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Habitat selection is the process by which an individual makes an active decision to make use of a particular habitat when others are available. The ability to infer habitat selection therefore requires observations of movement through space and time which can be particularly challenging for marine species that are cryptic and do not regularly visit the sea surface. Rajidae (skates) are benthic mesopredators that inhabit turbid coastal waters and exhibit site fidelity, making them an ideal group for studying habitat selection and resource partitioning using a fixed, passive acoustic receiver array in the western English Channel, UK. Using network analysis, significant differences were found in the way 4 species of Rajidae occupied different parts of the array; for example, Raja microocellata and R. clavata occupied shallower habitat than R. brachyura and R. montagui. R. montagui and R. brachyura were further separated, with each species detected more frequently at different receivers. Males and females of all 4 species were also detected at different receivers and at different times. These results demonstrate habitat selection, resource partitioning among species and sexual segregation in these 4 species of Rajidae. Our findings are important evidence for management of fisheries, such as the designation of marine protected areas, and further highlight the potential of this method for tracking other mobile marine species in temperate, open coastal regions.

Marine traffic is increasing globally yet collisions with endangered megafauna such as whales, sea turtles, and planktivorous sharks go largely undetected or unreported. Collisions leading to mortality can have population-level consequences for endangered species. Hence, identifying simultaneous space use of megafauna and shipping throughout ranges may reveal as-yet-unknown spatial targets requiring conservation. However, global studies tracking megafauna and shipping occurrences are lacking. Here we combine satellite-tracked movements of the whale shark, Rhincodon typus, and vessel activity to show that 92% of sharks’ horizontal space use and nearly 50% of vertical space use overlap with persistent large vessel (>300 gross tons) traffic. Collision-risk estimates correlated with reported whale shark mortality from ship strikes, indicating higher mortality in areas with greatest overlap. Hotspots of potential collision risk were evident in all major oceans, predominantly from overlap with cargo and tanker vessels, and were concentrated in gulf regions, where dense traffic co-occurred with seasonal shark movements. Nearly a third of whale shark hotspots overlapped with the highest collision-risk areas, with the last known locations of tracked sharks coinciding with busier shipping routes more often than expected. Depth-recording tags provided evidence for sinking, likely dead, whale sharks, suggesting substantial “cryptic” lethal ship strikes are possible, which could explain why whale shark population declines continue despite international protection and low fishing-induced mortality. Mitigation measures to reduce ship-strike risk should be considered to conserve this species and other ocean giants that are likely experiencing similar impacts from growing global vessel traffic., [Significance] Global vessel traffic is increasing alongside world economic growth. The potential for rising lethal ship strikes on endangered species of marine megafauna, such as the plankton-feeding whale shark, remains poorly understood since areas of highest overlap are seldom determined across an entire species range. Here we show how satellite tracking whale sharks and large vessel movements globally provides a means to localize high-overlap areas and to determine how collision risk changes in time. Our results point to potential high levels of undetected or unreported ship strikes, which may explain why whale shark populations continue to decline despite protection and low fishing-induced mortality. Collision mitigations in high-collision-risk areas appear necessary to help conserve this iconic species., Funding for data analysis was provided by the UK Natural Environment Research Council (NERC) through a University of Southampton INSPIRE DTP PhD Studentship to F.C.W. Additional funding for data analysis was provided by NERC Discovery Science (NE/R00997/X/1) and the European Research Council (ERC-AdG-2019 883583 OCEAN DEOXYFISH) to D.W.S., Fundação para a Ciência e a Tecnologia (FCT) under PTDC/BIA/28855/2017 and COMPETE POCI-01–0145-FEDER-028855, and MARINFO–NORTE-01–0145-FEDER-000031 (funded by Norte Portugal Regional Operational Program [NORTE2020] under the PORTUGAL 2020 Partnership Agreement, through the European Regional Development Fund–ERDF) to N.Q. FCT also supported N.Q. (CEECIND/02857/2018) and M.V. (PTDC/BIA-COM/28855/2017). D.W.S. was supported by a Marine Biological Association Senior Research Fellowship.

Groups of basking sharks engaged in circling behaviour are rarely observed, and their function remains enigmatic in the absence of detailed observations. Here, underwater and aerial video recordings of multiple circling groups of basking sharks during late summer (August and September 2016–2021) in the eastern North Atlantic Ocean showed groups numbering between 6 and 23 non-feeding individuals of both sexes. Sharks swam slowly in a rotating “torus” (diameter range: 17–39 m), with individuals layered vertically from the surface to a maximum depth of 16 m. Within a torus, sharks engaged in close-following, echelon, close-flank approach or parallel-swimming behaviours. Measured shark total body lengths were 5.4–9.5 m (mean LT: 7.3 m ± 0.9 s.d.; median: 7.2 m, n = 27), overlapping known lengths of sexually mature males and females. Males possessed large claspers with abrasions that were also observed on female pectoral fins. Female body colouration was paler than that of males, similar to colour changes observed during courtship and mating in other shark species. Individuals associated with most other members rapidly (within minutes), indicating toroidal behaviours facilitate multiple interactions. Sharks interacted through fin–fin and fin–body contacts, rolling to expose the ventral surfaces to following sharks, and breaching behaviour. Toruses formed in late summer when feeding aggregations in zooplankton-rich thermal fronts switched to non-feeding following and circling behaviours. Collectively, the observations explain a courtship function for toruses. This study highlights northeast Atlantic coastal waters as a critical habitat supporting courtship reproductive behaviour of endangered basking sharks, the first such habitat identified for this species globally. (https://www.youtube.com/watch?v=7i2H1wVOmb4) yes

REPLYING TO H. Murua et al. Nature https://doi.org/10.1038/s41586-021-03396-4 (2021) Our previously published paper1 provided global fine-scale spatiotemporal estimates (1° × 1°; monthly) of overlap and fishing exposure risk (FEI) between satellite-tracked shark space use and automatic identification system (AIS) longline fishing effort. We did not assess shark mortality directly, but in addition to replying to the Comment by Murua et al.2, we confirm—using regression analysis of spatially matched data—that fishing-induced pelagic shark mortality (catch per unit effort (CPUE)) is greater where FEI is higher. We focused on assessing shark horizontal spatiotemporal overlap and exposure risk with fisheries because spatial overlap is a major driver of fishing capture susceptibility and previous shark ecological risk assessments (ERAs) assumed a homogenous shark density within species-range distributions3,4,5 or used coarse-scale modelled occurrence data, rather than more ecologically realistic risk estimates in heterogeneous habitats that were selected by sharks over time. Furthermore, our shark spatial exposure risk implicitly accounts for other susceptibility factors with equal or similar probabilities to those commonly used in shark ERAs3,5.

