Iverson, Sara
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Item Open Access Polar Bear Diets and Arctic Marine Food Webs: Insights from Fatty Acid Analysis(2008-11) Thiemann, Gregory W.; Iverson, Sara J.; Stirling, IanWe used quantitative fatty acid signature analysis (QFASA) to examine the diets of 1738 individual polar bears (Ursus maritimus) sampled across the Canadian Arctic over a 30-year span. Polar bear foraging varied over large and small spatial and temporal scales, and between demographic groups. Diets in every subpopulation were dominated by ringed seals (Phoca hispida) and, in the eastern Arctic, secondarily by harp seals (Pagophilus groenlandica). Beluga whales (Delphinapterus leucas) were an important food source for bears in the High Arctic, which is consistent with previous anecdotal reports. Foraging patterns were most similar among neighboring subpopulations with similar prey assemblages, but also differed geographically within Western Hudson Bay. The sexual size dimorphism of polar bears had an important effect on foraging, as large bearded seals (Erignathus barbatus) and walruses (Odobenus rosmarus) were consumed most often by older, male bears, whereas ringed seals and, where available, harbor seals (P. vitulina) were most important to younger age classes. Larger, older bears also had the greatest dietary diversity, apparently because of their ability to include larger-bodied prey. During spring and summer, polar bears in some areas increased predation on migratory harp seals and beluga whales. In Western Hudson Bay, bearded seal consumption declined between 1995 and 2001 for both male and female bears and continued to decline among females up to the most recent sampling (2004). Ringed seal consumption in Western Hudson Bay increased between 1998 and 2001, perhaps in response to increased ringed seal productivity, but was not significantly affected by date of sea-ice breakup. Overall, our data indicate that polar bears are capable of opportunistically altering their foraging to take advantage of locally abundant prey, or to some degree compensating for a decline in a dominant prey species. However, in other areas polar bears are dependent on the availability of ringed and bearded seals. Recent population data suggest that polar bears with the most specialized diets may be most vulnerable to climate-related changes in ice conditions. The results of this large-scale, ecosystem-based study indicate a complex relationship between sea-ice conditions, prey population dynamics, and polar bear foraging.Item Open Access Development of the blood and muscle oxygen stores in gray seals (Halichoerus grypus): Implications for juvenile diving capacity and the necessity of a terrestrial postweaning fast(2005-07) Noren, SR; Iverson, Sara J.; Boness, DJTo successfully transition from nursing to foraging, phocid seal pups must develop adequate diving physiology within the limited time between birth and their first independent foraging trip to sea. We studied the postpartum development of oxygen stores in gray seals (Halichoerus grypus, n = 40) to better understand the ontogeny of diving capacity in phocids. Hemoglobin (Hb), hematocrit (Hct), blood volume (BV), and myoglobin (Mb) levels in newborn (3 d postpartum [DPP]) and newly weaned (DPP) pups were among the lowest 17 +/- 0.4 measured across age classes. During the pups' terrestrial postweaning fast (PWF), Hb, Hct, mass-specific BV, and Mb increased by 28%, 21%, 13%, and 29%, respectively, resulting in a 35% increase in total body mass-specific oxygen stores and a 23% increase in calculated aerobic dive limit (CADL). Although Hb and Hct levels at the end of the PWF were nearly identical to those of yearlings, total body mass-specific oxygen stores and CADL of weaned pups departing for sea were only 66% - 67% and 32% - 62%, respectively, of those for yearlings and adult females. The PWF represents an integral component of the physiological development of diving capacity in phocids; however, newly independent phocids still appear to have limited diving capabilities at the onset of foraging.Item Open Access Seasonal changes in buoyancy and diving behaviour of adult grey seals(2000-08) Beck, Carrie A.; Bowen, W. Don; Iverson, Sara J.Phocid seals go through dramatic seasonal changes in body mass and composition as a result of the spatial and temporal separation of foraging, reproduction and moulting. These changes in body fat content and body mass result in seasonal changes in buoyancy, which in turn may influence diving behaviour. We examined the longitudinal changes in buoyancy and diving behaviour of 14 adult grey seals (Halichoerus grypus) during two periods that represent maximal contrast in body mass and composition. During both the post-moulting (PM) and pre-breeding (PB) periods, grey seals were negatively buoyant. However, buoyancy increased by 47.9% between the PM and PB periods. Descent rate was significantly faster during the PM period (1.0+-0.07 m s-1) than during the PB period (0.7+-0.06 m s-1), suggesting that seals were aided by negative buoyancy during the downward portion of dives. Ascent rate was also significantly faster during the PM period (0.8+-0.06 m s-1) than during the PB period (0.6+-0.05 m s-1), contradicting the prediction that more buoyant animals should ascend faster. The effects of drag could not account for this discrepancy. Dive depth and surface interval between dives did not differ significantly between the two periods. Similarly, the distribution of dive shapes used by individuals did not differ between the two periods. However, dive duration was significantly longer during the PB period than during the PM period (5.5+-0.25 min compared with 4.4+-0.24 min, respectively) as was time spent at the bottom of the dive (3.1+-0.22 min compared with 2.5+-0.15 min, respectively).Item Open Access Weaning mass affects changes in body composition and food intake in harbour seal pups during the first month of independence(2003-05) Muelbert, MMC; Bowen, WD; Iverson, Sara J.In phocid seals, the transition to nutritional independence is abrupt, with females abandoning their offspring after weaning and returning to sea. We hypothesized that body size at weaning may play an important role in the nature of this transition. We studied the changes in body composition and water flux of newly weaning harbour seals over the first 4-6 wk postweaning. Thirty-three pups were dosed with deuterium oxide to estimate total body water (TBW) and a subset of 24 was dosed twice to estimate changes in body composition and water flux. All pups lost body mass over the study period, but TBW increased during the period of mass loss, indicating continued lean tissue growth. Combined data from this and our early study indicated that heavy (>median mass) pups were relatively fatter (41.0% vs. 37.1%) and had significantly greater total body energy at weaning than did light (less than or equal tomedian mass) pups. Percentage TBW declined linearly over time in light pups but was constant in heavy pups for the first 19 d postweaning and then declined linearly. Both the temporal pattern and composition of mass loss differed between light and heavy pups. Estimated food intake increased in the second 2 wk of study compared to the first 2 wk, in both light and heavy pups, reflecting increased foraging success but at levels still insufficient to meet daily expenditures of most individuals.Item Open Access Body condition at weaning affects the duration of the postweaning fast in gray seal pups (Halichoerus grypus)(2008-05) Noren, Shawn R.; Boness, Daryl J.; Iverson, Sara J.; McMillan, Jim; Bowen, W. DonGray seals (Halichoerus grypus) undergo a terrestrial postweaning fast (PWF) that depletes energy reserves acquired during the suckling interval. Plasticity in PWF duration may ensure that pups of variable body condition depart for sea with adequate energy reserves. To test this hypothesis, we examined body condition of 30 gray seal pups at weaning and monitored their PWF duration. On average, fat accounted for 47.3% +/- 0.7% of their 53.2 +/- 1.3-kg weaning mass. Although fasting duration averaged 21 +/- 1.1 d (n = 28), there was considerable variation in fasting duration (9 to >31 d) and the resulting age when pups departed to sea (26 to >49 d). Percent fat at weaning (38.6%-54.6%) was positively correlated with fasting duration (n = 28, r = 0.376, P = 0.0489). In contrast, total body gross energy (735.3-1,447.4 MJ) and body mass (39.0-66.0 kg) were not correlated with fasting duration. Thus, body composition, not overall body reserves, predicted fasting duration, but the effect was weak, indicating that other factors also account for the observed variation in fasting duration. We speculate that pups with greater percent fat more effectively utilized lipid and conserved protein while meeting metabolic costs throughout the PWF. As a result, fatter pups extended the PWF duration, which may be critical for development of diving physiology and may have facilitated their survivorship to age 1.Item Open Access Primiparous and multiparous females differ in mammary gland alveolar development: implications for milk production(2012-08) Lang, Shelley L. C.; Iverson, Sara J.; Bowen, W. DonMammary gland capacity is influenced by the number of secretory cells in the gland, the activity of those cells and the size and arrangement of the alveoli that they form. Although reproductive experience has been shown to affect the total number of secretory cells in the gland, its potential effect on the structural development of lobulo-alveolar tissues has not been directly investigated. To examine whether reproductive experience affects lobulo-alveolar development, we took mammary gland biopsies at early and peak lactation from primiparous and multiparous grey seal (Halichoerus grypus) females and used histological techniques to compare cell density, alveolar density and alveolar size within secretory lobules. Primiparous females had a significantly higher cell density compared