Hypoxia exposure blunts angiogenic signaling and upregulates the antioxidant system in endothelial cells derived from elephant seals.

BACKGROUND: Elephant seals exhibit extreme hypoxemic tolerance derived from repetitive hypoxia/reoxygenation episodes they experience during diving bouts. Real-time assessment of the molecular changes underlying protection against hypoxic injury in seals remains restricted by their at-sea inaccessibility. Hence, we developed a proliferative arterial endothelial cell culture model from elephant seals and used RNA-seq, functional assays, and confocal microscopy to assess the molecular response to prolonged hypoxia. RESULTS: Seal and human endothelial cells exposed to 1% O2 for up to 6 h respond differently to acute and prolonged hypoxia. Seal cells decouple stabilization of the hypoxia-sensitive transcriptional regulator HIF-1α from angiogenic signaling. Rapid upregulation of genes involved in glutathione (GSH) metabolism supports the maintenance of GSH pools, and intracellular succinate increases in seal but not human cells. High maximal and spare respiratory capacity in seal cells after hypoxia exposure occurs in concert with increasing mitochondrial branch length and independent from major changes in extracellular acidification rate, suggesting that seal cells recover oxidative metabolism without significant glycolytic dependency after hypoxia exposure. CONCLUSIONS: We found that the glutathione antioxidant system is upregulated in seal endothelial cells during hypoxia, while this system remains static in comparable human cells. Furthermore, we found that in contrast to human cells, hypoxia exposure rapidly activates HIF-1 in seal cells, but this response is decoupled from the canonical angiogenesis pathway. These results highlight the unique mechanisms that confer extraordinary tolerance to limited oxygen availability in a champion diving mammal.

Heterocheilus floridensis sp. n. (Nematoda: Heterocheilidae) from the West Indian manatee Trichechus manatus (Trichechidae, Sirenia) in Florida, USA.

Morphological data are used to describe a new nematode species, Heterocheilus floridensis sp. n. (Heterocheilidae) from the digestive tract of the Florida manatee Trichechus manatus latirostris (Harlan) (Trichechidae, Sirenia) from Florida, USA. Examination by light and scanning electron microscopy revealed that the new species differs from the related Heterocheilus tunicatus Diesing, 1839 mainly by having dentigerous ridges on the inner surface of the lips, a median unpaired papilla located anterior to the cloaca, and a considerably larger body size. Sequence data for subunits I and II of mitochondrial cytochrome oxidase gene, 18S small subunit and 28S ribosomal RNA genes were provided for molecular characterisation of the new species. However, the current unavailability of homologous sequence data for congeneric specimens precluded a molecular assessment of the morphological species hypothesis, and ascaridoid phylogenetic hypotheses could not be advanced. Specimens of Heterocheilus sp. collected from the Antillean manatee Trichechus manatus manatus Linnaeus in Puerto Rico, on loan from the US National Museum of Natural History, were morphologically consistent with the new species, so apparently all congeneric nematodes reported from both subspecies of the West Indian manatee Trichechus manatus Linnaeus and previously identified as H. tunicatus belong rather to H. floridensis sp. n. Heterocheilus hagenbecki (Khalil et Vogelsang, 1932) Sprent 1980 is here considered to be a species inquirenda. A key to valid species of Heterocheilus Diesing, 1839 is provided.

Faecal bacterial communities differ amongst discrete foraging populations of dugongs along the east Australian coast.

Gut bacterial communities play a vital role in a host's digestion, fermentation of complex carbohydrates, absorption of nutrients and energy harvest/storage. Dugongs are obligate seagrass grazers with an expanded hindgut and associated microbiome. Here, we characterised and compared the faecal bacterial communities of dugongs from genetically distinct populations along the east coast of Australia, between subtropical Moreton Bay and tropical Cleveland Bay. Amplicon sequencing of fresh dugong faecal samples (n=47) revealed Firmicutes (62%) dominating the faecal bacterial communities across all populations. Several bacterial genera (Bacteroides, Clostridium sensu stricto 1, Blautia and Polaribacter) were detected in samples from all locations, suggesting their importance in seagrass digestion. Principal coordinate analysis showed the three southern-most dugong populations having different faecal bacterial community compositions from northern populations. The relative abundances of the genera Clostridium sensu stricto 13 and dgA-11 gut group were higher, but Bacteroides was lower, in the southern dugong populations, compared to the northern populations, suggesting potential adaptive changes associated with location. This study contributes to our knowledge of the faecal bacterial communities of dugongs inhabiting Australian coastal waters. Future studies of diet selection in relation to seagrass availability throughout the dugong's range will help to advance our understanding of the roles that seagrass species may play in affecting the dugong's faecal bacterial community composition.

