Acoustic Telemetry Validates a Citizen Science Approach for Monitoring Sharks on Coral Reefs

Citizen science is promoted as a simple and cost-effective alternative to traditional approaches for the monitoring of populations of marine megafauna. However, the reliability of datasets collected by these initiatives often remains poorly quantified. We compared datasets of shark counts collected by professional dive guides with acoustic telemetry data from tagged sharks collected at the same coral reef sites over a period of five years. There was a strong correlation between the number of grey reef sharks (Carcharhinus amblyrhynchos) observed by dive guides and the telemetry data at both daily and monthly intervals, suggesting that variation in relative abundance of sharks was detectable in datasets collected by dive guides in a similar manner to data derived from telemetry at these time scales. There was no correlation between the number or mean depth of sharks recorded by telemetry and the presence of tourist divers, suggesting that the behaviour of sharks was not affected by the presence of divers during our study. Data recorded by dive guides showed that current strength and temperature were important drivers of the relative abundance of sharks at monitored sites. Our study validates the use of datasets of shark abundance collected by professional dive guides in frequently-visited dive sites in Palau, and supports the participation of experienced recreational divers as contributors to long-term monitoring programs of shark populations.     

Environmental Influences on Patterns of Vertical Movement and Site Fidelity of Grey Reef Sharks (Carcharhinus amblyrhynchos) at Aggregation Sites

We used acoustic telemetry to describe the patterns of vertical movement, site fidelity and residency of grey reef sharks (Carcharhinus amblyrhynchos) on the outer slope of coral reefs in Palau, Micronesia, over a period of two years and nine months. We tagged 39 sharks (mostly adult females) of which 31 were detected regularly throughout the study. Sharks displayed strong inter-annual residency with greater attendance at monitored sites during summer than winter months. More individuals were detected during the day than at night. Mean depths of tagged sharks increased from 35 m in winter to 60 m in spring following an increase in water temperature at 60 m, with maximum mean depths attained when water temperatures at 60 m stabilised around 29°C. Sharks descended to greater depths and used a wider range of depths around the time of the full moon. There were also crepuscular cycles in mean depth, with sharks moving into shallower waters at dawn and dusk each day. We suggest that daily, lunar and seasonal cycles in vertical movement and residency are strategies for optimising both energetic budgets and foraging behaviour. Cyclical patterns of movement in response to environmental variables might affect the susceptibility of reef sharks to fishing, a consideration that should be taken into account in the implementation of conservation strategies.     

Elevated body temperatures of adult female leopard sharks, Triakis semifasciata, while aggregating in shallow nearshore embayments: Evidence for behavioral thermoregulation?

The leopard shark, Triakis semifasciata, is an abundant nearshore elasmobranch, ranging from Baja California, Mexico to Oregon, USA. Mature female leopard sharks have been observed aggregating in shallow embayments throughout California; however, it is unclear why only females aggregate in these shallow areas. The goal of this study was to determine if mature female leopard sharks selectively occupy the warmest areas of a shallow embayment, if free-ranging leopard sharks' body temperatures are significantly warmer during the day than at night, and to quantify temporal use of these shallow habitats. Visual observations of sharks' fine-scale movements within the shallows of Big Fisherman's Cove Marine Life Refuge (Santa Catalina Island) aggregation site indicated that sharks preferred the warmest areas of the embayment and moved to warmer locations over the course of the day (p < 0.05). Active and passive acoustic tracking, along with archival transponder technology (Vemco: V13, V13-R256, VX32TP-CHAT tags respectively) of 16 sharks caught and tagged within this aggregation were used to monitor core body temperature, swimming depth, and movements. Sharks had significantly higher core body temperatures in the late afternoon (1700 h-2000 h) during the summer, showed increased fidelity to thermal refuges during the day and increased movement away from these refuges at night (χ², p < 0.05). Seasonal variations in warm, shallow water usage were also observed. Elevated core body temperature of mature female leopard sharks using warm shallow embayments will likely augment metabolic and physiological functions such as digestion, somatic growth, and possibly reproduction.     

