In the last thirty years, the emergence and progression of biologging technology has led to great advances in marine predator ecology. Large databases of location and dive observations from biologging devices have been compiled for an increasing number of diving predator species (such as pinnipeds, sea turtles, seabirds and cetaceans), enabling complex questions about animal activity budgets and habitat use to be addressed. Central to answering these questions is our ability to correctly identify and quantify the frequency of essential behaviours, such as foraging. Despite technological advances that have increased the quality and resolution of location and dive data, accurately interpreting behaviour from such data remains a challenge, and analytical methods are only beginning to unlock the full potential of existing datasets. This review evaluates both traditional and emerging methods and presents a starting platform of options for future studies of marine predator foraging ecology, particularly from location and two-dimensional (time-depth) dive data. We outline the different devices and data types available, discuss the limitations and advantages of commonly-used analytical techniques, and highlight key areas for future research. We focus our review on pinnipeds - one of the most studied taxa of marine predators - but offer insights that will be applicable to other air-breathing marine predator tracking studies. We highlight that traditionally-used methods for inferring foraging from location and dive data, such as first-passage time and dive shape analysis, have important caveats and limitations depending on the nature of the data and the research question. We suggest that more holistic statistical techniques, such as state-space models, which can synthesise multiple track, dive and environmental metrics whilst simultaneously accounting for measurement error, offer more robust alternatives. Finally, we identify a need for more research to elucidate the role of physical oceanography, device effects, study animal selection, and developmental stages in predator behaviour and data interpretation.

Young animals must learn to forage effectively to survive the transition from parental provisioning to independent feeding. Rapid development of successful foraging strategies is particularly important for capital breeders that do not receive parental guidance after weaning. The intrinsic and extrinsic drivers of variation in ontogeny of foraging are poorly understood for many species. Grey seals (Halichoerus grypus) are typical capital breeders; pups are abandoned on the natal site after a brief suckling phase, and must develop foraging skills without external input. We collected location and dive data from recently-weaned grey seal pups from two regions of the United Kingdom (the North Sea and the Celtic and Irish Seas) using animal-borne telemetry devices during their first months of independence at sea. Dive duration, depth, bottom time, and benthic diving increased over the first 40 days. The shape and magnitude of changes differed between regions. Females consistently had longer bottom times, and in the Celtic and Irish Seas they used shallower water than males. Regional sex differences suggest that extrinsic factors, such as water depth, contribute to behavioural sexual segregation. We recommend that conservation strategies consider movements of young naïve animals in addition to those of adults to account for developmental behavioural changes.

Rapid development of a successful foraging strategy is critical for juvenile survival, especially for naïve animals that receive no parental guidance. However, this process is poorly understood for many species. Although observation of early‐life movements is increasingly possible with miniaturisation of animal‐borne telemetry devices, analytical limitations remain. Here, we tracked 29 recently‐weaned grey seal Halichoerus grypus pups from colonies in two geographically distinct regions of the United Kingdom. We analysed at‐sea movements of pups throughout their initial months of nutritional independence to investigate the ontogeny of behaviour‐specific (foraging and travelling) movement patterns. Using generalized hidden Markov models (HMMs), we extended the conventional HMM framework to account for temporal changes in putative foraging and travelling movement characteristics, and investigate the effects of intrinsic (sex) and extrinsic (environment) factors on this process. Putative foraging behaviour became more tortuous with time, and travelling became faster and more directed, suggesting a reduction in search scale and an increase in travel efficiency as pups shifted from exploration to an adult‐like repeatable foraging strategy. Sex differences in movement characteristics were evident from colony departure, but sex‐specific activity budgets were only detected in one region. We show that sex‐specific behavioural strategies emerge before sexual size dimorphism in grey seals, and suggest that this phenomenon may occur in other long‐lived species. Our results also indicate that environmental variation may affect the emergence of sex‐specific foraging behaviour, highlighting the need to consider interacting intrinsic and extrinsic factors in shaping movement strategies of long‐lived vertebrates. Moreover, comparing the behavioural state estimations to those of a conventional HMM (no variation in state‐specific movement parameters) revealed differences in the amount and location of foraging activity, with implications for spatial conservation management. Overlooking intrinsic and extrinsic variation in movement processes could distort our understanding of foraging ecology, population dynamics and conservation requirements.

