Contrasting offspring dependence periods and diving development rates in two closely related marine mammal species.

Understanding the ontogeny of diving behaviour in marine megafauna is crucial owing to its influence on foraging success, energy budgets, and mortality. We compared the ontogeny of diving behaviour in two closely related species-northern elephant seals (Mirounga angustirostris, n = 4) and southern elephant seals (Mirounga leonina, n = 9)-to shed light on the ecological processes underlying migration. Although both species have similar sizes and behaviours as adults, we discovered that juvenile northern elephant seals have superior diving development, reaching 260 m diving depth in just 30 days, while southern elephant seals require 160 days. Similarly, northern elephant seals achieve dive durations of approximately 11 min on their first day of migration, while southern elephant seals take 125 days. The faster physiological maturation of northern elephant seals could be related to longer offspring dependency and post-weaning fast durations, allowing them to develop their endogenous oxygen stores. Comparison across both species suggests that weaned seal pups face a trade-off between leaving early with higher energy stores but poorer physiological abilities or leaving later with improved physiology but reduced fat stores. This trade-off might be influenced by their evolutionary history, which shapes their migration behaviours in changing environments over time.

Climate change impacts on seabirds and marine mammals: The importance of study duration, thermal tolerance and generation time.

Understanding climate change impacts on top predators is fundamental to marine biodiversity conservation, due to their increasingly threatened populations and their importance in marine ecosystems. We conducted a systematic review of the effects of climate change (prolonged, directional change) and climate variability on seabirds and marine mammals. We extracted data from 484 studies (4808 published studies were reviewed), comprising 2215 observations on demography, phenology, distribution, diet, behaviour, body condition and physiology. The likelihood of concluding that climate change had an impact increased with study duration. However, the temporal thresholds for the effects of climate change to be discernibly varied from 10 to 29 years depending on the species, the biological response and the oceanic study region. Species with narrow thermal ranges and relatively long generation times were more often reported to be affected by climate change. This provides an important framework for future assessments, with guidance on response- and region-specific temporal dimensions that need to be considered when reporting effects of climate change. Finally, we found that tropical regions and non-breeding life stages were poorly covered in the literature, a concern that should be addressed to enable a better understanding of the vulnerability of marine predators to climate change.

Spatial segregation in a sexually dimorphic central place forager: Competitive exclusion or niche divergence?

Sexual competition is increasingly recognized as an important selective pressure driving species distributions. However, few studies have investigated the relative importance of interpopulation versus intrapopulation competition in relation to habitat availability and selection. To explain spatial segregation between sexes that often occurs in non-territorial and central place foragers, such as seabirds, two hypotheses are commonly used. The 'competitive exclusion' hypothesis states that dominant individuals should exclude subordinate individuals through direct competition, whereas the 'niche divergence' hypothesis states that segregation occurs due to past competition and habitat specialization. We tested these hypotheses in two populations of an extreme wide-ranging and sexually dimorphic seabird, investigating the relative role of intrapopulation and interpopulation competition in influencing sex-specific distribution and habitat preferences. Using GPS loggers, we tracked 192 wandering albatrosses Diomedea exulans during four consecutive years (2016-2019), from two neighbouring populations in the Southern Ocean (Prince Edward and Crozet archipelagos). We simulated pseudo-tracks to create a null spatial distribution and used Kernel Density Estimates (KDE) and Resource Selection Functions (RSF) to distinguish the relative importance of within- versus between-population competition. Kernel Density Estimates showed that only intrapopulation sexual segregation was significant for each monitoring year, and that tracks between the two colonies resulted in greater overlap than expected from the null distribution, especially for the females. RSF confirmed these results and highlighted key at-sea foraging areas, even if the estimated of at-sea densities were extremely low. These differences in selected areas between sites and sexes were, however, associated with high interannual variability in habitat preferences, with no clear specific preferences per site and sex. Our results suggest that even with low at-sea population densities, historic intrapopulation competition in wide-ranging seabirds may have led to sexual dimorphism and niche specialization, favouring the 'niche divergence' hypothesis. In this study, we provide a protocol to study competition within as well as between populations of central place foragers. This is relevant for understanding their distribution patterns and population regulation, which could potentially improve management of threatened populations.

