Tracking of marine predators to protect Southern Ocean ecosystems.

Southern Ocean ecosystems are under pressure from resource exploitation and climate change1,2. Mitigation requires the identification and protection of Areas of Ecological Significance (AESs), which have so far not been determined at the ocean-basin scale. Here, using assemblage-level tracking of marine predators, we identify AESs for this globally important region and assess current threats and protection levels. Integration of more than 4,000 tracks from 17 bird and mammal species reveals AESs around sub-Antarctic islands in the Atlantic and Indian Oceans and over the Antarctic continental shelf. Fishing pressure is disproportionately concentrated inside AESs, and climate change over the next century is predicted to impose pressure on these areas, particularly around the Antarctic continent. At present, 7.1% of the ocean south of 40°S is under formal protection, including 29% of the total AESs. The establishment and regular revision of networks of protection that encompass AESs are needed to provide long-term mitigation of growing pressures on Southern Ocean ecosystems.

Long-term stability in the circumpolar foraging range of a Southern Ocean predator between the eras of whaling and rapid climate change.

Assessing environmental changes in Southern Ocean ecosystems is difficult due to its remoteness and data sparsity. Monitoring marine predators that respond rapidly to environmental variation may enable us to track anthropogenic effects on ecosystems. Yet, many long-term datasets of marine predators are incomplete because they are spatially constrained and/or track ecosystems already modified by industrial fishing and whaling in the latter half of the 20th century. Here, we assess the contemporary offshore distribution of a wide-ranging marine predator, the southern right whale (SRW, Eubalaena australis), that forages on copepods and krill from ~30°S to the Antarctic ice edge (>60°S). We analyzed carbon and nitrogen isotope values of 1,002 skin samples from six genetically distinct SRW populations using a customized assignment approach that accounts for temporal and spatial variation in the Southern Ocean phytoplankton isoscape. Over the past three decades, SRWs increased their use of mid-latitude foraging grounds in the south Atlantic and southwest (SW) Indian oceans in the late austral summer and autumn and slightly increased their use of high-latitude (>60°S) foraging grounds in the SW Pacific, coincident with observed changes in prey distribution and abundance on a circumpolar scale. Comparing foraging assignments with whaling records since the 18th century showed remarkable stability in use of mid-latitude foraging areas. We attribute this consistency across four centuries to the physical stability of ocean fronts and resulting productivity in mid-latitude ecosystems of the Southern Ocean compared with polar regions that may be more influenced by recent climate change.

Potential for redistribution of post-moult habitat for Eudyptes penguins in the Southern Ocean under future climate conditions.

Anthropogenic climate change is resulting in spatial redistributions of many species. We assessed the potential effects of climate change on an abundant and widely distributed group of diving birds, Eudyptes penguins, which are the main avian consumers in the Southern Ocean in terms of biomass consumption. Despite their abundance, several of these species have undergone population declines over the past century, potentially due to changing oceanography and prey availability over the important winter months. We used light-based geolocation tracking data for 485 individuals deployed between 2006 and 2020 across 10 of the major breeding locations for five taxa of Eudyptes penguins. We used boosted regression tree modelling to quantify post-moult habitat preference for southern rockhopper (E. chrysocome), eastern rockhopper (E. filholi), northern rockhopper (E. moseleyi) and macaroni/royal (E. chrysolophus and E. schlegeli) penguins. We then modelled their redistribution under two climate change scenarios, representative concentration pathways RCP4.5 and RCP8.5 (for the end of the century, 2071-2100). As climate forcings differ regionally, we quantified redistribution in the Atlantic, Central Indian, East Indian, West Pacific and East Pacific regions. We found sea surface temperature and sea surface height to be the most important predictors of current habitat for these penguins; physical features that are changing rapidly in the Southern Ocean. Our results indicated that the less severe RCP4.5 would lead to less habitat loss than the more severe RCP8.5. The five taxa of penguin may experience a general poleward redistribution of their preferred habitat, but with contrasting effects in the (i) change in total area of preferred habitat under climate change (ii) according to geographic region and (iii) the species (macaroni/royal vs. rockhopper populations). Our results provide further understanding on the regional impacts and vulnerability of species to climate change.

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.

Elephant seal foraging success is enhanced in Antarctic coastal polynyas.

