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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.

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.

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.).

Advances in satellite tracking and archival technologies now allow marine animal movements and behaviour to be recorded at much finer temporal scales, providing a more detailed ecological understanding that can potentially be applicable to conservation and management strategies. Pelagic sharks are commercially exploited worldwide with current concerns that populations are declining, however, how pelagic sharks use exploited environments remains enigmatic for most species. Here we analysed high resolution dive depth profiles of two pelagic shark species with contrasting feeding strategies to investigate movement patterns in relation to environmental heterogeneity. Seven macropredatory blue (Prionace glauca) and six plankton-feeding basking (Cetorhinus maximus) sharks were tagged with pop-off satellite-linked archival tags in the North Atlantic Ocean to examine habitat use and investigate the function of dives. We grouped dives of both species into five major categories based on the two dimensional dive profile shape. Each dive-shape class presented similar frequency and characteristics among the two species with U- and V-shaped dives predominating. We tested the spatial occurrence of different U- and V-shape dive parameters in response to environmental field gradients and found that mean depth and mean depth range decreased with increasing levels of primary productivity (chlorophyll ‘a’), whereas ascent velocities displayed a positive correlation. The results suggest that a planktivore and a macropredator responded behaviourally in similar ways to encountered environmental heterogeneity. This indicates fine-scale dive profiles of shark species with different feeding strategies can be used to identify key marine habitats, such as foraging areas where sharks aggregate and which may represent target areas for conservation.

The use of archival depth telemetry as a means of remotely assessing the reproductive rates of free-ranging fishes is explored. This is achieved by electronically tracking the vertical movements of individual female small spotted catsharks Scyliorhinus canicula in the natural environment, whilst simultaneously evaluating the temporal and vertical distributions of egg-laying in this species. Distinctive patterns of short-term (0·3-3·7 h), shallow-water activity are documented in the time-depth profiles of female S. canicula that occur at an appropriate depth (1·0-2·3 m) and periodicity (every 10-12 days) to represent egg-laying behaviour. Putative egg-laying behaviour was exhibited simultaneously by two individually tracked female S. canicula during late-spring and early-summer. The results highlight that, provided species behaviour is suitable and complementary methods such as previous data, laboratory experiments and field surveys can be used to validate the patterns observed, archival depth telemetry offers an unobtrusive means by which egg production and egg-laying behaviour of free-living fishes can be estimated. As precise information regarding life-history parameters is difficult to obtain for free-ranging fish species, this technique could be used to improve the parameterization of species demographic models that are relevant to the management of wild fish populations.

Patterns of vertical movement in pelagic predators can be highly complex, reflecting behaviours such as foraging, thermoregulatory excursions and spawning. Here we used fast Fourier analysis to identify periodicity in the vertical movements of 6 basking sharks Cetorhinus maximus from archival tagging data that totalled 595 d. We analysed quantitatively fine-scale vertical movements of basking sharks over seasonal scales (May to February) and detected predominant periodicities related to the vertical movements of the sharks' zooplankton prey. Normal and reverse diel vertical migration (DVM) represented the main periodic dive behaviour, occurring for 11 to 72% of individual track times. A tidal pattern of vertical movement, previously unreported for sharks, was also identified. A possible mechanism for this behaviour appears related to the shark exploiting tidally-induced aggregations of zooplankton prey at depth. The youngest shark tagged showed a markedly different pattern of vertical behaviour. Long-term data sets of swimming depth are becom- ing increasingly available for pelagic predators from pressure-sensitive data loggers. This study demonstrates the utility of signal processing techniques in objectively identifying both expected and unexpected periodicity in these continuous, high-resolution tracks.