Matching-to-Sample Task Training of a Killer Whale (Orcinus orca).

Matching-to-sample tasks have been a useful method in visual cognitive studies on non-human animals. The use of touch panels in matching-to-sample tasks has contributed to cognitive studies on terrestrial animals; however, there has been a difficulty in using these devices underwater, which is one of the factors that has slowed the progress of visual studies on underwater animals. Cetaceans (e.g., dolphins and whales) are highly adapted to underwater environments, and further studies on their cognitive abilities are needed to advance our understanding of the interactions between environmental factors and the evolution of cognitive abilities. In this study, we aimed to develop a new experimental method in which a captive killer whale performed a matching-to-sample task using a monitor shown through an underwater window as if a touch panel were used. In order to confirm the usefulness of this method, one simple experiment on mirror image discrimination was conducted, and the pairs with mirror images were shown to be more difficult to identify than the pairs with other normal images. The advantages of using this method include (1) simplicity in the devices and stimuli used in the experiments, (2) appropriate and rigorous experimental control, (3) the possibility of increasing the number of individuals to be tested and interspecies comparisons, and (4) contributions to animal welfare. The use of this method solves some of the problems in previous visual cognitive studies on cetaceans, and it suggests the further possibility of future comparative cognitive studies. It is also expected to contribute to animal welfare in terms of cognitive enrichment, and it could help with the proposal of new exhibition methods in zoos and aquariums.

Training the Concept of Innovate in Dolphins (Tursiops truncatus) Is Both Creative and Cognitively Stimulating.

Creative or novel behaviors in bottlenose dolphins (Tursiops truncatus) can be indicators of flexible thinking and problem solving. Over 50 years ago, two rough-tooth dolphins demonstrated creative novel behaviors acquired through reinforcement training in human care. Since this novel training, a variety of species have been trained to respond to this conceptual cue. The current study assessed the creativity of 12 bottlenose dolphins (5 females, 7 males) housed at the Roatan Institute for Marine Sciences (RIMS) in Roatan, Honduras. Individual differences were found across four constructs measured for creativity: fluency, flexibility, elaboration, and originality. Variability in performance occurred across test sessions. Animals with less experience with this task performed fewer "innovative" behaviors as compared to more experienced animals. Despite errors, dolphins continued to attempt the task during test sessions, suggesting the concept of "innovate" was intrinsically rewarding and cognitively engaging. This task may be utilized across species to promote the comparative study of innovative or creative behavior as well as to promote cognitive welfare.

Aerial single target acuity of harbor seals (Phoca vitulina) for stationary and moving targets of varying contrast.

Harbor seals (Phoca vitulina) need to detect single objects for example when orienting to landmarks or hunting prey. The detection of single objects, described by the single target acuity (STA), cannot be deduced from formerly determined grating acuity (GA) as different mechanisms underlie STA and GA. Thus, we assessed STA for stationary and moving single targets with varying contrast in two harbor seals in a first approach in air. In a two-alternative-forced-choice discrimination task, the seals had to indicate whether the single target was presented in a left or right stimulus field on a monitor. The STA for full-contrast stationary targets was determined as 0.27 deg of visual angle for both experimental animals. Contrary to our expectations, neither adding motion nor reducing contrast had a strong impact on STA. Additionally, we also determined GA in the two harbor seals (1.2 and 1.1 cycles/deg or 0.42 and 0.45 deg for a single stripe of the grating at threshold) to be slightly inferior to STA. Our results are in good correspondence with contrast sensitivity and allow calculating viewing distances in the context of for example visual orientation.

Rerouting of a major shipping lane through important harbour porpoise habitat caused no detectable change in annual occurrence or foraging patterns.

Shipping is one of the largest industries globally, with well-known negative impacts on the marine environment. Despite the known negative short-term (minutes to hours) impact of shipping on individual animal behavioural responses, very little is understood about the long-term (months to years) impact on marine species presence and area use. This study took advantage of a planned rerouting of a major shipping lane leading into the Baltic Sea, to investigate the impact on the presence and foraging behaviour of a marine species known to be sensitive to underwater noise, the harbour porpoise (Phocoena phocoena). Passive acoustic monitoring data were collected from 15 stations over two years. Against predictions, no clear change occurred in monthly presence or foraging behaviour of the porpoises, despite the observed changes in noise and vessel traffic. However, long-term heightened noise levels may still impact communication, echolocation, or stress levels of individuals, and needs further investigation.

