Area-restricted Search Research Articles

Staying Close to Home: Horizontal Movements of Satellite-Tracked Reef Manta Rays Mobula alfredi (Krefft, 1868) in the World’s Largest Manta Sanctuary

Indonesia is home to significant populations of globally vulnerable reef manta rays (Mobula alfredi) in at least four key regions: Berau, Nusa Penida, Komodo, and Raja Ampat. Despite detailed population studies in each of these regions, little is known about their horizontal movement patterns. Our study used satellite telemetry to investigate reef manta rays’ habitat use and home ranges. A total of 33 manta rays were tagged with SPLASH10F-321A satellite tags across the four regions: Berau (n = 5), Nusa Penida (n = 8), Komodo (n = 6), and Raja Ampat (n = 14), yielding usable data from 25 tags. The rays were tracked for 7 to 118 days (mean ± SD = 50 ± 30) from July 2014 to July 2022. The results showed localized movements, strong residency near tagging sites, and high site fidelity as evidenced by area-restricted search (ARS) behaviors and frequent revisitations. Most manta rays showed restricted home ranges in each region, with no connectivity between regions. Across 25 individuals, the home range (95% utilization distributions) varied significantly, ranging from 19 to 48,294 km2 (mean ± SD = 4667 ± 10,354). These findings offer important insights into the spatial movement patterns of reef manta rays in Indonesia, allowing the formulation of more effective management strategies.

Humpback whales in the Southwest Atlantic Ocean: investigating their breeding movements by satellite tracking

AbstractUnderstanding animals' movements is essential to assess habitat use, life‐history strategies, and population dynamics. Here, we investigate the movement and behavior patterns of 153 humpback whales in the Southwest Atlantic Ocean (SWA) using data obtained by satellite telemetry between 2003 and 2019 during the species' breeding season (August–December) off the Brazilian coast. Switching state space models were applied to estimate behavior states (bmode) classified as Area Restricted Search (ARS), Transiting (TRANS), or uncertain. Whales were distributed from 4°S to 24°S, and five clusters of ARS behavior were identified along the Brazilian coast. Generalized linear mixed modeling revealed three main results: (1) a transition towards more sinuous behavioral states with increasing latitude; (2) more sinuous movement behavior around new moons; (3) movement behavior was temporally dynamic throughout the breeding season over the years, particularly in 2019. The results then revealed important regions where humpback whales cluster to engage in mating and nursing behaviors, highlighting the influence of spatial location and environmental cycles on their behavior. Estimated movement behavior presented here improves the knowledge about the habitat use and movement patterns of SWA humpback whales in their breeding ground and can be used to mitigate potential human‐related impacts.

Search patterns, resource regeneration, and ambush locations impact the competition between active and ambush predators.

Many predators ambush prey rather than pursue them or shift between foraging modes. Active predators typically encounter prey more frequently than ambush predators. I designed a simulation model to examine whether this always holds and how active and ambush predators fare in capturing mobile prey. Prey foraged for clumped resources using area-restricted search, shifting from directional movement before resource encounter to less directional movement afterward. While active predators succeeded more than ambush predators, the advantage of active predators diminished when ambush predators were positioned inside resource patches rather than outside. I investigated the impact of eight treatments and their interactions. For example, regeneration of prey resources increased the difference between ambush predators inside and outside patches, and uncertain prey capture by predators decreased this difference. Several interactions resulted in outcomes different from each factor in isolation. For instance, reducing the directionality level of active predators impacted moderately when applied alone, but when combined with resource regeneration it led to the worst success of active predators against ambush predators inside patches. Ambush predators may not always be inferior to active predators, and one should consider the key traits of the studied system to predict the relative success of these two foraging modes.