Residency and spatial use by reef sharks of an isolated seamount and its implications for conservation

Although marine protected areas (MPAs) are a common conservation strategy, these areas are often designed with little prior knowledge of the spatial behaviour of the species they are designed to protect. Currently, the Coral Sea area and its seamounts (north-east Australia) are under review to determine if MPAs are warranted. The protection of sharks at these seamounts should be an integral component of conservation plans. Therefore, knowledge on the spatial ecology of sharks at the Coral Sea seamounts is essential for the appropriate implementation of management and conservation plans. Acoustic telemetry was used to determine residency, site fidelity and spatial use of three shark species at Osprey Reef: whitetip reef sharks Triaenodon obesus, grey reef sharks Carcharhinus amblyrhynchos and silvertip sharks Carcharhinus albimarginatus. Most individuals showed year round residency at Osprey Reef, although five of the 49 individuals tagged moved to the neighbouring Shark Reef (~14 km away) and one grey reef shark completed a round trip of ~250 km to the Great Barrier Reef. Additionally, individuals of white tip and grey reef sharks showed strong site fidelity to the areas they were tagged, and there was low spatial overlap between groups of sharks tagged at different locations. Spatial use at Osprey Reef by adult sharks is generally restricted to the north-west corner. The high residency and limited spatial use of Osprey Reef suggests that reef sharks would be highly vulnerable to targeted fishing pressure and that MPAs incorporating no-take of sharks would be effective in protecting reef shark populations at Osprey and Shark Reef.     

Fine scale movements and activity areas of white sharks (Carcharodon carcharias) in Mossel Bay, South Africa

Previous work on white sharks indicate the species show seasonally limited movement patters, at certain aggregation sites small areas may play vital roles in the life history of a large amount of the population. Acoustic telemetry was used to estimate habitat use of white sharks, Carcharodon carcharias, while aggregating at Mossel Bay, South Africa. Total range of all shark tracks combined accumulated 782 h and covered an area of 93.5 km² however, within this range, sharks were found to highly utilise a core habitat (50 % Kernel, K50) of just 1.05 km² over a reef system adjacent to a river mouth. Individual tracks revealed additional core habitats, some of which were previously undocumented and one adjacent to a commercial harbor. Much was found to be dependent on the size of the shark, with larger sharks (>400 cm) occupying smaller activity areas than subadult (300–399 cm) and juvenile (<300 cm) conspecifics, while Index of Reuse (IOR) and Index of Shared Space (IOSS) were both found to increase with shark size. Such results provide evidence that larger white sharks are more selective in habitat use, which indicates they have greater experience within aggregation sites. Furthermore, the focused nature of foraging means spatially restricted management strategies would offer a powerful tool to aid enforcement of current protective legislation for the white shark in similar environments of limited resources and capacity.     

Regulation of body temperature in the white shark, Carcharodon carcharias

Stomach temperatures of three white sharks, Carcharodoncarcharias, (one reported previously and two new individuals) were intermittently recorded by acoustic telemetry at the South Farallon Islands, central California. Temperature profiles of the water column were obtained concurrently. Stomach temperatures were elevated over ambient water temperatures by as much as 14.3 °C. Stomach temperatures varied within a narrow range while ambient water temperature fluctuated over a much larger range, showing that this species regulates its body temperature. These data, in combination with previous work on the physiology and anatomy of white sharks, indicate that the white shark is endothermic. It appears that the heat retention system in lamnid sharks has allowed them to inhabit cold water and remain active predators of swift and agile prey. Accepted: 17 February 1997     

Quantifying movement patterns for shark conservation at remote coral atolls in the Indian Ocean

Grey reef sharks (Carcharhinus amblyrhynchos) are apex predators found on many Indo-Pacific coral reefs, but little is known about their movement patterns and habitat requirements. We used acoustic telemetry to determine movements and habitat use of these sharks at the isolated Rowley Shoals atolls, 250 km off the coast of north-western Australia. We equipped 12 male and 14 female sharks ranging from 0.79 to 1.69 m in total length with transmitters that were detected by an array of 11 strategically placed receivers on two atoll reefs. Over 26,000 detections were recorded over the 325 days of receiver deployment. No sharks were observed to move between reefs. Receivers on the outer slopes of reefs provided nearly all (99%) of the detections. We found no differences in general attendance parameters due to size, sex or reef, except for maximum period of detection where larger sharks were detected over a longer period than smaller sharks. Male and female sharks were often detected at separate receivers at the outer slope habitat of one reef, suggesting sexual segregation, but this pattern did not occur at the second reef where males and females were detected at similar frequencies. We identified two patterns of daily behaviour: (1) sharks were present at the reef both day and night or (2) sharks spent more time in attendance during day than at night. Fast Fourier transforms identified 24-h cycles of attendance at the reef and a secondary peak of attendance at 12 h for most sharks, although no individuals shared the same attendance patterns. Our study provides baseline data that can be used to optimise the minimum area and habitat requirements for conservation of these apex predators.     

