Determining energy expenditure in a large seabird using accelerometry.

The trade off between energy gained and expended is the foundation of understanding how, why and when animals perform any activity. Based on the concept that animal movements have an energetic cost, accelerometry is increasingly being used to estimate energy expenditure. However, validation of accelerometry as an accurate proxy for field metabolic rate in free-ranging species is limited. In the present study, Australasian gannets (Morus serrator) from the Pope's Eye colony (38°16'42″S 144°41'48″E), south-eastern Australia, were equipped with GPS and tri-axial accelerometers and dosed with doubly labelled water (DLW) to measure energy expenditure during normal behaviour for 3-5 days. The correlation between daily energy expenditure from the DLW and vectorial dynamic body acceleration (VeDBA) was high for both a simple correlation and activity-specific approaches (R2=0.75 and 0.80, respectively). Varying degrees of success were observed for estimating at-sea metabolic rate from accelerometry when removing time on land using published energy expenditure constants (R2=0.02) or activity-specific approaches (R2=0.42). The predictive capacity of energy expenditure models for total and at-sea periods was improved by the addition of total distance travelled and proportion of the sampling period spent at sea during the night, respectively (R2=0.61-0.82). These results indicate that accelerometry can be used to estimate daily energy expenditure in free-ranging gannets and its accuracy may depend on the inclusion of movement parameters not detected by accelerometry.

Increasing numbers of killer whale individuals use fisheries as feeding opportunities within subantarctic populations.

Fisheries can generate feeding opportunities for large marine predators in the form of discards or accessible catch. How the use of this anthropogenic food may spread as a new behaviour, across individuals within populations over time, is poorly understood. This study used a 16-year (2003-2018) monitoring of two killer whale Orcinus orca subantarctic populations (regular and Type-D at Crozet), and Bayesian multistate capture-mark-recapture models, to assess temporal changes in the number of individuals feeding on fish caught on hooks ('depredation' behaviour) of a fishery started in 1996. For both populations, the number of depredating individuals increased during the study period (34 to 94 for regular; 17 to 43 for Type-D). Increasing abundance is unlikely to account for this and, rather, the results suggest depredation was acquired by increasing numbers of existing individuals. For regular killer whales, a plateau reached from 2014 suggests that it took 18 years for the behaviour to spread across the whole population. A more recent plateau was apparent for Type-Ds but additional years are needed to confirm this. These findings show how changes in prey availability caused by human activities lead to rapid, yet progressive, innovations in killer whales, likely altering the ecological role of this top-predator.

Stage-dependent niche segregation: insights from a multi-dimensional approach of two sympatric sibling seabirds.

Niche theory predicts that to reduce competition for the same resource, sympatric ecologically similar species should exploit divergent niches and segregate in one or more dimensions. Seasonal variations in environmental conditions and energy requirements can influence the mechanisms and the degree of niche segregation. However, studies have overlooked the multi-dimensional aspect of niche segregation over the whole annual cycle, and key facets of species co-existence still remain ambiguous. The present study provides insights into the niche use and partitioning of two morphologically and ecologically similar seabirds, the common (CDP, Pelecanoides urinatrix) and the South Georgian diving petrel (SGDP, Pelecanoides georgicus). Using phenology, at-sea distribution, diving behavior and isotopic data (during the incubation, chick-rearing and non-breeding periods), we show that the degree of partitioning was highly stage-dependent. During the breeding season, the greater niche segregation during chick-rearing than incubation supported the hypothesis that resource partitioning increases during energetically demanding periods. During the post breeding period, while species-specific latitudinal differences were expected (species specific water mass preference), CDP and SGDP also migrated in divergent directions. This segregation in migration area may not be only a response to the selective pressure arising from competition avoidance between sympatric species, but instead, could reflect past evolutionary divergence. Such stage-dependent and context-dependent niche segregation demonstrates the importance of integrative approaches combining techniques from different fields, throughout the entire annual cycle, to better understand the co-existence of ecologically similar species. This is particularly relevant in order to fully understand the short and long-term effects of ongoing environmental changes on species distributions and communities.This work demonstrates the need of integrative multi-dimensional approaches combining concepts and techniques from different fields to understand the mechanism and causal factors of niche segregation.