29 pages, 9 figures, 4 tables.-- This article is distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use and redistribution provided that the original author and source are credite, Climate-driven expansions of ocean hypoxic zones are predicted to concentrate pelagic fish in oxygenated surface layers, but how expanding hypoxia and fisheries will interact to affect threatened pelagic sharks remains unknown. Here, analysis of satellite-tracked blue sharks and environmental modelling in the eastern tropical Atlantic oxygen minimum zone (OMZ) shows shark maximum dive depths decreased due to combined effects of decreasing dissolved oxygen (DO) at depth, high sea surface temperatures, and increased surface-layer net primary production. Multiple factors associated with climate-driven deoxygenation contributed to blue shark vertical habitat compression, potentially increasing their vulnerability to surface fisheries. Greater intensity of longline fishing effort occurred above the OMZ compared to adjacent waters. Higher shark catches were associated with strong DO gradients, suggesting potential aggregation along suitable DO gradients contributed to habitat compression and higher fishing-induced mortality. Fisheries controls to counteract deoxygenation effects on shark catches will be needed as oceans continue warming, Funding was provided by the UK Natural Environment Research Council (NERC) (NE/R00997/X/1), European Research Council (ERC-AdG-2019 883583 OCEAN DEOXYFISH), NERC Oceans 2025 Strategic Programme (all to DWS), the Save Our Seas Foundation (DWS, NQ), Fundacao para a Ciencia e a Tecnologia (FCT) under PTDC/BIA/28855/2017 and COMPETE POCI-01–0145-FEDER-028855 (NQ, DWS), and MARINFO–NORTE-01–0145-FEDER-000031 (funded by Norte Portugal Regional Research article Ecology Operational Program [NORTE2020], under the PORTUGAL 2020 Partnership Agreement, through the European Regional Development Fund-RDF), and Xunta de Galicia Spain under the Isabel Barreto Program 2009–2012 (GM). FCT supported NQ (CEECIND/02857/2018), MV (PTDC/BIA-COM/28855/2017), GM (PTDC/MAR-BIO/4458/2012), FV (CEECIND/03469/2017), and JF (through the strategic project UID/MAR/04292/2013 to MARE). Direccao Regional de Cieˆ ncia e Tecnologia (DRCT) also supported JF (M3.1a/F/062/2016). DWS was supported by a Marine Biological Association Senior Research Fellowship

REPLYING TO A. V. Harry & J. M. Braccini Nature https://doi.org/10.1038/s41586-021-03463-w (2021) Our global analysis1 estimated the overlap and fishing exposure risk (FEI) using the space use of satellite-tracked sharks and longline fishing effort monitored by the automatic identification system (AIS). In the accompanying Comment, Harry and Braccini2 draw attention to two localized shark–longline vessel overlap hotspots in Australian waters, stating that 47 fishing vessels were misclassified as longline and purse seine vessels in the Global Fishing Watch (GFW)3 2012–2016 AIS fishing effort data product that we used. This, they propose2, results in misidentifications that highlight fishing exposure hotspots that are subject to an unexpected level of sensitivity in the analysis and they suggest that misidentifications could broadly affect the calculations of fishing exposure and the central conclusions of our study1. We acknowledged in our previously published paper1 that gear reclassifications were likely to occur for a small percentage of the more than 70,000 vessels studied, however, here we demonstrate that even using much larger numbers of vessel reclassifications than those proposed by Harry and Braccini2, the central results and conclusions of our paper1 do not change.

Additional co-authors: G. Peirson, K. K. Richter, M. I. Taylor,C. N. Trueman, B. Hayden, J. R. Stevens

Management and conservation of marine predator species relies on a fundamental knowledge of their movements and behaviour. Pop‐up satellite archival tags were used to investigate the vertical movement patterns of five blue sharks (Prionace glauca) and one thresher shark (Alopias vulpinus) within the southeastern Indian Ocean. Sections of similar depth distribution, identified using a split moving window analysis, were investigated in relation to the thermal structure of the water column and activity rates. Minimum horizontal displacement of between 66 and 5,187 km for blue sharks and 16 km for the thresher shark were recorded over 863 tracking days. Maximum depths ranged from 540 to 807 m for blue sharks and 144 m for the thresher shark. All sharks displayed plasticity in their depth distribution, with diel vertical movements and surface‐oriented movements the two most common patterns. Diel movement of prey is the most likely explanation for diel vertical movements of thresher and blue sharks. This study has improved our understanding of the vertical movement patterns of these pelagic predators and the relationship between their depth distribution, temperature, and activity.

Diel vertical migration (DVM) is a widespread behaviour among many pelagic species, from zooplankton to sharks and has been widely studied in both marine and freshwater environments. Usually, DVM comprises repeated daily vertical movements through the water column, from shallower at night to deeper during the day. Consequently, DVM is perhaps unexpected in benthic predators. Nonetheless, DVM has been observed in benthic sharks and freshwater teleosts, where it comprises inshore-offshore migrations over the substrate. However, there is no clear evidence of this behaviour in large temperate benthic predators, such as skates. Here we present new observations of DVM in 4 species of skate (Raja brachyura, R. clavata, R. microocellata and R. montagui) that identify it as a general behaviour in this clade. Analysis of 89 depth recording archival tags yielded 674 clear DVM events where skates left daytime deeper waters for shallower nighttime areas before returning to within 2.5 m of starting depths. Interestingly, these events closely resemble those of central place foragers, where shallow areas are foraging and deeper areas are refuging locations. Behaviour such as this has not been previously recorded in marine benthic predators, and the findings suggest that DVM might occur in many other benthic species. A broader understanding of DVM in benthic animals will be important in the design of effective boundaries for marine protected areas. These findings also have implications for trophic coupling between deep and shallow benthic zones. Further characteristics of this unexpected behaviour are presented and hypotheses for its occurrence are discussed.

