Validation of quantitative fatty acid signature analysis for estimating the diet composition of free-ranging killer whales.

Accurate diet estimates are necessary to assess trophic interactions and food web dynamics in ecosystems, particularly for apex predators like cetaceans, which can regulate entire food webs. Quantitative fatty acid analysis (QFASA) has been used to estimate the diets of marine predators in the last decade but has yet to be implemented on free-ranging cetaceans, from which typically only biopsy samples containing outer blubber are available, due to a lack of empirically determined calibration coefficients (CCs) that account for fatty acid (FA) metabolism. Here, we develop and validate QFASA for killer whales using full blubber from managed-care and free-ranging individuals. First, we compute full, inner, and outer blubber CCs from the FA signatures across the blubber layers of managed-care killer whales and their long-term diet items. We then run cross-validating simulations on the managed-care individuals to evaluate the accuracy of diet estimates by comparing full-depth and depth-specific estimates to true diets. Finally, we apply these approaches to subsistence-harvested killer whales from Greenland to test the utility of the method for free-ranging killer whales, particularly for the outer blubber. Accurate diet estimates for the managed-care killer whales were only achieved using killer whale-specific and blubber-layer-specific CCs. Modeled diets for the Greenlandic killer whales largely consisted of seals (75.9 ± 4.7%) and/or fish (20.4 ± 2.4%), mainly mackerel, which was consistent with stomach content data and limited literature on this population. Given the remote habitats and below surface feeding of most cetaceans, this newly developed cetacean-specific QFASA method, which can be applied to outer-layer biopsies, offers promise to provide a significant new understanding of diet dynamics of free-ranging odontocetes and perhaps other cetacean species throughout the world's oceans.

Exploring causal components of plasticity in grey seal birthdates: Effects of intrinsic traits, demography, and climate.

Change in breeding phenology is often a response to environmental forcing, but less is known of the mechanism underlying such changes and their fitness consequences. Here, we report on changes in the breeding phenology from a 27-year longitudinal study (1991-2017) of individually marked, known-aged grey seals (Halichoerus grypus) on Sable Island, Nova Scotia, Canada. We used generalized linear mixed models and a 3-step process to develop a model that includes interactions between intrinsic and extrinsic covariates and to test hypotheses about the influence of fixed factors (maternal age, parity, previous reproductive success, pup sex, colony density, Atlantic Multidecal Oscillation (AMO), North Atlantic Oscillation (NAO), and Sea Surface Temperature) and a random factor (female identity) on parturition dates. We also examined the consequences of the shift in birthdates on maternal energy allocation in offspring as measured by pup weaning mass. Birthdates were known for 2,768 pups of 660 known-age females. For 494 females with ≥2 parturition dates, repeatability as measured by the intraclass correlation was high (mean = 0.66). 87% of the variation in birthdates was explained by a mixed-effects model that included intrinsic and extrinsic fixed effects. Most of the explained variation was associated with the random effect of female identity. Parity was the most important intrinsic fixed effect, with inexperienced mothers giving birth later in the season than multiparous females. Over almost 3 decades, mean birthdates advanced by 15 days. The mixed model with intrinsic effects and population size, the detrended AMO from the previous year and mean NAO in the previous 3 years explained 80% of the variation with 21% of variation from the fixed effects. Both primiparous and multiparous individuals responded to the climate forcing, and there was strong evidence for heterogeneity in the response. Nevertheless, the shift in birthdates did not impact pup weaning mass.

Seasonal occupancy and connectivity amongst nearshore flats and reef habitats by permit Trachinotus falcatus: considerations for fisheries management.

