Climate influences on female survival in a declining population of southern elephant seals (Mirounga leonina).

The Southern Ocean has been disproportionately affected by climate change and is therefore an ideal place to study the influence of changing environmental conditions on ecosystems. Changes in the demography of predator populations are indicators of broader shifts in food web structure, but long-term data are required to study these effects. Southern elephant seals (Mirounga leonina) from Macquarie Island have consistently decreased in population size while all other major populations across the Southern Ocean have recently stabilized or are increasing. Two long-term mark-recapture studies (1956-1967 and 1993-2009) have monitored this population, which provides an opportunity to investigate demographic performance over a range of climatic conditions. Using a 9-state matrix population model, we estimated climate influences on female survival by incorporating two major climatic indices into our model: The Southern Annular Mode (SAM) and the Southern Oscillation Index (SOI). Our best model included a 1 year lagged effect of SAM and an unlagged SOI as covariates. A positive relationship with SAM1 (lagged) related the previous year's SAM with juvenile survival, potentially due to changes in local prey availability surrounding Macquarie Island. The unlagged SOI had a negative effect on both juvenile and adult seals, indicating that sea ice dynamics and access to foraging grounds on the East Antarctic continental shelf could explain the different contributions of ENSO events on the survival of females in this population.

Seal mothers expend more on offspring under favourable conditions and less when resources are limited.

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.

Bottom-up regulation of a pole-ward migratory predator population.

As the effects of regional climate change are most pronounced at polar latitudes, we might expect polar-ward migratory populations to respond as habitat suitability changes. The southern elephant seal (Mirounga leonina L.) is a pole-ward migratory species whose populations have mostly stabilized or increased in the past decade, the one exception being the Macquarie Island population which has decreased continuously over the past 50 years. To explore probable causes of this anomalous trend, we counted breeding female seals annually between 1988 and 2011 in order to relate annual rates of population change (r) to foraging habitat changes that have known connections with atmospheric variability. We found r (i) varied annually from -0.016 to 0.021 over the study period, (ii) was most effected by anomalous atmospheric variability after a 3 year time lag was introduced (R = 0.51) and (iii) was associated with sea-ice duration (SID) within the seals' foraging range at the same temporal lag. Negative r years may be extrapolated to explain, at least partially, the overall trend in seal abundance at Macquarie Island; specifically, increasing SID within the seals foraging range has a negative influence on their abundance at the island. Evidence is accruing that suggests southern elephant seal populations may respond positively to a reduced sea-ice field.

Climate change and seal survival: evidence for environmentally mediated changes in elephant seal, Mirounga leonina, pup survival.

Maternal and physical factors play a significant role in animal life-history variability, which means that large scale climate change has the potential to affect the size and dynamics of animal populations indirectly through maternal investment and directly through conditions that animals are exposed to. However, little is known about the effects of large-scale oceanographic events such as the El-Niño southern oscillation (ENSO) that influence productivity in the Southern Ocean and the abundance, quality and distribution of prey. The possible mechanisms by which physical factors and primary productivity could influence life-history traits, such as survival of apex predators, includes direct influences such as food availability and foraging success and indirect influences such as stored maternal investment and resource transfer during lactation. Here, we quantify the relative contribution of maternal investment and climate conditions at remote foraging sites to survival in the first year of life for southern elephant seals. We present evidence linking climate (ENSO) and variations in a key demographic parameter--first-year survival--and demonstrate that survival was highest during ENSO events and that the ability of mothers to store and acquire resources, which is typically related to ocean productivity, is the most important determinant of survival in the first year. This functional link provides valuable insights that can be used to model the responses of the seal populations to climate change scenarios.

Juvenile southern elephant seals exhibit seasonal differences in energetic requirements and use of lipids and protein stores.

Growing juvenile animals undergo many morphological, physiological, and behavioural changes that influence their energetic requirements, patterns of energy use, and ultimately, their survival and reproductive success. We examined changes in mass loss and body composition of juvenile southern elephant seals (1- and 2-yr-olds) during their two annual haul-outs. At the start and end of the midyear and molt haul-outs, we caught, weighed, and measured 41 and 14 seals, respectively. We measured blubber depth using ultrasound to estimate body composition (lean and adipose tissue mass). Using energy densities of the adipose and lean tissue, we calculated total, lean, and adipose mass changes and energy expenditure. While molting, juvenile seals used more energy than during the midyear, which is related to the increased use of lean tissue for hair and skin regeneration. The amount of energy used increases with mass as individuals mature. We found sexual differences in energy use where females retained greater fat reserves than males by utilizing more lean tissue. These differences are most likely related to haul-out function and behavior, growth, and earlier development of females toward sexual maturity.

Resource partitioning through oceanic segregation of foraging juvenile southern elephant seals ( Mirounga leonina).

In highly dynamic and unpredictable environments such as the Southern Ocean, species that have evolved behaviors that reduce the effects of intra-specific competition may have a selective advantage. This is particularly true when juveniles face disadvantages when foraging due to morphological or physiological limitation, which is the case for many marine mammals. We tracked the at-sea movements of 48 juvenile southern elephant seals ( Mirounga leonina) between the ages of 1 and 4 years from the population at Macquarie Island using locations derived from recorded light levels. There were significant differences in the total amount of the Southern Ocean covered by the different age-groups. The younger seals used a smaller area than the older seals. On average, the younger individuals also made more trips to sea than the older seals and did not travel as far on each trip. Females spent more time at sea than males and there were no significant differences between the total areas used by male and females. In summary, younger seals remained closer to the island at all times, and they spent more time in more northerly regions that older seals. These differences in behavior created temporal and spatial segregation between juveniles of different ages. Therefore, we suggest that these temporal and spatial separations help to avoid intra-specific competition for resources on land, space on beaches, and at-sea foraging areas. Such modifications of haul-out timing and behavior enable them to exploit a patchy and unpredictable environment.

The immunocompetence handicap hypothesis in two sexually dimorphic pinniped species--is there a sex difference in immunity during early development?

The 'immunocompetence handicap hypothesis' predicts that highly sexually dimorphic and polygynous species will exhibit sex differences in immunity. We tested this hypothesis in southern elephant and grey seals during their early development by measuring the following parameters: leucocyte counts, serum IgG levels, erythrocyte sedimentation rate and haematocrit. We failed to find any differences due to sex as assessed by the parameters investigated. Animals were sampled longitudinally during their development and there were significant age effects from birth to weaning in both species. Total and differential leucocyte counts and erythrocyte sedimentation rates increased just prior to weaning then decreased. Haematocrits declined whilst total circulating immunoglobulin G concentrations increased. Body temperatures remained constant throughout the postnatal period. Differences between the species were seen in total leucocyte counts and in polymorphonuclear cells and eosinophils. Southern elephant seals had higher concentrations than grey seals and total leucocyte counts in the former were among the highest reported for mammals.