A sympatric assemblage of morphologically similar predators is expected to exhibit fine-scale habitat segregation, or resource partitioning, to reduce the effects of direct competition. This principle has been well studied for predators in terrestrial ecosystems. In the marine environment, the fine-scale spatial segregation of sympatric species of large predators is poorly understood because detailed movement and behavioural data are often not available across multiple species within the same timeframe. The ways in which co-occurring congeneric predators separate spatially is even less well understood. Medium-sized species of skates (genus Raja) co-occur in temperate habitats of the north-east Atlantic Ocean, share similar morphologies and have distributional ranges that overlap significantly in the western English Channel ecosystem. In the present study, detailed depth time series retrieved from 89 electronic data storage tags attached to 4 species of skate were analysed to determine preferred depth ranges. The 4 species were found to segregate spatially into 2 groups, with one group having a significantly shallower core annual depth range than the other. To our knowledge, fine-scale segregation by depth has not been observed previously. Interestingly, the members of each species group appeared complementary, each group comprising species with different dietary preferences and with a larger and smaller body size. An understanding of how core depth ranges differ and how these species utilise vertical habitat could potentially enable geographic ranges around the coast to be predicted, with important implications for how these species interact with fisheries and Marine Protected Areas.

Efficient searching for resources such as food by animals is key to their survival. It has been proposed that diverse animals from insects to sharks and humans adopt searching patterns that resemble a simple Lévy random walk, which is theoretically optimal for ‘blind foragers’ to locate sparse, patchy resources. To test if such patterns are generated intrinsically, or arise via environmental interactions, we tracked free-moving Drosophila larvae with (and without) blocked synaptic activity in the brain, suboesophageal ganglion (SOG) and sensory neurons. In brain- blocked larvae, we found that extended substrate exploration emerges as multi-scale movement paths similar to truncated Lévy walks. Strikingly, power-law exponents of brain/SOG/sensory- blocked larvae averaged 1.96, close to a theoretical optimum (m ffi 2.0) for locating sparse resources. Thus, efficient spatial exploration can emerge from autonomous patterns in neural activity. Our results provide the strongest evidence so far for the intrinsic generation of Lévy-like movement patterns. Fil: Sims, David W. The Marine Biological Association; Reino Unido. University of Southampton; Reino Unido Fil: Humphries, Nicolas E. The Marine Biological Association; Reino Unido Fil: Hu, Nan. University of Cambridge; Reino Unido Fil: Medan, Violeta. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Fisiología, Biología Molecular y Neurociencias. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Fisiología, Biología Molecular y Neurociencias; Argentina. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Departamento de Fisiología, Biología Molecular y Celular; Argentina Fil: Berni, Jimena. University of Cambridge; Estados Unidos

There have been efforts around the globe to track individuals of many marine species and assess their movements and distribution, with the putative goal of supporting their conservation and management. Determining whether, and how, tracking data have been successfully applied to address real-world conservation issues is, however, difficult. Here, we compile a broad range of case studies from diverse marine taxa to show how tracking data have helped inform conservation policy and management, including reductions in fisheries bycatch and vessel strikes, and the design and administration of marine protected areas and important habitats. Using these examples, we highlight pathways through which the past and future investment in collecting animal tracking data might be better used to achieve tangible conservation benefits.

Effective ocean management and the conservation of highly migratory species depend on resolving the overlap between animal movements and distributions, and fishing effort. However, this information is lacking at a global scale. Here we show, using a big-data approach that combines satellite-tracked movements of pelagic sharks and global fishing fleets, that 24% of the mean monthly space used by sharks falls under the footprint of pelagic longline fisheries. Space-use hotspots of commercially valuable sharks and of internationally protected species had the highest overlap with longlines (up to 76% and 64%, respectively), and were also associated with significant increases in fishing effort. We conclude that pelagic sharks have limited spatial refuge from current levels of fishing effort in marine areas beyond national jurisdictions (the high seas). Our results demonstrate an urgent need for conservation and management measures at high-seas hotspots of shark space use, and highlight the potential of simultaneous satellite surveillance of megafauna and fishers as a tool for near-real-time, dynamic management., Data analysis was funded in part by the Marine Biological Association (MBA) and the UK Natural Environment Research Council (NERC) (NE/R00997X/1) (to D.W.S.) with additional research support from the Save Our Seas Foundation and the NERC Oceans 2025 Strategic Research Programme, in which D.W.S. was a principal investigator. D.W.S. was supported by an MBA Senior Research Fellowship, N.Q. by European Regional Development Fund (FEDER) via the Programa Operacional Competitividade e Internacionalização (COMPETE), National Funds via Fundação para a Ciência e a Tecnologia (FCT) under PTDC/MAR/100345/2008 and COMPETE FCOMP-01-0124-FEDER-010580 (to N.Q. and D.W.S.), and Norte Portugal Regional Operational Programme (NORTE 2020) under the PORTUGAL 2020 Partnership Agreement through the European Regional Development Fund (ERDF) in project MarInfo (NORTE-01-0145-FEDER-000031). Additional support was provided by an FCT Investigator Fellowship IF/01611/2013 (N.Q.), FCT Doctoral Fellowship PD/BD/52603/2014 (M.V.), PTDC/MAR-BIO/4458/2012, Xunta de Galicia - Isabel Barreto Program 2009-2012 (G.M.), Australian Research Council (ARC) grant DE170100841 and operational funds from the Australian Institute for Marine Science (AIMS) (both to A.M.M.S.).

Rhythmic activity patterns are ubiquitous in animals and in the marine environment a dominant rhythmic activity is the diel vertical migration (DVM) of pelagic organisms, moving or ‘migrating’ from deep waters during the day to shallower waters at night. While this overall pattern of movement is well understood, the cryptic nature of the marine environment has limited the study of fine-scale movements within each phase. Active pelagic predators, such as tuna, perform consistent, predictable large-scale vertical movements; however, the fine-scale movements nested within these larger movements have not previously been investigated in detail. Further, the prey field densities are known to differ significantly between day and night, presenting an opportunity to study differences in foraging patterns between these two phases. Here, using long-term depth time series recorded from 93 bigeye tuna, Thunnus obesus , with electronic tags (18 003 days of data), fine-scale changes in vertical movement patterns between day and night time phases were investigated in the context of the Levy foraging hypothesis, which predicts a Levy distribution of move steps during foraging when prey is scarce, but an exponential distribution when prey is abundant and searching is not required. During the day, T. obesus were found to exhibit scale-free movements well fitted by a Levy distribution indicating optimized searching for sparsely distributed prey. During night-time hours, however, exponentially distributed scale-dependent move step lengths were found to be dominant, supporting a simple, Brownian, movement pattern sufficient where prey is abundant. This study not only confirms the predictions of the Levy foraging hypothesis but suggests that the identification of Levy patterns in movement data can be a useful indicator of foraging activity in animals that are difficult to observe directly.

