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Understanding why sharks bite humans is essential for developing strategies to prevent these incidents. Here, we use bite wound characteristics and eye witness descriptions of shark behavior to determine the likely motivation for several bites perpetrated by an oceanic whitetip (OWT) shark Carcharhinus longimanus on an adult female snorkeler off Moorea island (French Polynesia) in October 2019. The victim was snorkeling with others in pelagic waters as part of an organized whale-watching tour when the shark-without any warning behavior-bit her at least three times resulting in severe injuries with substantial loss of soft tissue from the chest and both forearms. The victim survived these injuries thanks to rapid and effective first aid provided by her companions. The sudden, unprovoked and repeated bites with substantial tissue removal are consistent with predatory behavior although the dominance hypothesis cannot be fully ruled out. This would be the first case of a predatory shark bite ever documented in French Polynesia in over 70 years of data collection. Given the routine association of OWT sharks with cetaceans, in-water whale watching activities should adopt appropriate risk management strategies in regions hosting this species of shark.

20 pages, 3 tables, 5 figures.-- Samantha Andrzejaczek ... et al.-- Distributed under a Creative Commons Attribution NonCommercial License 4.0 (CC BY-NC), Knowledge of the three-dimensional movement patterns of elasmobranchs is vital to understand their ecological roles and exposure to anthropogenic pressures. To date, comparative studies among species at global scales have mostly focused on horizontal movements. Our study addresses the knowledge gap of vertical movements by compiling the first global synthesis of vertical habitat use by elasmobranchs from data obtained by deployment of 989 biotelemetry tags on 38 elasmobranch species. Elasmobranchs displayed high intra- and interspecific variability in vertical movement patterns. Substantial vertical overlap was observed for many epipelagic elasmobranchs, indicating an increased likelihood to display spatial overlap, biologically interact, and share similar risk to anthropogenic threats that vary on a vertical gradient. We highlight the critical next steps toward incorporating vertical movement into global management and monitoring strategies for elasmobranchs, emphasizing the need to address geographic and taxonomic biases in deployments and to concurrently consider both horizontal and vertical movements, Data analysis was funded by the Bertarelli Foundation through the Marine Science program through grants to D.J.C., B.A.B., and S.A. D.J.C. is also funded through Research England, UK. S.A. and B.A.B. thank the Moore Foundation and the Packard Foundation. F.G.-M. thanks the Instituto Politecnico Nacional for fellowships (COFAA, EDI). S.B.W. thanks funding from the Darwin Initiative (DPLUS046). A.D.M.D. acknowledges funding from the Research and Conservation Budget at Georgia Aquarium, including philanthropic gifts from several anonymous donors. K.F. acknowledges funding from the Rolex Awards for Enterprise and the Whitley Fund for Nature

BackgroundGreat hammerhead sharks (Sphyrna mokarran) routinely swim on their sides and periodically roll from side to side. A previous study used wind tunnel tests with a rigid model hammerhead shark to demonstrate that the rolling behavior could improve swimming efficiency using the tall first dorsal fin as a lift-generating surface. Scalloped hammerhead sharks (Sphyrna lewini) also have proportionally taller dorsal fins compared to pectoral fins than most shark species and similar to that of great hammerhead sharks, and thus might exhibit similar rolling behavior. This was assessed by deploying multi-sensor accelerometer instrument packages on free-swimming adult scalloped hammerhead sharks to directly measure swimming depth, body orientation and swimming performance. Specific objectives were to (1) determine whether scalloped hammerhead sharks exhibit side swimming and rolling behavior, (2) characterize the patterns of these behaviors, and (3) evaluate the purpose of these behaviors.ResultsWe obtained 196.7 total days (4720 h) of data from 9 free-swimming adult scalloped hammerhead sharks equipped with multi-instrument biologgers with deployment durations ranging from 7 to 29 days. All sharks exhibited rolling behavior throughout the entire period of observation. The roll angle magnitude and periodicity of rolling showed a clear diel pattern. During daytime, the sharks spent an average of 48% of the time swimming at a roll angle > 30°, with an average roll angle of 41° and rolling periodicity of around 4 min. At night, the sharks spent an average 82% of their time at an angle > 30°, with an average roll angle of 60° and rolling periodicity of around 13 min. In addition to an increase in degree of roll and roll duration, overall dynamic body acceleration (ODBA) also increased at night, and tailbeat frequency was more regular and consistent than during daytime.ConclusionWe observed rolling behavior in scalloped hammerhead sharks similar to that observed in great hammerhead sharks. The diel changes in roll angle and periodicity were accompanied by other changes in swimming behavior. These changes are possibly due to interplay between reducing cost of transport and social interactions with conspecifics.

Acoustic telemetry is widely used to investigate aquatic animal movement. Pulse position modulation (PPM) is an acoustic telemetry method that allows multiple unique identification codes to be transmitted at a single acoustic frequency, typically in the 69 kHz range. However, because the potential number of unique identification codes (i.e. tags) is ultimately limited by the number of pulses in the PPM signal, this poses a practical limitation. In addition, different manufacturers have developed different approaches to encoding the transmitted data, hampering compatibility across brands. A lack of broad compatibility across telemetry systems restricts users to a single manufacturer and operating system, reduces market competition and limits innovation. As the aquatic animal tracking research community organises towards networks of devices and data, incompatibility becomes more problematic and jeopardizes the unique scientific benefits offered by the networking approach. Here, we make a plea for collaboration among the manufacturers globally and propose a set of open protocols to ensure equipment interoperability as a medium-term solution.

DNA evidence is routinely used to identify individual predators responsible for attacks on people and livestock in terrestrial settings. However, the use of transfer DNA techniques in aquatic environments20 for similar purposes is a recent development. To date, DNA barcoding has been used successfully to identify shark species depredating fish catches and biting surfboards and neoprene surfaces. In this study we demonstrate the successful DNA barcoding and fingerprinting of individual sharks from transfer DNA collected directly from the wounds of two shark bite victims. The successful use of DNA techniques to identify both species and specific individuals responsible for shark bites opens the door to selective removal of these individuals as an innovative shark bite risk management strategy. This selective approach would be a more effective, eco-responsible, cost-effective and ethical solution for vulnerable taxa than ongoing non-selective culling campaigns.

