The Seasonal Aggregation of Whale Sharks at Ningaloo Reef, Western Australia: Currents, Migrations and the El Niño/ Southern Oscillation

Whale sharks, Rhincodon typus, seasonally aggregate in coastal waters off Ningaloo Reef, Western Australia. We review the oceanographic setting of the region and present evidence that such aggregations form as a result of migratory behavior associated with climatic and oceanographic processes. We utilise records of whale shark abundance collected at Ningaloo Reef from dedicated searches by boat and aircraft and from log sheets recorded by the tourism industry. Measures of whale shark abundance derived from log sheet data sets were moderately correlated with the Southern Oscillation Index and weakly correlated with coastal sea level, an index of the strength of the Leeuwin Current, and sea surface temperature over the period 1993 to 1998. Abundances of whale sharks derived from boat searches from 1983 to 1992 were also correlated with fluctuations in the Southern Oscillation Index, except during a three year period from 1988 to 1990. We conclude that, at least in some years, there appears to be a link between the abundance of aggregating whale sharks and the physical and biological oceanography of the region, with greater whale shark numbers in La Niña years. The lack of correlation in other years may be due to a combination of uneven quality of data and/or aggregations occurring in response to a complex interaction between the physical and biological oceanography of the region.     

Shark detection and classification with machine learning

Suitable shark conservation depends on well-informed population assessments. Direct methods such as scientific surveys and fisheries monitoring are adequate for defining population statuses, but species-specific indices of abundance and distribution coming from these sources are rare for most shark species. We can rapidly fill these information gaps by boosting media-based remote monitoring efforts with machine learning and automation.We created a database of 53,345 shark images covering 219 species of sharks, and packaged object-detection and image classification models into a Shark Detector bundle. The Shark Detector recognizes and classifies sharks from videos and images using transfer learning and convolutional neural networks (CNNs). We applied these models to common data-generation approaches of sharks: collecting occurrence records from photographs taken by the public or citizen scientists, processing baited remote camera footage and online videos, and data-mining Instagram. We examined the accuracy of each model and tested genus and species prediction correctness as a result of training data quantity.The Shark Detector can classify 47 species pertaining to 26 genera. It sorted heterogeneous datasets of images sourced from Instagram with 91% accuracy and classified species with 70% accuracy. It located sharks in baited remote footage and YouTube videos with 89% accuracy, and classified located subjects to the species level with 69% accuracy. All data-generation methods were processed without manual interaction.As media-based remote monitoring appears to dominate methods for observing sharks in nature, we developed an open-source Shark Detector to facilitate common identification applications. Prediction accuracy of the software pipeline increases as more images are added to the training dataset. We provide public access to the software on our GitHub page.     

Influence of teleost abundance on the distribution and abundance of sharks in Florida Bay, USA

Understanding the factors that influence the distribution and abundance of predators, including sharks, is important for predicting the impacts of human changes to the environment. Such studies are particularly important in Florida Bay, USA where there are planned large-scale changes to patterns of freshwater input from the Everglades ecosystem. Studies of many marine predators suggest that links between predator and prey habitat use may vary with spatial scale, but there have been few studies of the role of prey distribution in shaping habitat use and abundance of sharks. We used longline catches of sharks and trawls for potential teleost prey to determine the influence of teleost abundance on shark abundance at the scale of regions and habitats in Florida Bay. We found that shark catch per unit effort (CPUE) was not linked to CPUE ofteleosts at the scale of sampling sites, but shark CPUE was positively correlated with the mean CPUE for teleosts within a region. Although there does not appear to be a strong match between the abundance of teleosts and sharks at small spatial scales, regional shark abundance is likely driven, at least partially, by the availability of prey. Management strategies that influence teleost abundance will have cascading effects to higher trophic levels in Florida Bay. Electronic Supplementary Material is available for this article at     

Seasonal and long-term changes in relative abundance of bull sharks from a tourist shark feeding site in Fiji

Shark tourism has become increasingly popular, but remains controversial because of major concerns originating from the need of tour operators to use bait or chum to reliably attract sharks. We used direct underwater sampling to document changes in bull shark Carcharhinus leucas relative abundance at the Shark Reef Marine Reserve, a shark feeding site in Fiji, and the reproductive cycle of the species in Fijian waters. Between 2003 and 2009, the total number of C. leucas counted on each day ranged from 0 to 40. Whereas the number of C. leucas counted at the feeding site increased over the years, shark numbers decreased over the course of a calendar year with fewest animals counted in November. Externally visible reproductive status information indicates that the species' seasonal departure from the feeding site may be related to reproductive activity.     