Often the ecology of a commercially fished species and the behaviour of the fishers are inextricably linked. An understanding of both species ecology and fisher behaviour are therefore fundamental for effective fisheries management. In Europe, Rajidae are commercially fished species that are vulnerable to overfishing, have historically been grouped in landings data and declines have been reported to the point of local extinction in some species. Management of these species is therefore important, however an understanding of species ecology and the way they interact with the fishery is lacking. Here we investigate how the differing ecology of four species affects interactions with the fishery by considering gear types, Marine Protected Areas, location of capture and geographic displacements. Our results demonstrate that the four species interacted differently with the fishery, for example Raja clavata were more commonly captured by static nets (69 % of recaptures) and Raja brachyura more commonly by trawlers (77 %). There were clear differences in the location of capture, depths and displacements of the four species resulting in differing interactions with existing management in the form of Marine Protected Areas. For example 53 % of R. clavata recaptures were within the boundaries of an MPA compared to 18 % for R. brachyura. These results provide important differences among species that are generally grouped by current management, emphasising the need for further species specific research to ensure effective management.

The extent of increasing anthropogenic impacts on large marine vertebrates partly depends on the animals’ movement patterns. Effective conservation requires identification of the key drivers of movement including intrinsic properties and extrinsic constraints associated with the dynamic nature of the environments the animals inhabit. However, the relative importance of intrinsic versus extrinsic factors remains elusive. We analyze a global dataset of ∼2.8 million locations from >2,600 tracked individuals across 50 marine vertebrates evolutionarily separated by millions of years and using different locomotion modes (fly, swim, walk/paddle). Strikingly, movement patterns show a remarkable convergence, being strongly conserved across species and independent of body length and mass, despite these traits ranging over 10 orders of magnitude among the species studied. This represents a fundamental difference between marine and terrestrial vertebrates not previously identified, likely linked to the reduced costs of locomotion in water. Movement patterns were primarily explained by the interaction between species-specific traits and the habitat(s) they move through, resulting in complex movement patterns when moving close to coasts compared with more predictable patterns when moving in open oceans. This distinct difference may be associated with greater complexity within coastal microhabitats, highlighting a critical role of preferred habitat in shaping marine vertebrate global movements. Efforts to develop understanding of the characteristics of vertebrate movement should consider the habitat(s) through which they move to identify how movement patterns will alter with forecasted severe ocean changes, such as reduced Arctic sea ice cover, sea level rise, and declining oxygen content., Workshop funding was granted by the University of Western Australia (UWA) Oceans Institute, the Australian Institute of Marine Science (AIMS), and King Abdullah University of Science and Technology (KAUST). A.M.M.S. was supported by Australian Research Council Grant DE170100841 and an Indian Ocean Ocean Marine Research Centre (UWA, AIMS, Commonwealth of Scientific and Industrial Research Organisation) fellowship. J.P.R., V.M.E., and J.F.G. were supported by Agencia Estatal de Investigación (AEI, Spain) and Fondo Europeo de Desarrollo Regional (FEDER) through project Spatiotemporality in Sociobological Interactions, Models and Methods (SPASIMM) (FIS2016-80067-P AEI/FEDER, European Union), and by research funding from KAUST. J.P.R. was supported by Ministerio de Educación, Cultura y Deporte (Formación de Profesorado Universitario Grant, Spain). D.W.S. was supported by the UK Natural Environment Research Council and Save Our Seas Foundation. N.Q. was supported by Fundação para a Ciência e Tecnologia (Portugal). M.M.C.M. was supported by a Coordenação de Aperfeiçoamento de pessoal de Nível Superior fellowship (Ministry of Education).

A large, pregnant, female bull shark Carcharhinus leucas was tracked migrating from Seychelles across open ocean to south-east Madagascar, c. 2000 km away, and back again. In Madagascar, the shark spent a prolonged period shallower than 5 m, consistent with entering estuarine habitat to pup, and upon return to Seychelles the shark was slender and no longer gravid. This represents an unprecedented return migration across the open ocean for a C. leucas and highlights the need for international collaboration to manage the regional C. leucas population sustainably.

Eight whale sharks tagged with pop-up satellite archival tags off the Gulf of California, Mexico, were tracked for periods of 14–134 days. Five of these sharks were adults, with four females visually assessed to be pregnant. At least for the periods they were tracked, juveniles remained in the Gulf of California while adults moved offshore into the eastern Pacific Ocean. We propose that parturition occurs in these offshore waters. Excluding two juveniles that remained in the shallow tagging area for the duration of tracking, all sharks spent 65 ± 20.7% (SD) of their time near the surface, even over deep water, often in association with frontal zones characterized by cool-water upwelling. While these six sharks all made dives into the meso- or bathypelagic zones, with two sharks reaching the maximum depth recordable by the tags (1285.8 m), time spent at these depths represented a small proportion of the overall tracks. Most deep dives (72.7%) took place during the day, particularly during the early morning and late afternoon. Pronounced habitat differences by ontogenetic stage suggest that adult whale sharks are less likely to frequent coastal waters after the onset of maturity.

