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Open ocean predator‐prey interactions are often difficult to interpret because of a lack of information on prey fields at scales relevant to predator behaviour. Hence, there is strong interest in identifying the biological and physical factors influencing the distribution and abundance of prey species, which may be of broad predictive use for conservation planning and evaluating effects of environmental change. This study focuses on a key Southern Ocean prey species, Antarctic krill Euphausia superba, using acoustic observations of individual swarms (aggregations) from a large‐scale survey off East Antarctica. We developed two sets of statistical models describing swarm characteristics, one set using underway survey data for the explanatory variables, and the other using their satellite remotely sensed analogues. While survey data are in situ and contemporaneous with the swarm data, remotely sensed data are all that is available for prediction and inference about prey distribution in other areas or at other times. The fitted models showed that the primary biophysical influences on krill swarm characteristics included daylight (solar elevation/radiation) and proximity to the Antarctic continental slope, but there were also complex relationships with current velocities and gradients. Overall model performance was similar regardless of whether underway or remotely sensed predictors were used. We applied the latter models to generate regional‐scale spatial predictions using a 10‐yr remotely‐sensed time series. This retrospective modelling identified areas off east Antarctica where relatively dense krill swarms were consistently predicted during austral mid‐summers, which may underpin key foraging areas for marine predators. Spatiotemporal predictions along Antarctic predator satellite tracks, from independent studies, illustrate the potential for uptake into further quantitative modelling of predator movements and foraging. The approach is widely applicable to other krill‐dependent ecosystems, and our findings are relevant to similar efforts examining biophysical linkages elsewhere in the Southern Ocean and beyond. [ABSTRACT FROM AUTHOR]

State-space models have recently been shown to effectively model animal movement. In this paper we illustrate how such models can be used to improve our knowledge of animal navigation ability, something which is poorly understood. This work is of great interest when modeling the behavior of animals that are migrating, often over tremendously large distances. We use the term circle of confusion, first proposed by Kendall (1974), to describe the general inability of an animal to know its location precisely. Our modeling strategy enables us to statistically describe the circle of confusion associated with any animal movements where departure and destination points are known. For illustration, we use ARGOS satellite telemetry of leatherback turtles migrating over a distance of approximately 4000 km in the Atlantic Ocean. Robust features of the model enable one to deal with outlying observations, highly characteristic of these types of data. Although specifically designed for data obtained using satellite telemetry, our approach is generalizable to other common kinds of movement data such as archival tag data. [ABSTRACT FROM AUTHOR]

We explore the effect of differences in landscape structure, arisingfrom habitat loss, on the fine-scale movement behaviors of two congeneric damselflies - Calopteryx aequabilis and C. maculata. Both species require streams for breeding and naiad development and both often use forest for foraging. We compare movement behaviors across three types of landscape: forested landscapes, where stream and forest habitat are adjacent; partially forested landscapes, where streams and forest habitat are disjunct, and non-forested landscapes, where little to no forest habitat is available. We employ a reciprocal transplant experiment to determine the extent to which movement along and away from streams is influenced by landscape structure and historical behavior or morphological adaptations. For both species, we show that both the propensity to move away from streams and rates of net displacement differ among landscape types. Both species move away from streams on landscapes with high or moderate levels of forest cover but neithermoves away from streams on landscapes with little or no forest. Furthermore, C. maculata native to predominantly forested landscapes are more likely to move away from streams, regardless of the landscape structure they encounter, than are individuals native to moderately forested or non-forested landscapes. There was no effect of natal landscape on C. aequabilis. Comparisons with microlandscape studies suggestthat there may be some general similarities among the different systems but these are clouded by uncertainty regarding the similarity of the underlying processes responsible for observed behavioral responses to landscape structure. Despite this uncertainty, animal movement behaviors are contingent upon the structure of the broader landscape, regardless of the absolute scale of the landscape. [ABSTRACT FROM AUTHOR]