Bioenergetic modelling of a marine top predator's responses to changes in prey structure.

Determining how animals allocate energy, and how external factors influence this allocation, is crucial to understand species' life history requirements and response to disturbance. This response is driven in part by individuals' energy balance, prey characteristics, foraging behaviour and energy required for essential functions. We developed a bioenergetic model to estimate minimum foraging success rate (FSR), that is, the lowest possible prey capture rate for individuals to obtain the minimum energy intake needed to meet daily metabolic requirements, for female sperm whale (Physeter macrocephalus). The model was based on whales' theoretical energetic requirements using foraging and prey characteristics from animal-borne tags and stomach contents, respectively. We used this model to simulate two prey structure change scenarios: (1) decrease in mean prey size, thus lower prey energy content and (2) decrease in prey size variability, reducing the variability in prey energy content. We estimate the whales need minimum of ~14% FSR to meet their energetic requirements, and energy intake is more sensitive to energy content changes than a decrease in energy variability. To estimate vulnerability to prey structure changes, we evaluated the compensation level required to meet bioenergetic demands. Considering a minimum 14% FSR, whales would need to increase energy intake by 21% (5-35%) and 49% (27-67%) to compensate for a 15% and 30% decrease in energy content, respectively. For a 30% and 50% decrease in energy variability, whales would need to increase energy intake by 13% (0-23%) and 24% (10-35%) to meet energetic demands, respectively. Our model demonstrates how foraging and prey characteristics can be used to estimate impact of changing prey structure in top predator energetics, which can help inform bottom-up effects on marine ecosystems. We showed the importance of considering different FSR in bioenergetics models, as it can have decisive implications on estimates of energy acquired and affect the conclusions about top predator's vulnerability to possible environmental fluctuations.

120-years of ecological monitoring data shows that the risk of overhunting is increased by environmental degradation for an isolated marine mammal population: The Baltic grey seal.

The Baltic Sea is home to a genetically isolated and morphologically distinct grey seal population. This population has been the subject of 120-years of careful documentation, from detailed records of bounty statistics to annual monitoring of health and abundance. It has also been exposed to a range of well-documented stressors, including hunting, pollution and climate change. To investigate the vulnerability of marine mammal populations to multiple stressors, data series relating to the Baltic grey seal population size, hunt and health were compiled, vital demographic rates were estimated, and a detailed population model was constructed. The Baltic grey seal population fell from approximately 90,000 to as few as 3000 individuals during the 1900s as the result of hunting and pollution. Subsequently, the population has recovered to approximately 55,000 individuals. Fertility levels for mature females have increased from 9% in the 1970s to 86% at present. The recovery of the population has led to demands for increased hunting, resulting in a sudden increase in annual quotas from a few hundred to 3550 in 2020. Simultaneously, environmental changes, such as warmer winters and reduced prey availability due to overfishing, are likely impacting fecundity and health. Future population development is projected for a range of hunting and environmental stress scenarios, illustrating how hunting, in combination with environmental degradation, can lead to population collapse. The current combined hunting quotas of all Baltic Nations caused a 10% population decline within three generations in 100% of simulations. To enable continued recovery of the population, combined annual quotas of less than 1900 are needed, although this quota should be re-evaluated annually as monitoring of population size and seal health continues. Sustainable management of long-lived slowly growing species requires an understanding of the drivers of population growth and the repercussions of management decisions over many decades. The case of the Baltic grey seal illustrates how long-term ecological time series are pivotal in establishing historical baselines in population abundance and demography to inform sustainable management.

Management and research efforts are failing dolphins, porpoises, and other toothed whales.

Despite being subject to intensive research and public interest many populations of dolphins, porpoises, and other toothed whales (small cetaceans) continue to decline, and several species are on the verge of extinction. We examine small cetacean status, human activities driving extinction risk, and whether research efforts are addressing priority threats. We estimate that 22% of small cetaceans are threatened with extinction, with little signs of improvement in nearly thirty years. Fisheries and coastal habitat degradation are the main predictors of extinction risk. Contrary to popular belief, we show that the causal impact of small-scale fisheries on extinction risk is greater than from large-scale fisheries. Fisheries management strength had little influence on extinction risk, suggesting that the implementation of existing measures have been largely ineffective. Alarmingly, we find research efforts for priority threats to be vastly underrepresented and so a major shift in research focus is required. Small cetaceans are among the lower hanging fruits of marine conservation; continued failure to halt their decline bodes poorly for tackling marine biodiversity loss and avoiding an Anthropocene mass extinction.