Migratory movements of fin whales from the Gulf of St. Lawrence challenge our understanding of the Northwest Atlantic stock structure

Fin whales, Balenoptera physalus, are capital breeders, having the potential to separate breeding and feeding both spatially and temporally. Fin whales occur throughout the Northwest Atlantic, but stock structure and seasonal movements remain unclear. By deploying satellite transmitters on 28 individuals, we examine movement patterns within and beyond the Gulf of St. Lawrence (GSL), Canada, and challenge the current understanding of stock structure. Eight individuals left the GSL in autumn, with five tags persisting into January. Migration patterns of these whales showed considerable variation in timing and trajectory, with movements extending south to 24°N, and thus beyond the assumed distribution limit of the species in the Northwest Atlantic. A rapid return to the Scotian Shelf or Gulf of Maine was observed from several whales after incursions in southern waters, suggesting that fin whales in the Northwest Atlantic may not have a common winter destination that fits the definition of a breeding ground. Area-restricted search (ARS) behavior dominated fin whale activities during summer (92%) and fall (72%), with persistence into the winter (56%); ARS occurred at multiple locations in the GSL, Scotian Shelf and Shelf edge, and near seamounts of the North Atlantic, having characteristics consistent with foraging areas.

Evolution of explorative and exploitative search strategies in collective foraging

Evolutionary theories of foraging hypothesize that foraging strategies evolve to maximize search efficiency. Many studies have investigated the central trade-off between explore–exploit and how individual foragers manage it under various conditions. For foragers in groups, this trade-off can be affected by the social environment, influencing the evolution of individual search strategies. Previous work has shown that when learning socially, explorative search strategies can optimize group search efficiency. However, social learning can cause discrepancies in strategies that benefit the group versus an individual. We model the evolution of explorative and exploitative strategies using Lévy exponents under different levels of social learning and investigate their effect on individual and group search efficiencies. We show that reliance on social learning can lead to the evolution of mixed groups that are not optimally efficient. Exploiters can have a selective advantage in scrounging findings by explorers, but too many exploiters can diminish group efficiencies. However, greater opportunities for social learning can increase the benefits of explorative strategies. Finally, we show that area-restricted search can help individuals balance exploration and exploitation, and make groups more efficient. Our results demonstrate how exploration and exploitation must be balanced at both individual and collective levels for efficient search.

Distributional and behavioral responses of the wintering Oriental Storks to drought in China's largest freshwater lake

Extreme droughts are increasing in frequency and severity globally as a result of climate change. Developing understanding of species' responses to drought is crucial for their conservation, especially in regions experiencing increased aridity. Although numerous studies have investigated birds' responses to drought, the emphasis has primarily been on landbirds. Drought can significantly alter the wetland environments that waterbirds inhabit, but the response of waterbirds to drought remains understudied. In this study, we surveyed the distribution and behavior of Oriental Storks (Ciconia boyciana) in Poyang Lake, which is the largest freshwater lake in China. Results indicate that drought-induced catchment areas at the lowest water level limited the total population size of Oriental Storks in the sub-lakes. Sub-lakes with large catchment areas at the lowest water level demonstrated a capacity to support a larger population of wintering Oriental Storks. Over time, Oriental Storks exhibited a gradual concentration in Changhu Lake, characterized by larger catchments, after resource depletion in sub-lakes with smaller catchments. Additionally, the duration of Oriental Storks’ vigilance and moving behaviors decreased significantly compared with that observed before the drought. After the drought, Oriental Storks increased their foraging efforts, as evidenced by increased presence in deeper water and reaching their heads and necks into deeper water to forage, higher search rates, but lower foraging rates. In accordance with area-restricted search theory, reductions in habitat quality resulting from drought, including extensive fish die-offs, forced Oriental Storks to increase their foraging efforts. Sustaining a specific water area in sub-lakes during droughts can preserve resource availability, which is crucial for the conservation of Oriental Storks. Implementing measures such as water level control and micro-modification of lake bottoms in sub-lakes might mitigate the impact of drought on the piscivorous Oriental Storks.