The purpose of this study was to investigate the effect of changes in landscape pattern on generalist and specialist insects. We did thisby comparing the species richness and abundance of generalist and specialist herbivorous insects in alfalfa (Medicago sativa, L.) fields on 26 agricultural landscapes that differed in spatial structure. Theinsects were from the families Curculionidae (Coleoptera), weevils, and Cicadellidae (Auchennorhyncha), leafhoppers. We hypothesized that: (1) generalist richness and abundance would be highest in landscapes with high diversity (Shannon-Wiener); (2) specialist richness and abundance would be highest in landscapes with (i) high percent cover alfalfa and (ii) low mean inter-patch distance. We tested for these effects after controlling for the patch-level effects of field size, field age, frequency of disturbance and vegetation texture. The important findings of the study are: (1) generalist richness and abundance increased with increasing landscape diversity and (2) isolation (percent cover alfalfa in the landscape and/or mean inter-patch distance) does not affect specialist insects. These results are significant because they indicate that both generalist and specialist insects may move over much larger distances than previously thought. This is one of the first studies to demonstrate a large scale effect of spatial structure on insects across a broad range of landscapes. [ABSTRACT FROM AUTHOR]

For similar species to co-occur in places where resources are limited, they need to adopt strategies that partition resources to reduce competition. Our understanding of the mechanisms behind resource partitioning among sympatric marine predators is evolving, but we lack a clear understanding of how environmental change is impacting these dynamics. We investigated spatial and trophic resource partitioning among three sympatric seabirds with contrasting biological characteristics: greater crested terns Thalasseus bergii (efficient flyer, limited diver, and preference for high quality forage fish), little penguins Eudyptula minor (flightless, efficient diver, and preference for high quality forage fish) and silver gulls Chroicocephalus novaehollandiae (efficient flyer, limited diver and generalist diet). We investigated interannual variability in resource partitioning in relation to environmental variability in a climate change hotspot influenced by the warm and intensifying East Australian Current (EAC). Sampling was conducted from 2012 to 2014 during the austral summer breeding season of seabirds at Montague Island, Australia. Daily seabird movements were monitored using GPS trackers and feather tissues were collected and processed for stable isotope analysis (δ 15 N and δ 13 C). Generalised Linear Mixed Models were used to assess how changes in oceanographic conditions influenced space use for each species. Schoener's D and Bayesian mixing models were used to respectively investigate the levels of yearly inter-specific environmental and trophic niche overlaps. Crested terns and little penguins were less likely to be observed in warm, saline EAC waters and crested terns and silver gulls had smaller foraging areas on days when more than 30% of available habitat was classified as EAC origin. All species preferred areas with low variability in sea surface temperature (<0.5°C). Terns and penguins occupied similar marine trophic levels, with penguins having larger isotopic niche spaces in 2014 when the EAC was more dominant in the study area. Gulls occupied the lowest trophic level, with the widest niche and lowest interannual variability in niche area. As the EAC intensifies along the southeast coast of Australia under climate change, interspecific competition for resources may increase, with the greatest impacts on species like little penguins that have relatively restricted foraging ranges. This study suggests that species-specific biological traits and behavioural plasticity should be accounted for when predicting the effects of climate change on marine species., (© 2025 The Author(s). Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Ecological theory predicts niche partitioning between high-level predators living in sympatry as a mechanism to minimise the selective pressure of competition. Accordingly, male Australian fur seals Arctocephalus pusillus doriferus and New Zealand fur seals A. forsteri that live in sympatry should exhibit partitioning in their broad niches (in habitat and trophic dimensions) in order to coexist. However, at the northern end of their distributions in Australia, both are recolonising their historic range after a long absence due to over-exploitation, and their small population sizes suggest competition should be weak and may allow overlap in niche space. We found some niche overlap, yet clear partitioning in diet trophic level (δ 15 N values from vibrissae), spatial niche space (horizontal and vertical telemetry data) and circadian activity patterns (timing of dives) between males of each species, suggesting competition may remain an active driver of niche partitioning amongst individuals even in small, peripheral populations. Consistent with individual specialisation theory, broad niches of populations were associated with high levels of individual specialisation for both species, despite putative low competition. Specialists in isotopic space were not necessarily specialists in spatial niche space, further emphasising their diverse individual strategies for niche partitioning. Males of each species displayed distinct foraging modes, with Australian fur seals primarily benthic and New Zealand fur seals primarily epipelagic, though unexpectedly high individual specialisation for New Zealand fur seals might suggest marginal populations provide exceptions to the pattern generally observed amongst other fur seals., (© 2024. Crown.)