Identifying the species involved in shark bite incidents is an ongoing challenge but is important to mitigate risk. We developed a sampling protocol to identify shark species from DNA transferred to inanimate objects during bite incidents. To develop and refine the technique, we swabbed shark bite impressions on surfboards and wetsuit neoprene collected under semicontrolled conditions. Methods were tested experimentally and then successfully used to identify the species involved in a real-world shark bite incident. Thirty-two of 33 bite impressions yielded sufficient DNA sequences for species identification, producing barcodes from five test species, including dusky, Galapagos, bull, tiger, and white shark. The latter three species collectively account for a majority of shark bites worldwide. Our method successfully identified the species (Galeocerdo cuvier) responsible for a fatal shark bite on December 8th, 2020 on the island of Maui, from swab samples collected from the victim's surfboard 49 h after the bite incident. Our experimental results demonstrate that shark species can be accurately identified from transfer DNA recovered from bite impressions on surfboards and wetsuit neoprene. The successful use of our method in the real-world incident shows great potential for the practicality of this tool. We recommend DNA swabbing as a routine part of the forensic analysis of shark bites to help identify the species involved in human-shark interactions.

International audience; Advances in biologging technology have enabled 3D dead-reckoning reconstruction of marine animal movements at spatiotemporal scales of meters and seconds. Examining high-resolution 3D movements of sharks (Galeocerdo cuvier, N = 4; Rhincodon typus, N = 1), sea turtles (Chelonia mydas, N = 3), penguins (Aptenodytes patagonicus, N = 6), and marine mammals (Arctocephalus gazella, N = 4; Ziphius cavirostris, N = 1), we report the discovery of circling events where animals consecutively circled more than twice at relatively constant angular speeds. Similar circling behaviors were observed across a wide variety of marine megafauna, suggesting these behaviors might serve several similar purposes across taxa including foraging, social interactions, and navigation

International audience; We used long-term observations of 55 individually identified tiger sharks (Galeocerdo cuvier) at a shark eco-tourism site off Tahiti to estimate the prevalence, retention times and impacts of residual hooks and trailing line resulting from fisheries interactions. Thirty-eight percent of tiger sharks had at least one fisheries interaction resulting in retained hooks during the 8-year observation period. Both stainless-steel and corrodible hooks had a retention half-life of less than 1year, and all corrodible hooks were shed within 2.5 years whereas firmly embedded stainless-steel hooks persisted for at least 7.6 years and are potentially retained for the lifetime of the shark. The presence of residual hooks and trailing line did not impact tiger shark growth suggesting that sharks were able to feed normally despite trailing gear. However, only a single internally-hooked sharks trailing line was observed suggesting their survival may perhaps be lower than externally-hooked individuals. Hook-retention data obtained from this study could be extrapolated to other warm water shark species that are difficult to observe in their natural habitats (e.g. oceanic whitetip shark, Carcharhinus longimanus).

The application of genome-wide cytonuclear molecular data to identify management and adaptive units at various spatio-temporal levels is particularly important for overharvested large predatory organisms, often characterized by smaller, localized populations. Despite being “near threatened”, current understanding of habitat use and population structure of Carcharhinus galapagensis is limited to specific areas within its distribution. We evaluated population structure and connectivity across the Pacific Ocean using genome-wide single-nucleotide polymorphisms (~7200 SNPs) and mitochondrial control region sequences (945 bp) for 229 individuals. Neutral SNPs defined at least two genetically discrete geographic groups: an East Tropical Pacific (Mexico, east and west Galapagos Islands), and another central-west Pacific (Lord Howe Island, Middleton Reef, Norfolk Island, Elizabeth Reef, Kermadec, Hawaii and Southern Africa). More fine-grade population structure was suggested using outlier SNPs: west Pacific, Hawaii, Mexico, and Galapagos. Consistently, mtDNA pairwise Φ(ST) defined three regional stocks: east, central and west Pacific. Compared to neutral SNPs (F(ST) = 0.023–0.035), mtDNA exhibited more divergence (Φ(ST) = 0.258–0.539) and high overall genetic diversity (h = 0.794 ± 0.014; π = 0.004 ± 0.000), consistent with the longstanding eastern Pacific barrier between the east and central–west Pacific. Hawaiian and Southern African populations group within the west Pacific cluster. Effective population sizes were moderate/high for east/west populations (738 and 3421, respectively). Insights into the biology, connectivity, genetic diversity, and population demographics informs for improved conservation of this species, by delineating three to four conservation units across their Pacific distribution. Implementing such conservation management may be challenging, but is necessary to achieve long-term population resilience at basin and regional scales.

In aquatic environments, what one observes during the day can differ substantially by night. The species composition and associated ecological processes that occur during the day are often different than night. In polar seas and at great depths, "night" can span, months, years, and beyond. Teleosts and elasmobranchs have evolved unique sensory and behavioral modalities for living in darkness. As a consequence, fishers have adopted unique strategies for exploiting fish at night or in dark systems. We propose that neglecting the night has led to an incomplete understanding of aquatic organismal ecology, population/community dynamics, and ecosystem function with consequences for fisheries conservation management. To address this knowledge gap and stimulate the exchange of new data and ideas on behaviors, patterns, and processes relating to fish and fisheries in darkness, Fish at Night: an international symposium was held in Miami, Florida (USA), from 18 to 20 November, 2015. Here, we synthesize the findings from the symposium, providing an overview on the state-of-knowledge of fish studies in the dark, identifying critical information gaps, and charting a course for future research. We focus our commentary and synthesis on six areas: (1) nocturnal fish behavior and ecology; (2) fishing, fisheries, and enforcement; (3) deep and polar seas; (4) diel fish distribution and abundance comparisons; (5) methods for studying fish in darkness; (6) human threats to fish at night; and (7) larval fish at night. Taken together, we attempt to "shine a light" on fish at night, generating a greater interest and understanding of fishes and fisheries during darkness.