Decoding the internesting movements of marine turtles using a fine-scale behavioral state approach

IntroductionAn understanding of animal behavior is critical to determine their ecological role and to inform conservation efforts. However, observing hidden behaviors can be challenging, especially for animals that spend most of their time underwater. Animal-borne devices are valuable tools to estimate hidden behavioral states.MethodsWe investigated the fine-scale behavior of internesting hawksbill turtles using the mixed-membership method for movement (M4) which integrated dive variables with spatial components and estimated latent behavioral states.ResultsFive latent behavioral states were identified: 1) pre-nesting, 2) transit, 3) quiescence, and 4) area restricted search within and 5) near the residence of turtles. The last three states associated with a residency period, showed lower activity levels. Notably, when compared to other behaviors the pre-nesting exhibited shallower and remarkably long dives of up to 292 minutes. We noted high fidelity to residence core areas and nesting beaches, within and between nesting seasons, with residence areas decreasing within a season.DiscussionThe latent behaviors identified provide the most detailed breakdown of turtle movement behaviors during the internesting period to date, providing valuable insights into their ecology and behavior. This information can inform marine turtle conservation and management efforts since utilization distributions of individual behavioral states can be used to determine spatially-explicit susceptibility of turtles to various threats based on their behavior. The analyses of utilization distribution revealed a minimal overlap with existing marine protected areas (0.4%), and we show how a new proposal would expand protection to 30%. In short, this study provides valuable guidance for conservation and management of internesting marine turtles at a fine spatiotemporal resolution and can be used to enhance national action plans for endangered species, including the expansion of existing Marine Protected Areas. By flexibly incorporating biologically informative parameters, this approach can be used to study behavior outside of the hawksbill breeding season or even beyond this species.

Ontogeny of movement patterns in naïve grey seal pups inhabiting a complex continental shelf ecosystem.

Most vertebrate offspring must transition from the relative security of parental care (nutrition and protection) to independent foraging. Offspring face many challenges during this critical period, particularly in species where parental care ends at weaning, such as the grey seal (Halichoerus grypus). We studied the development of movement behaviour in naïve grey seal pups from their first trips to sea to about five months of age. Twenty-five (12 males and 13 females) newly-weaned pups were fitted with satellite-linked GPS tags on Sable Island, Nova Scotia, Canada in January 2016. The influence of fixed effects (pup size, sex, week) and the random effect of pup identity on trip characteristics were examined. Movement behaviour was analyzed using a move persistence mixed-effects model. Habitat use was highly variable among individuals and covered much of the geographic distribution of the population. Unlike older juveniles, subadults, and adults in this population, most naïve pups used multiple haulout sites to begin and end trips. There was little evidence of area-restricted search behaviour during trips, suggesting that naïve pups were using an opportunistic foraging tactic that may result in more variable foraging success than that of older, experienced animals. Naïve pups made longer trips with longer haulout durations between them than observed for older greys seals. Males and females differed in some trip characteristics, but sex effects were small over the first few months of life. Offspring size at weaning was not a useful predictor of trip characteristics. Move persistence of grey seal pups was initially high and then decreased over time as individuals gained experience. Both intrinsic and extrinsic factors were influential on the movements of grey seal pups. Greater body length at weaning, longer duration spent on shore after weaning, shallower water column depth, and farther distance from shore were all associated with lower move persistence. Female grey seal pups had lower move persistence than males. Overall, the movements of naïve grey seal pups during the first few months of life were characterized by extensive exploration, but move persistence decreased over time suggesting they may be using an exploration-refinement foraging tactic.

Do beluga whales truly migrate? Testing a key trait of the classical migration syndrome