Marine ecosystems in southeastern Australia are responding rapidly to climate change. We monitored the diet of the Australian fur seal ( Arctocephalus pusillus doriferus ), a key marine predator, over 17 years (1998-2014) to examine temporal changes. Frequency of occurrence (FO) of prey was used as a proxy for ecosystem change. Hard part analysis identified 71 prey taxa, with eight dominant taxa in greater than 70% of samples and predominantly included benthic and small pelagic fish. FO changed over time, e.g. redbait ( Emmelichthys nitidus ) reduced after 2005 when jack mackerel ( Trachurus declivis ) increased, and pilchard ( Sardinops sajax ) increased after 2009. Using generalized additive models, correlations between FO and environmental variables were evident at both the local (e.g. wind, sea surface temperature (SST)) and regional (e.g. El Niño-Southern Oscillation Index (SOI), Southern Annular Mode (SAM)) scales, with redbait and pilchard showing the best model fits (greater than 75% deviance explained). Positive SAM was correlated to FO for both species, and wind and season were important for redbait, while SOI and SST were important for pilchard. Both large-scale and regional processes influenced prey taxa in variable ways. We predict that the diverse and adaptable diet of the Australian fur seal will be advantageous in a rapidly changing ecosystem., (© 2022 The Authors.)

Understanding how marine predators encounter prey across patchy landscapes remains challenging due to difficulties in measuring the three-dimensional structure of pelagic prey fields at scales relevant to animal movement. We measured at-sea behaviour of a central-place forager, the little penguin ( Eudyptula minor ), over 5 years (2015-2019) using GPS and dive loggers. We made contemporaneous measurements of the prey field within the penguins' foraging range via boat-based acoustic surveys. We developed a prey encounter index by comparing estimates of acoustic prey density encountered along actual penguin tracks to those encountered along simulated penguin tracks with the same characteristics as real tracks but that moved randomly through the prey field. In most years, penguin tracks encountered prey better than simulated random movements greater than 99% of the time, and penguin dive depths matched peaks in the vertical distribution of prey. However, when prey was unusually sparse and/or deep, penguins had worse than random prey encounter indices, exhibited dives that mismatched depth of maximum prey density, and females had abnormally low body mass (5.3% lower than average). Reductions in prey encounters owing to decreases in the density or accessibility of prey may ultimately lead to reduced fitness and population declines in central-place foraging marine predators., (© 2022 The Authors.)