Movement Patterns of Juvenile Whale Sharks Tagged at an Aggregation Site in the Red Sea

Conservation efforts aimed at the whale shark, Rhincodon typus, remain limited by a lack of basic information on most aspects of its ecology, including global population structure, population sizes and movement patterns. Here we report on the movements of 47 Red Sea whale sharks fitted with three types of satellite transmitting tags from 2009–2011. Most of these sharks were tagged at a single aggregation site near Al-Lith, on the central coast of the Saudi Arabian Red Sea. Individuals encountered at this site were all juveniles based on size estimates ranging from 2.5–7 m total length with a sex ratio of approximately 1∶1. All other known aggregation sites for juvenile whale sharks are dominated by males. Results from tagging efforts showed that most individuals remained in the southern Red Sea and that some sharks returned to the same location in subsequent years. Diving data were recorded by 37 tags, revealing frequent deep dives to at least 500 m and as deep as 1360 m. The unique temperature-depth profiles of the Red Sea confirmed that several whale sharks moved out of the Red Sea while tagged. The wide-ranging horizontal movements of these individuals highlight the need for multinational, cooperative efforts to conserve R. typus populations in the Red Sea and Indian Ocean.     

Size frequency,dispersal distances and variable growth rates of young sharks in a multi-species aggregation

During a mark–recapture survey from November 2014 until April 2017, 333 neonatal and juvenile blacktip reef sharks Carcharhinus melanopterus and 302 neonatal and juvenile sicklefin lemon sharks Negaprion acutidens were tagged and measured at the uninhabited and isolated St. Joseph Atoll (Republic of Seychelles). Both species demonstrated seasonal reproductive synchronicity and relatively large sizes at birth. Despite the extended times at liberty > 2.5 years, the majority of recaptures were found in close proximity to the initial tagging location (< 500 m). Annual growth rates of C. melanopterus (n = 24) and N. acutidens (n = 62) ranged from 6.6 to 31.7 cm year⁻¹ (mean ± SE; 16.2 ± 1.2 cm year⁻¹) and 0.2 to 32.2 cm year⁻¹ (11.8 ± 1 cm year⁻¹), respectively and are to date the most variable ever recorded in wild juvenile sharks. High abundances of both species coupled with long-term and repeated recaptures are indicative of a habitat where juveniles can reside for their first years of life. However, large variability in annual growth rates in both species may suggest high intra and interspecific competition induced by a possibly resource limited, isolated habitat.     

Reef-Fidelity and Migration of Tiger Sharks,Galeocerdo cuvier,across the Coral Sea

Knowledge of the habitat use and migration patterns of large sharks is important for assessing the effectiveness of large predator Marine Protected Areas (MPAs), vulnerability to fisheries and environmental influences, and management of shark–human interactions. Here we compare movement, reef-fidelity, and ocean migration for tiger sharks, Galeocerdo cuvier, across the Coral Sea, with an emphasis on New Caledonia. Thirty-three tiger sharks (1.54 to 3.9 m total length) were tagged with passive acoustic transmitters and their localised movements monitored on receiver arrays in New Caledonia, the Chesterfield and Lord Howe Islands in the Coral Sea, and the east coast of Queensland, Australia. Satellite tags were also used to determine habitat use and movements among habitats across the Coral Sea. Sub-adults and one male adult tiger shark displayed year-round residency in the Chesterfields with two females tagged in the Chesterfields and detected on the Great Barrier Reef, Australia, after 591 and 842 days respectively. In coastal barrier reefs, tiger sharks were transient at acoustic arrays and each individual demonstrated a unique pattern of occurrence. From 2009 to 2013, fourteen sharks with satellite and acoustic tags undertook wide-ranging movements up to 1114 km across the Coral Sea with eight detected back on acoustic arrays up to 405 days after being tagged. Tiger sharks dove 1136 m and utilised three-dimensional activity spaces averaged at 2360 km³. The Chesterfield Islands appear to be important habitat for sub-adults and adult male tiger sharks. Management strategies need to consider the wide-ranging movements of large (sub-adult and adult) male and female tiger sharks at the individual level, whereas fidelity to specific coastal reefs may be consistent across groups of individuals. Coastal barrier reef MPAs, however, only afford brief protection for large tiger sharks, therefore determining the importance of other oceanic Coral Sea reefs should be a priority for future research.     