BackgroundMigration enables organisms to access resources in separate regions that have predictable but asynchronous spatiotemporal variability in habitat quality. The classical migration syndrome is defined by key traits including directionally persistent long-distance movements during which maintenance activities are suppressed. But recently, seasonal round-trip movements have frequently been considered to constitute migration irrespective of the traits required to meet this movement type, conflating common outcomes with common traits required for a mechanistic understanding of long-distance movements. We aimed to test whether a cetacean ceases foraging during so-called migratory movements, conforming to a trait that defines classical migration.MethodsWe used location and dive data collected by satellite tags deployed on beluga whales (Delphinapterus leucas) from the Eastern Beaufort Sea population, which undertake long-distance directed movements between summer and winter areas. To identify phases of directionally persistent travel, behavioural states (area-restricted search, ARS; or Transit) were decoded using a hidden-Markov model, based on step length and turning angle. Established dive profiles were then used as a proxy for foraging, to test the hypothesis that belugas cease foraging during these long-distance transiting movements, i.e., they suppress maintenance activities.ResultsBelugas principally made directed horizontal movements when moving between summer and winter residency areas, remaining in a Transit state for an average of 75.4% (range = 58.5–87.2%) of the time. All individuals, however, exhibited persistent foraging during Transit movements (75.8% of hours decoded as the Transit state had ≥ 1 foraging dive). These data indicate that belugas actively search for and/or respond to resources during these long-distance movements that are typically called a migration.ConclusionsThe long-distance movements of belugas do not conform to the traits defining the classical migration syndrome, but instead have characteristics of both migratory and nomadic behaviour, which may prove adaptive in the face of unpredictable environmental change. Such patterns are likely present in other cetaceans that have been labeled as migratory. Examination of not only horizontal movement state, but also the vertical behaviour of aquatic animals during directed movements is essential for identifying whether a species exhibits traits of the classical migration syndrome or another long-distance movement strategy, enabling improved ecological inference.

Foraging movements of humpback whales relate to the lateral and vertical distribution of capelin in the Barents Sea

Understanding how individual animals modulate their behaviour and movement patterns in response to environmental variability plays a central role in behavioural ecology. Marine mammal tracking studies typically use physical environmental characteristics that vary, and/or proxies of prey distribution, to explain predator movements. Studies linking predator movements and the actual distributions of prey are rare. Here we analysed satellite tag data from ten humpback whales in the Barents Sea (north-east Atlantic) to examine how their spatial movement and dive patterns are influenced by the geographic and vertical distribution of capelin, which is a key prey species for humpback whales. We used capelin density estimates based on direct observations from a trawl-acoustic survey and sun elevation to explore the drivers of changes in movement patterns. We found that the humpback whales’ exhibited characteristic area restricted search movement where capelin density was the highest. While horizontal movements showed both positive and negative individual relationships with sun elevation, humpback whale dive depth was positively correlated with diurnal variations in the vertical distribution of capelin. This suggests that in addition to whales foraging in regions of high capelin density, they also target the densest shoals of capelin at a range of depths, throughout the day and night. Overall, our findings suggest that regions of high capelin density are important foraging grounds for humpback whales, highlighting the central role capelin plays in the Barents Sea marine ecosystem.

How generalization relates to the exploration-exploitation tradeoff

It is known that animals foraging in the wild must balance their levels of exploitation and exploration so as to maximize resource consumption. This usually manifests as an area-restricted search strategy, such that animals tend to exploit environmental patches and make long excursions between patches. This optimal foraging strategy, however, relies on an underlying assumption: nearby locations yield similar resources. Here, we offer an explanation as to how animals utilize this assumption, which implicitly involves generalization. We also describe the computational mechanisms hypothesized to incorporate factors of exploitation, exploration, and generalization, thus, providing a more holistic picture of animal search strategies. Moreover, we connect this foraging behavior to cognition in general. As such, we suggest that cognitive processes, particularly those involved in sequential decision-making, reuse the computational principles grafted into neural activity by the evolution of optimal foraging. We speculate as to what neurobiological substrates may be using area-restricted search, as well as how a model of exploitation, exploration, and generalization can inform psychopathology.

Variation in foraging activity influences area-restricted search behaviour by bottlenose dolphins.

Area-restricted search (ARS) behaviour is commonly used to characterize spatio-temporal variation in foraging activity of predators, but evidence of the drivers underlying this behaviour in marine systems is sparse. Advances in underwater sound recording techniques and automated processing of acoustic data now provide opportunities to investigate these questions where species use different vocalizations when encountering prey. Here, we used passive acoustics to investigate drivers of ARS behaviour in a population of dolphins and determined if residency in key foraging areas increased following encounters with prey. Analyses were based on two independent proxies of foraging: echolocation buzzes (widely used as foraging proxies) and bray calls (vocalizations linked to salmon predation attempts). Echolocation buzzes were extracted from echolocation data loggers and bray calls from broadband recordings by a convolutional neural network. We found a strong positive relationship between the duration of encounters and the frequency of both foraging proxies, supporting the theory that bottlenose dolphins engage in ARS behaviour in response to higher prey encounter rates. This study provides empirical evidence for one driver of ARS behaviour and demonstrates the potential for applying passive acoustic monitoring in combination with deep learning-based techniques to investigate the behaviour of vocal animals.