Partitioning resources is a key mechanism for avoiding intraspecific competition and maximizing individual energy gain. However, in sexually dimorphic species it is difficult to discern if partitioning is due to competition or the different resource needs of morphologically distinct individuals. In the highly dimorphic southern elephant seal, there are intersexual differences in habitat use; at Iles Kerguelen, males predominantly use shelf waters, while females use deeper oceanic waters. There are equally marked intrasexual differences, with some males using the nearby Kerguelen Plateau, and others using the much more distant Antarctic continental shelf (~2,000 km away). We used this combination of inter and intrasexual behavior to test two hypotheses regarding habitat partitioning in highly dimorphic species. (a) that intersexual differences in habitat use will not appear until the seals diverge in body size and (b) that some habitats have higher rates of energy return than others. In particular, that the Antarctic shelf would provide higher energy returns than the Kerguelen Shelf, to offset the greater cost of travel. We quantified the habitat use of 187 southern elephant seals (102 adult females and 85 subadult males). The seals in the two groups were the same size (~2.4 m) removing the confounding effect of body size. We found that the intersexual differences in habitat use existed before the divergence in body size. Also, we found that the amount of energy gained was the same in all of the major habitats. This suggests that the use of shelf habitats by males is innate, and a trade-off between the need to access the large benthic prey available on shelf waters, against the higher risk of predation there. Intrasexual differences in habitat use are another trade-off; although there are fewer predators on the Antarctic shelf, it is subject to considerable interannual fluctuations in sea-ice extent. In contrast, the Kerguelen Plateau presents more consistent foraging opportunities, but contains higher levels of predation. Habitat partitioning in this highly dimorphic species is therefore the result of complex interplay of life history strategies, environmental conditions and predation pressure., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Knowledge about the movement ecology of endangered species is needed to identify biologically important areas and the spatio-temporal scale of potential human impacts on species. Blue whales (Balaenoptera musculus) are endangered due to twentieth century whaling and currently threatened by human activities. In Australia, they feed in the Great Southern Australian Coastal Upwelling System (GSACUS) during the austral summer. We investigate their movements, occupancy, behaviour, and environmental drivers to inform conservation management. Thirteen whales were satellite tagged, biopsy sampled and photo-identified in 2015. All were genetically confirmed to be of the pygmy subspecies (B. m. brevicauda). In the GSACUS, whales spent most of their time over the continental shelf and likely foraging in association with several seascape variables (sea surface temperature variability, depth, wind speed, sea surface height anomaly, and chlorophyll a). When whales left the region, they migrated west and then north along the Australian coast until they reached West Timor and Indonesia, where their movements indicated breeding or foraging behaviour. These results highlight the importance of the GSACUS as a foraging ground for pygmy blue whales inhabiting the eastern Indian Ocean and indicate the whales' migratory route to proposed breeding grounds off Indonesia. Information about the spatio-temporal scale of potential human impacts can now be used to protect this little-known subspecies of blue whale.

Globally, fisheries bycatch threatens the survival of many whale and dolphin species. Strategies for reducing bycatch can be expensive. Management is inclined to prioritize investment in actions that are inexpensive, but these may not be the most effective. We used an economic tool, return-on-investment, to identify cost-effective measures to reduce cetacean bycatch in the trawl, net, and line fisheries of Australia. We examined 3 management actions: spatial closures, acoustic deterrents, and gear modifications. We compared an approach for which the primary goal was to reduce the cost of bycatch reduction to fisheries with an approach that aims solely to protect whale and dolphin species. Based on cost-effectiveness and at a fine spatial resolution, we identified the management strategies across Australia that most effectively abated dolphin and whale bycatch. Although trawl-net modifications were the cheapest strategy overall, there were many locations where spatial closures were the most cost-effective solution, despite their high costs to fisheries, due to their effectiveness in reducing all fisheries interactions. Our method can be used to delineate strategies to reduce bycatch threats to mobile marine species across diverse fisheries at relevant spatial scales to improve conservation outcomes., (© 2019 Society for Conservation Biology.)

Mesopelagic fish and squid occupy ocean depths extending below the photic zone and their vertical migrations represent a massive pathway moving energy and carbon through the water column. Their spatio-temporal distribution is however, difficult to map across remote regions particularly the vast Southern Ocean. This represents a key gap in understanding biogeochemical processes, marine ecosystem structure, and how changing ocean conditions will affect marine predators, which depend upon mesopelagic prey. We infer mesopelagic prey vertical distribution and relative abundance in the Indian sector of the Southern Ocean (20° to 130°E) with a novel approach using predator-derived indices. Fourteen years of southern elephant seal tracking and dive data, from the open ocean between the Antarctic Polar Front and the southern Antarctic Circumpolar Current front, clearly show that the vertical distribution of mesopelagic prey is influenced by the physical hydrographic processes that structure their habitat. Mesopelagic prey have a more restricted vertical migration and higher relative abundance closer to the surface where Circumpolar Deep Water rises to shallower depths. Combining these observations with a future projection of Southern Ocean conditions we show that changes in the coupling of surface and deep waters will potentially redistribute mesopelagic prey. These changes are small overall, but show important spatial variability: prey will increase in relative abundance to the east of the Kerguelen Plateau but decrease to the west. The consequences for deep-diving specialists such as elephant seals and whales over this time scale will likely be minor, but the changes in mesoscale vertical energy flow have implications for predators that forage within the mesopelagic zone as well as the broader pelagic ecosystem.