The past six decades have seen the emergence of new electronic tag technologies enabling scientists to remotely study the behavior of fishes in a wide variety of aquatic habitats. This revolution began in the 1950s and 1960s with the first studies utilizing acoustic and radio transmitters to actively track fish movements. Subsequent decades saw the development of passive monitoring systems enabling researchers to scale up the number of individuals followed, and the duration and spatial extent of tracking studies. Recent decades also have witnessed the increasingly-widespread use of sophisticated satellite transmitters to quantify fish movements in remote parts of the ocean beyond the range of fixed listening arrays. Most recently, high-frequency sampling devices incorporating accelerometers and other novel sensors have given us unparalleled new and detailed insights into fish behavior. All of these technologies have yielded important insights into fish spatial behaviors, and shown that many species are most active at night, or in deep waters, but we need better ground-truthing to reveal the full context and purpose of these spatial patterns. Recent advances in animal-mounted cameras have provided important ground-truthing breakthroughs, allowing us to see behavior and habitat use from the fishes' own perspective. However, current generations of animal-mounted cameras are reliant on natural light, or use artificial light overlapping the visual spectrum of most fishes. To see what fishes are doing at night, we need to develop new animal-mounted cameras using far red spectrum light, or high-frequency sound, to illuminate the view field without impacting natural behavior.

International audience; Selective removal of problem individuals following shark bite incidents would be consistent with current management practices for terrestrial predators, and would be more effective and more environmentally responsible than current mass-culling programs. In parallel, and in addition to traditional forensics analysis, we recommend the routine collection of shark DNA from wounds or devices following shark bite incidents in order to genetically identify the individual responsible. This approach would require creating an extensive database of shark identities in high-risk areas against which to compare DNA forensically recovered from shark bite incidents. At a local and regional scale, we propose utilizing existing shark tagging programs and artificial sharkaggregation sites to collect DNA, behavioural and morphological data for the database, and to facilitate removal of problem individuals. In several places around the world, selective removal of problem individuals would not be significantly more expensive and definitely less environmentally-destructive than traditional approaches and would also help reconcile people and sharks by underlining individuality in shark behaviour.

This “Perspectives” paper identifies aspects of tiger shark (Galeocerdo cuvier) biology that are currently unknown or for which additional data are needed to improve interpretive power. Some of these data gaps may be regional. Technical or methodological approaches to acquiring these data are suggested. Some of these technologies already exist, some are in development and some exist in concept only. Reproductive biology and behavior, social interactions and the behavioral ecology of sub-adults are among the areas identified as deserving of future research effort.

We compared tiger shark (Galeocerdo cuvier) spatial behaviour among 4 Hawaiian Islands to evaluate whether local patterns of movement could explain higher numbers of shark bites seen around Maui than other islands. Our sample consisted of 96 electronically-tagged (satellite and acoustic transmitters) tiger sharks, individually tracked for up to 6 years. Most individuals showed fidelity to a specific ‘home’ island, but also swam between islands and sometimes ranged far (up to 1,400 km) offshore. Movements were primarily oriented to insular shelf habitat (0–200 m depth) in coastal waters, and individual sharks utilized core-structured home ranges within this habitat. Core utilization areas of large tiger sharks were closer to high-use ocean recreation sites around Maui, than around Oahu. Tiger sharks routinely visited shallow ocean recreation sites around Maui and were detected on more days overall at ocean recreation sites around Maui (62–80%) than Oahu (

Aim Broadly distributed reef fishes tend to have high gene flow mediated by a pelagic larval phase. Here, we survey a reef-associated fish distributed across half the tropical oceans, from the Red Sea to the central Pacific. Our goal is to determine whether genetic structure of the broadly distributed Yellowstripe Goatfish (Mulloidichthys flavolineatus) is defined by biogeographical barriers, or facilitated via larval dispersal. Location Red Sea, Indian Ocean, Pacific Ocean Methods Specimens were obtained at 19 locations from the Red Sea to Hawai'i. Genetic data include mtDNA cytochrome b (n = 217) and 12 microsatellite loci (n = 185). Analysis of molecular variance (AMOVA), structure, a parsimony network and coalescence analyses were used to resolve recent population history and connectivity. Results Population structure was significant (mtDNA ϕST = 0.68, P

Marine predators will often perform diel and seasonal movements associated with specific habitats. In tropical areas, mesophotic coral reefs may be an important habitat type for many predators but their use of these areas has rarely been investigated. We used results of acoustic telemetry and stable isotope analyses to investigate the diel and seasonal movements of Galapagos sharks Carcharhinus galapagensis and giant trevally Caranx ignobilis captured from a mesophotic reef (depth: 50–70 m) at an uninhabited Pacific atoll. All predators performed horizontal and vertical movements over seasonal and diel time frames associated with mesophotic reefs. Galapagos sharks performed reverse diel vertical movements diving deeper during the night than during the day, while giant trevally displayed a mix, with some individuals performing regular diel movements (deep during the day, shallow at night) while others performed reverse vertical diel movements. Trevally used very shallow water during the spawning periods in the summer. The isotopic compositions of predators suggest they primarily forage in shallow reefs, although approximately 35% of resources came from mesophotic reefs. Similar to variability in vertical movement strategies, giant trevally occupied a wide range of trophic positions, potentially due to individual specialization in diet and high levels of intra-specific competition. Mesophotic reefs provide some prey to upper level predators but may primarily serve as a refuge habitat. The frequent movements between habitats suggest that marine predators may function as significant transporters of nutrients from shallow to mesophotic reefs. Publisher PDF

Blooms of alien invasive marine algae have become common, greatly altering the health and stability of nearshore marine ecosystems. Concurrently, herbivorous fishes have been severely overfished in many locations worldwide, contributing to increases in macroalgal cover. We used a multi-pronged, interdisciplinary approach to test if higher biomass of herbivorous fishes inside a no-take marine reserve makes this area more resistant to invasive algal overgrowth. Over a two year time period, we (1) compared fish biomass and algal cover between two fished and one unfished patch reef in Hawai’i, (2) used acoustic telemetry to determine fidelity of herbivorous fishes to the unfished reef, and (3) used metabarcoding and next-generation sequencing to determine diet composition of herbivorous fishes. Herbivore fish biomass was significantly higher in the marine reserve compared to adjacent fished reefs, whereas invasive algal cover differed by species. Herbivorous fish movements were largely confined to the unfished patch reef where they were captured. Diet analysis indicated that the consumption of invasive algae varied among fish species, with a high prevalence of comparatively rare native algal species. Together these findings demonstrate that the contribution of herbivores to coral reef resilience, via resistance to invasive algae invasion, is complex and species-specific.