Animal lifestyle affects acceptable mass limits for attached tags.

Animal-attached devices have transformed our understanding of vertebrate ecology. To minimize any associated harm, researchers have long advocated that tag masses should not exceed 3% of carrier body mass. However, this ignores tag forces resulting from animal movement. Using data from collar-attached accelerometers on 10 diverse free-ranging terrestrial species from koalas to cheetahs, we detail a tag-based acceleration method to clarify acceptable tag mass limits. We quantify animal athleticism in terms of fractions of animal movement time devoted to different collar-recorded accelerations and convert those accelerations to forces (acceleration × tag mass) to allow derivation of any defined force limits for specified fractions of any animal's active time. Specifying that tags should exert forces that are less than 3% of the gravitational force exerted on the animal's body for 95% of the time led to corrected tag masses that should constitute between 1.6% and 2.98% of carrier mass, depending on athleticism. Strikingly, in four carnivore species encompassing two orders of magnitude in mass (ca 2-200 kg), forces exerted by '3%' tags were equivalent to 4-19% of carrier body mass during moving, with a maximum of 54% in a hunting cheetah. This fundamentally changes how acceptable tag mass limits should be determined by ethics bodies, irrespective of the force and time limits specified.

Consistent sociality but flexible social associations across temporal and spatial foraging contexts in a colonial breeder.

When the consequences of sociality differ depending on the state of individual animals and the experienced environment, individuals may benefit from altering their social behaviours in a context-dependent manner. Thus, to fully address the hypotheses about the role of social associations it is imperative to consider the multidimensional nature of sociality by explicitly examining social associations across multiple scales and contexts. We simultaneously recorded > 8000 associations from 85% of breeding individuals from a colony of Australasian gannets (Morus serrator) over a 2-week period, and examined gregariousness across four foraging states using multilayer social network analysis. We found that social associations varied in a context-dependent manner, highlighting that social associations are most prevalent during foraging (local enhancement) and in regions expected to provide clustered resources. We also provide evidence of individual consistency in gregariousness, but flexibility in social associates, demonstrating that individuals can adjust their social behaviours to match experienced conditions.

Evidence of sociality in the timing and location of foraging in a colonial seabird.

Social foraging behaviours, which range from cooperative hunting to local enhancement, can result in increased prey capture and access to information, which may significantly reduce time and energy costs of acquiring prey. In colonial species, it has been proposed that the colony itself may act as a site of social information transfer and group formation. However, conclusive evidence from empirical studies is lacking. In particular, most studies in colonial species have generally focussed on behaviours either at the colony or at foraging sites in isolation, and have failed to directly connect social associations at the colony to social foraging. In this study, we simultaneously tracked 85% of a population of Australasian gannets (Morus serrator) over multiple foraging trips, to study social associations at the colony and test whether these associations influence the location of foraging sites. We found that gannets positively associate with conspecifics while departing from the colony and that co-departing gannets have more similar initial foraging patches than individuals that did not associate at the colony. These results provide strong evidence for the theory that the colony may provide a source of information that influences foraging location.

Regional variability in diving physiology and behavior in a widely distributed air-breathing marine predator, the South American sea lion (Otaria byronia).

Our understanding of how air-breathing marine predators cope with environmental variability is limited by our inadequate knowledge of their ecological and physiological parameters. Because of their wide distribution along both coasts of the sub-continent, South American sea lions (Otaria byronia) provide a valuable opportunity to study the behavioral and physiological plasticity of a marine predator in different environments. We measured the oxygen stores and diving behavior of South American sea lions throughout most of its range, allowing us to demonstrate that diving ability and behavior vary across its range. We found no significant differences in mass-specific blood volumes of sea lions among field sites and a negative relationship between mass-specific oxygen storage and size, which suggests that exposure to different habitats and geographical locations better explains oxygen storage capacities and diving capability in South American sea lions than body size alone. The largest animals in our study (individuals from Uruguay) were the shallowest and shortest duration divers, and had the lowest mass-specific total body oxygen stores, while the deepest and longest duration divers (individuals from southern Chile) had significantly larger mass-specific oxygen stores, despite being much smaller animals. Our study suggests that the physiology of air-breathing diving predators is not fixed, but that it can be adjusted, to a certain extent, depending on the ecological setting and or habitat. These adjustments can be thought of as a 'training effect': as the animal continues to push its physiological capacity through greater hypoxic exposure, its breath-holding capacity increases.