Foraging site fidelity allows animals to increase their efficiency by returning to profitable feeding areas. However, the mechanisms underpinning why animals 'stay' or 'switch' sites have rarely been investigated. Here, we explore how habitat quality and prior prey capture experience influence short-term site fidelity by the little penguin ( Eudyptula minor ). Using 88 consecutive foraging trips by 20 brooding penguins, we found that site fidelity was higher after foraging trips where environmental conditions were favourable, and after trips where prey capture success was high. When penguins exhibited lower site fidelity, the number of prey captures relative to the previous trip increased, suggesting that switches in foraging location were an adaptive strategy in response to low prey capture rates. Penguins foraged closer to where other penguins foraged on the same day than they did to the location of their own previous foraging site, and caught more prey when they foraged close together. This suggests that penguins aggregated flexibly when prey was abundant and accessible. Our results illustrate how foraging predators can integrate information about prior experience with contemporary information such as social cues. This gives insight into how animals combine information adaptively to exploit changing prey distribution in a dynamic environment., (© 2018 The Author(s).)

Animal movement research relies on biotelemetry, and telemetry-based locations are increasingly augmented with ancillary information. This presents an underutilized opportunity to enhance movement process models. Given tags designed to record specific behaviors, efforts are increasing to update movement models beyond reliance solely upon horizontal movement information to improve inference of space use and activity budgets. We present two state-space models adapted to incorporate ancillary data to inform three discrete movement states: directed, resident, and an activity state. These were developed for two case studies: (1) a "haulout" model for Weddell seals, and (2) an "activity" model for Antarctic fur seals which intersperse periods of diving activity and inactivity. The methodology is easily implementable with any ancillary data that can be expressed as a proportion (or binary) indicator. A comparison of the models augmented with ancillary information and unaugmented models confirmed that many behavioral states appeared mischaracterized in the latter. Important changes in subsequent activity budgets occurred. Haulout accounted for 0.17 of the overall Weddell seal time budget, with the estimated proportion of time spent in a resident state reduced from a posterior median of 0.69 (0.65-0.73; 95% HPDI) to 0.54 (0.50-0.58 HPDI). The drop was more dramatic in the Antarctic fur seal case, from 0.57 (0.52-0.63 HPDI) to 0.22 (0.20-0.25 HPDI), with 0.35 (0.31-0.39 HPDI) of time spent in the inactive (nondiving) state. These findings reinforce previously raised contentions about the drawbacks of behavioral states inferred solely from horizontal movements. Our findings have implications for assessing habitat requirements; estimating energetics and consumption; and management efforts such as mitigating fisheries interactions. Combining multiple sources of information within integrated frameworks should improve inference of relationships between movement decisions and fitness, the interplay between resource and habitat dependencies, and their changes at the population and landscape level.

It is a golden age for animal movement studies and so an opportune time to assess priorities for future work. We assembled 40 experts to identify key questions in this field, focussing on marine megafauna, which include a broad range of birds, mammals, reptiles, and fish. Research on these taxa has both underpinned many of the recent technical developments and led to fundamental discoveries in the field. We show that the questions have broad applicability to other taxa, including terrestrial animals, flying insects, and swimming invertebrates, and, as such, this exercise provides a useful roadmap for targeted deployments and data syntheses that should advance the field of movement ecology., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