Halocercus lagenorhynchi infection in a stranded striped dolphin Stenella coeruleoalba (Meyen, 1833) on the Southwest coastline of India.

Necropsy on a striped dolphin Stenella coeruleoalba (Meyen, 1833) entangled in ghost fishing net and dead while rescuing yielded some helminth parasites, later identified as Halocercus lagenorhynchi. DNA barcoding of the host and parasite and the phylogenetic analysis of the parasite was conducted. This study provides valuable information towards establishing basal data of marine mammal parasite diversity and distribution in the Indian waters. We believe this is the first report of the occurrence of Halocercus lagenorhynchi in marine mammals in India.

Population genomics of harbour seal Phoca vitulina from northern British Columbia through California and comparison to the Atlantic subspecies.

The harbour seal Phoca vitulina is a ubiquitous pinniped species found throughout coastal waters of the Northern Hemisphere. Harbour seal impacts on ecosystem dynamics may be significant due to their high abundance and food web position. Two subspecies exist in North America, P. v. richardii in the Pacific Ocean and P. v. vitulina in the Atlantic. Strong natal philopatry of harbour seals can result in fine-scale genetic structure and isolation by distance. Management of harbour seals is expected to benefit from improved resolution of seal population structure and dynamics. Here, we use genotyping-by-sequencing to genotype 146 harbour seals from the eastern Pacific Ocean (i.e. British Columbia (BC), Oregon and California) and the western Atlantic Ocean (i.e. Québec, Newfoundland and Labrador). Using 12,742 identified variants, we confirm the recently identified elevated genetic diversity in the eastern Pacific relative to the western Atlantic and greatest differentiation between the subspecies. Further, we demonstrate that this is independent of reference genome bias or other potential technical artefacts. Coast-specific analyses with 8933 and 3828 variants in Pacific and Atlantic subspecies, respectively, identify divergence between BC and Oregon-California, and between Québec and Newfoundland-Labrador. Unexpected PCA outlier clusters were observed in two populations due to cryptic relatedness of individuals; subsequently, closely related samples were removed. Admixture analysis indicates an isolation-by-distance signature where Oregon seals contained some of the BC signature, whereas California did not. Additional sampling is needed in the central and north coast of BC to determine whether a discrete separation of populations exists within the region.

Heartworm and seal louse: Trends in prevalence, characterisation of impact and transmission pathways in a unique parasite assembly on seals in the North and Baltic Sea.

The ectoparasitic seal louse, Echinophthirius horridus infects harbour (Phoca vitulina) and grey seals (Halichoerus grypus) in the North and Baltic Sea. The endoparasitic heartworm Acanthocheilonema spirocauda parasitizes the right heart and blood vessels of harbour seals. The complete lifecycle of the heartworm is not entirely understood although the seal louse is assumed to serve as vector for its transmission. Knowledge about the impact of both parasite species on host health are scarce. In this study, necropsy data and archived parasites of harbour and grey seals in German waters were analysed to determine long-term seal louse (SLP) and heartworm prevalence (HWP) from 2014 to 2021. Histology, microbiology and scanning electron microscopy (SEM) were applied on seal louse infected and uninfected skin to investigate associated lesions and the health impact. During the study period, HWP in harbour seals was 13%, the SLP in harbour seals was 4% and in grey seals 10%. HWP of harbour seals was significantly higher during the winter months compared to the summer. SLP in adults was significantly higher in comparison to juvenile harbour seals. SLP varied significantly between grey seals from the North and Baltic Sea. Filarial nematodes were detected in the haemocoel, pharynx, and intestine of E. horridus highlighting the seal louse as vector for heartworms. Alopecia and folliculitis were associated with the attachment posture of E. horridus and microbiological investigations isolated bacteria commonly associated with folliculitis.

Cardiorespiratory adaptations in small cetaceans and marine mammals.