From video recordings to whisker stable isotopes: a critical evaluation of timescale in assessing individual foraging specialisation in Australian fur seals.

Estimating the degree of individual specialisation is likely to be sensitive to the methods used, as they record individuals' resource use over different time-periods. We combined animal-borne video cameras, GPS/TDR loggers and stable isotope values of plasma, red cells and sub-sampled whiskers to investigate individual foraging specialisation in female Australian fur seals (Arctocephalus pusillus doriferus) over various timescales. Combining these methods enabled us to (1) provide quantitative information on individuals' diet, allowing the identification of prey, (2) infer the temporal consistency of individual specialisation, and (3) assess how different methods and timescales affect our estimation of the degree of specialisation. Short-term inter-individual variation in diet was observed in the video data (mean pairwise overlap = 0.60), with the sampled population being composed of both generalist and specialist individuals (nested network). However, the brevity of the temporal window is likely to artificially increase the level of specialisation by not recording the entire diet of seals. Indeed, the correlation in isotopic values was tighter between the red cells and whiskers (mid- to long-term foraging ecology) than between plasma and red cells (short- to mid-term) (R(2) = 0.93-0.73 vs. 0.55-0.41). δ(13)C and δ(15)N values of whiskers confirmed the temporal consistency of individual specialisation. Variation in isotopic niche was consistent across seasons and years, indicating long-term habitat (WIC/TNW = 0.28) and dietary (WIC/TNW = 0.39) specialisation. The results also highlight time-averaging issues (under-estimation of the degree of specialisation) when calculating individual specialisation indices over long time-periods, so that no single timescale may provide a complete and accurate picture, emphasising the benefits of using complementary methods.

Disentangling the cause of a catastrophic population decline in a large marine mammal.

Considerable uncertainties often surround the causes of long-term changes in population abundance. One striking example is the precipitous decline of southern sea lions (SSL; Otariaflavescens) at the Falkland Islands, from 80 555 pups in the mid 1930s to just 5506 pups in 1965. Despite an increase in SSL abundance over the past two decades, the population has not recovered, with the number of pups born in 2014 (minimum 4443 pups) less than 6% of the 1930s estimate. The order-of-magnitude decline is primarily attributed to commercial sealing in Argentina. Here, we test this established paradigm and alternative hypotheses by assessing (1) commercial sealing at the Falkland Islands, (2) winter migration of SSL from the Falkland Islands to Argentina, (3) whether the number of SSL in Argentina could have sustained the reported level of exploitation, and (4) environmental change. The most parsimonious hypothesis explaining the SSL population decline was environmental change. Specifically, analysis of 160 years of winter sea surface temperatures revealed marked changes, including a period of warming between 1930 and 1950 that was consistent with the period of SSL decline. Sea surface temperature changes likely influenced the distribution or availability of SSL prey and impacted its population dynamics. We suggest that historical harvesting may not always be the "smoking gun" as is often purported. Rather, our conclusions support the growing evidence for bottom-up forcing on the abundance of species at lower trophic levels (e.g., plankton and fish) and resulting impacts on higher trophic levels across a broad range of ecosystems.

Determinants of individual foraging specialization in large marine vertebrates, the Antarctic and subantarctic fur seals.

The degree of individual specialization in resource use differs widely among wild populations where individuals range from fully generalized to highly specialized. This interindividual variation has profound implications in many ecological and evolutionary processes. A recent review proposed four main ecological causes of individual specialization: interspecific and intraspecific competition, ecological opportunity and predation. Using the isotopic signature of subsampled whiskers, we investigated to what degree three of these factors (interspecific and intraspecific competition and ecological opportunity) affect the population niche width and the level of individual foraging specialization in two fur seal species, the Antarctic and subantarctic fur seals (Arctocephalus gazella and Arctocephalus tropicalis), over several years. Population niche width was greater when the two seal species bred in allopatry (low interspecific competition) than in sympatry or when seals bred in high-density stabilized colonies (high intraspecific competition). In agreement with the niche variation hypothesis (NVH), higher population niche width was associated with higher interindividual niche variation. However, in contrast to the NVH, all Antarctic females increased their niche width during the interbreeding period when they had potential access to a wider diversity of foraging grounds and associated prey (high ecological opportunities), suggesting they all dispersed to a similar productive area. The degree of individual specialization varied among populations and within the annual cycle. Highest levels of interindividual variation were found in a context of lower interspecific or higher intraspecific competition. Contrasted results were found concerning the effect of ecological opportunity. Depending on seal species, females exhibited either a greater or lower degree of individual specialization during the interbreeding period, reflecting species-specific biological constraints during that period. These results suggest a significant impact of ecological interactions on the population niche width and degree of individual specialization. Such variation at the individual level may be an important factor in the species plasticity with significant consequences on how it may respond to environmental variability.