Antarctic polynyas are persistent open water areas which enable early and large seasonal phytoplankton blooms. This high primary productivity, boosted by iron supply from coastal glaciers, attracts organisms from all trophic levels to form a rich and diverse community. How the ecological benefit of polynya productivity is translated to the highest trophic levels remains poorly resolved. We studied 119 southern elephant seals feeding over the Antarctic shelf and demonstrated that: (i) 96% of seals foraging here used polynyas, with individuals spending on average 62% of their time there; (ii) the seals exhibited more area-restricted search behaviour when in polynyas; and (iii) these seals gained more energy (indicated by increased buoyancy from greater fat stores) when inside polynyas. This higher-quality foraging existed even when ice was not present in the study area, indicating that these are important and predictable foraging grounds year-round. Despite these energetic advantages from using polynyas, not all the seals used them extensively. Factors other than food supply may influence an individual's choice in their use of feeding grounds, such as exposure to predation or the probability of being able to return to distant sub-Antarctic breeding sites.

Predicting the distribution of foraging seabirds during a period of heightened environmental variability.

Quantifying the links between the marine environment, prey occurrence, and predator distribution is the first step towards identifying areas of biological importance for marine spatial planning. Events such as marine heatwaves result in an anomalous change in the physical environment, which can lead to shifts in the structure, biomass, and distribution of lower trophic levels. As central-place foragers, seabirds are vulnerable to changes in their foraging grounds during the breeding season. We first quantified spatiotemporal variability in the occurrence and biomass of prey in response to an abrupt change in oceanography as a result of a marine heatwave event. Secondly, using multivariate techniques and machine learning, we investigated if differences in the foraging technique and prey of seabirds resulted in varying responses to changes in prey occurrence and the environment over a 2.5-yr period. We found that the main variables correlated with seabird distribution were also important in structuring the occurrence and biomass of prey; sea-surface temperature (SST), current speed, mixed-layer depth, and bathymetry. Both zooplankton biomass and the occurrence of fish schools exhibited negative relationships with temperature, and temperature was subsequently an important variable in determining seabird distribution. We were able to establish correlations between the distribution of prey and the spatiotemporal distribution of albatross, little penguins and common-diving petrels. We were unable to find a correlation between the distribution of prey and that of short-tailed shearwaters and fairy prions. For high-use coastal areas, the delineation of important foraging regions is essential to balance human use of an area with the needs of marine predators, particularly seabirds.

Decadal changes in blood δ13C values, at-sea distribution, and weaning mass of southern elephant seals from Kerguelen Islands.

Changes in the foraging environment and at-sea distribution of southern elephant seals from Kerguelen Islands were investigated over a decade (2004-2018) using tracking, weaning mass, and blood δ13C values. Females showed either a sub-Antarctic or an Antarctic foraging strategy, and no significant shift in their at-sea distribution was detected between 2004 and 2017. The proportion of females foraging in sub-Antarctic versus Antarctic habitats did not change over the 2006-2018 period. Pup weaning mass varied according to the foraging habitat of their mothers. The weaning mass of sub-Antarctic foraging mothers' pups decreased by 11.7 kg over the study period, but they were on average 5.8 kg heavier than pups from Antarctic foraging mothers. Pup blood δ13C values decreased by 1.1‰ over the study period regardless of their sex and the presumed foraging habitat of their mothers. Together, these results suggest an ecological change is occurring within the Indian sector of the Southern Ocean with possible consequences on the foraging performance of southern elephant seals. We hypothesize that this shift in δ13C is related to a change in primary production and/or in the composition of phytoplankton communities, but this requires further multidisciplinary investigations.

Preferred, small-scale foraging areas of two Southern Ocean fur seal species are not determined by habitat characteristics.

BACKGROUND: To understand and predict the distribution of foragers, it is crucial to identify the factors that affect individual movement decisions at different scales. Individuals are expected to adjust their foraging movements to the hierarchical spatial distribution of resources. At a small local scale, spatial segregation in foraging habitat happens among individuals of closely situated colonies. If foraging segregation is due to differences in distribution of resources, we would expect segregated foraging areas to have divergent habitat characteristics. RESULTS: We investigated how environmental characteristics of preferred foraging areas differ between two closely situated Subantarctic fur seal (Arctocephalus tropicalis) colonies and a single Antarctic fur seal (A. gazella) colony that forage in different pelagic areas even though they are located well within each other's foraging range. We further investigated the influence of the seasonal cycle on those environmental factors. This study used tracking data from 121 adult female Subantarctic and Antarctic fur seals, collected during summer and winter (2009-2015), from three different colonies. Boosted Regression Tree species distribution models were used to determine key environmental variables associated with areas of fur seal restricted search behaviour. There were no differences in the relative influence of key environmental variables between colonies and seasons. The variables with the most influence for each colony and season were latitude, longitude and magnitude of sea-currents. The influence of latitude and longitude is a by-product of the species' distinct foraging areas, despite the close proximity (< 25 km) of the colonies. The predicted potential foraging areas for each colony changed from summer to winter, reflecting the seasonal cycle of the Southern Ocean. The model predicted that the potential foraging areas of females from the three colonies should overlap, and the fact they do not in reality indicates that factors other than environmental are influencing the location of each colony's foraging area. CONCLUSIONS: The results indicated that small scale spatial segregation of foraging habitats is not driven by bottom-up processes. It is therefore important to also consider other potential drivers, e.g. competition, information transfer, and memory, to understand animal foraging decisions and movements.