Capsule Three quarters of tracked Northern Gannets (Morus bassanus) at Grassholm gathered in rafts around the colony, concentrated within a recently designated at-sea Special Protection Area (SPA), but rafting was not correlated with foraging effort. Aims To investigate the incidence, distribution and foraging implications of Northern Gannet rafting behaviour in waters adjacent to a large colony. Methods Using bird-borne global positioning system (GPS) loggers we reconstructed at-sea movement and used a speed filter to identify rafting behaviour within 10 km of the colony. We mapped the spatial distribution of rafting events from 160 breeding individuals over 5 years, and investigated the relationship between foraging effort (trip duration and total distance travelled) and the presence/absence of rafting. Results On average, 74% of tracked birds engaged in rafting. Of the 381 foraging trips analysed, rafting was recorded on 237 (62%). Birds were more likely to raft on outbound (224 trips, 59%), than inbound journeys (38 trips, 10%). Presence/absence of rafting did not correlate significantly with foraging trip distance or duration nor with duration of nest attendance. The majority of rafting was concentrated in a 2-km radius around the colony within a recently designated seaward SPA extension. Birds showed low individual repeatability in rafting, although there was lower variation within, than among, individuals. Conclusion Our results show that rafting is important for breeding gannets on Grassholm, and a recently designated at-sea SPA encapsulates the core distribution of rafting. Rafting did not appear to be correlated with foraging behaviour. Given the dearth of literature on rafting and the wealth of GPS tracking data for seabirds, we suggest that similar research be conducted elsewhere to further elucidate the ecological and applied significance of this behaviour.

Marine predator populations are crucial to the structure and functioning of ecosystems. Like many predator taxa, pinnipeds face an increasingly complex array of natural and anthropogenic threats. Understanding the relationship between at-sea processes and trends in abundance at land-based monitoring sites requires robust estimates of at-sea distribution, often on multi-region scales. Such an understanding is critical for effective conservation management, but estimates are often limited in spatial extent by spatial coverage of animal-borne tracking data. Grey (Halichoerus grypus) and harbour seals (Phoca vitulina) are sympatric predators in North Atlantic shelf seas. The United Kingdom (UK) and Ireland represents an important population centre for both species, and Special Areas of Conservation (SACs) are designated for their monitoring and protection. Here we use an extensive high-resolution GPS tracking dataset, unprecedented in both size (114 grey and 239 harbour seals) and spatial coverage, to model habitat preference and generate at-sea distribution estimates for the entire UK and Ireland populations of both species. We found regional differences in environmental drivers of distribution for both species which likely relate to regional variation in diet and population trends. Moreover, we provide SAC-specific estimates of at-sea distribution for use in marine spatial planning, demonstrating that hotspots of at-sea density in UK and Ireland-wide maps cannot always be apportioned to the nearest SAC. We show that for grey seals, colonial capital breeders, there is a mismatch between SACs (where impacts are likely to be detected) and areas where impacts are most likely to occur (at sea). We highlight an urgent need for further research to elucidate the links between at-sea distribution during the foraging season and population trends observed in SACs. More generally, we highlight that the potential for such a disconnect needs to be considered when designating and managing protected sites, particularly for species that aggregate to breed and exhibit partial migration (e.g. grey seals), or spatial variation in migration strategies. We demonstrate the use of strategic tracking efforts to predict distribution across multiple regions, but caution that such efforts should be mindful of the potential for differences in species-environment relationships despite similar accessible habitats.

Marine megafauna species are affected by a wide range of anthropogenic threats. To evaluate the risk of such threats, species’ vulnerability to each threat must first be determined. We build on the existing threats classification scheme and ranking system of the International Union for Conservation of Nature (IUCN) Red List of Threatened Species by assessing the vulnerability of 256 marine megafauna species to 23 at‐sea threats. The threats we considered included individual fishing gear types, climate‐change‐related subthreats not previously assessed, and threats associated with coastal impacts and maritime disturbances. Our ratings resulted in 70 species having high vulnerability ( v > 0.778 out of 1) to at least 1 threat, primarily drifting longlines, temperature extremes, or fixed gear. These 3 threats were also considered to have the most severe effects (i.e., steepest population declines). Overall, temperature extremes and plastics and other solid waste were rated as affecting the largest proportion of populations. Penguins, pinnipeds, and polar bears had the highest vulnerability to temperature extremes. Bony fishes had the highest vulnerability to drifting longlines and plastics and other solid waste; pelagic cetaceans to 4 maritime disturbance threats; elasmobranchs to 5 fishing threats; and flying birds to drifting longlines and 2 maritime disturbance threats. Sirenians and turtles had the highest vulnerability to at least one threat from all 4 categories. Despite not necessarily having severe effects for most taxonomic groups, temperature extremes were rated among the top threats for all taxa except bony fishes. The vulnerability scores we provide are an important first step in estimating the risk of threats to marine megafauna. Importantly, they help differentiate scope from severity, which is key to identifying threats that should be prioritized for mitigation.