hematology, marine mammal, serum chemistry, Steller sea lion
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Trade-off between foraging efficiency and pup feeding rate of lactating northern fur seals in a declining population.
Jeanniard-du-Dot, T., A.W. Trites, J.P.Y. Arnould, J.R. Speakman, and C. Guinet. 2018.
Marine Ecology Progress Series 600:207-222.
Foraging strategies and their resulting efficiency (energy gain to cost ratio) affect animals' survival and reproductive success and can be linked to population dynamics. However, they have rarely been studied quantitatively in free-ranging animals. We investigated foraging strategies and efficiencies of wild northern fur seals Callorhinus ursinus during their breeding season to understand potential links to the observed population decline in the Bering Sea. We equipped 20 lactating females with biologgers to determine at-sea foraging behaviors. We measured energy expenditure while foraging using the doubly labeled water method, and energy gained using (1) the types and energy densities of prey consumed, and (2) the number of prey capture attempts (from acceleration data). Our results show that seals employed 2 foraging strategies. One group (40%) fed mostly in oceanic waters on small high energy-density prey, while the other (60%) stayed over the shallow continental shelf feeding mostly on larger, lower quality fish. Females foraging in oceanic waters captured 3 times more prey, and had double the foraging efficiencies of females that foraged on-shelf in neritic waters. However, neritic seals made comparatively shorter trips, and likely fed their pups ~20-25% more frequently. The presence of these strategies which either favor foraging efficiency (energy) or frequency of nursing (time) might be maintained in the population because they have similar net fitness outcomes. However, neither strategy appears to simultaneously maximise time and energy allocated to nursing, with potential impacts on the survival of pups during their first year at sea.
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Telemetry tags increase the costs of swimming in northern fur seals, Callorhinus ursinus.
Rosen, D. A. S., C. D. Gerlinsky and A. W. Trites. 2018.
Marine Mammal Science 34 (2):385-402. doi:10.1111/mms.12460
Animal-borne instruments have become a standard tool for collecting important data from marine mammals. However, few studies have examined whether placement of these data loggers affects the behavior and energetics of individual animals, potentially leading to biasing data. We measured the effect of two types of relatively small data loggers (<1% of animals

keywords     northern fur seals, Callorhinus ursinus, telemetry, bioenergetics, biologging, diving, swimming, marine mammal
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Accelerometers can measure total and activity-specific energy expenditure in free-ranging marine mammals only if linked to time-activity budgets.
Jeanniard du Dot, T., C. Guinet, J. P. Y. Arnould, J. R. Speakman and A. W. Trites. 2017.
Functional Ecology 31:377-386.
1-Energy expenditure is an important component of foraging ecology, but is extremely difficult to estimate in free-ranging animals and depends on how animals partition their time between different activities during foraging. Acceleration data has emerged as a new way to determine energy expenditure at a fine scale but needs to be tested and validated in wild animals. 2-This study investigated whether vectorial dynamic body acceleration (VeDBA) could accurately predict the energy expended by marine predators during a full foraging trip. We also aimed to determine whether the accuracy of predictions of energy expenditure derived from acceleration increased when partitioned by different types of at-sea activities (i.e., diving, transiting, resting and surface activities) vs calculated activity-specific metabolic rates. 3-To do so, we equipped 20 lactating northern (Callorhinus ursinus) and 20 Antarctic fur seals (Arctocephalus gazella) with GPS, time-depth recorders and tri-axial accelerometers, and obtained estimates of field metabolic rates using the doubly-labelled water (DLW) method. VeDBA was derived from tri-axial acceleration, and at-sea activities (diving, transiting, resting and surface activities) were determined using dive depth, tri-axial acceleration and traveling speed. 4-We found that VeDBA did not accurately predict the total energy expended by fur seals during their full foraging trips (R2 = 0.36). However, the accuracy of VeDBA as a predictor of total energy expenditure increased significantly when foraging trips were partitioned by activity and used activity-specific VeDBA paired with time activity budgets (R2 = 0.70). Activity-specific VeDBA also accurately predicted the energy expenditures of each activity independent of each other (R2 > 0.85). 5-Our study confirms that acceleration is a promising way to estimate energy expenditures of free-ranging marine mammals at a fine scale never attained before. However, it shows that it needs to be based on the time-activity budget that make up foraging trips rather than being derived as a single measure of VeDBA applied to entire foraging trips. Our activity-based method provides a cost-effective means to accurately calculate energy expenditures of fur seals using acceleration and time-activity budgets, a stepping stone for numerous other research fields.