Archival Tagging Of School Shark, Galeorhinus Galeus, in Australia: Initial Results

Archival tags were used to study the movement and depth behaviour of school sharks, Galeorhinus galeus, in southern australia. Thirty fish were tagged in late 1997, and to date there have been nine recaptures (30% recapture rate). Periods at liberty varied from 8 days to 18 months. The sharks spent about 80% of their time on the continental shelf, and appeared to swim close to the bottom during the day. At night they often ascended for periods of up to several hours, except at times around the full moon. When in deep water, the sharks typically descended at dawn to depths of up to 600m, before ascending at dusk. It was not possible to use the light data from the tags to estimate position when the sharks were in deep water, because they were often at depths beyond the sensitivity of the tag. In shallower water, longitude was estimated from the light data but latitude was estimated from the maximum daily depth, assuming the fish were on the bottom. The timing of the dives in deepwater appeared sufficiently regular to offer the prospect of using it to estimate longitude. We propose future research using archival tags on this species should address questions about female reproductive migrations, pelagic behaviour and vertical movements.     

Crossing Latitudes—Long-Distance Tracking of an Apex Predator

Tiger sharks (Galeocerdo cuvier) are apex predators occurring in most tropical and warm temperate marine ecosystems, but we know relatively little of their patterns of residency and movement over large spatial and temporal scales. We deployed satellite tags on eleven tiger sharks off the north-western coast of Western Australia and used the Brownian Bridge kernel method to calculate home ranges and analyse movement behaviour. One individual recorded one of the largest geographical ranges of movement ever reported for the species, travelling over 4000 km during 517 days of monitoring. Tags on the remainder of the sharks reported for shorter periods (7-191 days). Most of these sharks had restricted movements and long-term (30-188 days) residency in coastal waters in the vicinity of the area where they were tagged. Core home range areas of sharks varied greatly from 1166.9 to 634,944 km². Tiger sharks spent most of their time in water temperatures between 23°-26°C but experienced temperatures ranging from 6°C to 33°C. One shark displayed seasonal movements among three distinct home range cores spread along most of the coast of Western Australia and generalized linear models showed that this individual had different patterns of temperature and depth occupancy in each region of the coast, with the highest probability of residency occurring in the shallowest areas of the coast with water temperatures above 23°C. These results suggest that tiger sharks can migrate over very large distances and across latitudes ranging from tropical to the cool temperate waters. Such extensive long-term movements may be a key element influencing the connectivity of populations within and among ocean basins.     

Acoustic telemetry reveals cryptic residency of whale sharks

Although whale sharks (Rhincodon typus) have been documented to move thousands of kilometres, they are most frequently observed at a few predictable seasonal aggregation sites. The absence of sharks at the surface during visual surveys has led to the assumption that sharks disperse to places unknown during the long ‘off-seasons’ at most of these locations. Here we compare 2 years of R. typus visual sighting records from Mafia Island in Tanzania to concurrent acoustic telemetry of tagged individuals. Sightings revealed a clear seasonal pattern with a peak between October and February and no sharks observed at other times. By contrast, acoustic telemetry demonstrated year-round residency of R. typus. The sharks use a different habitat in the off-season, swimming deeper and further away from shore, presumably in response to prey distributions. This behavioural change reduces the sharks'' visibility, giving the false impression that they have left the area. We demonstrate, for the first time to our knowledge, year-round residency of unprovisioned, individual R. typus at an aggregation site, and highlight the importance of using multiple techniques to study the movement ecology of marine megafauna.     