The redistribution of species has emerged as one of the most pervasive impacts of anthropogenic climate warming, and presents many societal challenges. Understanding how temperature regulates species distributions is particularly important for mobile marine fauna such as sharks given their seemingly rapid responses to warming, and the socio-political implications of human encounters with some dangerous species. The predictability of species distributions can potentially be improved by accounting for temperature's influence on performance, an elusive relationship for most large animals. We combined multi-decadal catch data and bio-logging to show that coastal abundance and swimming performance of tiger sharks Galeocerdo cuvier are both highest at ~22°C, suggesting thermal constraints on performance may regulate this species' distribution. Tiger sharks are responsible for a large proportion of shark bites on humans, and a focus of controversial control measures in several countries. The combination of distribution and performance data moves towards a mechanistic understanding of tiger shark's thermal niche, and delivers a simple yet powerful indicator for predicting the location and timing of their occurrences throughout coastlines. For example, tiger sharks are mostly caught at Australia's popular New South Wales beaches (i.e. near Sydney) in the warmest months, but our data suggest similar abundances will occur in winter and summer if annual sea surface temperatures increase by a further 1-2°C.

Diel vertical migration is a widespread behavioral phenomenon where organisms migrate through the water column and may modify behavior relative to changing environmental conditions based on physiological tolerances. Here, we combined a novel suite of biologging technologies to examine the thermal physiology (intramuscular temperature), fine-scale swimming behavior and activity (overall dynamic body acceleration as a proxy for energy expenditure) of bluntnose sixgill sharks (Hexanchus griseus) in response to environmental changes (depth, water temperature, dissolved oxygen) experienced during diel vertical migrations. In the subtropical waters off Hawai‘i, sixgill sharks undertook pronounced diel vertical migrations and spent considerable amounts of time in cold (5–7°C), low oxygen conditions (10–25% saturation) during their deeper daytime distribution. Further, sixgill sharks spent the majority of their deeper daytime distribution with intramuscular temperatures warmer than ambient water temperatures, thereby providing them with a significant thermal advantage over non-vertically migrating and smaller-sized prey. Sixgill sharks exhibited relatively high rates of activity during both shallow (night) and deep (day) phases and contrary to our predictions, did not reduce activity levels during their deeper daytime distribution while experiencing low temperature and dissolved oxygen levels. This demonstrates an ability to tolerate the low oxygen conditions occurring within the local oxygen minimum zone. The novel combination of biologging technologies used here enabled innovative in situ deep-sea natural experiments and provided significant insight into the behavioral and physiological ecology of an ecologically important deepwater species.

It has been hypothesized that protecting herbivorous fishes within Marine Reserves (MRs) will help these areas to resist algal overgrowth of corals. However, we lack empirical studies demonstrating the validity of key assumptions underpinning this concept, including that herbivorous fishes (1) are permanently resident within MR boundaries, (2) routinely consume the invasive algae of concern at significant levels and (3) are not significant mobile vectors for propagules of the invasive algae. A 46-year-old MR (Hawaii Marine Laboratory Refuge, 21°27′35″N, 157°48′15″W) in waters off Oahu provided a natural setting to examine these factors with the herbivorous bluespine unicornfish (Naso unicornis) and the invasive rhodophyte, Gracilaria salicornia. We acoustically tracked five unicornfish to quantify their residency and habitat use within the MR, conducted field and laboratory observations to confirm N. unicornis consume G. salicornia, and tested the viability of G. salicornia fragments recovered from unicornfish feces. Unicornfish were resident within the MR where they spent most time in reef crest habitat with occasional, brief forays into reef flat habitat. We confirmed unicornfish consume G. salicornia and found a significant positive correlation between algal canopy height and distance from the reef crest, presumably because grazing intensity is lower in less frequently utilized reef flat habitat. We demonstrated that unicornfish egest viable fragments of G. salicornia that resume vegetative growth after several weeks. We conclude that N. unicornis may act as both a natural control agent and a mobile vector for invasive alien algae.

We monitored the long-term residency of reef-associated ballan wrasse and sand-dwelling rays captured at the site of a potential future Marine Protected Area (MPA: Portelet Bay, Jersey) by implanting them with small transmitters and deploying underwater receivers inside the bay. Individual fish were detected at Portelet Bay for up to 618 days, but there were species-specific differences in residency and detection patterns. Ballan wrasse were year-round residents at the study site where they exhibited distinct, rhythmic, diel, tidal and seasonal patterns of behaviour, whereas rays were occasional visitors to Portelet Bay with no discernible pattern to their visits. Results indicate relatively small MPAs (2) that with suitable habitat could provide effective, long-term protection for ballan wrasse, but would likely be of little conservation benefit for rays. Our findings emphasize the importance of quantifying fish movements when planning MPAs which intend to protect multi-species assemblages of coastal fishes.

Sharks zigzag vertically through the water in a series of alternating ascending and descending segments, changing depth by a few tens of meters over a period of a few hundred seconds. This 'yo-yo' like behavior has several characteristic patterns, identifiable by the way the swimming and vertical velocities vary along the dive. We suggest that these patterns represent different optimal strategies minimizing the cost of locomotion under different constraints; moreover, these constraints can be inferred by matching the pattern of a dive with a (standard) optimal swimming strategy for which the constraints are known. We used three sets of constraints and two definitions of the 'cost of locomotion' to analytically generate four standard optimal strategies; we have used high resolution tracking data from four tiger sharks to identify two different yo-yo diving patterns. These patterns seem to match two of the standard strategies: one that maximizes range, given an alternating power supply (e.g., swimming actively on ascents and lazily on descents); and the other that maximizes range, given an alternating vertical velocity (implying an 'intentional' up-and-down motion).

Summary 1. Animal search patterns reflect sensory perception ranges combined with memory and knowledge of the surrounding environment. 2. Random walks are used when the locations of resources are unknown, whereas directed walks should be optimal when the location of favourable habitats is known. However, directed walks have been quantified for very few species. 3. We re-analysed tracking data from three shark species to determine whether they were using directed walks, and if so, over which spatial scales. Fractal analysis was used to quantify how movement structure varied with spatial scale and determine whether the sharks were using patches. 4. Tiger sharks performed directed walks at large spatial scales (at least 6–8 km). Thresher sharks also showed directed movement (at scales of 400–1900 m), and adult threshers were able to orient at greater scales than juveniles, which may suggest that learning improves the ability to perform directed walks. Blacktip reef sharks had small home ranges, high site fidelity and showed no evidence of oriented movements at large scales. 5. There were inter- and intraspecific differences in path structure and patch size, although most individuals showed scale-dependent movements. Furthermore, some individuals of each species performed movements similar to a correlated random walk. 6. Sharks can perform directed walks over large spatial scales, with scales of movements reflecting site fidelity and home range size. Understanding when and where directed walks occur is crucial for developing more accurate population-level dispersal models.