keywords     Antarctic fur seal, Arctocephalus gazella, Callorhinus ursinus, diving, energy expenditure, foraging, metabolic rate, northern fur seal, time-activity budget
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Reproductive success is energetically linked to foraging efficiency in Antarctic fur seals.
Jeanniard-du-Dot, T., A.W. Trites, J.P. Arnould, and C. Guinet. 2017.
PLoS ONE. 12:e0174001
The efficiency with which individuals extract energy from their environment defines their survival and reproductive success, and thus their selective contribution to the population. Individuals that forage more efficiently (i.e., when energy gained exceeds energy expended) are likely to be more successful at raising viable offspring than individuals that forage less efficiently. Our goal was to test this prediction in large long-lived mammals under free-ranging conditions. To do so, we equipped 20 lactating Antarctic fur seals (Arctocephalus gazella) breeding on Kerguelen Island in the Southern Ocean with tags that recorded GPS locations, depth and tri-axial acceleration to determine at-sea behaviours and detailed time-activity budgets during their foraging trips. We also simultaneously measured energy spent at sea using the doubly-labeled water (DLW) method, and estimated the energy acquired while foraging from 1) type and energy content of prey species present in scat remains, and 2) numbers of prey capture attempts determined from head acceleration. Finally, we followed the growth of 36 pups from birth until weaning (of which 20 were the offspring of our 20 tracked mothers), and used the relative differences in body mass of pups at weaning as an index of first year survival and thus the reproductive success of their mothers. Our results show that females with greater foraging efficiencies produced relatively bigger pups at weaning. These mothers achieved greater foraging efficiency by extracting more energy per minute of diving rather than by reducing energy expenditure. This strategy also resulted in the females spending less time diving and less time overall at sea, which allowed them to deliver higher quality milk to their pups, or allowed their pups to suckle more frequently, or both. The linkage we demonstrate between reproductive success and the quality of individuals as foragers provides an individual-based quantitative framework to investigate how changes in the availability and accessibility of prey can affect fitness of animals.

keywords     foraging, energetics, pups, growth, biologging, doubly-labeled water, Antarctic fur seal, Arctocephalus gazella
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Activity-specific metabolic rates for diving, transiting and resting at sea can be estimated from time-activity budgets in free-ranging marine mammals.
Jeanniard-du-Dot, T., A.W. Trites, J.P.Y. Arnould, and C. Guinet. 2017.
Ecology and Evolution 2017:1-8.
Time and energy are the two most important currencies in animal bioenergetics. How much time animals spend engaged in different activities with specific energetic costs ultimately defines their likelihood of surviving and successfully reproducing. However, it is extremely difficult to determine the energetic costs of independent activities for free-ranging animals. In this study, we developed a new method to calculate activity-specific metabolic rates, and applied it to female fur seals. We attached biologgers (that recorded GPS locations, depth profiles, and triaxial acceleration) to 12 northern (Callorhinus ursinus) and 13 Antarctic fur seals (Arctocephalus gazella), and used a hierarchical decision tree algorithm to determine time allocation between diving, transiting, resting, and performing slow movements at the surface (grooming, etc.). We concomitantly measured the total energy expenditure using the doubly-labelled water method. We used a general least-square model to establish the relationship between time-activity budgets and the total energy spent by each individual during their foraging trip to predict activity-specific metabolic rates. Results show that both species allocated similar time to diving (~29%), transiting to and from their foraging grounds (~26-30%), and resting (~8-11%). However, Antarctic fur seals spent significantly more time grooming and moving slowly at the surface than northern fur seals (36% vs. 29%). Diving was the most expensive activity (~30 MJ/day if done non-stop for 24 hr), followed by transiting at the surface (~21 MJ/day). Interestingly, metabolic rates were similar between species while on land or while slowly moving at the surface (~13 MJ/day). Overall, the average field metabolic rate was ~20 MJ/day (for all activities combined). The method we developed to calculate activity-specific metabolic rates can be applied to terrestrial and marine species to determine the energetic costs of daily activities, as well as to predict the energetic consequences for animals forced to change their time allocations in response to environmental shifts.