with multiparous females throughout lactation, suggesting that primiparous females have smaller, less-developed secretory cells. Primiparous females had a significantly smaller average alveolar size compared with multiparous females throughout lactation. Although alveolar density was higher in primiparous females compared with multiparous females at early lactation, there was no significant difference between the groups at peak lactation. These results suggest that the mammary gland of primiparous females may have both a lower secretory capacity and a lower storage capacity on a relative basis than those of multiparous females and demonstrate, for the first time, that reproductive experience has a significant effect on both the rate and pattern of mammary gland alveolar development and, potentially, on a female's capacity for milk production.Item Open Access The Influence of Reproductive Experience on Milk Energy Output and Lactation Performance in the Grey Seal (Halichoerus grypus)(2011-05) Lang, Shelley L. C.; Iverson, Sara J.; Bowen, W. DonAlthough evidence from domestic and laboratory species suggests that reproductive experience plays a critical role in the development of aspects of lactation performance, whether reproductive experience may have a significant influence on milk energy transfer to neonates in wild populations has not been directly investigated. We compared maternal energy expenditures and pup growth and energy deposition over the course of lactation between primiparous and fully-grown, multiparous grey seal (Halichoerus grypus) females to test whether reproductive experience has a significant influence on lactation performance. Although there was no difference between primiparous females in milk composition and, thus, milk energy content at either early or peak lactation primiparous females had a significantly lower daily milk energy output than multiparous females indicating a reduced physiological capacity for milk secretion. Primiparous females appeared to effectively compensate for lower rates of milk production through an increased nursing effort and, thus, achieved the same relative rate of milk energy transfer to pups as multiparous females. There was no difference between primiparous and multiparous females in the proportion of initial body energy stores mobilised to support the costs of lactation. Although primiparous females allocated a greater proportion of energy stores to maternal maintenance versus milk production than multiparous females, the difference was not sufficient to result in significant differences in the efficiency of energy transfer to pups. Thus, despite a lower physiological capacity for milk production, primiparous females weaned pups of the same relative size and condition as multiparous females without expending proportionally more energy. Although reproductive experience does not significantly affect the overall lactation performance of grey seals, our results suggest that increases in mammary gland capacity with reproductive experience may play a significant role in the age-related increases in neonatal growth rates and weaning masses observed in other free-ranging mammals.Item Open Access Repeatability in lactation performance and the consequences for maternal reproductive success in gray seals(2009-09) Lang, S. L. C.; Iverson, Sara J.; Bowen, W. D.In mammals, the most significant maternal effect on offspring growth during lactation is the ability of females to efficiently transfer milk energy to their neonates. However, despite the importance of the transfer of milk energy to both maternal and offspring fitness, nothing is known about the extent to which variation among females may be attributed to differences in individual quality or environmental variation in natural populations. We measured repeatability over multiple lactation periods in components of lactation performance in free-ranging, multiparous gray seal (Halichoerus grypus) females to examine to what extent variation among females in pup weaning mass may be attributed to inherent differences in their physiological capacity to deliver milk energy. Levels of repeatability were high for milk composition (r = 0.38-0.50), daily milk output (r = 0.46), and the duration of lactation (r = 0.57), demonstrating that there are consistent differences among females in these characters across lactations and that the overall capacity of gray seal females to deliver milk energy to their pups is characteristic of individuals. The repeatability in pup weaning mass (r = 0.48) was consistent with the values for the components of total milk energy output and suggests that, over a large proportion of their reproductive life, individual gray seal females will consistently wean pups with greater or lesser probabilities of survival. Our results suggest that inherent differences among females in their physiological capacity to deliver milk energy may be an important component of variation in individual quality and, thus, lifetime reproductive success in