Variation in body condition during the post-moult foraging trip of southern elephant seals and its consequences on diving behaviour.

Mature female southern elephant seals (Mirounga leonina) come ashore only in October to breed and in January to moult, spending the rest of the year foraging at sea. Mature females may lose as much as 50% of their body mass, mostly in lipid stores, during the breeding season due to fasting and lactation. When departing to sea, post-breeding females are negatively buoyant, and the relative change in body condition (i.e. density) during the foraging trip has previously been assessed by monitoring the descent rate during drift dives. However, relatively few drift dives are performed, resulting in low resolution of the temporal reconstruction of body condition change. In this study, six post-breeding females were equipped with time-depth recorders and accelerometers to investigate whether changes in active swimming effort and speed could be used as an alternative method of monitoring density variations throughout the foraging trip. In addition, we assessed the consequences of density change on the swimming efforts of individuals while diving and investigated the effects on dive duration. Both descent swimming speed and ascent swimming effort were found to be strongly correlated to descent rate during drift dives, enabling the fine-scale monitoring of seal density change over the whole trip. Negatively buoyant seals minimized swimming effort during descents, gliding down at slower speeds, and reduced their ascent swimming effort to maintain a nearly constant swimming speed as their buoyancy increased. One per cent of seal density variation over time was found to induce a 20% variation in swimming effort during dives with direct consequences on dive duration.

Foraging behaviour and habitat use during chick-rearing in the Australian endemic black-faced cormorant (Phalacrocorax fuscescens).

Despite its wide distribution, relatively little is known of the foraging ecology and habitat use of the black-faced cormorant (Phalacrocorax fuscescens), an Australian endemic seabird. Such information is urgently required in view of the rapid oceanic warming of south-eastern Australia, the stronghold of the species. The present study used a combination of opportunistically collected regurgitates and GPS/dive behaviour data loggers to investigate diet, foraging behaviour and habitat-use of black-faced cormorants during four chick-rearing periods (2020-2023) on Notch Island, northern Bass Strait. Observed prey species were almost exclusively benthic (95%), which is consistent with the predominantly benthic diving behaviour recorded. Males foraged at deeper depths than females (median depth males: 18 m; median depth females: 8 m), presumably due to a greater physiological diving capacity derived from their larger body size. This difference in dive depths was associated with sexual segregation of foraging locations, with females predominantly frequenting shallower areas closer to the coastline. These findings have strong implications for the management of the species, as impacts of environmental change may disproportionally affect the foraging range of one sex and, thereby, reproductive success.

Morphometric and molecular characterization of the species of Uncinaria Frölich, 1789 (Nematoda) parasitic in the Australian fur seal Arctocephalus pusillus doriferus (Schreber), with notes on hookworms in three other pinniped hosts.

This study presents morphological and molecular data on hookworms from the Australian fur seal Arctocephalus pusillus doriferus (Schreber) currently identified in Australian waters as Uncinaria hamiltoni Baylis, 1933. Additional specimens from the Australian sea lion Neophoca cinerea (Péron) and the New Zealand fur seal Arctocephalus forsteri (Lesson) from Australia, and the Southern elephant seal Mirounga leonina (Linnaeus) from Antarctica, were included. Using the internal transcribed spacer (ITS), hookworms from A. p. doriferus, N. cinerea and A. forsteri were found to be genetically similar but distinct from Uncinaria spp. found in M. leonina from Antarctica, as well as from Zalophus californianus (Lesson) and Callorhinus ursinus (Linnaeus) from California. Few morphological differences were detected between these taxa.