The basking shark Cetorhinus maximus is listed as Vulnerable (A1a,d, + 2d) worldwide, and Endangered (EN A1a,d) in the north-east Atlantic in the IUCN Red List. However, protection for this species is limited in European waters and varies spatially. Without information on the amount of time individuals spend in different areas, any potential risks to population levels from incidental capture, possible future exploitation and climate change will be difficult to assess. To quantify the annual space-use patterns within political–economic zones in the north-east Atlantic we used geolocation data from seven transmitter-tagged basking sharks tracked for 964 days (16,754 km). Basking sharks tagged within the UK protection zone off south-west England and north-west Scotland spent subsequently only about 22% (range, 2.4–47.7%) of time at liberty within this zone, and a further 30% in the UK fishing zone. Although only about 6% of time was spent in the territorial waters of Ireland and France, basking sharks remained in UK, Irish and French fishing and EEZ zones for over 71% of track time (range, 51.4–89.2%). Sharks did not occupy International waters away from the European shelf at any time. These results indicate basking sharks move between different economic zones and were not afforded statutory protection for the major part of the time (78%) they spent within preferred habitat on the European shelf. This demonstrates the limited capacity of the British protection zone for encompassing the greater part of shark space utilisation. Tracked basking sharks regularly crossed national zone boundaries suggesting that conservation measures for this species need to be framed on a European, rather than national, basis. This study highlights the need for better information about the movements and habitat use by marine animals if conservation strategies are to be truly effective.

Movements of wide-ranging top predators can now be studied effectively using satellite and archival telemetry. However, the motivations underlying movements remain difficult to determine because trajectories are seldom related to key biological gradients, such as changing prey distributions. Here, we use a dynamic prey landscape of zooplankton biomass in the north-east Atlantic Ocean to examine active habitat selection in the plankton-feeding basking shark Cetorhinus maximus . The relative success of shark searches across this landscape was examined by comparing prey biomass encountered by sharks with encounters by random-walk simulations of ‘model’ sharks. Movements of transmitter-tagged sharks monitored for 964 days (16 754 km estimated minimum distance) were concentrated on the European continental shelf in areas characterized by high seasonal productivity and complex prey distributions. We show movements by adult and sub-adult sharks yielded consistently higher prey encounter rates than 90% of random-walk simulations. Behavioural patterns were consistent with basking sharks using search tactics structured across multiple scales to exploit the richest prey areas available in preferred habitats. Simple behavioural rules based on learned responses to previously encountered prey distributions may explain the high performances. This study highlights how dynamic prey landscapes enable active habitat selection in large predators to be investigated from a trophic perspective, an approach that may inform conservation by identifying critical habitat of vulnerable species.

current concerns about the population levels of the basking shark (cetorhinus maximus) in the north-east atlantic have prompted a need to understand population distribution, habitat preference and centres of abundance. in this study, spatial distribution maps derived from satellite-tag geolocations, boat surveys and public sightings data were compared. the broad distribution patterns revealed by these different methods are similar, but there are considerable differences in density distributions. surface sightings data show high densities, or ‘hotspots’ in the hebridean sea, clyde sea, irish sea and close inshore around devon and cornwall. tag geolocations, in contrast, identified two areas where individuals spent considerable time outside the distributions indicated by surveys and public sightings: the celtic sea and western approaches of the english channel. the reason for this disparity and its implications for population estimates for the species are discussed.

the tendency for marine fish to refuge in natural and artificial structures underwater is well documented, but the incidence of this behaviour among predatory sharks is not widely known. a sub-adult male nursehound scyliorhinus stellaris was tracked by intermittent acoustic telemetry over 168 days in a tidal sea lough. this individual, in-between undertaking nocturnal foraging excursions, refuged in at least five different narrow-entrance holes. refuge location was validated by underwater and surface observations during which time other nursehounds (sub-adult male and female), were also seen refuging, sometimes together. this indicates nursehound display philopatric behaviour centred on aggregation in ‘home’ refuges that, in this case, were labyrinthine rock systems.