We used acoustic telemetry to quantify permit Trachinotus falcatus habitat use and connectivity in proximity to the Florida Keys, USA, and assessed these patterns relative to current habitat and fisheries management practices. From March 2017 to June 2018, 45 permit tagged within 16 km of the lower Florida Keys were detected at stationary acoustic receivers throughout the south Florida region, the majority of which remained within the Special Permit Zone, where more extensive fisheries harvest regulations are implemented. There was a high level of connectivity between nearshore flats (i.e., <3 m water depth) and the Florida reef tract (FRT; 15-40 m water depth), with 75% of individuals detected in both habitats. These locations probably function primarily as foraging and spawning habitats, respectively. Permit occupancy on the FRT peaked during the months of March-September, with the highest number of individuals occurring there in April and May. Specific sites on the FRT were identified as potentially important spawning locations, as they attracted a high proportion of individuals that exhibited frequent visits with high residency durations. There were also significant positive relationships between seasonal habitat-use metrics on the FRT and an empirical permit gonadosomatic index. Large aggregations of permit at spawning sites on the FRT are potentially vulnerable to the effects of fishing (including predation during catch and release) at a critical point in their life cycle. These data on permit space use and movement, coupled with knowledge of stressors on their ecology, provide insights for implementing science-based strategic management plans.

Egg size is independent of variation in pre-breeding feather corticosterone in Cassin's auklets during favorable oceanographic conditions.

The measurement of corticosterone levels in feathers (fCort) is gaining recognition as an effective means for describing links between stages of the annual cycle in birds. Many seabirds are especially good models for exploring these links, or carryover effects, due to their migratory behavior and reproductive investment in a single-egg clutch. Here, we measure fCort in Cassin's auklet (Ptychoramphus aleuticus) breast feathers at two colonies in British Columbia during 2011, a year of favorable oceanographic conditions, and examine its relationship with egg size. These feathers are grown at sea during the late winter period, 1-2 months prior to egg laying. Assuming that fCort provides some measure of nutritional stress and hypothalamic-pituitary-adrenal axis activity during feather growth, we predicted that fCort would be positively correlated with egg size via increased support for foraging activity during the nutritionally demanding molt stage. We also analyzed the δ13C and δ15N stable isotope content of breast feathers, measures commonly used to characterize seabird diet composition. Contrary to prediction, neither fCort nor stable isotope ratios were good predictors of egg size. Our results appear to conflict with two previous studies on alcids in which fCort and stable isotopes showed clear links with egg size; however, both studies were conducted in years when oceanographic and foraging conditions were poor. Under these conditions, upregulation of corticosterone may be needed to mediate the energetic tradeoffs between self-maintenance and reproduction, supporting increased foraging effort and thus increasing both the likelihood of reproduction and large egg size. However, when foraging conditions are favorable, we suggest that such tradeoffs and associated physiological constraints are minimized and regulation of egg size may be effectively independent of circulating corticosterone levels and diet type.

A novel approach to quantifying the spatiotemporal behavior of instrumented grey seals used to sample the environment.

BACKGROUND: Paired with satellite location telemetry, animal-borne instruments can collect spatiotemporal data describing the animal's movement and environment at a scale relevant to its behavior. Ecologists have developed methods for identifying the area(s) used by an animal (e.g., home range) and those used most intensely (utilization distribution) based on location data. However, few have extended these models beyond their traditional roles as descriptive 2D summaries of point data. Here we demonstrate how the home range method, T-LoCoH, can be expanded to quantify collective sampling coverage by multiple instrumented animals using grey seals (Halichoerus grypus) equipped with GPS tags and acoustic transceivers on the Scotian Shelf (Atlantic Canada) as a case study. At the individual level, we illustrate how time and space-use metrics quantifying individual sampling coverage may be used to determine the rate of acoustic transmissions received. RESULTS: Grey seals collectively sampled an area of 11,308 km (2) and intensely sampled an area of 31 km (2) from June-December. The largest area sampled was in July (2094.56 km (2)) and the smallest area sampled occurred in August (1259.80 km (2)), with changes in sampling coverage observed through time. CONCLUSIONS: T-LoCoH provides an effective means to quantify changes in collective sampling effort by multiple instrumented animals and to compare these changes across time. We also illustrate how time and space-use metrics of individual instrumented seal movement calculated using T-LoCoH can be used to account for differences in the amount of time a bioprobe (biological sampling platform) spends in an area.