Satellite tag derived data from two Antarctic blue whales (Balaenopteramusculusintermedia) tagged in the east Antarctic sector of the Southern Ocean.

Satellite tags were deployed on two Antarctic blue whales (Balaenopteramusculusintermedia) in the east Antarctic sector of the Southern Ocean as part of the International Whaling Commission's Southern Ocean Research Partnership initiative. The satellite tracks generated are the first and currently, the only, satellite telemetry data that exist for this critically endangered species. These data provide valuable insights into the movements of Antarctic blue whales on their Antarctic feeding ground. The data were collected between February and April 2013 and span a 110° longitudinal range. This dataset is the first and only detailed movement data that exist for this critically endangered species. As such, this dataset provides the first measures of movement rates (distances travelled, speeds) and movement behaviour (distinguishing transit behaviour from area restricted search behaviour) within the Southern Ocean. These movement-based measures are critical to the ongoing management of Antarctic blue whales as they recover from commercial whaling as they provide insight into foraging behaviour, habitat use, population structure and overlap with anthropogenic threats.

Drivers of polar bear behavior and the possible effects of prey availability on foraging strategy

BackgroundChange in behavior is one of the earliest responses to variation in habitat suitability. It is therefore important to understand the conditions that promote different behaviors, particularly in areas undergoing environmental change. Animal movement is tightly linked to behavior and remote tracking can be used to study ethology when direct observation is not possible.MethodsWe used movement data from 14 polar bears (Ursus maritimus) in Hudson Bay, Canada, during the foraging season (January–June), when bears inhabit the sea ice. We developed an error-tolerant method to correct for sea ice drift in tracking data. Next, we used hidden Markov models with movement and orientation relative to wind to study three behaviors (stationary, area-restricted search, and olfactory search) and examine effects of 11 covariates on behavior.ResultsPolar bears spent approximately 47% of their time in the stationary drift state, 29% in olfactory search, and 24% in area-restricted search. High energy behaviors occurred later in the day (around 20:00) compared to other populations. Second, olfactory search increased as the season progressed, which may reflect a shift in foraging strategy from still-hunting to active search linked to a shift in seal availability (i.e., increase in haul-outs from winter to the spring pupping and molting seasons). Last, we found spatial patterns of distribution linked to season, ice concentration, and bear age that may be tied to habitat quality and competitive exclusion.ConclusionsOur observations were generally consistent with predictions of the marginal value theorem, and differences between our findings and other populations could be explained by regional or temporal variation in resource availability. Our novel movement analyses and finding can help identify periods, regions, and conditions of critical habitat.

Dynamic body acceleration improves mapping of at‐sea foraging behavior in black‐tailed gulls Larus crassirostris

Areas at which seabirds forage intensively can be discriminated by tracking the individuals' at‐sea movements. However, such tracking data may not accurately reflect the birds' exact foraging locations. In addition to tracking data, gathering information on the dynamic body acceleration of individual birds may refine inferences on their foraging activity. Our aim was to classify the foraging behaviors of surface‐feeding seabirds using data on their body acceleration and use this signal to discriminate areas where they forage intensively. Accordingly, we recorded the foraging movements and body acceleration data from seven and ten black‐tailed gulls Larus crassirostris in 2017 and 2018, respectively, using GPS loggers and accelerometers. By referring to video footage of flying and foraging individuals, we were able to classify flying (flapping flight, gliding and hovering), foraging (surface plunging, hop plunging and swimming) and maintenance (drifting, preening, etc.) behaviors using the speed, body angle and cycle and amplitude of body acceleration of the birds. Foraging areas determined from acceleration data corresponded roughly with sections of low speed and area‐restricted searching (ARS) identified from the GPS tracks. However, this study suggests that the occurrence of foraging behaviors may be overestimated based on low‐speed trip sections, because birds may exhibit long periods of reduced movement devoted to maintenance. Opposite, the ARS‐based approach may underestimate foraging behaviors since birds can forage without conducting an ARS. Therefore, our results show that the combined use of accelerometers and GPS tracking helps to adequately determine the important foraging areas of black‐tailed gulls. Our approach may contribute to better discriminate ecologically or biologically significant areas in marine environments.