Energetic consequences of prey type in little penguins (Eudyptula minor).

Investigation of foraging decisions can help understand how animals efficiently gather and exploit food. Prey chase and handling times are important aspects of foraging efficiency, influencing the net energy gain derived from a prey item. However, these metrics are often overlooked in studies of foraging behaviour due to the difficulty in observing them. The present study used animal-borne cameras to investigate the type, duration and energetic consequences of predator-prey interactions in little penguins (Eudyptula minor) (n = 32) from two colonies in Bass Strait, south-eastern Australia. A total of seven main prey items were observed and consumed by little penguins. Penguins were observed to consume prey types and use strategies that have not been previously documented. These included consumption of bellowsfish (Macroramphosus scolopax) and other fish species captured sheltering around jellyfish or extracted dead from the tentacles. Chase and handling time varied with prey type and lasted approximately 2 s for most prey. Profitability varied among prey types, with a greater amount of low profitable prey being consumed, suggesting a trade-off between minimizing energetic costs, and increasing capture rates. These results highlight the use of animal-borne video data loggers to further understand the foraging adaptations of important predators in the marine environment.

Using ecotourism boats for estimating the abundance of a bottlenose dolphin population in south-eastern Australia.

It is challenging to collect robust, long-term datasets to properly monitor the viability and social structure of large, long-lived animals, especially marine mammals. The present study used a unique long-term dataset to investigate the population parameters and social structure of a poorly studied population of bottlenose dolphins (Tursiops sp.) in southern Port Phillip Bay, south-eastern Australia. Photo-identification images have been collected between 2012-2022 both opportunistically and following a protocol by patrons, staff, and volunteers of ecotourism companies using their vessels as platforms. The resulting large dataset was available to be processed through the online platform Flukebook and used in capture recapture models to estimate abundance and demographic parameters. In addition, the social structure of the population and the reproductive parameters were investigated. The marked adult population abundance (45.2 ± 2.7 individuals) was found to be stable over the last decade and the calving rate ranged between 0.06-0.19 new calves per identified individuals per year, while the inter-birth interval was 3.7 ± 0.8 years. Social analysis suggested the population has a fission-fusion structure with no apparent clusters. The stability of the population over the study period suggests no deleterious effect of anthropogenic or environmental factors during the last decade. This study is the outcome of the effort of the ecotourism organisations and the results obtained, along with their similarity to those of other dolphin populations worldwide, highlight the importance of such data sources for long-term information that would otherwise be too expensive or logistically difficult to obtain.

Pacific Gulls (Larus pacificus) as Potential Vectors of Coxiella burnetii in an Australian Fur Seal Breeding Colony.

Recently, Coxiella burnetii has been described as a novel pathogen potentially contributing to decreased pup production in Australian fur seals (AusFS, Arctocephalus pusillus doriferus). Pacific gulls (PGs, Larus pacificus) are known to scavenge AusFS placental material during the fur seal breeding season. It is hypothesized that PGs may act as vectors for this pathogen. In the present study, cloacal swabs, oral swabs and serum were collected from PGs on Kanowna Island (KI, an AusFS breeding colony) and a nearby island, Seal Island (SI), not occupied by pinnipeds. All sample sets were evaluated with qPCR for the com1, htpAB and IS1111 markers. Most oral and cloacal swabs from KI tested positive on both the com1 (94.1%; 88.2%) and htpAB targets (76.5%; 76.5%). Amplification was very low from the SI oral swabs and cloacal swabs. Only the KI serum samples had amplification (17.7% for both com1 and htpAB). There was no IS1111 amplification in either colony. The results demonstrate that PGs can potentially act as vectors for the spread of C. burnetii. In some birds, C. burnetii was detectable in the serum, indicating that gulls can experience bacteraemia. It appears that different feeding strategies in the same species within the same ecosystem can have profound effects on the prevalence of pathogens. Further studies are required to better understand the epidemiology and potential risks of this organism.

A Novel Marine Mammal Coxiella burnetii-Genome Sequencing Identifies a New Genotype with Potential Virulence.