ECOLOGY. Aquatic animal telemetry: A panoramic window into the underwater world.

The distribution and interactions of aquatic organisms across space and time structure our marine, freshwater, and estuarine ecosystems. Over the past decade, technological advances in telemetry have transformed our ability to observe aquatic animal behavior and movement. These advances are now providing unprecedented ecological insights by connecting animal movements with measures of their physiology and environment. These developments are revolutionizing the scope and scale of questions that can be asked about the causes and consequences of movement and are redefining how we view and manage individuals, populations, and entire ecosystems. The next advance in aquatic telemetry will be the development of a global collaborative effort to facilitate infrastructure and data sharing and management over scales not previously possible.

Offspring size at weaning affects survival to recruitment and reproductive performance of primiparous gray seals.

Offspring size affects survival and subsequent reproduction in many organisms. However, studies of offspring size in large mammals are often limited to effects on juveniles because of the difficulty of following individuals to maturity. We used data from a long-term study of individually marked gray seals (Halichoerus grypus; Fabricius, 1791) to test the hypothesis that larger offspring have higher survival to recruitment and are larger and more successful primiparous mothers than smaller offspring. Between 1998 and 2002, 1182 newly weaned female pups were branded with unique permanent marks on Sable Island, Canada. Each year through 2012, all branded females returning to the breeding colony were identified in weekly censuses and a subset were captured and measured. Females that survived were significantly longer offspring than those not sighted, indicating size-selective mortality between weaning and recruitment. The probability of female survival to recruitment varied among cohorts and increased nonlinearly with body mass at weaning. Beyond 51.5 kg (mean population weaning mass) weaning mass did not influence the probability of survival. The probability of female survival to recruitment increased monotonically with body length at weaning. Body length at primiparity was positively related to her body length and mass at weaning. Three-day postpartum mass (proxy for birth mass) of firstborn pups was also positively related to body length of females when they were weaned. However, females that were longer or heavier when they were weaned did not wean heavier firstborn offspring.

Local contamination, and not feeding preferences, explains elevated PCB concentrations in Labrador ringed seals (Pusa hispida).

Polychlorinated biphenyls (PCBs) in high trophic level species typically reflect the contributions of myriad sources, such that source apportionment is rarely possible. The release of PCBs by a military radar station into Saglek Bay, Labrador contaminated the local marine food web. For instance, while heavier (higher chlorinated) PCB profiles in some ringed seals (Pusa hispida) were previously attributed to this local source, differences in feeding preferences among seals could not be ruled out as a contributing factor. Herein, similar fatty acid profiles between those seals with 'local' PCB profiles and those with 'long-range' or background profiles indicate little support for the possibility that differential feeding ecologies underlay the divergent PCB profiles. Ringed seals appeared to feed predominantly on zooplankton (Mysis oculata and Themisto libellula), followed by the dusky snailfish (Liparis gibbus), arctic cod (Boreogadus saida), and shorthorn sculpin (Myoxocephalus scorpius). Principal components analysis (PCA) and PCB homolog profiles illustrated the extent of contamination of the Saglek food web, which had very different (and much heavier) PCB profiles than those food web members contaminated by 'long-range' sources. Locally contaminated prey had PCB levels that were higher (2- to 544-fold) than prey contaminated by 'long-range' sources and exceeded wildlife consumption guidelines for PCBs. The application of multivariate analyses to two distinct datasets, including PCB congeners (n=50) and fatty acids (n=65), afforded the opportunity to clearly distinguish the contribution of locally-released PCBs to a ringed seal food web from those delivered via long-ranged transport. Results from the present study strongly suggest that habitat use rather than differences in prey selection is the primary mechanism explaining the divergent PCB patterns in Labrador ringed seals.