Reef sharks exhibit site-fidelity and higher relative abundance in marine reserves on the Mesoamerican Barrier Reef

Carcharhinid sharks can make up a large fraction of the top predators inhabiting tropical marine ecosystems and have declined in many regions due to intense fishing pressure. There is some support for the hypothesis that carcharhinid species that complete their life-cycle within coral reef ecosystems, hereafter referred to as “reef sharks”, are more abundant inside no-take marine reserves due to a reduction in fishing pressure (i.e., they benefit from marine reserves). Key predictions of this hypothesis are that (a) individual reef sharks exhibit high site-fidelity to these protected areas and (b) their relative abundance will generally be higher in these areas compared to fished reefs. To test this hypothesis for the first time in Caribbean coral reef ecosystems we combined acoustic monitoring and baited remote underwater video (BRUV) surveys to measure reef shark site-fidelity and relative abundance, respectively. We focused on the Caribbean reef shark (Carcharhinus perezi), the most common reef shark in the Western Atlantic, at Glover''s Reef Marine Reserve (GRMR), Belize. Acoustically tagged sharks (N = 34) were detected throughout the year at this location and exhibited strong site-fidelity. Shark presence or absence on 200 BRUVs deployed at GRMR and three other sites (another reserve site and two fished reefs) showed that the factor “marine reserve” had a significant positive effect on reef shark presence. We rejected environmental factors or site-environment interactions as predominant drivers of this pattern. These results are consistent with the hypothesis that marine reserves can benefit reef shark populations and we suggest new hypotheses to determine the underlying mechanism(s) involved: reduced fishing mortality or enhanced prey availability.     

Ontogenetic changes in the diet of the sandbar shark, <Emphasis Type="Italic">Carcharhinus plumbeus</Emphasis>, in lower Chesapeake Bay and Virginia (USA) coastal waters

This study describes the diet of the sandbar shark, Carcharhinus plumbeus, highlighting differences in diet within various regions of the Virginia (USA) nursery area, as well as ontogenetic changes in diet. Stomach samples were obtained in 2001 and 2002 from 232 sharks caught by gillnets or longlines. Historical data from the Virginia Institute of Marine Science (VIMS) Shark Ecology Program were also analyzed. Ontogenetic changes in diet were evident, with crustacean prey decreasing in frequency with increasing shark size, and elasmobranch prey importance increasing with increasing shark size. Whereas previous research in Chincoteague Bay, VA showed the blue crab, Callinectes sapidus, was the dominant crustacean in sandbar shark diet, the mantis shrimp, Squilla empusa, dominated the crustacean portion of the diet in this study. Differences in diet of sharks were observed among locations within the study area. Small juveniles (≤80 cm precaudal length) in the lower Chesapeake Bay ate more fishes, whereas Eastern Shore juveniles ate more crustaceans. Crustacean prey items varied among locations along the Eastern Shore, with more portunid crabs consumed in waters near Wachapreague and more mantis shrimp consumed near Sand Shoal Inlet. Our study showed that Carcharhinus plumbeus is a generalist predator and is thus unlikely to strongly impact the population of any particular prey species, and in turn is not likely to be strongly affected by fluctuations in abundance of a single prey species.     