The dive response, or the 'master switch of life', is probably the most studied physiological trait in marine mammals and is thought to conserve the available O2 for the heart and brain. Although generally thought to be an autonomic reflex, several studies indicate that the cardiovascular changes during diving are anticipatory and can be conditioned. The respiratory adaptations, where the aquatic breathing pattern resembles intermittent breathing in land mammals, with expiratory flow exceeding 160 litres s-1 has been measured in cetaceans, and where exposure to extreme pressures results in alveolar collapse (atelectasis) and recruitment upon ascent. Cardiorespiratory coupling, where breathing results in changes in heart rate, has been proposed to improve gas exchange. Cardiorespiratory coupling has also been reported in marine mammals, and in the bottlenose dolphin, where it alters both heart rate and stroke volume. When accounting for this respiratory dependence on cardiac function, several studies have reported an absence of a diving-related bradycardia except during dives that exceed the duration that is fuelled by aerobic metabolism. This review summarizes what is known about the respiratory physiology in marine mammals, with a special focus on cetaceans. The cardiorespiratory coupling is reviewed, and the selective gas exchange hypothesis is summarized, which provides a testable mechanism for how breath-hold diving vertebrates may actively prevent uptake of N2 during routine dives, and how stress results in failure of this mechanism, which results in diving-related gas emboli.

Fecal and skin microbiota of two rescued Mediterranean monk seal pups during rehabilitation.

IMPORTANCE: This study showed that during the rehabilitation of two rescued Mediterranean monk seal pups (Monachus monachus), the skin and fecal bacterial communities showed similar succession patterns between the two individuals. This finding means that co-housed pups share their microbiomes, and this needs to be considered in cases of infection outbreaks and their treatment. The housing conditions, along with the feeding scheme and care protocols, including the admission of antibiotics as prophylaxis, probiotics, and essential food supplements, resulted in bacterial communities with no apparent pathogenic bacteria. This is the first contribution to the microbiome of the protected seal species of M. monachus and contributes to the animal's conservation practices through its microbiome.

Improving the knowledge management of marine megafauna strandings.

Although the marine megafauna often strands on beaches around the world, such as sea turtles and whales, stranding data are poorly managed and incorporated into management and conservation strategies. Here we use a knowledge value chain framework to call attention for the urgent need to improve our data architecture and knowledge management on marine megafauna strandings. We use Brazil, a continental megadiverse federative republic, as study model. After describing the main components and identifying the strengths and weaknesses of the current Brazilian data architecture, we propose 10 practical measures for its improvement involving researchers, companies, non-governmental organizations, legislators, policy makers, public agents, citizen scientists, and local communities. Although Brazil has notable strengths such as comprehensive environmental legislation, hundreds of scientists and dozens of prestigious research institutions, stranding data is not translated into technical-scientific knowledge; technical-scientific knowledge is not transformed into effective public regulations; deficient regulations lead to bad decisions and limited actions, which in turn result in ineffective management and conservation strategies. In light of the UN Decade of Ocean Science for Sustainable Development (2021-2030), we propose (1) expanding standardized beach monitoring projects to the entire Brazilian coast; (2) creating a governmental database with FAIR principles; (3) encouraging the development of broad citizen science initiatives; (4) funding scientists and research institutions; (5) boosting outreach activities among researchers to popularize the scientific knowledge; (6) raising awareness among legislators and policy makers on the problem of strandings; (7) updating the existing legal provisions on the environmental licensing of activities developed at sea; (8) hiring new environmental analysts and inspectors and improving the infrastructure of executing environmental agencies; (9) strengthening existing conservation networks with multiple stakeholders; and (10) making the results of the management and conservation strategies broadly accessible to society. These recommendations may also apply to other coastal countries around the world.

Imitation of Novel Intransitive Body Actions in a Beluga Whale (Delphinapterus leucas): A "Do as Other Does" Study.

Cetaceans are well known for their unique behavioral habits, such as calls and tactics. The possibility that these are acquired through social learning continues to be explored. This study investigates the ability of a young beluga whale to imitate novel behaviors. Using a do-as-other-does paradigm, the subject observed the performance of a conspecific demonstrator involving familiar and novel behaviors. The subject: (1) learned a specific 'copy' command; (2) copied 100% of the demonstrator's familiar behaviors and accurately reproduced two out of three novel actions; (3) achieved full matches on the first trial for a subset of familiar behaviors; and (4) demonstrated proficiency in coping with each familiar behavior as well as the two novel behaviors. This study provides the first experimental evidence of a beluga whale's ability to imitate novel intransitive (non-object-oriented) body movements on command. These results contribute to our understanding of the remarkable ability of cetaceans, including dolphins, orcas, and now beluga whales, to engage in multimodal imitation involving sounds and movements. This ability, rarely documented in non-human animals, has significant implications for the development of survival strategies, such as the acquisition of knowledge about natal philopatry, migration routes, and traditional feeding areas, among these marine mammals.