The world's oceans are undergoing rapid, regionally specific warming. Strengthening western boundary currents play a role in this phenomenon, with sea surface temperatures (SST) in their paths rising faster than the global average. To understand how dynamic oceanography influences food availability in these ocean warming "hotspots", we use a novel prey capture signature derived from accelerometry to understand how the warm East Australian Current shapes foraging success by a meso-predator, the little penguin. This seabird feeds on low trophic level species that are sensitive to environmental change. We found that in 2012, prey capture success by penguins was high when SST was low relative to the long-term mean. In 2013 prey capture success was low, coincident with an unusually strong penetration of warm water. Overall there was an optimal temperature range for prey capture around 19-21 °C, with lower success at both lower and higher temperatures, mirroring published relationships between commercial sardine catch and SST. Spatially, higher SSTs corresponded to a lower probability of penguins using an area, and lower prey capture success. These links between high SST and reduced prey capture success by penguins suggest negative implications for future resource availability in a system dominated by a strengthening western boundary current.

State-space models provide a powerful way to scale up inference of movement behaviours from individuals to populations when the inference is made across multiple individuals. Here, I show how a joint estimation approach that assumes individuals share identical movement parameters can lead to improved inference of behavioural states associated with different movement processes. I use simulated movement paths with known behavioural states to compare estimation error between nonhierarchical and joint estimation formulations of an otherwise identical state-space model. Behavioural state estimation error was strongly affected by the degree of similarity between movement patterns characterising the behavioural states, with less error when movements were strongly dissimilar between states. The joint estimation model improved behavioural state estimation relative to the nonhierarchical model for simulated data with heavy-tailed Argos location errors. When applied to Argos telemetry datasets from 10 Weddell seals, the nonhierarchical model estimated highly uncertain behavioural state switching probabilities for most individuals whereas the joint estimation model yielded substantially less uncertainty. The joint estimation model better resolved the behavioural state sequences across all seals. Hierarchical or joint estimation models should be the preferred choice for estimating behavioural states from animal movement data, especially when location data are error-prone.

Strategies employed by wide-ranging foraging animals involve consideration of habitat quality and predictability and should maximise net energy gain. Fidelity to foraging sites is common in areas of high resource availability or where predictable changes in resource availability occur. However, if resource availability is heterogeneous or unpredictable, as it often is in marine environments, then habitat familiarity may also present ecological benefits to individuals. We examined the winter foraging distribution of female Antarctic fur seals, Arctocephalus gazelle, over four years to assess the degree of foraging site fidelity at two scales; within and between years. On average, between-year fidelity was strong, with most individuals utilising more than half of their annual foraging home range over multiple years. However, fidelity was a bimodal strategy among individuals, with five out of eight animals recording between-year overlap values of greater than 50%, while three animals recorded values of less than 5%. High long-term variance in sea surface temperature, a potential proxy for elevated long-term productivity and prey availability, typified areas of overlap. Within-year foraging site fidelity was weak, indicating that successive trips over the winter target different geographic areas. We suggest that over a season, changes in prey availability are predictable enough for individuals to shift foraging area in response, with limited associated energetic costs. Conversely, over multiple years, the availability of prey resources is less spatially and temporally predictable, increasing the potential costs of shifting foraging area and favouring long-term site fidelity. In a dynamic and patchy environment, multi-year foraging site fidelity may confer a long-term energetic advantage to the individual. Such behaviours that operate at the individual level have evolutionary and ecological implications and are potential drivers of niche specialization and modifiers of intra-specific competition.