The obligate intracellular bacterial pathogen Coxiella burnetii has been identified in a few species of marine mammals, some of which are showing population declines. It has been hypothesized that C. burnetii in marine mammals is a distinct genotype that varies significantly from the typical terrestrial genotypes. It appears to lack an IS1111. Isolates originating from Australian marine animals have a distinctly non-Australian profile of multiple-locus variable-number tandem-repeat analysis (MLVA). Extracted Coxiella DNA of Australian fur seal placental origin was sequenced using the Novaseq platform. Illumina 150 bp paired-end reads were filtered and trimmed with Trimgalore. The microbial community present in the sequenced genome was evaluated with Kraken and Bracken software using the NCBI database. A phylogenetic analysis was performed using 1131 core genes. Core genes were identified using Panaroo and inputted into Iqtree to determine the maximum-likelihood tree. A second phylogenetic tree was created using Rickettsiella grylii and using seven housekeeping genes. Results were compared with the C. burnetii Nine Mile RSA439 virulent genome. This new Australian marine mammal isolate of Coxiella (PG457) appears to be a novel genotype that lacks IS1111 and has a distinct MLVA signature (ms26, ms27, ms28, ms30, and ms31). The presence of genes for multiple virulence factors appears to give this genotype sufficient pathogenicity for it to be considered a possible causative agent of abortion in Australian fur seals as well as a potential zoonotic risk.

Geographical, temporal, and individual-based differences in the trophic ecology of female Cape fur seals.

Information on resource use and trophic dynamics of marine predators is important for understanding their role in ecosystem functioning and predicting population-level responses to environmental change. Where separate populations experience different local environmental conditions, geographic variability in their foraging ecology is often expected. Within populations, individuals also vary in morphology, physiology, and experience, resulting in specialization in resource use. In this context, isotopic compositions of incrementally grown tissues such as keratinous hairs offer a valuable opportunity to study long-term variation in resource and habitat use. We investigated the trophic ecology of female Cape fur seals (Arctocephalus pusillus pusillus) using carbon and nitrogen isotopic compositions of serially sampled whiskers collected at four breeding sites along the coast of South Africa. Drawing on over 900 isotopic measurements, we assessed geographic variability in isotopic niche width between colonies and the degree of individual specialization. We found slight, but clear geographic differences in isotopic ratios and isotopic niche widths, seemingly related to ecological setting, with niche widths being proportional to the area of available shelf and shelf-slope habitat surrounding the colony. We further identified periodic oscillations in isotopic ratios, which likely reflect temporal patterns in foraging distribution and prey type, linked to shifts in the availability of prey resources and their interaction with constraints on individual females throughout their breeding cycle. Finally, individual specialization indices revealed that each of the study populations contain specialist individuals that utilize only a small subset of the total population niche width. The degree of individual specialization was, however, not consistent across colonies and may reflect an interactive influence between density-dependent effects and habitat heterogeneity. Overall, this study provides important information on the trophic ecology of Cape fur seals breeding in South Africa and highlights the need to consider geographic and individual variability when assessing the foraging ecology of marine predators.

Global assessment of marine plastic exposure risk for oceanic birds.

Plastic pollution is distributed patchily around the world's oceans. Likewise, marine organisms that are vulnerable to plastic ingestion or entanglement have uneven distributions. Understanding where wildlife encounters plastic is crucial for targeting research and mitigation. Oceanic seabirds, particularly petrels, frequently ingest plastic, are highly threatened, and cover vast distances during foraging and migration. However, the spatial overlap between petrels and plastics is poorly understood. Here we combine marine plastic density estimates with individual movement data for 7137 birds of 77 petrel species to estimate relative exposure risk. We identify high exposure risk areas in the Mediterranean and Black seas, and the northeast Pacific, northwest Pacific, South Atlantic and southwest Indian oceans. Plastic exposure risk varies greatly among species and populations, and between breeding and non-breeding seasons. Exposure risk is disproportionately high for Threatened species. Outside the Mediterranean and Black seas, exposure risk is highest in the high seas and Exclusive Economic Zones (EEZs) of the USA, Japan, and the UK. Birds generally had higher plastic exposure risk outside the EEZ of the country where they breed. We identify conservation and research priorities, and highlight that international collaboration is key to addressing the impacts of marine plastic on wide-ranging species.