mammals. High levels of repeatability also suggest that components of milk energy transfer may have a significant heritable genetic basis.Item Open Access Animal-Borne Acoustic Transceivers Reveal Patterns of at-Sea Associations in an Upper-Trophic Level Predator(2012-11) Lidgard, Damian C.; Bowen, W. Don; Jonsen, Ian D.; Iverson, Sara J.Satellite telemetry data have substantially increased our understanding of habitat use and foraging behaviour of upper-trophic marine predators, but fall short of providing an understanding of their social behaviour. We sought to determine whether novel acoustic and archival GPS data could be used to examine at-sea associations among grey seals (Halichoerus grypus) during the fall foraging period. Fifteen grey seals from Sable Island, Canada were deployed with Vemco Mobile Transceivers and Satellite-GPS transmitters in October 2009, 13 of which were recaptured and units retrieved 79 +/- 2.3 days later during the following breeding season, December 2009-January 2010. An association between two individuals was defined as a cluster of acoustic detections where the time between detections was <30 min. Bathymetry, travel rate, and behavioural state (slow and fast movement) were determined for each GPS archival point (3.7 +/- 0.1 locations recorded per hour). Behavioural state was estimated using a hidden Markov model. All seals had been involved in associations with other instrumented seals while at sea, with a total of 1,872 acoustic detections recorded in 201 associations. The median number of detections per association was 3 (range: 1-151) and the median duration of an association was 0.17 h (range: <0.1-11.3 h). Linear mixed-effects models showed that associations occurred when seals were exhibiting slow movement (0.24 +/- 0.01 ms(-1)) on shallow (53.4 +/- 3.7 m) offshore banks where dominant prey is known to occur. These results suggest the occurrence of short-term associations among multiple individuals at foraging grounds and provide new insights into the foraging ecology of this upper-trophic marine predator.Item Open Access Variation in milk production and lactation performance in grey seals and consequences for pup growth and weaning characteristics(1999-12) Mellish, JAE; Iverson, Sara J.; Bowen, WDPhocid seals are one of the few groups of mammals capable of sustaining the energetic demands of lactation entirely through body nutrient stores while fasting. Lactation performance of the female in turn influences the rate and pattern of pup growth. We examined variation in and patterns of milk composition and production, maternal energy output, and pup growth and energy deposition over the entire lactation period in 18 grey seal mother-pup pairs using hydrogen isotope ((H2O)-H-3 and D2O) dilution. Milk composition was independent of maternal mass and nutrient stores, indicating dependence on other physiological and genetic factors. Heavier females lactated longer (r(2) = 0.653, P < 0.001), had higher total milk outputs (r(2) = 0.652, P < 0.001), and produced larger pups at weaning (r(2) = 0.417, P = 0.005). While fatter females lactated for longer periods of time (r(2) = 0.595, P < 0.001), females with a larger lean body mass at parturition produced more milk (r(2) = 0.579, P < 0.001). Total milk energy output was the strongest predictor of pup weaning mass, which, along with the pup's efficiency of energy storage, accounted for 91% of the variation in weaning mass. Nevertheless, there was sufficient plasticity in milk composition and energy output that some smaller females produced relatively large pups. Few females appeared to deplete body nutrients to the point where it might limit the duration of lactation.Item Open Access Effect of a low-fat diet on body composition and blubber fatty acids of captive juvenile harp seals (Phoca groenlandica)(2000-01) Kirsch, PE; Iverson, Sara J.; Bowen, WDWe investigated the effects of a change from a high-fat diet to a low-fat diet of differing fatty acid (FA) composition on the body composition and blubber FA of five captive juvenile harp seals. Seals that had been maintained for 1 yr on a diet of Atlantic herring (greater than or equal to 9% fat) were switched to a diet of Atlantic pollock (1.7% fat) for 30 d. On days 0, 14, and 30, mass and body composition (using isotope dilution) were measured, and blubber biopsies (5 cm x 6 mm) were taken for FA analysis. Fat accounted for 38%-49% of body mass at the start of the experiment. When switched to the pollock diet, and despite food intakes averaging 6.5 kg/d (32.3 MJ/d), body fat: declined by an average of 6.4 kg or by 32% over the 30-d experiment. In contrast, body protein increased in direct relation to protein intake (r(2) = 0.836, P = 0.030). Despite substantial loss of body fat, blubber FA signature changed significantly to reflect the changes in dietary intake of FA, and the deposition of FA was quantifiably predictable. Our results suggest that young growing phocids are unable to maintain body fat stores on low-fat diets