Foraging in a dynamic environment: Response of four sympatric sub-Antarctic albatross species to interannual environmental variability.

Seasonal and annual climate variations are linked to fluctuations in the abundance and distribution of resources, posing a significant challenge to animals that need to adjust their foraging behavior accordingly. Particularly during adverse conditions, and while energetically constrained when breeding, animals ideally need to be flexible in their foraging behavior. Such behavioral plasticity may separate "winners" from "losers" in light of rapid environmental changes due to climate change. Here, the foraging behavior of four sub-Antarctic albatross species was investigated from 2015/16 to 2017/18, a period characterized by pronounced environmental variability. Over three breeding seasons on Marion Island, Prince Edward Archipelago, incubating wandering (WA, Diomedea exulans; n = 45), grey-headed (GHA, Thalassarche chrysostoma; n = 26), sooty (SA, Phoebetria fusca; n = 23), and light-mantled (LMSA, P. palpebrata; n = 22) albatrosses were tracked with GPS loggers. The response of birds to environmental variability was investigated by quantifying interannual changes in their foraging behavior along two axes: spatial distribution, using kernel density analysis, and foraging habitat preference, using generalized additive mixed models and Bayesian mixed models. All four species were shown to respond behaviorally to environmental variability, but with substantial differences in their foraging strategies. WA was most general in its habitat use defined by sea surface height, eddy kinetic energy, wind speed, ocean floor slope, and sea-level anomaly, with individuals foraging in a range of habitats. In contrast, the three smaller albatrosses exploited two main foraging habitats, with habitat use varying between years. Generalist habitat use by WA and interannually variable use of habitats by GHA, SA, and LMSA would likely offer these species some resilience to predicted changes in climate such as warming seas and strengthening of westerly winds. However, future investigations need to consider other life-history stages coupled with demographic studies, to better understand the link between behavioral plasticity and population responses.

High mortality rates in a juvenile free-ranging marine predator and links to dive and forage ability.

High juvenile mortality rates are typical of many long-lived marine vertebrate predators. Insufficient development in dive and forage ability is considered a key driver of this. However, direct links to survival outcome are sparse, particularly in free-ranging marine animals that may not return to land.In this study, we conduct exploratory investigations toward early mortality in juvenile southern elephant seals Mirounga leonina. Twenty postweaning pups were equipped with (a) a new-generation satellite relay data tag, capable of remotely transmitting fine-scale behavioral movements from accelerometers, and (b) a location transmitting only tag (so that mortality events could be distinguished from device failures). Individuals were followed during their first trip at sea (until mortality or return to land). Two analyses were conducted. First, the behavioral movements and encountered environmental conditions of nonsurviving pups were individually compared to temporally concurrent observations from grouped survivors. Second, common causes of mortality were investigated using Cox's proportional hazard regression and penalized shrinkage techniques.Nine individuals died (two females and seven males) and 11 survived (eight females and three males). All but one individual died before the return phase of their first trip at sea, and all but one were negatively buoyant. Causes of death were variable, although common factors included increased horizontal travel speeds and distances, decreased development in dive and forage ability, and habitat type visited (lower sea surface temperatures and decreased total [eddy] kinetic energy).For long-lived marine vertebrate predators, such as the southern elephant seal, the first few months of life following independence represent a critical period, when small deviations in behavior from the norm appear sufficient to increase mortality risk. Survival rates may subsequently be particularly vulnerable to changes in climate and environment, which will have concomitant consequences on the demography and dynamics of populations.

Body condition influences ontogeny of foraging behavior in juvenile southern elephant seals.