Intracellular negative feedback mechanisms in blubber and muscle moderate acute stress responses in fasting seals.

Animals may limit the cost of stress responses during key life history stages such as breeding and molting by reducing tissue sensitivity to energy-mobilizing stress hormones (e.g. cortisol). We measured expression of genes encoding glucocorticoid receptor (GR, NR3C1), GR inhibitor (FKBP5) and cortisol-inactivating enzyme (HSD11B2) in blubber and muscle of northern elephant seals before and after stress axis stimulation by adrenocorticotropic hormone (ACTH) early and late in a fasting period associated with molting. ACTH elevated cortisol levels for >24 h and increased FKBP5 and HSD11B2 expression while downregulating NR3C1 expression in blubber and muscle, suggesting robust intracellular negative feedback in peripheral tissues. This feedback was maintained over prolonged fasting, despite differences in baseline cortisol and gene expression levels between early and late molt, suggesting that fasting-adapted animals use multiple tissue-specific, intracellular negative feedback mechanisms to modulate downstream impacts of acute stress responses during key life history stages.

Infection with Pythium flevoense in a harbour porpoise (Phocoena phocoena) as a novel cause of dermatitis in marine mammals.

The oomycete Pythium flevoense was diagnosed as the cause of dermatitis in a young adult female harbour porpoise (Phocoena phocoena) that had been trapped in a pound net in a temperate saltwater environment. Disease from Pythium sp. infection-pythiosis-is infrequently diagnosed in humans, horses, dogs, cattle, and few other mammalian species. Pythiosis is typically associated with exposure to tropical or subtropical freshwater conditions, and typically caused by Pythium insidiosum. However, until now, pythiosis has been reported in neither marine mammals nor temperate saltwater conditions, and P. flevoense is not known as a cause of pythiosis in mammals. This porpoise developed generalised dermatitis despite treatment and euthanasia was necessary. Histopathological evaluation revealed a chronic active erosive dermatitis, with intralesional hyphae morphologically consistent with a Pythium sp. PCR analysis and sequencing of affected skin matched Pythium flevoense with a 100% similarity to the reference strain. Additional diagnostics excluded other pathogens. Based on this case report, P. flevoense needs to be considered as a mammalian pathogen. Furthermore, harbour porpoises and possibly other marine mammals may be at risk of infection with P. flevoense, and pythiosis should be included in the differential diagnosis of dermatitis in marine mammals.

Age-specific reproduction in female Steller sea lions in Southeast Alaska.

Age-, region-, and year-specific estimates of reproduction are needed for monitoring wildlife populations during periods of ecosystem change. Population dynamics of Steller sea lions (Eumetopias jubatus) in Southeast Alaska varied regionally (with high population growth and survival in the north vs. the south) and annually (with reduced adult female survival observed following a severe marine heatwave event), but reproductive performance is currently unknown. We used mark-resighting data from 1006 Steller sea lion females marked as pups at ~3 weeks of age from 1994 to 1995 and from 2001 to 2005 and resighted from 2002 to 2019 (to a maximum age of 25) to examine age-, region-, and year-specific reproduction. In the north versus the south, age of first reproduction was earlier (beginning at age 4 vs. age 5, respectively) but annual birth probabilities of parous females were reduced by 0.05. In an average year pre-heatwave, the proportion of females with pup at the end of the pupping season peaked at ages 12-13 with ~0.60/0.65 (north/south) with pup, ~0.30/0.25 with juvenile, and ~0.10 (both regions) without a dependent. In both regions, reproductive senescence was gradual after age 12: ~0.40, 0.40, and 0.20 of females were in these reproductive states, respectively, by age 20. Correcting for neonatal mortality, true birth probabilities at peak ages were 0.66/0.72 (north/south). No cost of reproduction on female survival was detected, but pup production remained lower (-0.06) after the heatwave event, which if sustained could result in population decline in the south. Reduced pup production and greater retention of juveniles during periods of poor prey conditions may be an important strategy for Steller sea lions in Southeast Alaska, where fine-tuning reproduction based on nutritional status may improve the lifetime probability of producing pups under good conditions in a variable and less productive environment.