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Age structure is a fundamental aspect of animal population biology. Age is strongly related to individual physiological condition, reproductive potential and mortality rate. Currently, there are no robust molecular methods for age estimation in birds. Instead, individuals must be ringed as chicks to establish known-age populations, which is a labour-intensive and expensive process. The estimation of chronological age using DNA methylation (DNAm) is emerging as a robust approach in mammals including humans, mice and some non-model species. Here, we quantified DNAm in whole blood samples from a total of 71 known-age Short-tailed shearwaters (Ardenna tenuirostris) using digital restriction enzyme analysis of methylation (DREAM). The DREAM method measures DNAm levels at thousands of CpG dinucleotides throughout the genome. We identified seven CpG sites with DNAm levels that correlated with age. A model based on these relationships estimated age with a mean difference of 2.8 years to known age, based on validation estimates from models created by repeated sampling of training and validation data subsets. Longitudinal observation of individuals re-sampled over 1 or 2 years generally showed an increase in estimated age (6/7 cases). For the first time, we have shown that epigenetic changes with age can be detected in a wild bird. This approach should be of broad interest to researchers studying age biomarkers in non-model species and will allow identification of markers that can be assessed using targeted techniques for accurate age estimation in large population studies., (© 2018 John Wiley & Sons Ltd.)

In iteroparous species, intermittent breeding is an important life-history tactic that can greatly affect animal population growth and viability. Despite its importance, few studies have quantified the consequences of breeding pauses on lifetime reproductive output, principally because calculating lifetime reproductive output requires knowledge of each individual's entire reproductive history. This information is extremely difficult to obtain in wild populations. We applied novel statistical approaches that account for uncertainty in state assessment and individual heterogeneity to an 18-year capture-recapture dataset of 6,631 female southern elephant seals from Macquarie Island. We estimated survival and breeding probabilities, and investigated the consequences of intermittent breeding on lifetime reproductive output. We found consistent differences in females' demographic performance between two heterogeneity classes. In particular, breeding imbued a high cost on survival in the females from the heterogeneity class 2, assumed to be females of lower quality. Individual quality also appeared to play a major role in a female's decision to skip reproduction with females of poorer quality more likely to skip breeding events than females of higher quality. Skipping some breeding events allowed females from both heterogeneity classes to increase lifetime reproductive output over females that bred annually. However, females of lower quality produced less offspring over their lifetime. Intermittent breeding seems to be used by female southern elephant seals as a tactic to offset reproductive costs on survival and enhance lifetime reproductive output but remains unavoidable and driven by individual-specific constraints in some other females., (© 2017 The Authors. Journal of Animal Ecology © 2017 British Ecological Society.)

In mammals, maternal expenditure on offspring is a complex mix of several factors including the species' mating system, offspring sex and the condition and age of the mother. While theory suggests that in polygynous species mothers should wean larger male offspring than females when resources and maternal conditions allow, the evidence for this remains equivocal. Southern elephant seals are highly dimorphic, polygynous capital breeders existing in an environment with highly variable resources and should therefore provide clear evidence to support the theoretical expectations of differential maternal expenditure in male and female pups. We quantified maternal size (mass and length) and pup size at birth and weaning for 342 elephant seal mothers at Macquarie Island. The study was conducted over 11 years of contrasting sea-ice and Southern Annular Mode values, both indices of maternal prey resources. Overall, large females weaned male pups that weighed 17 kg (15·5%) more than female pups. Maternal condition varied by as much as 59 kg among years, and was positively related to Southern Annular Mode, and negatively to maximum sea-ice extent. Smaller mothers weaned relatively larger male pups under favourable conditions, this effect was less apparent for larger mothers. We developed a simple model linking environmental variation to maternal masses post-partum, followed by maternal masses post-partum to weaning masses and then weaning masses to pup survival and demonstrated that environmental conditions affected predicted survival so that the pups of small mothers had an estimated 7% increase in first year survival in 'good' vs. 'bad' years compared to 1% for female pups of large mothers. Co-occurrence of environmental quality and conservative reproductive tactics suggests that mothers retain substantial plasticity in maternal care, enhancing their lifetime reproductive success by adjusting reproductive expenditure relative to both prevailing environmental conditions and their own capabilities., (© 2016 The Authors. Journal of Animal Ecology © 2016 British Ecological Society.)

1. Body condition plays a fundamental role in many ecological and evolutionary processes at a variety of scales and across a broad range of animal taxa. An understanding of how body condition changes at fine spatial and temporal scales as a result of interaction with the environment provides necessary information about how animals acquire resources. 2. However, comparatively little is known about intra- and interindividual variation of condition in marine systems. Where condition has been studied, changes typically are recorded at relatively coarse time-scales. By quantifying how fine-scale interaction with the environment influences condition, we can broaden our understanding of how animals acquire resources and allocate them to body stores. 3. Here we used a hierarchical Bayesian state-space model to estimate the body condition as measured by the size of an animal's lipid store in two closely related species of marine predator that occupy different hemispheres: northern elephant seals (Mirounga angustirostris) and southern elephant seals (Mirounga leonina). The observation model linked drift dives to lipid stores. The process model quantified daily changes in lipid stores as a function of the physiological condition of the seal (lipid:lean tissue ratio, departure lipid and departure mass), its foraging location, two measures of behaviour and environmental covariates. 4. We found that physiological condition significantly impacted lipid gain at two time-scales - daily and at departure from the colony - that foraging location was significantly associated with lipid gain in both species of elephant seals and that long-term behavioural phase was associated with positive lipid gain in northern and southern elephant seals. In northern elephant seals, the occurrence of short-term behavioural states assumed to represent foraging were correlated with lipid gain. Lipid gain was a function of covariates in both species. Southern elephant seals performed fewer drift dives than northern elephant seals and gained lipids at a lower rate. 5. We have demonstrated a new way to obtain time series of body condition estimates for a marine predator at fine spatial and temporal scales. This modelling approach accounts for uncertainty at many levels and has the potential to integrate physiological and movement ecology of top predators. The observation model we used was specific to elephant seals, but the process model can readily be applied to other species, providing an opportunity to understand how animals respond to their environment at a fine spatial scale., (© 2013 The Authors. Journal of Animal Ecology © 2013 British Ecological Society.)