Probability of detecting marine predator-prey and species interactions using novel hybrid acoustic transmitter-receiver tags.

Understanding the nature of inter-specific and conspecific interactions in the ocean is challenging because direct observation is usually impossible. The development of dual transmitter/receivers, Vemco Mobile Transceivers (VMT), and satellite-linked (e.g. GPS) tags provides a unique opportunity to better understand between and within species interactions in space and time. Quantifying the uncertainty associated with detecting a tagged animal, particularly under varying field conditions, is vital for making accurate biological inferences when using VMTs. We evaluated the detection efficiency of VMTs deployed on grey seals, Halichoerus grypus, off Sable Island (NS, Canada) in relation to environmental characteristics and seal behaviour using generalized linear models (GLM) to explore both post-processed detection data and summarized raw VMT data. When considering only post-processed detection data, only about half of expected detections were recorded at best even when two VMT-tagged seals were estimated to be within 50-200 m of one another. At a separation of 400 m, only about 15% of expected detections were recorded. In contrast, when incomplete transmissions from the summarized raw data were also considered, the ratio of complete transmission to complete and incomplete transmissions was about 70% for distances ranging from 50-1000 m, with a minimum of around 40% at 600 m and a maximum of about 85% at 50 m. Distance between seals, wind stress, and depth were the most important predictors of detection efficiency. Access to the raw VMT data allowed us to focus on the physical and environmental factors that limit a transceiver's ability to resolve a transmitter's identity.

The sun, moon, wind, and biological imperative-shaping contrasting wintertime migration and foraging strategies of adult male and female northern fur seals (Callorhinus ursinus).

Adult male and female northern fur seals (Callorhinus ursinus) are sexually segregated in different regions of the North Pacific Ocean and Bering Sea during their winter migration. Explanations for this involve interplay between physiology, predator-prey dynamics, and ecosystem characteristics, however possible mechanisms lack empirical support. To investigate factors influencing the winter ecology of both sexes, we deployed five satellite-linked conductivity, temperature, and depth data loggers on adult males, and six satellite-linked depth data loggers and four satellite transmitters on adult females from St. Paul Island (Bering Sea, Alaska, USA) in October 2009. Males and females migrated to different regions of the North Pacific Ocean: males wintered in the Bering Sea and northern North Pacific Ocean, while females migrated to the Gulf of Alaska and California Current. Horizontal and vertical movement behaviors of both sexes were influenced by wind speed, season, light (sun and moon), and the ecosystem they occupied, although the expression of the behaviors differed between sexes. Male dive depths were aligned with the depth of the mixed layer during daylight periods and we suspect this was the case for females upon their arrival to the California Current. We suggest that females, because of their smaller size and physiological limitations, must avoid severe winters typical of the northern North Pacific Ocean and Bering Sea and migrate long distances to areas of more benign environmental conditions and where prey is shallower and more accessible. In contrast, males can better tolerate often extreme winter ocean conditions and exploit prey at depth because of their greater size and physiological capabilities. We believe these contrasting winter behaviors 1) are a consequence of evolutionary selection for large size in males, important to the acquisition and defense of territories against rivals during the breeding season, and 2) ease environmental/physiological constraints imposed on smaller females.

Validation of adipose lipid content as a body condition index for polar bears.

Body condition is a key indicator of individual and population health. Yet, there is little consensus as to the most appropriate condition index (CI), and most of the currently used CIs have not been thoroughly validated and are logistically challenging. Adipose samples from large datasets of capture biopsied, remote biopsied, and harvested polar bears were used to validate adipose lipid content as a CI via tests of accuracy, precision, sensitivity, biopsy depth, and storage conditions and comparisons to established CIs, to measures of health and to demographic and ecological parameters. The lipid content analyses of even very small biopsy samples were highly accurate and precise, but results were influenced by tissue depth at which the sample was taken. Lipid content of capture biopsies and samples from harvested adult females was correlated with established CIs and/or conformed to expected biological variation and ecological changes. However, lipid content of remote biopsies was lower than capture biopsies and harvested samples, possibly due to lipid loss during dart retrieval. Lipid content CI is a biologically relevant, relatively inexpensive and rapidly assessed CI and can be determined routinely for individuals and populations in order to infer large-scale spatial and long-term temporal trends. As it is possible to collect samples during routine harvesting or remotely using biopsy darts, monitoring and assessment of body condition can be accomplished without capture and handling procedures or noninvasively, which are methods that are preferred by local communities. However, further work is needed to apply the method to remote biopsies.

