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Evaluating physiological responses in the context of a species’ life history, demographics, and ecology is essential to understanding the health of individuals and populations. Here, we measured the main mammalian glucocorticoid, cortisol, in an elusive Antarctic apex predator, the leopard seal ( Hydrurga leptonyx ). We also examined intraspecific variation in cortisol based on life history (sex), morphometrics (body mass, body condition), and ecological traits ( δ 15 N, δ 13 C). To do this, blood samples, life history traits, and morphometric data were collected from 19 individual leopard seals off the Western Antarctic Peninsula. We found that adult leopard seals have remarkably high cortisol concentrations (100.35 ± 16.72 μg/dL), showing the highest circulating cortisol concentration ever reported for a pinniped: 147 μg/dL in an adult male. Leopard seal cortisol concentrations varied with sex, body mass, and diet. Large adult females had significantly lower cortisol (94.49 ± 10.12 μg/dL) than adult males (120.85 ± 6.20 μg/dL). Similarly, leopard seals with higher isotope values (i.e., adult females, δ 15 N: 11.35 ± 0.69‰) had lower cortisol concentrations than seals with lower isotope values (i.e., adult males, δ 15 N: 10.14 ± 1.65‰). Furthermore, we compared cortisol concentrations across 26 closely related Arctoid taxa (i.e., mustelids, bears, and pinnipeds) with comparable data. Leopard seals had the highest mean cortisol concentrations that were 1.25 to 50 times higher than other Arctoids. More broadly, Antarctic ice seals (Lobodontini: leopard seal, Ross seal, Weddell seal, crabeater seal) had higher cortisol concentrations compared to other pinnipeds and Arctoid species. Therefore, high cortisol is a characteristic of all lobodontines and may be a specialized adaptation within this Antarctic-dwelling clade. Together, our results highlight exceptionally high cortisol concentrations in leopard seals (and across lobodontines) and reveal high variability in cortisol concentrations among individuals from a single location. This information provides the context for understanding how leopard seal physiology changes with life history, ecology, and morphology and sets the foundation for assessing their physiology in the context of a rapidly changing Antarctic environment. 

Elephant seals sleep 2 hours a day while diving for months at sea, rivaling the record for the least sleep among mammals. 

Animals that display plasticity in behavioral, ecological, and morphological traits are better poised to cope with environmental disturbances. Here, we examined individual plasticity and intraspecific variation in the morphometrics, movement patterns, and dive behavior of an enigmatic apex predator, the leopard seal ( Hydrurga leptonyx ). Satellite/GPS tags and time-depth recorders were deployed on 22 leopard seals off the Western Antarctic Peninsula. Adult female leopard seals were significantly larger (454±59 kg) and longer (302±11 cm) than adult males (302±22 kg, 276±11 cm). As females were 50% larger than their male counterparts, leopard seals are therefore one of the most extreme examples of female-biased sexual size dimorphism in marine mammals. Female leopard seals also spent more time hauled-out on land and ice than males. In the austral spring/summer, three adult female leopard seals hauled-out on ice for 10+ days, which likely represent the first satellite tracks of parturition and lactation for the species. While we found sex-based differences in morphometrics and haul-out durations, other variables, including maximum distance traveled and dive parameters, did not vary by sex. Regardless of sex, some leopard seals remained in near-shore habitats, traveling less than 50 kilometers, while other leopard seals traveled up to 1,700 kilometers away from the tagging location. Overall, leopard seals were short (3.0±0.7 min) and shallow (29±8 m) divers. However, within this general pattern, some individual leopard seals primarily used short, shallow dives, while others switched between short, shallow dives and long, deep dives. We also recorded the single deepest and longest dive made by any leopard seal—1, 256 meters for 25 minutes. Together, our results showcased high plasticity among leopard seals tagged in a single location. These flexible behaviors and traits may offer leopard seals, an ice-associated apex predator, resilience to the rapidly changing Southern Ocean. 