article i nfo The increasing importance of the European common cuttlefish Sepia officinalis as a commercial fishery re- source raises concerns that our limited understanding of its behaviour and movements in natural habitats may impede future management for this species. Whilst information regarding the fine-scale movements and behaviour of small marine species such as cuttlefish can be difficult to obtain, the increasing miniaturisation of electronic tags, combined with novel attachment techniques, has now enabled tracking of this cephalopod. In this study we describe the first use of electronic tags attached to adult and sub-adult S. officinalis to monitor their movements and behaviour within natural environments. Eight adults (170- 205 mm dorsal mantle length (DML)) were fitted with continuous acoustic transmitters and ten sub-adults (132-180 mm DML) were fitted with coded acoustic transmitters. All transmitters were attached to the internal cuttlebone of the living animal. Tagged animals recovered rapidly from anaesthesia and surgical pro- cedures and were found to retain tags for periods of up to 11 weeks in this short-term study. Six of the eight adult cuttlefish released into a radio-acoustic positioning (VRAP) array left the area rapidly (≤45 min) whilst two remained for multiple days (≤9 days). Short-term site fidelity at a local level was observed in only two adults, whilst larger scale movements along the coastline (≤35 km) occurred in three adults, indicating that a complex range of spatial behaviour patterns could occur among spawning adults. Similar patterns were ob- served in the space use of sub-adults monitored using a passive acoustic-receiver array on the seabed. Three sub-adults were detected within the study area over an extended period of time (≥73 days), whilst the remaining sub-adults left the study area rapidly and without return. The capacity to monitor the movements of both adult and sub-adult cuttlefish in their natural environment will facilitate identification and under- standing of ontogenic changes in behaviours and migratory patterns of wild populations. Our results high- light the potential of electronic tagging methods as tools to aid conservation and management of this important commercial fishery species in the future.

The ocean sunfish (Mola mola) is the world’s heaviest bony fish reaching a body mass of up to 2.3 tonnes. However, the prey M. mola consumes to fuel this prodigious growth remains poorly known. Sunfish were thought to be obligate gelatinous plankton feeders, but recent studies suggest a more generalist diet. In this study, through molecular barcoding and for the first time, the diet of sunfish in the north-east Atlantic Ocean was characterised. Overall, DNA from the diet content of 57 individuals was successfully amplified, identifying 41 different prey items. Sunfish fed mainly on crustaceans and teleosts, with cnidarians comprising only 16% of the consumed prey. Although no adult fishes were sampled, we found evidence for an ontogenetic shift in the diet, with smaller individuals feeding mainly on small crustaceans and teleost fish, whereas the diet of larger fish included more cnidarian species. Our results confirm that smaller sunfish feed predominantly on benthic and on coastal pelagic species, whereas larger fish depend on pelagic prey. Therefore, sunfish is a generalist predator with a greater diversity of links in coastal food webs than previously realised. Its removal as fisheries’ bycatch may have wider reaching ecological consequences, potentially disrupting coastal trophic interactions.

1. Search processes play an important role in physical, chemical and biological systems. In animal foraging, the search strategy predators should use to search optimally for prey is an enduring question. Some models demonstrate that when prey is sparsely distributed, an optimal search pattern is a specialised random walk known as a Levy flight, whereas when prey is abundant, simple Brownian motion is sufficiently efficient. These predictions form part of what has been termed the Levy flight foraging hypothesis (LFF) which states that as Levy flights optimise random searches, movements approximated by optimal Levy flights may have naturally evolved in organisms to enhance encounters with targets (e.g. prey) when knowledge of their locations is incomplete. 2. Whether free-ranging predators exhibit the movement patterns predicted in the LFF hypothesis in response to known prey types and distributions, however, has not been determined. We tested this using vertical and horizontal movement data from electronic tagging of an apex predator, the great white shark Carcharodon carcharias, across widely differing habitats reflecting different prey types. 3. Individual white sharks exhibited movement patterns that predicted well the prey types expected under the LFF hypothesis. Shark movements were best approximated by Brownian motion when hunting near abundant, predictable sources of prey (e.g. seal colonies, fish aggregations), whereas movements approximating truncated Levy flights were present when searching for sparsely distributed or potentially difficult-to-detect prey in oceanic or shelf environments, respectively. 4. That movement patterns approximated by truncated Levy flights and Brownian behaviour were present in the predicted prey fields indicates search strategies adopted by white sharks appear to be the most efficient ones for encountering prey in the habitats where such patterns are observed. This suggests that C. carcharias appears capable of exhibiting search patterns that are approximated as optimal in response to encountered changes in prey type and abundance, and across diverse marine habitats, from the surf zone to the deep ocean. 5. Our results provide some support for the LFF hypothesis. However, it is possible that the observed Levy patterns of white sharks may not arise from an adaptive behaviour but could be an emergent property arising from simple, straight-line movements between complex (e.g. fractal) distributions of prey. Experimental studies are needed in vertebrates to test for the presence of Levy behaviour patterns in the absence of complex prey distributions.

Over-fishing may lead to a decrease in fish abundance and a proliferation of jellyfish. Active movements and prey search might be thought to provide a competitive advantage for fish, but here we use data-loggers to show that the frequently occurring coastal jellyfish ( Rhizostoma octopus ) does not simply passively drift to encounter prey. Jellyfish (327 days of data from 25 jellyfish with depth collected every 1 min) showed very dynamic vertical movements, with their integrated vertical movement averaging 619.2 m d −1 , more than 60 times the water depth where they were tagged. The majority of movement patterns were best approximated by exponential models describing normal random walks. However, jellyfish also showed switching behaviour from exponential patterns to patterns best fitted by a truncated Lévy distribution with exponents (mean μ = 1.96, range 1.2–2.9) close to the theoretical optimum for searching for sparse prey ( μ opt ≈ 2.0). Complex movements in these ‘simple’ animals may help jellyfish to compete effectively with fish for plankton prey, which may enhance their ability to increase in dominance in perturbed ocean systems.