even when protein intakes are high. This may have significant implications for juvenile pinniped survival in the wild. In addition, turnover and deposition of dietary FA in blubber takes place in nonfattening seals.Item Open Access Jellyfish Support High Energy Intake of Leatherback Sea Turtles (Dermochelys coriacea): Video Evidence from Animal-Borne Cameras(2012-03) Heaslip, Susan G.; Iverson, Sara J.; Bowen, W. Don; James, Michael C.The endangered leatherback turtle is a large, highly migratory marine predator that inexplicably relies upon a diet of low-energy gelatinous zooplankton. The location of these prey may be predictable at large oceanographic scales, given that leatherback turtles perform long distance migrations (1000s of km) from nesting beaches to high latitude foraging grounds. However, little is known about the profitability of this migration and foraging strategy. We used GPS location data and video from animal-borne cameras to examine how prey characteristics (i.e., prey size, prey type, prey encounter rate) correlate with the daytime foraging behavior of leatherbacks (n = 19) in shelf waters off Cape Breton Island, NS, Canada, during August and September. Video was recorded continuously, averaged 1: 53 h per turtle (range 0: 08-3: 38 h), and documented a total of 601 prey captures. Lion's mane jellyfish (Cyanea capillata) was the dominant prey (83-100%), but moon jellyfish (Aurelia aurita) were also consumed. Turtles approached and attacked most jellyfish within the camera's field of view and appeared to consume prey completely. There was no significant relationship between encounter rate and dive duration (p = 0.74, linear mixed-effects models). Handling time increased with prey size regardless of prey species (p = 0.0001). Estimates of energy intake averaged 66,018 kJ.d(-1) but were as high as 167,797 kJ.d(-1) corresponding to turtles consuming an average of 330 kg wet mass.d(-1) (up to 840 kg.d(-1)) or approximately 261 (up to 664) jellyfish.d(-1). Assuming our turtles averaged 455 kg body mass, they consumed an average of 73% of their body mass.d(-1) equating to an average energy intake of 3-7 times their daily metabolic requirements, depending on estimates used. This study provides evidence that feeding tactics used by leatherbacks in Atlantic Canadian waters are highly profitable and our results are consistent with estimates of mass gain prior to southward migration.Item Open Access Metabolism of dietary cetoleic acid (22 : 1n-11) in mink (Mustela vison) and gray seals (Halichoerus grypus) studied using radiolabeled fatty acids(2006-07) Cooper, Margaret H.; Iverson, Sara J.; Rouvinen-Watt, KirstiCetoleic acid (22:1n-11) is a good indicator of diet in marine predators and has proven to be an important fatty acid (FA) when using adipose tissue FA composition to study diet in marine mammals and seabirds. Feeding studies have shown that 22:1 isomers are predictably underrepresented in adipose tissue relative to diet, implying that metabolism within the predator strongly influences the relationship between the level of these FAs in diet and adipose tissue. Fully understanding such metabolic processes for individual FAs is important for the quantitative estimation of predator diets. We employed a dual-label radioisotope tracer technique to investigate the potential modification of 22:1n-11 and its recovery in the blubber of gray seals (Halichoerus grypus) and in the adipose tissue and liver of mink (Mustela vison), a smaller model carnivore also accustomed to fish-based diets. In both seals and mink, H-3 radioactivity was found in the chain-shortened products of 22:1n-11, with 18:1 being the dominant product. We also found H-3 radioactivity in saturated FAs. The distribution patterns of H-3 radioactivity across the FAs isolated from seal blubber and mink subcutaneous adipose tissue were comparable, indicating that mink are a good model for the investigation of lipid metabolism in marine carnivores.Item Open Access Interspecific and intraspecific variation in proximate, mineral, and fatty acid composition of milk in old world fruit bats (Chiroptera : Pteropodidae)(2001-01) Hood, WR; Kunz, TH; Oftedal, OT; Iverson, Sara J.; LeBlanc, D.; Seyjagat, J.We examine the effect of body mass on milk composition among Old World fruit bats, including Pteropus pumilus (0.175 kg), Pteropus rodricensus (0.265 kg), Pteropus hypomelanus (0.571 kg), and Pteropus vampyrus (1.133 kg). We describe intra- and interspecific differences in the proximate composition of milk among these four species and the minerals and fatty acids in the milk of the latter two species. There were no differences between species in the concentrations of dry matter, fat, or lactose in milk. However, there were significant, although small, differences in the protein content of milk among species, with protein being significantly greater in P. rodricensus than in P. pumilus and P. hypomelanus and protein being significantly less in P. hypomelanus than in P. rodricensus and