keywords     Antarctic fur seal, Arctocephalus gazella, Callorhinus ursinus, diving, energy expenditure, foraging, metabolic rate, northern fur seal, time-activity budget
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Combining hard-part and DNA analyses in scats with biologging and stable isotopes can reveal different diet compositions and feeding strategies within a population.
Jeanniard-du-Dot, T., A. C. Thomas, Y. Cherel, A. W. Trites and C. Guinet. 2017.
Marine Ecology Progress Series 584:1-16.
Accurately estimating predators' diets at relevant spatial and temporal scales is key to understanding animals' energetics and fitness, particularly in populations whose decline might be related to their diet such as northern fur seals Callorhinus ursinus. Our goals were to improve the accuracy of diet estimates and extend understanding of feeding ecology by combining 2 scat-based methods of diet determination (hard-part identification and DNA-metabarcoding) with stable isotope measurements and individual behavioural data. We collected 98 scats on a northern fur seal breeding colony. We also tracked 20 females with biologgers, and took blood samples to determine δ13C and δ15N values as proxies for seal foraging habitat and diet. Results show that diet composition from hard-parts analysis corresponded well with DNA results, with DNA yielding a greater diversity of prey species at a finer taxonomic level. Overall, scat-based methods showed that seals mostly fed on neritic shelf-associated prey. Cluster analyses of combined hard-parts and DNA results however identified 2 diet groups, one mostly neritic and the other mostly pelagic. Stable isotopes and behavioural data revealed that 40% of seals fed in oceanic waters on pelagic prey. This is more than indicated by scat-based analyses, which are likely biased towards animals foraging closest to the colony and underestimate some dietary specializations within the population. Consequently, the combination of multiple methods for diet identification with at-sea tracking of individuals can help identify and quantify specialist groups within a population and provide a wider spatial and temporal ecological context for dietary analysis.
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Proxies of energy expenditure for marine mammals: an experimental test of the time trap.
Ladds, M. A., D. A. S. Rosen, D. J. Slip and R. G. Harcourt. 2017.
Scientific Reports 7:11815
Direct measures of energy expenditure are difficult to obtain in marine mammals, and accelerometry may be a useful proxy. Recently its utility has been questioned as some analyses derived their measure of activity level by calculating the sum of accelerometry-based values and then comparing this summation to summed (total) energy expenditure (the so-called 'time trap'). To test this hypothesis, we measured oxygen consumption of captive fur seals and sea lions wearing accelerometers during submerged swimming and calculated total and rate of energy expenditure. We compared these values with two potential proxies of energy expenditure derived from accelerometry data: flipper strokes and dynamic body acceleration (DBA). Total number of strokes, total DBA, and submergence time all predicted total oxygen consumption (sVO2 ml kg−1). However, both total DBA and total number of strokes were correlated with submergence time. Neither stroke rate nor mean DBA could predict the rate of oxygen consumption (sV.O2 ml min−1 kg−1). The relationship of total DBA and total strokes with total oxygen consumption is apparently a result of introducing a constant (time) into both sides of the relationship. This experimental evidence supports the conclusion that proxies derived from accelerometers cannot estimate the energy expenditure of marine mammals.

keywords     energy expenditure, accelerometers, Steller sea lions
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On the utility of accelerometers to predict stroke rate using captive fur seals and sea lions.
Ladds, M. A., D. A. S. Rosen, D. J. Slip and R. G. Harcourt. 2017.
Biology Open 6:1396-1400.
Energy expenditure of free-living fur seals and sea lions is difficult to measure directly, but may be indirectly derived from flipper stroke rate. We filmed 10 captive otariids swimming with accelerometers either attached to a harness (Daily Diary: sampling frequency 32Hz, N = 4) or taped to the fur (G6a+: 25Hz, N = 6). We used down sampling to derive four recording rates from each accelerometer (Daily Diary: 32, 16, 8, 4Hz; G6a+: 25, 20, 10, 5Hz). For each of these sampling frequencies we derived 20 combinations of two parameters (RMW - the window size used to calculate the running mean, and m – the minimum number of points smaller than the local maxima used to detect a peak), from the dynamic acceleration of x, z and x+z, to estimate stroke rate from the accelerometers. These estimates differed by up to ~20% in comparison to the actual number of foreflipper strokes counted from videos. RMW had little effect on the overall differences, nor did the choice of axis used to make the calculations (x, z or x+z), though the variability was reduced when using x+z. The best m varied depending on the axis used and the sampling frequency, where a larger m was needed for higher sampling frequencies. This study demonstrates that when parameters are appropriately tuned, accelerometers are a simple yet valid tool for estimating the stroke rates of swimming otariids.