Shifts in the movement, activity or behaviour of individual animals in relation to chang- ing environmental landscapes play an important role in determining re-distribution patterns of pop- ulations. Such spatial dynamics are poorly understood for pelagic sharks despite the decline of many species due to overfishing. Satellite-linked archival transmitters were used to record the movements of blue sharks Prionace glauca for the first time in the northeast Atlantic. A total of 256 tracking days were recorded, with sharks covering an estimated minimum distance of 11 432 km. Generally, blue sharks displayed wide-ranging southward movements away from the tagging areas (English Chan- nel, southern Portugal and the Azores Islands), with juvenile sharks displaying less site fidelity than previously suggested from other methods. Vertical movements ranged from the surface to a maxi- mum depth of 696 m, and water temperatures varied from 10.6 to 24.6°C. Shifts in both thermal and depth-use patterns were detected, with tracked sharks displaying behavioural variability correlated with the environment, both within and between individuals. Patterns of behaviour in coastal areas were generally linked to the thermal structure of the water column; sharks displayed some prefer- ence for surface layers in thermally stratified waters, but displayed less regular behaviour in well- mixed or weakly stratified waters. In offshore locations, sharks usually adopted either a surface- oriented or a depth-oriented diving pattern, often switching between both in response to probable changes in prey distribution or prey type when in deeper water. Fishing-induced mortality of tracked sharks was approximately 10% during the period of 13 to 70 d post-tagging. Our results indicate a variety of short and longer range movements of juvenile and sub-adult blue sharks with respect to time and environment in the northeast Atlantic, which suggests complex spatial dynamics and popu- lation structuring in this exploited species.

Understanding the habitat preferences of large marine vertebrates has only recently become tractable with the widespread availability of satellite telemetry for monitoring movements and behaviour. For many species with low population abundances, however, little progress has been made in identifying space use patterns. The endothermic porbeagle shark, Lamna nasus , has declined in the North Atlantic due to severe fishing pressure, with little evidence of recovery. One potential factor exacerbating population decline is area fidelity to coastal waters where fisheries are intensive. We tested for short-term area fidelity by attaching pop-up satellite-linked archival transmitters to four porbeagles in summer 2007, resulting in 175 days total tracking time covering an estimated 10,256 km distance. Throughout July and August the sharks occupied localised areas (8,602 – 90,153 km 2 ) within the Celtic Sea, between the south-west UK, south-west Wales and southern Ireland. Only one shark was tracked into the autumn, when it moved into deep water off the continental shelf, then north towards colder latitudes. Sharks occupied a broad vertical depth range (0 – 552 m) and water temperatures (9° - 19 °C). Dives were made frequently from the surface to near the seabed in shelf areas, however, in shelf edge habitats extended periods of time were spent at depths > 300 m. Porbeagles showed considerable plasticity in diel depth changes within and between individuals and as a function of habitat type. In addition to no obvious day-night difference in depth occupation, some sharks showed reverse diel vertical migration (DVM) (dawn ascent – dusk descent) in well-mixed coastal waters whereas normal DVM (dawn descent – dusk ascent) characterised movements into deeper, thermally well-stratified waters. The variable behaviours may reflect the need for different search strategies depending on habitat and prey types encountered. These results show porbeagles are potentially vulnerable to fisheries throughout the summer when they aggregate, and that large scale movement across national boundaries identifies the need for international conservation measures.

11 páginas, 5 figuras.-- James S. E. Lea ... et al.-- This work is licensed under a Creative Commons Attribution 4.0 International License, Long-distance movements of animals are an important driver of population spatial dynamics and determine the extent of overlap with area-focused human activities, such as fishing. Despite global concerns of declining shark populations, a major limitation in assessments of population trends or spatial management options is the lack of information on their long-term migratory behaviour. For a large marine predator, the tiger shark Galeocerdo cuvier, we show from individuals satellite-tracked for multiple years (up to 1101 days) that adult males undertake annually repeated, round-trip migrations of over 7,500 km in the northwest Atlantic. Notably, these migrations occurred between the highly disparate ecosystems of Caribbean coral reef regions in winter and high latitude oceanic areas in summer, with strong, repeated philopatry to specific overwintering insular habitat. Partial migration also occurred, with smaller, immature individuals displaying reduced migration propensity. Foraging may be a putative motivation for these oceanic migrations, with summer behaviour showing higher path tortuosity at the oceanic range extremes. The predictable migratory patterns and use of highly divergent ecosystems shown by male tiger sharks appear broadly similar to migrations seen in birds, reptiles and mammals, and highlight opportunities for dynamic spatial management and conservation measures of highly mobile sharks, Funding for this study was provided by the Guy Harvey Ocean Foundation, Guy Harvey Research Institute, the Shark Foundation (Hai Stiftung), and the Bermuda Shark Project. Funding for data analysis was provided by the UK Natural Environment Research Council (NERC) ‘Oceans 2025’ Strategic Research Programme in which D.W.S. is a principal investigator, a Fundação para a Ciência e a Tecnologia (FCT) grant (PTDC/MAR/100345/2008) to N.Q. and D.W.S., and an FCT grant (SFRH/BD/68717/2010) to L.L.S., an FCT Investigator Fellowship to N.Q. (IF/01611/2013). J.S.E.L. was supported by Danah Divers and the Marine Biological Association of the UK (MBA) and D.W.S. by an MBA Senior Research Fellowship.

Efficient searching is crucial for timely location of food and other resources. Recent studies show that diverse living animals use a theoretically optimal scale-free random search for sparse resources known as a Lévy walk, but little is known of the origins and evolution of foraging behavior and the search strategies of extinct organisms. Here, using simulations of self-avoiding trace fossil trails, we show that randomly introduced strophotaxis (U-turns)--initiated by obstructions such as self-trail avoidance or innate cueing--leads to random looping patterns with clustering across increasing scales that is consistent with the presence of Lévy walks. This predicts that optimal Lévy searches may emerge from simple behaviors observed in fossil trails. We then analyzed fossilized trails of benthic marine organisms by using a novel path analysis technique and find the first evidence, to our knowledge, of Lévy-like search strategies in extinct animals. Our results show that simple search behaviors of extinct animals in heterogeneous environments give rise to hierarchically nested Brownian walk clusters that converge to optimal Lévy patterns. Primary productivity collapse and large-scale food scarcity characterizing mass extinctions evident in the fossil record may have triggered adaptation of optimal Lévy-like searches. The findings suggest that Lévy-like behavior has been used by foragers since at least the Eocene but may have a more ancient origin, which might explain recent widespread observations of such patterns among modern taxa.