Many pelagic fishes exhibit ‘yo-yo’ diving behavior, which may serve several possible functions, including energy conservation, prey searching and navigation. We deployed accelerometers and digital still cameras on 4 free-ranging tiger sharks Galeocerdo cuvier, to test whether their vertical movements are most consistent with energy conservation or prey searching. All sharks swam continuously, with frequent vertical movements through the water column at mean swimming speeds of 0.5 to 0.9 m s–1. Tail-beating was continuous except for sporadic, powerless gliding during descents (from 0 to 18% of total descent time). At a given tailbeat frequency, swimming speeds were lower during ascent than descent (consistent with negative buoyancy). Burst swimming events, which might represent prey pursuits, were observed during all phases of vertical movements. Camera images showed a variety of potential prey and the possible capture of a unicornfish. Collectively, results suggest that yo-yo diving by tiger sharks is not primarily for energy conservation, but probably represents an effective search strategy for locating prey throughout the water column.

The main Hawaiian Islands (MHI) are the principal breeding ground for humpback whales in the North Pacific. Over the past 3 decades, population recovery from whaling-era losses has resulted in a steady increase in the number of whales wintering in Hawaiian waters and a geo- graphic expansion of their distribution in the MHI. Until recently, no existing evidence showed that this expansion included the islands, atolls, and banks of the Northwestern Hawaiian Islands (NWHI). To better understand the occurrence of humpback whales in the NWHI, 9 ecological acoustic recorders (EARs) were deployed at sites throughout the archipelago to record the occurrence of humpback whale song, an indicator of winter breeding activity. Song was found to be prevalent at Maro Reef, Lisianski Island, and French Frigate Shoals but was also recorded at Kure Atoll, Midway Atoll, and Pearl and Hermes Atoll. Both the timing and abundance of song at several locations closely followed trends observed on Oahu, which is one of the MHI, strongly suggesting that humpback whales use the NWHI as a wintering area. This finding is of particular relevance in light of recent suggestions that, based on photo-identification data, a yet undocumented wintering area exists somewhere in the central North Pacific. We propose that the NWHI could be that area.

We equipped individual tiger (Galeocerdo cuvier Peron and Lesueur, 1822) and Galapagos (Carcharhinus galapagensis Snodgrass and Heller, 1905) sharks with both acoustic and satellite transmitters to quantify their long-term movements in the Papahanaumokuakea Marine National Monument (Northwestern Hawaiian Islands). Tiger sharks exhibited two broad patterns of behavior. Some individuals were detected at French Frigate Shoals (FFS) year round, whereas others visited FFS atoll in summer to forage on fledging albatross, then swam thousands of kilometers along the Hawaiian chain, or out into open ocean to the North Pacific transition zone chlorophyll front, before returning to FFS in subsequent years. These patterns suggest tiger sharks may use cognitive maps to navigate between distant foraging areas. Different patterns of spatial behavior may arise because cognitive maps are built up through individual exploration, and each tiger shark learns a unique combination of foraging sites. Galapagos shark detections were all associated with FFS, suggesting these sharks may be more resident around oceanic islands. Both Galapagos and tiger sharks primarily used the mixed layer (

We tracked the long-term movements of 70 parrotfishes, surgeonfishes and goatfishes captured inside a small (1.3 km2) marine protected area (MPA: Kealakekua Bay Marine Life Conservation District, Hawaii) by implanting them with small transmitters and deploying underwater monitoring devices inside the bay and along 100 km of the adjacent west Hawaii coastline. Individual fish were detected inside Kealakekua Bay for up to 612 days but many were detected for much shorter periods (median = 52 days). There were species-specific differences in the scale of movements and habitats used, but most fish utilized between 0.2 and 1.6 km of coastline, and individuals of each species showed some degree of diel habitat shift. A wide variety of reef fishes captured inside the MPA swam back and forth across an MPA boundary intersecting continuous reef (i.e., this boundary was porous to reef fish movements), but only 1 of 11 species tagged crossed a wide sandy channel inside Kealakekua Bay suggesting that this feature may function as a natural barrier to movements. Results indicate relatively small MPAs (

Prototype acoustic 'business card' (BC) tags were deployed on 4 free-swimming Gala- pagos sharks Carcharhinus galapagensis associated with a shark ecotourism operation near Haleiwa, Hawaii, USA. These transmitter/receiver tags employed mobile peer-to-peer (MP2P) technology that allowed the tagged sharks to exchange codes among each other and to detect other sharks implanted with standard one-way coded acoustic transmitters. Two tags were recovered (after 20 and 132 d); both tags had multiple detections of all other BC tags, and a comparison of detections made by these tags to those made by a fixed array of standard VR2 receivers indicated that the BC tags accurately captured the 'presence-absence' patterns of the other tagged sharks. Importantly, the BC tags detected sharks that were beyond the range of the fixed receiver array. The results indicate that the BC tag/MP2P approach can elucidate important inter-and intra-specific interactions among individ- uals in areas remote from traditional fixed receiver arrays.

SUMMARYShark cage diving is both popular and controversial, with proponents citing educational value and non-extractive use of natural resources and opponents raising concerns about public safety and ecological impacts. Logbook data collected 2004–2008 from two Oahu (Hawaii) shark cage diving operations were analysed to determine whether such voluntary records provide useful insights into shark ecology or ecotourism impacts. Operators correctly identified common shark species and documented gross seasonal cycles and long-term trends in abundance of Galapagos (Carcharhinus galapagensis), sandbar (Carcharhinus plumbeus) and tiger sharks (Galeorcerdo cuvier). Annual cycles in shark abundance may indicate seasonal migrations, whereas long-term trends suggest gradual exclusion of smaller sandbar sharks from cage diving sites. Numerically dominant (> 98%) Galapagos and sandbar sharks are rarely implicated in attacks on humans. Negligible impact on public safety is supported by other factors such as: (1) remoteness of the sites, (2) conditioning stimuli that are specific to the tour operations and different from inshore recreational stimuli and (3) no increase in shark attacks on the north coast of Oahu since cage diving started. Tracking studies are required to validate logbook data and to determine whether sharks associated with offshore cage diving travel into inshore areas used for in-water recreation.