Ontogeny of diving and foraging behavior in marine top predators is poorly understood despite its importance in population recruitment. This lack of knowledge is partly due to the difficulties of monitoring juveniles in the wild, which is linked to high mortality early in life. Pinnipeds are good models for studying the development of foraging behaviors because juveniles are large enough to robustly carry tracking devices for many months. Moreover, parental assistance is absent after a juvenile departs for its first foraging trip, minimizing confounding effects of parental input on the development of foraging skills. In this study, we tracked 20 newly weaned juvenile southern elephant seals from Kerguelen Islands for up to 338 days during their first trip at sea following weaning. We used a new generation of satellite relay tags, which allow for the transmission of dive, accelerometer, and location data. We also monitored, at the same time, nine adult females from the colony during their post-breeding trips, in order to compare diving and foraging behaviors. Juveniles showed a gradual improvement through time in their foraging skills. Like adults females, they remarkably adjusted their swimming effort according to temporal changes in buoyancy (i.e., a proxy of their body condition). They also did not appear to exceed their aerobic physiological diving limits, although dives were constrained by their smaller size compared to adults. Changes in buoyancy appeared to also influence their decision to either keep foraging or return to land, alongside the duration of their haul outs and choice of foraging habitat (oceanic vs. plateau). Further studies are thus needed to better understand how patterns in juveniles survival, and therefore elephant seal populations, might be affected by their changes in foraging skills and changes in their environmental conditions.

Processing of acceleration and dive data on-board satellite relay tags to investigate diving and foraging behaviour in free-ranging marine predators.

Biologging technologies are changing the way in which the marine environment is observed and monitored. However, because device retrieval is typically required to access the high-resolution data they collect, their use is generally restricted to those animals that predictably return to land. Data abstraction and transmission techniques aim to address this, although currently these are limited in scope and do not incorporate, for example, acceleration measurements which can quantify animal behaviours and movement patterns over fine-scales.In this study, we present a new method for the collection, abstraction and transmission of accelerometer data from free-ranging marine predators via the Argos satellite system. We test run the technique on 20 juvenile southern elephant seals Mirounga leonina from the Kerguelen Islands during their first months at sea following weaning. Using retrieved archival data from nine individuals that returned to the colony, we compare and validate abstracted transmissions against outputs from established accelerometer processing procedures.Abstracted transmissions included estimates, across five segments of a dive profile, of time spent in prey catch attempt (PrCA) behaviours, swimming effort and pitch. These were then summarised and compared to archival outputs across three dive phases: descent, bottom and ascent. Correlations between the two datasets were variable but generally good (dependent on dive phase, marginal R2 values of between .45 and .6 to >.9) and consistent between individuals. Transmitted estimates of PrCA behaviours and swimming effort were positively biased to those from archival processing.Data from this study represent some of the first remotely transmitted quantifications from accelerometers. The methods presented and analysed can be used to provide novel insight towards the behaviours and movements of free-ranging marine predators, such as juvenile southern elephant seals, from whom logger retrieval is challenging. Future applications could however benefit from some adaption, particularly to reduce positive bias in transmitted PrCA behaviours and swimming effort, for which this study provides useful insight.

Early diving behaviour in juvenile penguins: improvement or selection processes.

The early life stage of long-lived species is critical to the viability of population, but is poorly understood. Longitudinal studies are needed to test whether juveniles are less efficient foragers than adults as has been hypothesized. We measured changes in the diving behaviour of 17 one-year-old king penguins Aptenodytes patagonicus at Crozet Islands (subantartic archipelago) during their first months at sea, using miniaturized tags that transmitted diving activity in real time. We also equipped five non-breeder adults with the same tags for comparison. The data on foraging performance revealed two groups of juveniles. The first group made shallower and shorter dives that may be indicative of early mortality while the second group progressively increased their diving depths and durations, and survived the first months at sea. This surviving group of juveniles required the same recovery durations as adults, but typically performed shallower and shorter dives. There is thereby a relationship between improved diving behaviour and survival in young penguins. This long period of improving diving performance in the juvenile life stage is potentially a critical period for the survival of deep avian divers and may have implications for their ability to adapt to environmental change.