Many optimal foraging models for diving animals examine strategies that maximize time spent in the foraging zone, assuming that prey acquisition increases linearly with search time. Other models have considered the effect of patch quality and predict a net energetic benefit if dives where no prey is encountered early in the dive are abandoned. For deep divers, however, the energetic benefit of giving up is reduced owing to the elevated energy costs associated with descending to physiologically hostile depths, so patch residence time should be invariant. Others consider an asymptotic gain function where the decision to leave a patch is driven by patch-depletion effects - the marginal value theorem. As predator behaviour is increasingly being used as an index of marine resource density and distribution, it is important to understand the nature of this gain function. We investigated the dive behaviour of the world's deepest-diving seal, the southern elephant seal Mirounga leonina, in response to patch quality. Testing these models has largely been limited to controlled experiments on captive animals. By integrating in situ measurements of the seal's relative lipid content obtained from drift rate data (a measure of foraging success) with area-restricted search behaviour identified from first-passage time analysis, we identified regions of high- and low-quality patches. Dive durations and bottom times were not invariant and did not increase in regions of high quality; rather, both were longer when patches were of relatively low quality. This is consistent with the predictions of the marginal value theorem and provides support for a nonlinear relationship between search time and prey acquisition. We also found higher descent and ascent rates in high-quality patches suggesting that seals minimized travel time to the foraging patch when quality was high; however, this was not achieved by increasing speed or dive angle. Relative body lipid content was an important predictor of dive behaviour. Seals did not schedule their diving to maximize time spent in the foraging zone in higher-quality patches, challenging the widely held view that maximizing time in the foraging zone translates to greater foraging success., (© 2012 The Authors. Journal of Animal Ecology © 2012 British Ecological Society.)

1. Seasonal long-distance migrations are often expected to be related to resource distribution, and foraging theory predicts that animals should spend more time in areas with relatively richer resources. Yet for highly migratory marine species, data on feeding success are difficult to obtain. We analysed the temporal feeding patterns of wild juvenile southern bluefin tuna from visceral warming patterns recorded by archival tags implanted within the body cavity. 2. Data collected during 1998-2000 totalled 6221 days, with individual time series (n = 19) varying from 141 to 496 days. These data span an annual migration circuit including a coastal summer residency within Australian waters and subsequent migration into the temperate south Indian Ocean. 3. Individual fish recommenced feeding between 5 and 38 days after tagging, and feeding events (n = 5194) were subsequently identified on 76.3 +/- 5.8% of days giving a mean estimated daily intake of 0.75 +/- 0.05 kg. 4. The number of feeding events varied significantly with time of day with the greatest number occurring around dawn (58.2 +/- 8.0%). Night feeding, although rare (5.7 +/- 1.3%), was linked to the full moon quarter. Southern bluefin tuna foraged in ambient water temperatures ranging from 4.9 degrees C to 22.9 degrees C and depths ranging from the surface to 672 m, with different targeting strategies evident between seasons. 5. No clear relationship was found between feeding success and time spent within an area. This was primarily due to high individual variability, with both positive and negative relationships observed at all spatial scales examined (grid ranges of 2 x 2 degrees to 10 x 10 degrees ). Assuming feeding success is proportional to forage density, our data do not support the hypothesis that these predators concentrate their activity in areas of higher resource availability. 6. Multiple-day fasting periods were recorded by most individuals. The majority of these (87.8%) occurred during periods of apparent residency within warmer waters (sea surface temperature > 15 degrees C) at the northern edge of the observed migratory range. These previously undocumented nonfeeding periods may indicate alternative motivations for residency. 7. Our results demonstrate the importance of obtaining information on feeding when interpreting habitat utilization from individual animal tracks.

1. Environmental variation influences food abundance and availability, which is reflected in the reproductive success of top predators. We examined maternal expenditure, offspring mass and condition for Weddell seals in 2 years when individuals exhibited marked differences in these traits. 2. For females weighing > or = 355 kg there was a positive relationship between maternal post-partum mass (MPPM) and lactation length, but below this there was no relationship, suggesting that heavier females were able to increase lactation length but lighter females were restricted to a minimum lactation period of 33 days. 3. Overall, females were heavier in 2002, but in 2003 shorter females were lighter than similar-sized females in 2002 suggesting that the effects of environmental variability on foraging success and condition are more pronounced in smaller individuals. 4. There was no relationship between MPPM and pup birth mass, indicating pre-partum investment did not differ between years. However, there was a positive relationship between MPPM and pup mass gain. Mass and energy transfer efficiency were 10.2 and 5.4% higher in 2002 than 2003, which suggests costs associated with a putatively poor-resource year were delayed until lactation. 5. Heavier females lost a higher proportion of mass during lactation in both years, so smaller females may not have been able to provide more to their offspring to wean a pup of similar size to larger females. 6. MPPM had only a small influence on total body lipid; therefore, regardless of mass, females had the same relative body composition. Females with male pups lost a higher percentage of lipid than those with female pups, but by the end of lactation female pups had 4.5% higher lipid content than males. 7. It appears that for Weddell seals the consequences of environmentally induced variation in food availability are manifested in differences in maternal mass and expenditure during lactation. These differences translate to changes in pup mass and condition at weaning with potential consequences for future survival and recruitment.