P. vampyrus. There were no differences in mineral content between P. hypomelanus and P. vampyrus in milk minerals, but minor differences were evident in fatty acids 12:0, 14:0, 18:0, 18:1n11, and 18:2n6. Our findings suggest that milk composition is relatively constant across lactation for most proximate, mineral, and fatty acid components. We found a significant increase in dry matter and energy across lactation in the concentration of dry matter and energy in P. pumilus and fat in P. hypomelanus. In P. hypomelanus, we found a significant increase in the concentration of fatty acids 10:0 and 20:1n9 and a significant decrease in Iso15 and 20:1n7. No other differences associated with day of lactation were found. These findings suggest that milk composition is generally similar within the genus Pteropus, despite a 6.5-fold difference in body mass between species that we evaluated.Item Open Access Quantitative fatty acid signature analysis: A new method of estimating predator diets(2004-05) Iverson, Sara J.; Field, Chris; Bowen, W. Don; Blanchard, WadeAccurate estimates of the diets of predators are required in many areas of ecology, but for many species current methods are imprecise, limited to the last meal, and often biased. The diversity of fatty acids and their patterns in organisms, coupled with the narrow limitations on their biosynthesis, properties of digestion in monogastric animals, and the prevalence of large storage reservoirs of lipid in many predators; led us to propose the use of quantitative fatty acid signature analysis (QFASA) to study predator diets. We present a statistical model that provides quantitative estimates of the proportions of prey specie's in the diets of individual predators using fatty acid signatures. We conducted simulation studies using a database of 28; prey species (n = 954 individuals) from the Scotian Shelf off eastern Canada to investigate properties of the model and to evaluate the reliability with which prey could be distinguished in the model. We then conducted experiments on. grey seals (Halichoerus grypus, n = 25) and harp seals (Phoca groenlandica, n = 5) to assess quantitative characteristics of fatty acid deposition and to develop calibration coefficients for individual fatty acids to account for predator lipid metabolism, We then tested the model and calibration coefficients by estimating the diets of. experimentally fed captive grey seals (n = 6, switched from herring to a mackerel/capelin diet) and mink kits (Mustela vison, n = 46, switched from milk to one of three oil-supplemented diets). The diets of all experimentally fed animals were generally well estimated using QFASA and were consistent with qualitative and quantitative expectations, provided that appropriate calibration coefficients were used. In a final case, we compared video data of foraging by individual free-ranging harbor seals (Phoca vitulina, n = 23) fitted with Crittercams and QFASA estimates of the diet of those same seals using a complex ecosystem-wide prey database. Among the 28 prey species in the database, QFASA estimated sandlance to be the dominant prey species in the diet of all seals (averaging 62% of diet), followed primarily by flounders, but also capelin and minor amounts of other species, although there was also considerable individual variability among seals. These estimates were consistent with video data showing sandlance to be the predominant prey, followed by flatfish. We conclude that QFASA provides estimates of diets for individuals at time scales that are relevant to the ecological.. processes affecting survival, and can be used to study diet variability within individuals over time, which will provide important opportunities rarely possible with other indirect methods. We propose that the QFASA model we have set forth will be applicable to a wide range of predators and ecosystems.Item Open Access Linking movement, diving, and habitat to foraging success in a large marine predator(2006-12) Austin, Deborah; Bowen, W. Don; McMillan, Jim I.; Iverson, Sara J.Establishing where and when predators forage is essential to understanding trophic interactions, yet foraging behavior remains poorly understood in large marine carnivores. We investigated the factors leading to foraging success in gray seals (Halichoerus grypus) in the Northwest Atlantic in the first study to use simultaneous deployments of satellite transmitters, time depth recorders, and stomach-temperature loggers on a free-ranging marine mammal. Thirty-two seals were each fitted with the three types of instrumentation; however, complete records from all three instruments were obtained from only 13 individuals, underscoring the difficulty of such a multi-instrument approach. Our goal was to determine the characteristics of diving, habitat, and movement that predict feeding. We linked diving behavior to foraging success at two temporal scales: trips ( days) and bouts ( hours) to test models of optimal diving, which indicate that feeding can be predicted by time