Optimizing lifetime reproductive output: Intermittent breeding as a tactic for females in a long-lived, multiparous mammal.

In iteroparous species, intermittent breeding is an important life-history tactic that can greatly affect animal population growth and viability. Despite its importance, few studies have quantified the consequences of breeding pauses on lifetime reproductive output, principally because calculating lifetime reproductive output requires knowledge of each individual's entire reproductive history. This information is extremely difficult to obtain in wild populations. We applied novel statistical approaches that account for uncertainty in state assessment and individual heterogeneity to an 18-year capture-recapture dataset of 6,631 female southern elephant seals from Macquarie Island. We estimated survival and breeding probabilities, and investigated the consequences of intermittent breeding on lifetime reproductive output. We found consistent differences in females' demographic performance between two heterogeneity classes. In particular, breeding imbued a high cost on survival in the females from the heterogeneity class 2, assumed to be females of lower quality. Individual quality also appeared to play a major role in a female's decision to skip reproduction with females of poorer quality more likely to skip breeding events than females of higher quality. Skipping some breeding events allowed females from both heterogeneity classes to increase lifetime reproductive output over females that bred annually. However, females of lower quality produced less offspring over their lifetime. Intermittent breeding seems to be used by female southern elephant seals as a tactic to offset reproductive costs on survival and enhance lifetime reproductive output but remains unavoidable and driven by individual-specific constraints in some other females.

Decadal changes in habitat characteristics influence population trajectories of southern elephant seals.

Understanding divergent biological responses to climate change is important for predicting ecosystem level consequences. We use species habitat models to predict the winter foraging habitats of female southern elephant seals and investigate how changes in environmental variables within these habitats may be related to observed decreases in the Macquarie Island population. There were three main groups of seals that specialized in different ocean realms (the sub-Antarctic, the Ross Sea and the Victoria Land Coast). The physical and climate attributes (e.g. wind strength, sea surface height, ocean current strength) varied amongst the realms and also displayed different temporal trends over the last two to four decades. Most notably, sea ice extent increased on average in the Victoria Land realm while it decreased overall in the Ross Sea realm. Using a species distribution model relating mean residence times (time spent in each 50 × 50 km grid cell) to 9 climate and physical co-variates, we developed spatial predictions of residence time to identify the core regions used by the seals across the Southern Ocean from 120°E to 120°W. Population size at Macquarie Island was negatively correlated with ice concentration within the core habitat of seals using the Victoria Land Coast and the Ross Sea. Sea ice extent and concentration is predicted to continue to change in the Southern Ocean, having unknown consequences for the biota of the region. The proportion of Macquarie Island females (40%) utilizing the relatively stable sub-Antarctic region, may buffer this population against longer-term regional changes in habitat quality, but the Macquarie Island population has persistently decreased (-1.45% per annum) over seven decades indicating that environmental changes in the Antarctic are acting on the remaining 60% of the population to impose a long-term population decline in a top Southern Ocean predator.

Seal mothers expend more on offspring under favourable conditions and less when resources are limited.

In mammals, maternal expenditure on offspring is a complex mix of several factors including the species' mating system, offspring sex and the condition and age of the mother. While theory suggests that in polygynous species mothers should wean larger male offspring than females when resources and maternal conditions allow, the evidence for this remains equivocal. Southern elephant seals are highly dimorphic, polygynous capital breeders existing in an environment with highly variable resources and should therefore provide clear evidence to support the theoretical expectations of differential maternal expenditure in male and female pups. We quantified maternal size (mass and length) and pup size at birth and weaning for 342 elephant seal mothers at Macquarie Island. The study was conducted over 11 years of contrasting sea-ice and Southern Annular Mode values, both indices of maternal prey resources. Overall, large females weaned male pups that weighed 17 kg (15·5%) more than female pups. Maternal condition varied by as much as 59 kg among years, and was positively related to Southern Annular Mode, and negatively to maximum sea-ice extent. Smaller mothers weaned relatively larger male pups under favourable conditions, this effect was less apparent for larger mothers. We developed a simple model linking environmental variation to maternal masses post-partum, followed by maternal masses post-partum to weaning masses and then weaning masses to pup survival and demonstrated that environmental conditions affected predicted survival so that the pups of small mothers had an estimated 7% increase in first year survival in 'good' vs. 'bad' years compared to 1% for female pups of large mothers. Co-occurrence of environmental quality and conservative reproductive tactics suggests that mothers retain substantial plasticity in maternal care, enhancing their lifetime reproductive success by adjusting reproductive expenditure relative to both prevailing environmental conditions and their own capabilities.