Understanding how invasive species spread is of particular concern in the current era of globalisation and rapid environmental change. The occurrence of super-diffusive movements within the context of Levy flights has been discussed with respect to particle physics, human movements, microzooplankton, disease spread in global epidemiology and animal foraging behaviour. Super-diffusive movements provide a theoretical explanation for the rapid spread of organisms and disease, but their applicability to empirical data on the historic spread of organisms has rarely been tested. This study focuses on the role of long-distance dispersal in the invasion dynamics of aquatic invasive species across three contrasting areas and spatial scales: open ocean (north-east Atlantic), enclosed sea (Mediterranean) and an island environment (Ireland). Study species included five freshwater plant species, Azolla filiculoides, Elodea canadensis, Lagarosiphon major, Elodea nuttallii and Lemna minuta; and ten species of marine algae, Asparagopsis armata, Antithamnionella elegans, Antithamnionella ternifolia, Codium fragile, Colpomenia peregrina, Caulerpa taxifolia, Dasysiphonia sp., Sargassum muticum, Undaria pinnatifida and Womersleyella setacea. A simulation model is constructed to show the validity of using historical data to reconstruct dispersal kernels. Levy movement patterns similar to those previously observed in humans and wild animals are evident in the re-constructed dispersal pattern of invasive aquatic species. Such patterns may be widespread among invasive species and could be exacerbated by further development of trade networks, human travel and environmental change. These findings have implications for our ability to predict and manage future invasions, and improve our understanding of the potential for spread of organisms including infectious diseases, plant pests and genetically modified organisms.

While evidence for optimal random search patterns, known as Lévy walks, in empirical movement data is mounting for a growing list of taxa spanning motile cells to humans, there is still much debate concerning the theoretical generality of Lévy walk optimisation. Here, using a new and robust simulation environment, we investigate in the most detailed study to date (24×10(6) simulations) the foraging and search efficiencies of 2-D Lévy walks with a range of exponents, target resource distributions and several competing models. We find strong and comprehensive support for the predictions of the Lévy flight foraging hypothesis and in particular for the optimality of inverse square distributions of move step-lengths across a much broader range of resource densities and distributions than previously realised. Further support for the evolutionary advantage of Lévy walk movement patterns is provided by an investigation into the 'feast and famine' effect, with Lévy foragers in heterogeneous environments experiencing fewer long 'famines' than other types of searchers. Therefore overall, optimal Lévy foraging results in more predictable resources in unpredictable environments.

International audience; The decisions animals make about how long to wait between activities can determine the success of diverse behaviours such as foraging, group formation or risk avoidance. Remarkably, for diverse animal species, including humans, spontaneous patterns of waiting times show random 'burstiness' that appears scale-invariant across a broad set of scales. However, a general theory linking this phenomenon across the animal kingdom currently lacks an ecological basis. Here, we demonstrate from tracking the activities of 15 sympatric predator species (cephalopods, sharks, skates and teleosts) under natural and controlled conditions that bursty waiting times are an intrinsic spontaneous behaviour well approximated by heavy-tailed (power-law) models over data ranges up to four orders of magnitude. Scaling exponents quantifying ratios of frequent short to rare very long waits are species-specific, being determined by traits such as foraging mode (active versus ambush predation), body size and prey preference. A stochastic-deterministic decision model reproduced the empirical waiting time scaling and species-specific exponents, indicating that apparently complex scaling can emerge from simple decisions. Results indicate temporal power-law scaling is a behavioural 'rule of thumb' that is tuned to species' ecological traits, implying a common pattern may have naturally evolved that optimizes move-wait decisions in less predictable natural environments.

Summary 1. A first step in the analysis of complex movement data often involves discretisation of the path into a series of step-lengths and turns, for example in the analysis of specialised random walks, such as L evy flights. However, the identification of turning points, and therefore step-lengths, in a tortuous path is dependent onad-hoc parameter choices. Consequently, studies testing for movement patterns in these data, such as L evy flights, have generated debate. However, studies focusing on one-dimensional (1D) data, as in the vertical displacements of marine pelagic predators, where turning points can be identified unambiguously have provided strong support for L evy flight movement patterns. 2. Here, we investigate how step-length distributions in 3D movement patterns would be interpreted by tags recording in 1D (i.e. depth) and demonstrate the dimensional symmetry previously shown mathematically for L evy-flight movements. We test the veracity of this symmetry by simulating several measurement errors common in empirical datasets and find L evy patterns and exponents to be robust to low-quality movement data. 3. We then consider exponential and composite Brownian random walks and show that these also project into 1D with sufficient symmetry to be clearly identifiable as such. 4. By extending the symmetry paradigm, we propose a new methodology for step-length identification in 2D or 3D movement data. The methodology is successfully demonstrated in a re-analysis of wandering albatross Global Positioning System (GPS) location data previously analysed using a complex methodology to determine bird-landing locations as turning points in a L evy walk. For this high-resolution GPS data, we show that there is strong evidence for albatross foraging patterns approximated by truncated L evy flights spanning over 3 5o rders of magnitude. 5. Our simple methodology and freely available software can be used with any 2D or 3D movement data at any scale or resolution and are robust to common empirical measurement errors. The method should find wide applicability in the field of movement ecology spanning the study of motile cells to humans.

Marine protected areas (MPAs) are commonly employed to protect ecosystems from threats like overfishing. Ideally, MPA design should incorporate movement data from multiple target species to ensure sufficient habitat is protected. We used long-term acoustic telemetry and network analysis to determine the fine-scale space use of five shark and one turtle species at a remote atoll in the Seychelles, Indian Ocean, and evaluate the efficacy of a proposed MPA. Results revealed strong, species-specific habitat use in both sharks and turtles, with corresponding variation in MPA use. Defining the MPA's boundary from the edge of the reef flat at low tide instead of the beach at high tide (the current best in Seychelles) significantly increased the MPA's coverage of predator movements by an average of 34%. Informed by these results, the larger MPA was adopted by the Seychelles government, demonstrating how telemetry data can improve shark spatial conservation by affecting policy directly.

It is an open question how animals find food in dynamic natural environments where they possess little or no knowledge of where resources are located. Foraging theory predicts that in environments with sparsely distributed target resources, where forager knowledge about resources’ locations is incomplete, Lévy flight movements optimize the success of random searches. However, the putative success of Lévy foraging has been demonstrated only in model simulations. Here, we use high-temporal-resolution Global Positioning System (GPS) tracking of wandering ( Diomedea exulans ) and black-browed albatrosses ( Thalassarche melanophrys ) with simultaneous recording of prey captures, to show that both species exhibit Lévy and Brownian movement patterns. We find that total prey masses captured by wandering albatrosses during Lévy movements exceed daily energy requirements by nearly fourfold, and approached yields by Brownian movements in other habitats. These results, together with our reanalysis of previously published albatross data, overturn the notion that albatrosses do not exhibit Lévy patterns during foraging, and demonstrate that Lévy flights of predators in dynamic natural environments present a beneficial alternative strategy to simple, spatially intensive behaviors. Our findings add support to the possibility that biological Lévy flight may have naturally evolved as a search strategy in response to sparse resources and scant information.