Little is known about the long-term movement patterns of most marine apex predators. A network of acoustic receivers was used to quantify the long-term movements of transmitter- equipped tiger sharks Galeocerdo cuvier Peron & Lesueur, 1822 in the Main Hawaiian Islands. Tiger sharks were wide-ranging, swam between islands and patrolled up to 109 km of contiguous coastline. Visits to specific acoustic receiver sites were typically brief (mean duration 3.3 min), unpredictable and interspersed by absences of weeks, months or years. This pattern may be an optimal foraging strategy for capturing risk-averse prey. Tiger sharks may have to move on soon after arriving in an area because the element of surprise is quickly lost and potential prey become wary and difficult to catch. Juvenile tiger sharks were significantly wider-ranging and less frequently detected than mature females. Juveniles may be avoiding predation by larger individuals, or explor- ing to find suitable home ranges. Tiger sharks may also switch movement patterns and foraging strategies to take advantage of different prey types, restricting their movements to exploit seasonally abundant and naive prey. Further empirical studies are required to directly link movement patterns with foraging.

We quantified spatial dynamics and substrate impacts of snorkelers and SCUBA divers within four Hawaiian MPAs to determine: (1) whether coral reefs in these areas are being damaged by recreational activities, and (2) how damage might be mitigated. Observers secretly followed snorkelers and SCUBA divers, and used handheld Global Positioning System (GPS) units to record their geographic tracks and substrate contacts. Most activities occurred within relatively small, well-defined areas associated with access points. Despite heavy use, recreation impact was low at Hawaiian MPAs because most fragile corals occurred below the maximum depth of the dominant recreational activity (snorkeling). SCUBA diving was only common at one MPA with physically durable benthic habitats. GPS tracking provided useful insights into how recreational impacts to MPAs could be reduced. General topography could be used to predict where visitors will go, and designated access points could be used to focus substrate contact away from fragile habitats.

In order to investigate ontogenetic changes in diet and diet overlap between rohu (Labeo rohita) and common carp (Cyprinus carpio) in polyculture ponds, food preferences of different size classes of these fishes were quantified. Rohu diet consisted of both phytoplankton and zooplankton, and there was a distinct ontogenetic shift in the relative importance of these food items. Zooplankton was the dominant food for rohu up to 20.6 cm total length (TL) and then gradually decreased in importance as fish grew. Phytoplankton was initially a minor component of rohu diet but gradually increased in importance and became the dominant food for rohu at 24.2 cm TL. Phytoplankton biovolume in rohu guts was positively correlated with fish size (TL). Chesson’s α indicated that rohu of all sizes preferentially selected Cladocera and avoided Cyanophyceae and Euglenophyceae. Young rohu initially preferred Rotifera and Copepoda but gradually switched to Bacillariophyceae and Chlorophyceae. Common carp diet consisted of phytoplankton, zooplankton, and benthic macroinvertebrates, but was dominated by benthic macroinvertebrates (63–92% of total diet). As common carp grew, the proportion of zooplankton ingested decreased and the proportion of benthic macroinvertebrates increased. Benthic macroinvertebrate biovolume in common carp guts was positively correlated with fish size. Common carp of up to 15.4 cm TL preferentially selected zooplankton, but common carp larger than 18.9 cm TL avoided this food item. Common carp of all sizes avoided phytoplankton. A low dietary overlap was found between rohu and common carp (Schoener overlap index: 0.08–0.35), probably due to ingestion of smaller quantities of zooplankton by the latter. Dietary overlap also decreased with increasing rohu and common carp size because of divergent ontogenetic shifts in dietary preferences of the two species.

Gaps in our knowledge of basic fish ecology have provided impetus for development of novel "ecology tags" to detect and quantify hard to observe behaviors such as spawning, schooling and feeding. The acoustic envi- ronment is one source of potentially useful information about these behaviors. We implanted an acoustic recording tag (Bioacoustic Probe) into the gut cavity of a blacktip reef shark to determine whether an implanted tag could success- fully record external and internal sounds. The tag successfully recorded reef fish vocalizations, boat engine noise, the sound of the shark feeding and unidentified rhythmic sounds that may derive from shark tail beats. Technical challenges remain, but sound recording tags have the potential to provide novel insights into shark and fish ecology.

Little is known about the feeding habits of large free ranging fish, due in large part to lack of an appropriate technique for quantifying feeding variables. A previous study demonstrated that changes in gastric pH can be used as a proxy for feeding events in free-ranging sharks. Here we describe the development of a new acoustic pH transmitter to remotely measure gastric pH in sharks in the field. The transmitter consists of a dual sensor (pH and temperature) continuous pinger, and was tested in captive adult blacktip reef sharks (Carcharhinus melanopterus). The transmitter was retained in the shark's stomach for between 5-12 days. The empty stomach had a low pH (1.6 ± 0.2) and feeding induced a rapid increase in gastric pH, which was clearly distinguishable from baseline levels. Meal size showed a significant linear relationship with the magnitude of the pH changes. Measurement accuracy of the pH transmitter ranged from 0.05-0.9, although resolution of the VR100 receiver that decoded the transmitter signal was 0.1 units. The pH transmitter can be used to determine when free-ranging sharks in the field are feeding and hence quantify feeding chronology, frequency and daily ration.

I used shoreline creel surveys to quantify fishing activities in and around a small (0.34 km 2 ) Marine Protected Area (MPA) in Hawaii (Waikiki Marine Life Conservation District). Spear fishing and shoreline pole & line fishing (angling) were the dominant fishing activities at Waikiki. Spear fishing had a greater overall impact on reef fishes than shoreline pole & line fishing, accounting for 70% of the total reef fish harvest at Waikiki, despite accounting for only 25% of fishing activities observed. Fishing activities at Waikiki were unevenly distributed in space and time. The MPA experienced minor illegal fishing and was located between an area of high diurnal spear fishing effort and an area of generally low fishing effort. This pattern of fishing activities allows jacks and goatfishes to evade capture despite nightly excursions from the MPA into fished areas, and may partly explain why these fishes remain more abundant and larger inside the Waikiki MPA than in surrounding fished areas. Quantifying fishing activities at MPA sites can provide valuable insight into how these areas function, and this information can be used to improve MPA design and effectiveness.