Intraspecific variation in vertical habitat use by tiger sharks (Galeocerdo cuvier) in the western North Atlantic

Tiger sharks (Galeocerdo cuvier) are a wide ranging, potentially keystone predator species that display a variety of horizontal movement patterns, making use of coastal and pelagic waters. Far less, however, is known about their vertical movements and use of the water column. We used pop‐up satellite archival tags with two data sampling rates (high rate and standard rate tags) to investigate the vertical habitat use and diving behavior of tiger sharks tagged on the Puerto Rico–Virgin Islands platform and off Bermuda between 2008 and 2009. Useable data were received from nine of 14 sharks tagged, tracked over a total of 529 days. Sharks spent the majority of their time making yo‐yo dives within the upper 50 m of the water column and considerable time within the upper 5 m of the water column. As a result, sharks typically occupied a narrow daily temperature range (~2°C). Dives to greater than 200 m were common, and all sharks made dives to at least 250 m, with one shark reaching a depth of 828 m. Despite some similarities among individuals, a great deal of intraspecific variability in vertical habit use was observed. Four distinct depth distributions that were not related to tagging location, horizontal movements, sex, or size were detected. In addition, similar depth distributions did not necessitate similar dive patterns among sharks. Recognition of intraspecific variability in habitat use of top predators can be crucial for effective management of these species and for understanding their influence on ecosystem dynamics.     

Fine‐scale oceanographic drivers of reef manta ray (Mobula alfredi) visitation patterns at a feeding aggregation site

Globally, reef manta rays (Mobula alfredi) are in decline and are particularly vulnerable to exploitation and disturbance at aggregation sites. Here, passive acoustic telemetry and a suite of advanced oceanographic technologies were used for the first time to investigate the fine‐scale (5‐min) influence of oceanographic drivers on the visitation patterns of 19 tagged M. alfredi to a feeding aggregation site at Egmont Atoll in the Chagos Archipelago. Boosted regression trees indicate that tag detection probability increased with the intrusion of cold‐water bores propagating up the atoll slope through the narrow lagoon inlet during flood tide, potentially transporting zooplankton from the thermocline. Tag detection probability also increased with warmer near‐surface temperature close to low tide, with near‐surface currents flowing offshore, and with high levels of backscatter (a proxy of zooplankton biomass). These combinations of processes support the proposition that zooplankton carried from the thermocline into the lagoon during the flood may be pumped back out through the narrow inlet during an ebb tide. These conditions provide temporally limited feeding opportunities for M. alfredi, which are tied on the tides. Results also provide some evidence of the presence of Langmuir Circulation, which transports and concentrates zooplankton, and may partly explain why M. alfredi occasionally remained at the feeding location for longer than that two hours. Identification of these correlations provides unique insight into the dynamic synthesis of fine‐scale oceanographic processes which are likely to influence the foraging ecology of M. alfredi at Egmont Atoll, and elsewhere throughout their range.     

Telemetry and Satellite Tracking of Whale Sharks, Rhincodon Typus, in the Sea of Cortez, Mexico, and the North Pacific Ocean

We used satellite-linked radio telemetry to document the geographic and vertical movements and thermal habitats of whale sharks in the Sea of Cortez and as they migrated into the north Pacific Ocean. Of 17 sharks tagged between 1994 and 1996, six dispersed widely in the Sea of Cortez during 12–39 days of tracking. Four others left the Sea of Cortez and ranged extensively in the north Pacific Ocean. Indeed, one whale shark migrated to the western north Pacific Ocean, covering over 13000km in 37 months of tracking. The sharks generally occupied areas where sea surface water temperatures were between 28 and 32°C, though several ranged to depths of 240m or deeper where water temperature reached 10°C or colder. Whale sharks may segregate by size and sex, and their movement patterns appear to be related to oceanographic features, such as sea mounts and boundary currents, where primary productivity may be enhanced. These results have important implications for the global conservation of the world's largest yet least known fish. We think that satellite telemetry is a exceptionally promising tool for learning more about the ecology of whale sharks, especially when combined with conventional methods of telemetry and molecular biology.authorship arranged alphabetically     