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.

Taking animal tracking to new depths: synthesizing horizontal--vertical movement relationships for four marine predators.

In animal ecology, a question of key interest for aquatic species is how changes in movement behavior are related in the horizontal and vertical dimensions when individuals forage. Alternative theoretical models and inconsistent empirical findings mean that this question remains unresolved. Here we tested expectations by incorporating the vertical dimension (dive information) when predicting switching between movement states ("resident" or "directed") within a state-space model. We integrated telemetry-based tracking and diving data available for four seal species (southern elephant, Weddell, antarctic fur, and crabeater) in East Antarctica. Where possible, we included dive variables derived from the relationships between (1) dive duration and depth (as a measure of effort), and (2) dive duration and the postdive surface interval (as a physiological measure of cost). Our results varied within and across species, but there was a general tendency for the probability of switching into "resident" state to be positively associated with shorter dive durations (for a given depth) and longer postdive surface intervals (for a given dive duration). Our results add to a growing body of literature suggesting that simplistic interpretations of optimal foraging theory based only on horizontal movements do not directly translate into the vertical dimension in dynamic marine environments. Analyses that incorporate at least two dimensions can test more sophisticated models of foraging behavior.

Bottom-up regulation of a pole-ward migratory predator population.

As the effects of regional climate change are most pronounced at polar latitudes, we might expect polar-ward migratory populations to respond as habitat suitability changes. The southern elephant seal (Mirounga leonina L.) is a pole-ward migratory species whose populations have mostly stabilized or increased in the past decade, the one exception being the Macquarie Island population which has decreased continuously over the past 50 years. To explore probable causes of this anomalous trend, we counted breeding female seals annually between 1988 and 2011 in order to relate annual rates of population change (r) to foraging habitat changes that have known connections with atmospheric variability. We found r (i) varied annually from -0.016 to 0.021 over the study period, (ii) was most effected by anomalous atmospheric variability after a 3 year time lag was introduced (R = 0.51) and (iii) was associated with sea-ice duration (SID) within the seals' foraging range at the same temporal lag. Negative r years may be extrapolated to explain, at least partially, the overall trend in seal abundance at Macquarie Island; specifically, increasing SID within the seals foraging range has a negative influence on their abundance at the island. Evidence is accruing that suggests southern elephant seal populations may respond positively to a reduced sea-ice field.

Climate change and Southern Ocean ecosystems I: how changes in physical habitats directly affect marine biota.

Antarctic and Southern Ocean (ASO) marine ecosystems have been changing for at least the last 30 years, including in response to increasing ocean temperatures and changes in the extent and seasonality of sea ice; the magnitude and direction of these changes differ between regions around Antarctica that could see populations of the same species changing differently in different regions. This article reviews current and expected changes in ASO physical habitats in response to climate change. It then reviews how these changes may impact the autecology of marine biota of this polar region: microbes, zooplankton, salps, Antarctic krill, fish, cephalopods, marine mammals, seabirds, and benthos. The general prognosis for ASO marine habitats is for an overall warming and freshening, strengthening of westerly winds, with a potential pole-ward movement of those winds and the frontal systems, and an increase in ocean eddy activity. Many habitat parameters will have regionally specific changes, particularly relating to sea ice characteristics and seasonal dynamics. Lower trophic levels are expected to move south as the ocean conditions in which they are currently found move pole-ward. For Antarctic krill and finfish, the latitudinal breadth of their range will depend on their tolerance of warming oceans and changes to productivity. Ocean acidification is a concern not only for calcifying organisms but also for crustaceans such as Antarctic krill; it is also likely to be the most important change in benthic habitats over the coming century. For marine mammals and birds, the expected changes primarily relate to their flexibility in moving to alternative locations for food and the energetic cost of longer or more complex foraging trips for those that are bound to breeding colonies. Few species are sufficiently well studied to make comprehensive species-specific vulnerability assessments possible. Priorities for future work are discussed.

Age-specific cost of first reproduction in female southern elephant seals.