Underwater visibility constrains the foraging behaviour of a diving pelagic seabird.

Understanding the sensory ecology of species is vital if we are to predict how they will function in a changing environment. Visual cues are fundamentally important for many predators when detecting and capturing prey. However, many marine areas have become more turbid through processes influenced by climate change, potentially affecting the ability of marine predators to detect prey. We performed the first study that directly relates a pelagic seabird species's foraging behaviour to oceanic turbidity. We collected biologging data from 79 foraging trips and 5472 dives of a visually dependent, pursuit-diving seabird, the Manx shearwater (Puffinus puffinus). Foraging behaviour was modelled against environmental variables affecting underwater visibility, including water turbidity, cloud cover and solar angle. Shearwaters were more likely to initiate area-restricted search and foraging dives in clearer waters. Underwater visibility also strongly predicted dive rate and depth, suggesting that fine-scale prey capture was constrained by the detectability of prey underwater. Our novel use of dynamic descriptors of underwater visibility suggests that visual cues are vital for underwater foraging. Our data indicate that climate change could negatively impact seabird populations by making prey more difficult to detect, compounded by the widely reported effects of reduced prey populations.

Ergodicity Breaking in Area-Restricted Search of Avian Predators

Quantifying and comparing patterns of dynamical ecological systems requires averaging over measurable quantities. For example, to infer variation in movement and behavior, metrics such as step length and velocity are averaged over large ensembles. Yet, in nonergodic systems, such averaging is inconsistent; thus, identifying ergodicity breaking is essential in ecology. Using rich, high-resolution, movement data sets (greater than 7×107 localizations) from 70 individuals and continuous-time random walk modeling, we find subdiffusive behavior and ergodicity breaking in the localized movement of three species of avian predators. Small-scale, within-patch movement was found to be qualitatively different, not inferrable and separated from large-scale interpatch movement. Local search is characterized by long, power-law-distributed waiting times with a diverging mean, giving rise to ergodicity breaking in the form of considerable variability uniquely observed at this scale. This implies that wild animal movement is scale specific, with no typical waiting time at the local scale.Received 15 November 2021Revised 13 February 2022Accepted 20 May 2022DOI:https://doi.org/10.1103/PhysRevX.12.031005Published by the American Physical Society under the terms of the Creative Commons Attribution 4.0 International license. Further distribution of this work must maintain attribution to the author(s) and the published article’s title, journal citation, and DOI.Published by the American Physical SocietyPhysics Subject Headings (PhySH)Research AreasAnomalous diffusionBiological movementContinuous-time random walkEcology & evolutionPhase transitionsTechniquesData analysisErgodic theoryFirst passage problemsStatistical PhysicsBiological PhysicsNonlinear Dynamics

Intra-seasonal variation in feeding rates and diel foraging behaviour in a seasonally fasting mammal, the humpback whale.

Antarctic humpback whales forage in summer, coincident with the seasonal abundance of their primary prey, the Antarctic krill. During the feeding season, humpback whales accumulate energy stores sufficient to fuel their fasting period lasting over six months. Previous animal movement modelling work (using area-restricted search as a proxy) suggests a hyperphagic period late in the feeding season, similar in timing to some terrestrial fasting mammals. However, no direct measures of seasonal foraging behaviour existed to corroborate this hypothesis. We attached high-resolution, motion-sensing biologging tags to 69 humpback whales along the Western Antarctic Peninsula throughout the feeding season from January to June to determine how foraging effort changes throughout the season. Our results did not support existing hypotheses: we found a significant reduction in foraging presence and feeding rates from the beginning to the end of the feeding season. During the early summer period, feeding occurred during all hours at high rates. As the season progressed, foraging occurred mostly at night and at lower rates. We provide novel information on seasonal changes in foraging of humpback whales and suggest that these animals, contrary to nearly all other animals that seasonally fast, exhibit high feeding rates soon after exiting the fasting period