spent at the bottom of a dive. Using an information-theoretic approach, a Generalized Linear Mixed Model with trip duration and accumulated bottom time per day best explained the number of feeding events per trip, whereas the best predictor of the number of feeding events per bout was accumulated bottom time. We then tested whether characteristics of movement were predictive of feeding. Significant predictors of the number of feeding events per trip were angular variance (i.e., path tortuosity) and distance traveled per day. Finally, we integrated measures of diving, movement, and habitat at four temporal scales to determine overall predictors of feeding. At the 3-h scale, mean bottom time and distance traveled were the most important predictors of feeding frequency, whereas at the 6-h and 24-h time scales, distance traveled alone was most important. Bathymetry was the most significant predictor of feeding at the 12-h interval, with feeding more likely to occur at deeper depths. Our findings indicate that several factors predict feeding in gray seals, but predictor variables differ across temporal scales such that environmental variation becomes important at some scales and not others. Overall, our results illustrate the value of simultaneously recording and integrating multiple types of information to better understand the circumstances leading to foraging success.Item Open Access The energetics of male reproduction in an acquatically mating pinniped, the harbour seal(1998-07) Coltman, DW; Bowen, WD; Iverson, Sara J.; Boness, DJThe energy expenditure of breeding male harbour seals, Phoca vitulina, on Sable Island, Nova Scotia, was investigated by measuring changes in body mass, body composition, and water flux using isotope dilution. Seals lost 0.47% +/- 0.04% (n = 34) of their initial mass per day during the breeding season (4 wk), and fat, water, and protein accounted for 64.3% +/- 4.8%, 27.8% +/- 3.3%, and 6.9% +/- 1.4% of this mass loss, respectively (n = 31). Total energy expenditure was estimated as 33.3 +/- 1.9 MJ d(-1), or 3.9 +/- 0.2 W kg(-1) (n = 17), similar to rates measured in terrestrially mating pinniped species. However, unlike terrestrially mating pinnipeds, male harbour seals did not fast during the breeding season, and energy intake from foraging accounted for 61.8% +/- 4.0% of the total energy expended. Males derived most of their expended energy from food intake early in. the breeding season. However, as oestrus females became increasingly available, reduced rates of food intake in males were coupled with increased rates of total energy expenditure. Larger males expended significantly more energy from body stores and more total energy than smaller males. Male harbour seals appeared to balance the energetic costs of reproduction against the constraints of small body size by foraging during deep-diving trips before the appearance of oestrus females and by opportunistic feeding throughout the breeding season while at sea. We suggest that size dimorphism may be less pronounced in aquatically mating pinnipeds partly because the temporal and spatial separation of foraging and reproduction is less distinct than it is for terrestrially breeding pinnipeds.Item Open Access Demonstration of the deposition and modification of dietary fatty acids in pinniped blubber using radiolabelled precursors(2004-07) Budge, SM; Cooper, MH; Iverson, Sara J.Radioisotopes are commonly used to study the in vivo metabolism and deposition of dietary fatty acids in adipose tissue. The application of this approach to pinnipeds is problematic because of their large mass and blubber fat content. We have developed a method where labelled lipids can be fed to seals at financially feasible levels, with the radioactivity in individual fatty acids isolated from blubber detected with standard laboratory equipment. A combination of techniques including argentation thin layer chromatography, high performance liquid chromatography with ultraviolet detection, and independent liquid scintillation counting were employed. Juvenile gray seals (Halichoerus grypus) were fed either 0.5 mCi H-3-labelled triolein (18:1n-9, n=2) or palmitic acid (16:0, n=2). Blubber samples were taken 12 h later, and the radioactivity in individual fatty acids was determined. Radioactivity was detected in only 18:1 from the animals fed H-3-labelled triolein, indicating direct deposition without modification. Both animals fed H-3-labelled palmitic acid showed clear peaks of radioactivity in 16:0; however, there was also significant activity (23%-29%) found in the desaturation product 16:1. Our results demonstrate that this method is sufficiently sensitive to track the deposition of labelled dietary lipids as well as modification products of ingested fatty acids and will be important in the application of fatty acid signatures to study predator diets.Item Open Access Estimation of total body water in pinnipeds using hydrogen-isotope dilution(1998-05) Bowen, WD; Iverson, Sara J.No abstract available.