When to commence breeding is a crucial life-history decision that may be the most important determinant of an individual's lifetime reproductive output and can have major consequences on population dynamics. The age at which individuals first reproduce is an important factor influencing the intensity of potential costs (e.g. reduced survival) involved in the first breeding event. However, quantifying age-related variation in the cost of first reproduction in wild animals remains challenging because of the difficulty in reliably recording the first breeding event. Here, using a multi-event capture-recapture model that accounts for both imperfect detection and uncertainty in the breeding status on an 18-year dataset involving 6637 individuals, we estimated age and state-specific survival of female elephant seals (Mirounga leonina) in the declining Macquarie Island population. We detected a clear cost of first reproduction on survival. This cost was higher for both younger first-time breeders and older first-time breeders compared with females recruiting at age four, the overall mean age at first reproduction. Neither earlier primiparity nor delaying primiparity appear to confer any evolutionary advantage, rather the optimal strategy seems to be to start breeding at a single age, 4 years.

Scaling laws of marine predator search behaviour.

Many free-ranging predators have to make foraging decisions with little, if any, knowledge of present resource distribution and availability. The optimal search strategy they should use to maximize encounter rates with prey in heterogeneous natural environments remains a largely unresolved issue in ecology. Lévy walks are specialized random walks giving rise to fractal movement trajectories that may represent an optimal solution for searching complex landscapes. However, the adaptive significance of this putative strategy in response to natural prey distributions remains untested. Here we analyse over a million movement displacements recorded from animal-attached electronic tags to show that diverse marine predators-sharks, bony fishes, sea turtles and penguins-exhibit Lévy-walk-like behaviour close to a theoretical optimum. Prey density distributions also display Lévy-like fractal patterns, suggesting response movements by predators to prey distributions. Simulations show that predators have higher encounter rates when adopting Lévy-type foraging in natural-like prey fields compared with purely random landscapes. This is consistent with the hypothesis that observed search patterns are adapted to observed statistical patterns of the landscape. This may explain why Lévy-like behaviour seems to be widespread among diverse organisms, from microbes to humans, as a 'rule' that evolved in response to patchy resource distributions.

Correction: Incorporating mesopelagic fish into the evaluation of marine protected areas under climate change scenarios.

[This corrects the article DOI: 10.1007/s42995-023-00188-9.].

Incorporating mesopelagic fish into the evaluation of conservation areas for marine living resources under climate change scenarios.

Mesopelagic fish (meso-fish) are central species within the Southern Ocean (SO). However, their ecosystem role and adaptive capacity to climate change are rarely integrated into protected areas assessments. This is a pity given their importance as crucial prey and predators in food webs, coupled with the impacts of climate change. Here, we estimate the habitat distribution of nine meso-fish using an ensemble model approach (MAXENT, random forest, and boosted regression tree). Four climate model simulations were used to project their distribution under two representative concentration pathways (RCP4.5 and RCP8.5) for short-term (2006-2055) and long-term (2050-2099) periods. In addition, we assess the ecological representativeness of protected areas under climate change scenarios using meso-fish as indicator species. Our models show that all species shift poleward in the future. Lanternfishes (family Myctophidae) are predicted to migrate poleward more than other families (Paralepididae, Nototheniidae, Bathylagidae, and Gonostomatidae). In comparison, lanternfishes were projected to increase habitat area in the eastern SO but lose area in the western SO; the opposite was projected for species in other families. Important areas (IAs) of meso-fish are mainly distributed near the Antarctic Peninsula and East Antarctica. Negotiated protected area cover 23% of IAs at present and 38% of IAs in the future (RCP8.5, long-term future). Many IAs of meso-fish still need to be included in protected areas, such as the Prydz Bay and the seas around the Antarctic Peninsula. Our results provide a framework for evaluating protected areas incorporating climate change adaptation strategies for protected areas management. Supplementary Information: The online version contains supplementary material available at 10.1007/s42995-023-00188-9.

Integrative modelling of animal movement: incorporating in situ habitat and behavioural information for a migratory marine predator.

A fundamental goal in animal ecology is to quantify how environmental (and other) factors influence individual movement, as this is key to understanding responsiveness of populations to future change. However, quantitative interpretation of individual-based telemetry data is hampered by the complexity of, and error within, these multi-dimensional data. Here, we present an integrative hierarchical Bayesian state-space modelling approach where, for the first time, the mechanistic process model for the movement state of animals directly incorporates both environmental and other behavioural information, and observation and process model parameters are estimated within a single model. When applied to a migratory marine predator, the southern elephant seal (Mirounga leonina), we find the switch from directed to resident movement state was associated with colder water temperatures, relatively short dive bottom time and rapid descent rates. The approach presented here can have widespread utility for quantifying movement-behaviour (diving or other)-environment relationships across species and systems.