ABSTRACT The hypothalamic–pituitary–adrenal (HPA) axis controls the release of glucocorticoids, which regulate immune and inflammatory function by modulating cytokines, white blood cells and oxidative stress via glucocorticoid receptor (GR) signaling. Although the response to HPA activation is well characterized in many species, little is known about the impacts of HPA activation during extreme physiological conditions. Hence, we challenged 18 simultaneously fasting and developing elephant seal pups with daily intramuscular injections of adrenocorticotropin (ACTH), a GR antagonist (RU486), or a combination of the two (ACTH+RU486) for 4 days. We collected blood at baseline, 2 h and 4 days after the beginning of treatment. ACTH and ACTH+RU486 elevated serum aldosterone and cortisol at 2 h, with effects diminishing at 4 days. RU486 alone induced a compensatory increase in aldosterone, but not cortisol, at 4 days. ACTH decreased neutrophils at 2 h, while decreasing lymphocytes and increasing the neutrophil:lymphocyte ratio at 4 days. These effects were abolished by RU486. Despite alterations in white blood cells, there was no effect of ACTH or RU486 on transforming growth factor-β or interleukin-6 levels; however, both cytokines decreased with the 4 day fasting progression. Similarly, ACTH did not impact protein oxidation, lipid peroxidation or antioxidant enzymes, but plasma isoprostanes and catalase activity decreased while glutathione peroxidase increased with fasting progression. These data demonstrate differential acute (2 h) and chronic (4 days) modulatory effects of HPA activation on white blood cells and that the chronic effect is mediated, at least in part, by GR. These results also underscore elephant seals' extraordinary resistance to oxidative stress derived from repeated HPA activation. 

Marine mammals such as northern elephant seals (NES) routinely experience hypoxemia and ischemia-reperfusion events to many tissues during deep dives with no apparent adverse effects. Adaptations to diving include increased antioxidants and elevated oxygen storage capacity associated with high hemoprotein content in blood and muscle. The natural turnover of heme by heme oxygenase enzymes (encoded by HMOX1 and HMOX2 ) produces endogenous carbon monoxide (CO), which is present at high levels in NES blood and has been shown to have cytoprotective effects in laboratory systems exposed to hypoxia. To understand how pathways associated with endogenous CO production and signaling change across ontogeny in diving mammals, we measured muscle CO and baseline expression of 17 CO-related genes in skeletal muscle and whole blood of three age classes of NES. Muscle CO levels approached those of animals exposed to high exogenous CO, increased with age, and were significantly correlated with gene expression levels. Muscle expression of genes associated with CO production and antioxidant defenses ( HMOX1 , BVR , GPX3 , PRDX1 ) increased with age and was highest in adult females, while that of genes associated with protection from lipid peroxidation ( GPX4 , PRDX6 , PRDX1 , SIRT1 ) was highest in adult males. In contrast, muscle expression of mitochondrial biogenesis regulators ( PGC1A , ESRRA , ESRRG ) was highest in pups, while genes associated with inflammation ( HMOX2 , NRF2 , IL1B ) did not vary with age or sex. Blood expression of genes involved in regulation of inflammation ( IL1B , NRF2 , BVR , IL10 ) was highest in pups, while HMOX1 , HMOX2 and pro-inflammatory markers ( TLR4 , CCL4 , PRDX1 , TNFA ) did not vary with age. We propose that ontogenetic upregulation of baseline HMOX1 expression in skeletal muscle of NES may, in part, underlie increases in CO levels and expression of genes encoding antioxidant enzymes. HMOX2 , in turn, may play a role in regulating inflammation related to ischemia and reperfusion in muscle and circulating immune cells. Our data suggest putative ontogenetic mechanisms that may enable phocid pups to transition to a deep-diving lifestyle, including high baseline expression of genes associated with mitochondrial biogenesis and immune system activation during postnatal development and increased expression of genes associated with protection from lipid peroxidation in adulthood.