Predicting the Foraging Habitats of Sympatrically Breeding Gadfly Petrels in the South Pacific Ocean

Gadfly petrels (genus Pterodroma) are one of the most threatened groups of birds. They are exceptionally well adapted to forage over enormous areas to maximize chances of encountering prey. Their wide-ranging travel, extensive use of oceanic habitats beyond national jurisdictions (the high seas), and limited information on their at-sea distributions and foraging ecology pose several management challenges. Here, we examined the foraging distributions and habitat preferences of three gadfly petrels that breed on Phillip Island (Norfolk Island Group), in the southwest Pacific Ocean, and tested the ability of species distribution models (SDMs) to predict important marine habitats. GPS loggers were deployed in 2018 and 2019 on chick-provisioning black-winged petrels (P. nigripennis) and white-necked petrels (P. cervicalis) and in 2020 on Kermadec petrels (P. neglecta), and hidden Markov models (HMMs) were used to estimate behavioral states across 387 foraging trips. SDMs were built using six algorithms and the predictive performance of models constructed using conventional random cross-validation (CV) was compared to those constructed with spatially independent CV. All three species demonstrated dual-foraging strategies with short trips closer to the colony and longer, presumably self-provisioning, trips with maximum distances from the colony of several thousand kilometers for black-winged and white-necked petrels. Foraging areas of each species were distinctly partitioned across the Tasman Sea during long trips, but there was high overlap during short trips. Black-winged and white-necked petrels exhibited area-restricted search foraging behavior throughout their foraging ranges which spanned almost the entire Tasman Sea and into the western Pacific, whereas the foraging range of Kermadec petrels was restricted closer to the colony. Approximately half of each species’ foraging range extended into the high seas. Response curves and variable importance between the two SDM CV approaches were similar, suggesting that model fitting was robust to the CV approach. However, evaluation using spatially independent CV indicated that generalizability of ensemble SDMs to new data ranged from poor to fair for all three species. This suggests that the maximal-area foraging strategy of gadfly petrels (whereby they search opportunistically for resources across expansive oceanic habitats) results in weak or wide associations with environmental features making predicting important habitats extremely challenging.

Mesoscale movement patterns of common bottlenose dolphins along Kuroshio Current and its bifurcation

AbstractIn the waters off Japan, the patterns of the Kuroshio Current (KC) and its bifurcations are highly variable, which significantly affect the physical and biological environments. Knowledge about species' habitats of marine apex predators such as cetaceans and their variability is fundamental to understand not only their population structures but also their ecosystem structures related through bottom‐up and top‐down trophic linkages. In dynamic ocean environments, however, such crucial information has not been provided adequately by snapshot‐based conventional approaches, such as sighting surveys. Spatiotemporal habitat utilization patterns of the common bottlenose dolphin (Tursiops truncatus) were investigated using fin‐mounted Advanced Research and Global Observation Satellite (ARGOS) tags. Behavioral states at each tracking location were categorized into transiting‐ or area‐restricted searching (ARS) modes based on Bayesian state‐space model. Twelve satellite‐tracked dolphins showed different movement patterns. Their habitat ranges were estimated by minimum convex polygons (MCPs), and movement patterns were classified into three types (four animals each of eastward, offshore, and westward movements) based on MCP overlapping among tagged dolphins. The animals that restricted their habitat to the coastward side of the KC (CSKC) tended to show ARS modes more frequently. Estimated locations of these animals corresponded well with cold coastal waters. On the other hand, offshore movements corresponded well with clockwise flow generated by the bifurcation of the KC, and those dolphins were likely straying into less productive waters. These patterns suggest that common bottlenose dolphins off the southern coast of Japan mainly utilize productive CSKC waters but occasionally move to offshore waters.