Seasonal foraging movements and migratory patterns of female Lamna ditropis tagged in Prince William Sound, Alaska

Conventional and electronic tags were used to investigate social segregation, distribution, movements and migrations of salmon sharks Lamna ditropis in Prince William Sound, Alaska. Sixteen salmon sharks were tagged with satellite transmitters and 246 with conventional tags following capture, and were then released in Prince William Sound during summer 1999 to 2001. Most salmon sharks sexed during the study were female (95%), suggesting a high degree of sexual segregation in the region. Salmon sharks congregated at adult Pacific salmon Oncorhynchus spp. migration routes and in bays near Pacific salmon spawning grounds in Prince William Sound during July and August. Adult Pacific salmon were the principal prey in 51 salmon shark stomachs collected during summer months in Prince William Sound, but the fish appeared to be opportunistic predators and consumed sablefish Anoplopoma fimbria, gadids, Pacific herring Clupea pallasi, rockfish Sebastes spp. and squid (Teuthoidea) even when adult Pacific salmon were locally abundant. As Pacific salmon migrations declined in late summer, the salmon sharks dispersed; some continued to forage in Prince William Sound and the Gulf of Alaska into autumn and winter months, while others rapidly moved south‐east thousands of kilometres toward the west coasts of Canada and the U.S. Three movement modes are proposed to explain the movement patterns observed in the Gulf of Alaska and eastern North Pacific Ocean: ‘focal foraging’ movements, ‘foraging dispersals’ and ‘direct migrations’. Patterns of salmon shark movement are possibly explained by spatio‐temporal changes in prey quality and density, an energetic trade‐off between prey availability and water temperature, intra‐specific competition for food and reproductive success. Transmissions from the electronic tags also provided data on depth and water temperatures experienced by the salmon sharks. The fish ranged from the surface to a depth of 668 m, encountered water temperatures from 4·0 to 16·8° C and generally spent the most time above 40 m depth and between 6 and 14° C (60 and 73%, respectively).     

Aggregations of juvenile whale sharks (<Emphasis Type="Italic">Rhincodon typus</Emphasis>) in the Gulf of Tadjoura,Djibouti

A total of 23 whale sharks were identified over a 5 d period in the Arta Bay region of the Gulf of Tadjora, Djibouti. Most of the sharks aggregating in this area were small (<4 m TL) males. Individuals were identified using photographs of distinctive scars and spot and stripe patterns on the sides of the animals. Of these, 65% had scarring that was attributable to boat or propeller strikes. Most of the whale sharks we encountered were feeding on dense accumulations of plankton in shallow water just off (10–200 m) the shoreline. This food source may account for the aggregation of sharks in this area. One 3 m male shark was tagged with an ARGOS (Splash) satellite tag for 9 d. During this time the shark traversed to the shoreline on the opposite side of the Gulf (a distance of 14 km) and then returned to the Arta Bay area before retracing his path to the other shore. The shark spent most of the daylight hours at the surface, while at night dives were more frequent, deeper and for longer durations.

Regional movements of satellite‐tagged whale sharks Rhincodon typus in the Gulf of Aden

To gain insight into whale shark (Rhincodon typus) movement patterns in the Western Indian Ocean, we deployed eight pop‐up satellite tags at an aggregation site in the Arta Bay region of the Gulf of Tadjoura, Djibouti in the winter months of 2012, 2016, and 2017. Tags revealed movements ranging from local‐scale around the Djibouti aggregation site, regional movements along the coastline of Somaliland, movements north into the Red Sea, and a large‐scale (>1,000 km) movement to the east coast of Somalia, outside of the Gulf of Aden. Vertical movement data revealed high occupation of the top ten meters of the water column, diel vertical movement patterns, and deep diving behavior. Long‐distance movements recorded both here and in previous studies suggest that connectivity between the whale sharks tagged at the Djibouti aggregation and other documented aggregations in the region are likely within annual timeframes. In addition, wide‐ranging movements through multiple nations, as well as the high use of surface waters recorded, likely exposes whale sharks in this region to several anthropogenic threats, including targeted and bycatch fisheries and ship‐strikes. Area‐based management approaches focusing on seasonal hotspots offer a way forward in the conservation of whale sharks in the Western Indian Ocean.