keywords     otariid, swim mechanics, stroke rate, accelerometer, energetics, biologger
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Physiological constraints and energetic costs of diving behaviour in marine mammals: a review of studies using trained Steller sea lions diving in the open ocean.
Rosen, D.A.S., A.G. Hindle, C. Gerlinsky, E. Goundie, G.D. Hastie and A.W. Trites. 2017.
Journal of Comparative Physiology B 187:29-50.
Marine mammals are characterized as having physiological specializations that maximize the use of oxygen stores to prolong time spent under water. However, it has been difficult to undertake the requisite controlled studies to determine the physiological limitations and trade-offs that marine mammals face while diving in the wild under varying environmental and nutritional conditions. For the past decade, Steller sea lions (Eumetopias jubatus) trained to swim and dive in the open ocean away from the physical confines of pools participated in studies that investigated the interactions between diving behaviour, energetic costs, physiological constraints, and prey availability. Many of these studies measured the cost of diving to understand how it varies with behaviour and environmental and physiological conditions. Collectively, these studies show that the type of diving (dive bouts or single dives), the level of underwater activity, the depth and duration of dives, and the n utritional status and physical condition of the animal affect the cost of diving and foraging. They show that dive depth, dive and surface duration, and the type of dive result in physiological adjustments (heart rate, gas exchange) that may be independent of energy expenditure. They also demonstrate that changes in prey abundance and nutritional status cause sea lions to alter the balance between time spent at the surface acquiring oxygen (and offloading CO2 and other metabolic by-products) and time spent at depth acquiring prey. These new insights into the physiological basis of diving behaviour further our understanding of the potential scope for behavioural responses of marine mammals to environmental changes, the energetic significance of these adjustments, and the consequences of approaching physiological limits.
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Marine mammals exploring the oceans pole to pole: a review of the MEOP consortium.
Treasure, A. M., F. Roquet, I. J. Ansorge, M. N. Bester, L. Boehme, H. Bornemann, J.-B. Charrassin, D. Chevallier, D. P. Costa, M. A. Fedak, C. Guinet, M. O. Hammill, R. G. Harcourt, M. A. Hindell, K. M. Kovacs, M.-A. Lea, P. Lovell, A. D. Lowther, C. Lydersen, T. McIntyre, C. R. McMahon, M. M. C. Muelbert, K. Nicholls, B. Picard, G. Reverdin, A. W. Trites, G. D. Williams and P.J. Nico de Bruyn. 2017.
Oceanography 30:132-138.
Polar oceans are poorly monitored despite the important role they play in regulating Earth's climate system. Marine mammals equipped with biologging devices are now being used to fill the data gaps in these logistically difficult to sample regions. Since 2002, instrumented animals have been generating exceptionally large data sets of oceanographic CTD casts (>500,000 profiles), which are now freely available to the scientific community through the MEOP data portal (http://meop.net). MEOP (Marine Mammals Exploring the Oceans Pole to Pole) is a consortium of international researchers dedicated to sharing animal-derived data and knowledge about the polar oceans. Collectively, MEOP demonstrates the power and cost-effectiveness of using marine mammals as data-collection platforms that can dramatically improve the ocean observing system for biological and physical oceanographers. Here, we review the MEOP program and database to bring it to the attention of the international community.
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Fine-scale foraging movements by fish-eating killer whales (Orcinus orca) relate to the vertical distributions and escape responses of salmonid prey (Oncorhynchus spp.).
Wright, B. M., J. K. B. Ford, G. M. Ellis, V. B. Deecke, A. D. Shapiro, B. C. Battaile and A. W. Trites. 2017.
Movement Ecology 5:1-18.
Background: We sought to quantitatively describe the fine-scale foraging behavior of northern resident killer whales (Orcinus orcas), a population of fish-eating killer whales that feeds almost exclusively on Pacific salmon (Oncorhynchus spp.). To reconstruct the underwater movements of these specialist predators, we deployed 34 biologging Dtags on 32 individuals and collected high-resolution, three-dimensional accelerometry and acoustic data. We used the resulting dive paths to compare killer whale foraging behavior to the distributions of different salmonid prey species. Understanding the foraging movements of these threatened predators is important from a conservation standpoint, since prey availability has been identified as a limiting factor in their population dynamics and recovery. Results: Three-dimensional dive tracks indicated that foraging (N = 701) and non-foraging dives (N = 10,618) were kinematically distinct (Wilks

keywords     Foraging, Movement, Diving behavior, Biologging, Dtag, Accelerometry, Killer whale, Orcinus orca, Pacific salmon
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The Consortium is conducting a long-term research program on Steller sea lions, northern fur seals and other marine mammals and their interactions with fisheries, other species and oceanographic conditions in the North Pacific Ocean and Eastern Bering Sea.

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