Characterization of blubber fatty acid signatures in northern elephant seals (Mirounga angustirostris) over the postweaning fast.

Phocids routinely fast for extended periods. During these fasts, energetic requirements are met primarily through the catabolism of blubber lipid. To assess whether fatty acid (FA) composition changes during the postweaning fast in northern elephant seals, blubber biopsies were acquired longitudinally from 43 pups at 2.3 ± 1.5 and 55.2 ± 3.7 days postweaning in 1999 and 2000. At weaning, short-chain monounsaturated FA (SC-MUFA, ≤18 carbons) dominated the blubber while saturated FA (SFA) were found in the next highest proportion. The major FA (all ≥1 % by mass) comprised approximately 91 % of total blubber FA. In both years, 18:1n-9 and 16:0 were the most prevalent FA. Major FA mobilized during the fast consisted of polyunsaturated FA (PUFA), SFA, and SC-MUFA. Long-chain MUFA (>18 carbons) tended to be conserved. The fractional mobilization value of 20:5n-3 was the highest, resulting in significant reductions of this PUFA. Although concentrations of some blubber FA changed significantly during the postweaning fast, the general FA signature of blubber was similar at weaning and near the end of the fast. Changes in some FA differed across years. For example, the concentration of 20:4n-6, a minor PUFA, was significantly reduced in 1999 but not in 2000. FA mobilization patterns in northern elephant seal pups are somewhat similar to those reported previously for other fasting phocids and terrestrial mammals, though there are some notable differences. Differences in FA mobilization patterns across mammalian species may be related to differences in diets, geographical distribution, environmental factors, physiological adaptations, and life history stage.

Global change effects on the long-term feeding ecology and contaminant exposures of East Greenland polar bears.

Rapid climate changes are occurring in the Arctic, with substantial repercussions for arctic ecosystems. It is challenging to assess ecosystem changes in remote polar environments, but one successful approach has entailed monitoring the diets of upper trophic level consumers. Quantitative fatty acid signature analysis (QFASA) and fatty acid carbon isotope (δ(13) C-FA) patterns were used to assess diets of East Greenland (EG) polar bears (Ursus maritimus) (n = 310) over the past three decades. QFASA-generated diet estimates indicated that, on average, EG bears mainly consumed arctic ringed seals (47.5 ± 2.1%), migratory subarctic harp (30.6 ± 1.5%) and hooded (16.7 ± 1.3%) seals and rarely, if ever, consumed bearded seals, narwhals or walruses. Ringed seal consumption declined by 14%/decade over 28 years (90.1 ± 2.5% in 1984 to 33.9 ± 11.1% in 2011). Hooded seal consumption increased by 9.5%/decade (0.0 ± 0.0% in 1984 to 25.9 ± 9.1% in 2011). This increase may include harp seal, since hooded and harp seal FA signatures were not as well differentiated relative to other prey species. Declining δ(13) C-FA ratios supported shifts from more nearshore/benthic/ice-associated prey to more offshore/pelagic/open-water-associated prey, consistent with diet estimates. Increased hooded seal and decreased ringed seal consumption occurred during years when the North Atlantic Oscillation (NAO) was lower. Thus, periods with warmer temperatures and less sea ice were associated with more subarctic and less arctic seal species consumption. These changes in the relative abundance, accessibility, or distribution of arctic and subarctic marine mammals may have health consequences for EG polar bears. For example, the diet change resulted in consistently slower temporal declines in adipose levels of legacy persistent organic pollutants, as the subarctic seals have higher contaminant burdens than arctic seals. Overall, considerable changes are occurring in the EG marine ecosystem, with consequences for contaminant dynamics.