Dramatic population declines among species of pelagic shark as a result of overfishing have been reported, with some species now at a fraction of their historical biomass. Advanced telemetry techniques enable tracking of spatial dynamics and behaviour, providing fundamental information on habitat preferences of threatened species to aid conservation. We tracked movements of the highest pelagic fisheries by-catch species, the blue shark Prionace glauca, in the North-east Atlantic using pop-off satellite-linked archival tags to determine the degree of space use linked to habitat and to examine vertical niche. Overall, blue sharks moved south-west of tagging sites (English Channel; southern Portugal), exhibiting pronounced site fidelity correlated with localized productive frontal areas, with estimated space-use patterns being significantly different from that of random walks. Tracked female sharks displayed behavioural variability in diel depth preferences, both within and between individuals. Diel depth use ranged from normal DVM (nDVM; dawn descent, dusk ascent), to reverse DVM (rDVM; dawn ascent, dusk descent), to behavioural patterns where no diel differences were apparent. Results showed that blue sharks occupy some of the most productive marine zones for extended periods and structure diel activity patterns across multiple spatio-temporal scales in response to particular habitat types. In so doing, sharks occupied an extraordinarily broad vertical depth range for their size (1.0–2.0 m fork length), from the surface into the bathypelagic realm (max. dive depth, 1160 m). The space-use patterns of blue sharks indicated they spend much of the time in areas where pelagic longlining activities are often highest, and in depth zones where these fisheries particularly target other species, which could account for the rapid declines recently reported for blue sharks in many parts of the world's oceans. Our results provide habitat targets for blue shark conservation that may also be relevant to other pelagic species.

An optimal search theory, the so-called Levy-flight foraging hypothesis1, predicts that predators should adopt search strategies known as Levy flights where prey is sparse and distributed unpredictably, but that Brownian movement is sufficiently efficient for locating abundant prey2, 3, 4. Empirical studies have generated controversy because the accuracy of statistical methods that have been used to identify Levy behaviour has recently been questioned5, 6. Consequently, whether foragers exhibit Levy flights in the wild remains unclear. Crucially, moreover, it has not been tested whether observed movement patterns across natural landscapes having different expected resource distributions conform to the theory’s central predictions. Here we use maximum-likelihood methods to test for Levy patterns in relation to environmental gradients in the largest animal movement data set assembled for this purpose. Strong support was found for Levy search patterns across 14 species of open-ocean predatory fish (sharks, tuna, billfish and ocean sunfish), with some individuals switching between Levy and Brownian movement as they traversed different habitat types. We tested the spatial occurrence of these two principal patterns and found Levy behaviour to be associated with less productive waters (sparser prey) and Brownian movements to be associated with productive shelf or convergence-front habitats (abundant prey). These results are consistent with the Levy-flight foraging hypothesis1, 7, supporting the contention8, 9 that organism search strategies naturally evolved in such a way that they exploit optimal Levy patterns.

Many free-ranging predators have to make foraging decisions with little, if any, knowledge of present resource distribution and availability1. The optimal search strategy they should use to maximize encounter rates with prey in heterogeneous natural environments remains a largely unresolved issue in ecology1, 2, 3. Lévy walks4 are specialized random walks giving rise to fractal movement trajectories that may represent an optimal solution for searching complex landscapes5. However, the adaptive significance of this putative strategy in response to natural prey distributions remains untested6, 7. 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 optimum2. 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 organisms3, from microbes8 to humans9, as a 'rule' that evolved in response to patchy resource distributions.

Satellite tracking of large pelagic fish provides insights on free-ranging behaviour, distributions and population structuring. Up to now, such fish have been tracked remotely using two principal methods: direct positioning of transmitters by Argos polar-orbiting satellites, and satellite relay of tag-derived light-level data for post hoc track reconstruction. Error fields associated with positions determined by these methods range from hundreds of metres to hundreds of kilometres. However, low spatial accuracy of tracks masks important details, such as foraging patterns. Here we use a fast-acquisition global positioning system (Fastloc GPS) tag with remote data retrieval to track long-term movements, in near real time and position accuracy of

Determining the habitat use of mobile marine species is important for understanding responses to climate change and aids the implementation of management and conservation measures. Inference of preferred habitat use has been greatly improved by combining satellite-based oceanographic data with animal tracking techniques. Although there have been several satellite-tracking studies on ocean sunfish Mola mola, limited information is available about either horizontal or vertical environmental preferences. In this study, both geographical movements and diving behaviour of ocean sunfish were explored together with the environmental factors influencing this species’ space use in the north-east Atlantic. Habitat selection of electronic-tagged sunfish (n = 22 individuals; 0.6–1.4 m total length, TL) was investigated using geolocations from Argos-linked and pop-up satellite archival transmitters. Satellite tracking (up to 171 days, mean 66 days) revealed seasonal movements: northward in spring–summer and southward in cooler months. Sunfish spent extended periods in three focal areas, the Gulf of Cadiz, north-east Iberia and the Alboran gyre, which are characterised by the presence of frontal features with elevated primary production. Habitat modelling revealed that sea surface temperature and thermal gradients significantly influenced sunfish distribution. Diving profiles, extending from the surface to a maximum depth of 704 m, revealed different depth-use patterns not linked to geographical region or water column stratification. Overall, a size-related movement pattern was detected with larger individuals (>0.92 m TL) travelling further, exploiting greater depth ranges and spending more time at depth than smaller fish. Ocean sunfish in the north-east Atlantic displayed seasonal movements, primarily driven by thermal preferences, extending into higher latitudes in summer. Moreover, fish also occupied productive frontal areas for long periods, presumably for improved foraging opportunities. Lastly, sunfish showed considerable variability in diving patterns which likely reflect the tracking of planktonic prey distributions.