Empirical data quantifying the long-term movement patterns of coral reef top predators are needed in order to design marine protected areas (MPAs) that will provide these fishes with effective, long-term protection. Acoustic telemetry was used to quantify the movements of a large coral reef top predator (Aprion virescens, Lutjanidae; Hawaiian name ‘uku’) at five atolls in the Northwestern Hawaiian Islands Marine National Monument (NWHIMNM) from May 2005 to September 2006. The study atolls were located between 23.8°N, 166.2°W and 28.5°N, 178.3°W, and were separated from their nearest receiver-equipped neighbor by distances ranging from 100 to 478 km. No inter-atoll movements by uku were detected but individuals were seasonally site-attached to core activity areas of up to 12 km in length, and ranged up to 19 km across atolls. Within their core areas, tagged uku exhibited diel and tidal habitat shifts, with the latter resulting in round trips of up to 24 km in 24 h. Seasonal uku migrations resulted in extended winter (October–April) absences from summer (May–September) core activity areas and may be linked to summer spawning. Large MPAs (i.e., entire islands, atolls or banks) would probably be required for full protection of resident populations of adult uku, but such ‘island-scale’ MPAs will not benefit fisheries unless there is significant larval supply from MPAs to neighboring fished areas, or adult emigration over time scales exceeding the 16-month monitoring period of this study. A mixed management strategy of combining smaller MPAs with conventional measures (e.g., minimum size limits, catch, and effort restrictions) may be the best approach for sustaining uku fisheries. However, this would still require relatively large MPAs (12 km in length) to contain uku short-term (diel and tidal) movements, and seasonal migrations would still take uku beyond the boundaries of MPAs of this size. These fluxes across MPA boundaries could supply fish to fisheries but, if high exploitation rates exist, fishing could eliminate key MPA benefits such as increased numbers of large, highly fecund individuals.

We need to understand the long-term movement patterns of coral reef top predators in order to design marine protected areas that will provide these animals with effective, long-term pro- tection. We used acoustic telemetry to quantify the movements of giant trevally, a large coral reef top predator, at 5 atolls in the Northwestern Hawaiian Islands Marine National Monument. We did not observe any inter-atoll movements but found that giant trevally were attached to core activity areas from which they made periodic atoll-wide excursions of up to 29 km. Within the core areas, the tagged fish exhibited diel habitat shifts. We identified Rapture Reef at French Frigate Shoals (FFS) atoll as a spawning site, where giant trevally form seasonal mating aggregations. Giant trevally that use Rapture Reef as their core daytime area live there year-round, whereas those that occupy other areas of FFS migrate to Rapture Reef in the summertime during specific phases of the moon. Although giant trevally were wide-ranging and would require large marine protected areas (e.g. entire atolls or islands) to protect their entire range, core activity areas and spawning sites for giant trevally could be contained within relatively small marine protected areas. Identification of spawning sites could assist in the establishment of focused marine protected areas that, while quite small in size, could have significant impact in preserving the spawning biomass of this species. Very large marine protected areas are not feasible in most heavily populated areas, hence effective alternative management strategies must be found for this fishery-targeted, wide-ranging top predator. We sug- gest using a combination of relatively small marine protected areas, to protect core activity areas and spawning sites for giant trevally, and conventional measures such as minimum size or 'slot' limits to further safeguard spawning stocks.

We do not expect non air-breathing aquatic animals to exhibit positive buoyancy. Sharks, for example, rely on oil-filled livers instead of gas-filled swim bladders to increase their buoyancy, but are nonetheless ubiquitously regarded as either negatively or neutrally buoyant. Deep-sea sharks have particularly large, oil-filled livers, and are believed to be neutrally buoyant in their natural habitat, but this has never been confirmed. To empirically determine the buoyancy status of two species of deep-sea sharks (bluntnose sixgill sharks, Hexanchus griseus, and a prickly shark, Echinorhinus cookei) in their natural habitat, we used accelerometer-magnetometer data loggers to measure their swimming performance. Both species of deep-sea sharks showed similar diel vertical migrations: they swam at depths of 200–300 m at night and deeper than 500 m during the day. Ambient water temperature was around 15°C at 200–300 m but below 7°C at depths greater than 500 m. During vertical movements, all deep-sea sharks showed higher swimming efforts during descent than ascent to maintain a given swimming speed, and were able to glide uphill for extended periods (several minutes), indicating that these deep-sea sharks are in fact positively buoyant in their natural habitats. This positive buoyancy may adaptive for stealthy hunting (i.e. upward gliding to surprise prey from underneath) or may facilitate evening upward migrations when muscle temperatures are coolest, and swimming most sluggish, after spending the day in deep, cold water. Positive buoyancy could potentially be widespread in fish conducting daily vertical migration in deep-sea habitats.

We quantified bluespine unicornfish, Naso unicornis, movement patterns, home range size and habitat preferences in a small Hawaiian marine reserve. Bluespine unicornfish were site-attached to home ranges situated within the reserve boundaries and their movements were aligned with topographic features. Two different diel movement patterns (‘commuting’ and ‘foraying’) were observed. Commuters made crepuscular migrations of several hundred meters between daytime foraging areas and nighttime refuge holes. Foraying fish did not partake in crepuscular migrations and utilized refuge holes both day and night. Two bluespine unicornfish were also nocturnally active. There was little direct evidence of dispersal from the reserve but differences in bluespine unicornfish abundance and size among reef habitat zones were consistent with ontogenetic habitat shifts. The influence of habitat topography on bluespine unicornfish movements suggests that gross habitat characteristics could be used to predict reef fish movements. This could provide a simple method for setting marine reserve boundaries at sites for which empirical fish movement data are unavailable.

The feasibility of long-term acoustic monitoring of coral reef fish movements was evaluated by quantifying transmitter retention times for captive bluefin trevally Caranx melampygus and detection patterns of transmitter-implanted free-ranging bluefin trevally. Wax-coated dummy transmitters (9 × 22 mm; 3.6 g) were surgically implanted into the gut cavity of four captive bluefin trevally. The fish retained the dummy transmitters for 17–22 months, but at the end of the study transintestinal expulsion was in progress in one individual. Five bluefin trevally captured at Honokohau, Hawaii, were surgically implanted with long-life (1-year) coded transmitters and released. Their movements were monitored with the use of 27 underwater receivers deployed along a 115-km stretch of coastline. The five bluefin trevally were detected by the receivers over periods ranging from 1 to 173 d (mean, 60 d), and their movements spanned up to 10.2 km of coastline. Daily movements were consistent with those observed by p...