Summary 1. Megaplanktivores such as filter-feeding sharks and baleen whales are at the apex of a short food chain (phytoplankton‐zooplankton‐vertebrate) and are sensitive indicators of sea-surface plankton availability. Even though they spend the majority of their time below the surface it is still not known how most of these species utilize vertical habitat and adapt to short-term changes in food availability. 2. A key factor likely to control vertical habitat selection by planktivorous sharks is the diel vertical migration (DVM) of zooplankton; however, no study has determined whether specific ocean-habitat type influences their behavioural strategy. Based on the first high-resolution dive data collected for a plankton-feeding fish species we show that DVM patterns of the basking shark Cetorhinus maximus reflect habitat type and zooplankton behaviour. 3. In deep, well-stratified waters sharks exhibited normal DVM (dusk ascent‐dawn descent) by tracking migrating sound-scattering layers characterized by Calanus and euphausiids. Sharks occupying shallow, inner-shelf areas near thermal fronts conducted reverse DVM (dusk descent‐dawn ascent) possibly due to zooplankton predator‐prey interactions that resulted in reverse DVM of Calanus . 4. These opposite DVM patterns resulted in the probability of daytime-surface sighting differing between these habitats by as much as two orders of magnitude. Ship-borne surveys undertaken at the same time as trackings reflected these behavioural differences. 5. The tendency of basking sharks to feed or rest for long periods at the surface has made them vulnerable to harpoon fisheries. Ship-borne and aerial surveys also use surface occurrence to assess distribution and abundance for conservation purposes. Our study indicates that without bias reduction for habitat-specific DVM patterns, current surveys could under- or overestimate shark abundance by at least 10-fold.

Habitat selection processes in highly migratory animals such as sharks and whales are important to understand because they influence patterns of distribution, availability and therefore catch rates. However, spatial strategies remain poorly understood over seasonal scales in most species, including, most notably, the plankton-feeding basking shark Cetorhinus maximus. It was proposed nearly 50 yr ago that this globally distributed species migrates from coastal summer- feeding areas of the northeast Atlantic to hibernate during winter in deep water on the bottom of con- tinental-shelf slopes. This view has perpetuated in the literature even though the 'hibernation theory' has not been tested directly. We have now tracked basking sharks for the first time over seasonal scales (1.7 to 6.5 mo) using 'pop-up' satellite archival transmitters. We show that they do not hiber- nate during winter but instead undertake extensive horizontal (up to 3400 km) and vertical (>750 m depth) movements to utilise productive continental-shelf and shelf-edge habitats during summer, autumn and winter. They travel long distances (390 to 460 km) to locate temporally discrete produc- tivity 'hotspots' at shelf-break fronts, but at no time were prolonged movements into open-ocean regions away from shelf waters observed. Basking sharks have a very broad vertical diving range and can dive beyond the known range of planktivorous whales. Our study suggests this species can exploit shelf and slope-associated zooplankton communities in mesopelagic (200 to 1000 m) as well as epipelagic habitat (0 to 200 m).

for many decades, scientists at the marine biological association (mba) in plymouth and the centre for environment, fisheries and aquaculture science (cefas), formerly the directorate of fisheries research, at lowestoft have undertaken considerable research into the ecology and life histories of elasmobranch fish around the british isles.

The first confirmed record of a smooth hammerhead shark (Sphyrna zygaena) in British waters for 140 years is described. The head section of the specimen was found washed ashore in relatively good condition at Portreath, south-west England, in December 2004. Evidence suggests it was not a fishery discard, and, like most other previous records of this species in British waters, probably came ashore moribund or dead.

Dermal denticles are unique tooth-like structures embedded in the skin of sharks and rays that protect them from predators and ectoparasites, reduce mechanical abrasion and possibly minimize swimming-induced drag. Here, we show that juvenile lesser spotted dogfish (Scyliorhinus canicula) also use this body armour to anchor food items near their tail so that bite-sized pieces can be torn away by rapid jaw and head movements. This scale-rasp behaviour is novel among fishes and suggests a new role for skin in the feeding ecology of sharks. Scale rasping may be important ecologically because it could function to increase the dietary breadth and growth potential of juveniles.