Determining where, when and how much animals eat is fundamental to understanding their ecology. We developed a technique to identify a prey capture signature for little penguins from accelerometry, in order to quantify food intake remotely. We categorised behaviour of captive penguins from HD video and matched this to time-series data from back-mounted accelerometers. We then trained a support vector machine (SVM) to classify the penguins' behaviour at 0.3 s intervals as either 'prey handling' or 'swimming'. We applied this model to accelerometer data collected from foraging wild penguins to identify prey capture events. We compared prey capture and non-prey capture dives to test the model predictions against foraging theory. The SVM had an accuracy of 84.95±0.26% (mean ± s.e.) and a false positive rate of 9.82±0.24% when tested on unseen captive data. For wild data, we defined three independent, consecutive prey handling observations as representing true prey capture, with a false positive rate of 0.09%. Dives with prey captures had longer duration and bottom times, were deeper, had faster ascent rates, and had more 'wiggles' and 'dashes' (proxies for prey encounter used in other studies). The mean (±s.e.) number of prey captures per foraging trip was 446.6±66.28. By recording the behaviour of captive animals on HD video and using a supervised machine learning approach, we show that accelerometry signatures can classify the behaviour of wild animals at unprecedentedly fine scales., (© 2014. Published by The Company of Biologists Ltd.)

Argos recently implemented a new algorithm to calculate locations of satellite-tracked animals that uses a Kalman filter (KF). The KF algorithm is reported to increase the number and accuracy of estimated positions over the traditional Least Squares (LS) algorithm, with potential advantages to the application of state-space methods to model animal movement data. We tested the performance of two Bayesian state-space models (SSMs) fitted to satellite tracking data processed with KF algorithm. Tracks from 7 harbour seals (Phoca vitulina) tagged with ARGOS satellite transmitters equipped with Fastloc GPS loggers were used to calculate the error of locations estimated from SSMs fitted to KF and LS data, by comparing those to "true" GPS locations. Data on 6 fin whales (Balaenoptera physalus) were used to investigate consistency in movement parameters, location and behavioural states estimated by switching state-space models (SSSM) fitted to data derived from KF and LS methods. The model fit to KF locations improved the accuracy of seal trips by 27% over the LS model. 82% of locations predicted from the KF model and 73% of locations from the LS model were <5 km from the corresponding interpolated GPS position. Uncertainty in KF model estimates (5.6 ± 5.6 km) was nearly half that of LS estimates (11.6 ± 8.4 km). Accuracy of KF and LS modelled locations was sensitive to precision but not to observation frequency or temporal resolution of raw Argos data. On average, 88% of whale locations estimated by KF models fell within the 95% probability ellipse of paired locations from LS models. Precision of KF locations for whales was generally higher. Whales' behavioural mode inferred by KF models matched the classification from LS models in 94% of the cases. State-space models fit to KF data can improve spatial accuracy of location estimates over LS models and produce equally reliable behavioural estimates.

The need to balance energy reserves during migration is a critical factor for most long-distance migrants and an important determinant of migratory strategies in birds, insects and land mammals. Large baleen whales migrate annually between foraging and breeding sites, crossing vast ocean areas where food is seldom abundant. How whales respond to the demands and constraints of such long migrations remains unknown. We applied a behaviour discriminating hierarchical state-space model to the satellite tracking data of 12 fin whales and 3 blue whales tagged off the Azores, to investigate their movements, behaviour (transiting and area-restricted search, ARS) and daily activity cycles during the spring migration. Fin and blue whales remained at middle latitudes for prolonged periods, spending most of their time there in ARS behaviour. While near the Azores, fin whale ARS behaviour occurred within a restricted area, with a high degree of overlap among whales. There were noticeable behavioural differences along the migratory pathway of fin whales tracked to higher latitudes: ARS occurred only in the Azores and north of 56°N, whereas in between these areas whales travelled at higher overall speeds while maintaining a nearly direct trajectory. This suggests fin whales may alternate periods of active migration with periods of extended use of specific habitats along the migratory route. ARS behaviour in blue whales occurred over a much wider area as whales slowly progressed northwards. The tracks of these whales terminated still at middle latitudes, before any behavioural switch was detected. Fin whales exhibited behavioural-specific diel rhythms in swimming speed but these varied significantly between geographic areas, possibly due to differences in the day-night cycle across areas. Finally, we show a link between fin whales seen in the Azores and those summering in eastern Greenland-western Iceland along a migratory corridor located in central Atlantic waters.