Animal-borne acoustic transceivers reveal patterns of at-sea associations in an upper-trophic level predator.

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.

Primiparous and multiparous females differ in mammary gland alveolar development: implications for milk production.

Mammary 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.

Jellyfish support high energy intake of leatherback sea turtles (Dermochelys coriacea): video evidence from animal-borne cameras.

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.

The influence of reproductive experience on milk energy output and lactation performance in the grey seal (Halichoerus grypus).

Although 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.

Validating quantitative fatty acid signature analysis to estimate diets of spectacled and Steller's eiders (Somateria fischeri and Polysticta stelleri).

Fatty acid (FA) signature analysis has been used to study foraging ecology and food webs in marine ecosystems. This powerful method provides information about diets over an extended time period (e.g., 2-4 weeks), rather than just the most recent meal as with most traditional approaches. Using consumer FA signatures, along with a comprehensive database of diet FA signatures, and accounting for consumer FA metabolism, it is possible to estimate the proportions of diet items in the consumer's diet using quantitative FA signature analysis (QFASA). However, before applying QFASA to free-ranging populations, ideally, controlled feeding studies are performed to determine FA deposition and turnover characteristics. We conducted feeding experiments to validate QFASA in captive spectacled eiders (Somateria fischeri) and Steller's eiders (Polysticta stelleri) as a minimally invasive method for studying the diets of these threatened species. We determined FA deposition in eider adipose tissue relative to long-term diet, and developed calibration coefficients (CCs) to account for eider lipid metabolism. Using these CCs with subsequent diet trials, QFASA accurately indicated diet and diet switches. QFASA estimates also indicated that turnover of dietary FAs was not complete by 21 or 29 days, and confirmed that diets could be estimated over an extended period of >29 days. Thus, our understanding of diet can be backtracked to more than a month in captive feeding eiders. We conclude that applying QFASA techniques to eiders and other birds in the wild has the potential to provide valuable information about their diets at various life history stages.

The effects of diet and caloric restriction on adipose tissue fatty acid signatures of tufted puffin (Fratercula cirrhata) nestlings.

Fatty acid (FA) signature analysis is a powerful tool to investigate foraging ecology and food web dynamics in marine ecosystems. However, use of FA signatures to qualitatively or quantitatively infer diets is potentially complicated by effects of nutritional state on lipid metabolism. Estimation of diets using the quantitative fatty acid signature analysis (QFASA) model requires the use of calibration coefficients to account for predator metabolism of individual FAs. We conducted a captive feeding experiment to determine the effects of a 50% reduction in food intake on growth rate and adipose tissue FA signatures of tufted puffin (Fratercula cirrhata) nestlings, a species that routinely experiences food restriction during growth. FA signatures of chicks fed low- and high-calorie diets both exhibited a change in composition in response to the dietary shift with the direction of change in the composition of individual FAs matching the direction of change in the dietary FAs. Despite a growth rate in the restricted nestlings that was 38% of those in the well-fed group, rates of FA turnover were not different between high and low-calorie treatments, and turnover was close to, but not entirely complete, after 27 days on both high-calorie and restricted diets. FA signatures of tufted puffin nestlings were significantly affected by caloric restriction, but these effects were much less pronounced than those of dietary turnover, and calibration coefficients of puffins fed low and high-calorie diets were highly correlated. Our results demonstrate that changes in physiological state can affect FA metabolism, but future research is required to better understand whether the size of these effects is sufficient to substantially alter diet estimation using the QFASA model.

Body condition at weaning affects the duration of the postweaning fast in gray seal pups (Halichoerus grypus).

Gray 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 = p28), 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.