Movement patterns, site fidelity and growth were studied for giant trevally ( Caranx ignobilis) inhabiting a marine reserve surrounding Coconut Island in Kaneohe Bay, off the Island of Oahu, Hawaii. Short-term movements of fish were determined by acoustic telemetry and long-term movements and growth were investigated with conventional tag and release methods. Giant trevally appear to move to the Coconut Island marine reserve from the adjacent bay floor upon attaining a size of approximately 20–25 cm fork length. After several years spent frequenting the reserve, giant trevally move out of the bay into deeper water. Tracked fish spent considerable time within the marine reserve moving along the reef slope, but frequently ventured outside of the reserve boundaries. The recapture rate for tagged fish was 11%, with an average time at liberty of 346 days. Nearly one-third of recaptured fish were caught at distances greater than 3 km from the tagging site with maximum values demonstrating long-term (more than 7 years) and long-distance (30 km) movements. The protective function of the Coconut Island marine reserve for giant trevally is limited because the reserve is utilized by only a portion of the population, and even these fish regularly move outside of the refuge. © 2003 Elsevier B.V. All rights reserved.

Stomach content, capture method and capture location data were collected for 401 carangids captured during three annual 1-day fishing tournaments held at a coastal bay in Hawaii. Blue jack (Caranx melampygus), white jack (Caranx ignobilis) and island jack (Carangoides orthogrammus) were the most common species, accounting for 83.5, 8.5 and 5.2% of tournament catches, respectively. Geographical area fished consisted of a sheltered bay and the adjacent seaward coastal reef beyond. Area of capture and fishing method influenced species and size of fish captured. Small ( 500 mm) C. melampygus were captured outside Kaneohe Bay. Trolling (towing a surface lure) accounted for 80% of C. melampygus, 76% of C. orthogrammus and 55% of C. ignobilis captured. Differential vulnerability to trolling may be related to interspecific differences in diet; captured C. melampygus had fed primarily on fish whereas C. orthogrammus had consumed both fish and benthic crustaceans, and C. ignobilis had eaten mainly benthic crustaceans. Differences in diet may indicate resource partitioning between these sympatric and closely related species. For C. melampygus there was a consistent relationship between prey size and predator size. When conducted under scientific scrutiny, fishing tournaments can provide synoptic data on diet and gear vulnerability that would otherwise be very difficult to obtain.

Suitability of small (< 1 km2) marine reserves for protecting a commercially important endemic Hawaiian goatfish, Parupeneus porphyreus, was examined by quantifying goatfish habitat use, home range size and site fidelity in an existing marine reserve (Coconut Island in Kaneohe Bay, Hawaii). Five goatfish equipped with acoustic transmitters were tracked for up to 93 h each over 3–14 days. Daytime habitat use patterns of two of these fish were continuously monitored for one month using a fixed hydrophone hardwired to an onshore computer. Acoustically tagged fish showed consistent diel patterns of behavior, refuging in holes in the reef by day and moving over extensive areas of sand and coral rubble habitat at night. Remote monitoring of daytime habitat use by two goatfish revealed that the same daytime refuge was used by both fish for at least one month (the battery life of the transmitters). Home ranges of all fish were within the boundaries of the Coconut Island reserve suggesting that even small areas containing suitable habitat can make effective reserves for this species. A relatively low abundance of reproductive size P. porphyreus at Coconut Island in comparison with deeper areas may indicate an ontogenetic shift to deeper habitat in this species.

Externally and internally implanted sonic transmitters were used to track the movements of eight tiger sharks (Galeocerdo cuvier) ranging between 200 and 417 cm total length (TL), captured by hook-and-line on the south coast of Oahu, Hawaii, between 1993 and 1997. Attachment of the transmitters was facilitated by the fact that captured sharks exhibited tonic immobility when restrained and inverted at the side of the tagging vessel. Three common themes emerged from the horizontal movements of the tracked sharks: (1) offshore movements away from the island, (2) extended periods of directed, “straight-line” swimming, (3) orientation to the Penguin Banks – a shallow bank located ≃35 km from the release point. In shallow water ( 300 m) they swam within the mixed layer at depths of ∼80 m. One shark dove briefly to 335 m. The average estimated swimming speed of sharks traversing open water was 0.29 body length (BL) s−1. Two sharks were recaptured after termination of the tracks; one of these sharks was recaptured twice, with a total time at liberty of 377 d. The data suggest that Hawaiian tiger sharks move within large home ranges and that they can efficiently navigate between distant parts of their range, even when this requires crossing open ocean waters.

Empirical data on the abundance and habitat preferences of coral reef top predators are needed to evaluate their ecological impacts and guide management decisions. We used longline surveys to quantify the shark assemblage at French Frigate Shoals (FFS) atoll from May to August 2009. Fishing effort consisted of 189 longline sets totaling 6,862 hook hours of soak time. A total of 221 sharks from 7 species were captured, among which Galapagos (Carcharhinus galapagensis, 36.2%), gray reef (Carcharhinus amblyrhynchos, 25.8%) and tiger (Galeocerdo cuvier, 20.4%) sharks were numerically dominant. A lack of blacktip reef sharks (Carcharhinus melanopterus) distinguished the FFS shark assemblage from those at many other atolls in the Indo-Pacific. Compared to prior underwater visual survey estimates, longline methods more accurately represented species abundance and composition for the majority of shark species. Sharks were significantly less abundant in the shallow lagoon than adjacent habitats. Recaptures of Galapagos sharks provided the first empirical estimate of population size for any Galapagos shark population. The overall recapture rate was 5.4%. Multiple closed population models were evaluated, with Chao M(h) ranking best in model performance and yielding a population estimate of 668 sharks with 95% confidence intervals ranging from 289-1720. Low shark abundance in the shallow lagoon habitats suggests removal of a small number of sharks from the immediate vicinity of lagoonal islets may reduce short-term predation on endangered monk seal (Monachus schauinslandi) pups, but considerable fishing effort would be required to catch even a small number of sharks. Additional data on long-term movements and habitat use of sharks at FFS are required to better assess the likely ecological impacts of shark culling.