SHARK ATTACKS ON BOTTLENOSE DOLPHINS (TURSIOPS ADUNCUS) IN SHARK BAY, WESTERN AUSTRALIA: ATTACK RATE, BITE SCAR FREQUENCIES, AND ATTACK SEASONALITY

Shark predation may have been a central factor influencing the evolution of sociality in dolphins, as well as a determinant of dolphin habitat use and behavior. To understand the role of predation in driving interpopulation differences in behavior and sociality, it is important to quantify differences in predation risk among populations. This study describes the frequency of shark-inflicted scars and estimates the shark attack rate on bottlenose dolphins ( Tursiops aduncus ) in Shark Bay, Western Australia. Shark bite scars were found on 74.2% (95 of 128) of non-calves, and most of these scars were inflicted by tiger sharks ( Galeocerdo cuvier ). Although there were no differences among age/sex classes in the frequency of scarring, significantly more adult males than adult females bore multiple scars. The rate of unsuccessful shark attack was estimated to be between 11% and 13% of dolphins attacked each year. Large sharks (>3 m) were responsible for a disproportionate number of attacks. However, bites from small carcharhinid sharks on 6.2% of dolphins suggest that some of these small sharks may be dolphin ectoparasites. Both the scar frequencies and attack rate suggest that Shark Bay dolphins face a greater risk of predation than bottlenose dolphins in other locations.     

Shark attacks in Dakar and the Cap Vert Peninsula, Senegal: low incidence despite high occurrence of potentially dangerous species

Background

The International Shark Attack File mentions only four unprovoked shark attacks on the coast of West Africa during the period 1828–2004, an area where high concentrations of sharks and 17 species potentially dangerous to man have been observed. To investigate if the frequency of shark attacks could be really low and not just under-reported and whether there are potentially sharks that might attack in the area, a study was carried out in Dakar and the Cap Vert peninsula, Senegal.

Methodology/Principal Findings

Personnel of health facilities, administrative services, traditional authorities and groups of fishermen from the region of Dakar were interviewed about the occurrence of shark attacks, and visual censuses were conducted along the coastline to investigate shark communities associated with the coasts of Dakar and the Cap Vert peninsula. Six attacks were documented for the period 1947–2005, including two fatal ones attributed to the tiger shark Galeocerdo cuvieri. All attacks concerned fishermen and only one occurred after 1970. Sharks were observed year round along the coastline in waters 3–15 m depth. Two species potentially dangerous for man, the nurse shark Ginglymostoma cirratum and the blacktip shark Carcharhinus limbatus, represented together 94% of 1,071 sharks enumerated during 1,459 hours of observations. Threatening behaviour from sharks was noted in 12 encounters (1.1%), including 8 encounters with C. limbatus, one with Galeocerdo cuvieri and 3 with unidentified sharks.

Conclusions/Significance

These findings suggest that the frequency of shark attacks on the coast of West Africa is underestimated. However, they also indicate that the risk is very low despite the abundance of sharks. In Dakar area, most encounters along the coastline with potentially dangerous species do not result in an attack. Compared to other causes of water related deaths, the incidence of shark attack appears negligible, at least one thousand fold lower.     

The shark nursery of Bulls Bay,South Carolina,with a review of the shark nurseries of the southeastern coast of the United States

Synopsis Shark nurseries, or nursery areas, are geographically discrete parts of a species range where the gravid females of most species of coastal sharks deliver their young or deposit their eggs, and where their young spend their first weeks, months, or years. These areas are usually located in shallow, energy rich coastal areas where the young find abundant food and have little predation by larger sharks. Nurseries are characterized by the presence of both gravid females and free swimming neonates. Neonates are young bearing fresh, unhealed umbilical scars in the case of placental species, or those at or near the birth size in aplacental species. Bulls Bay, South Carolina, is a nursery for the blacknose, spinner, finetooth, blacktip, sandbar, dusky, Atlantic sharpnose, scalloped hammerhead, and smooth dogfish sharks. The lemon shark has its nursery in shallow waters of south Florida and the Bahamas. The bull shark has its nursery in the lagoons of the east coast of central Florida.     

Fiji’s largest marine reserve benefits reef sharks

To provide more information about whether sharks benefit from no-take marine reserves, we quantified the relative abundance and biomass of reef sharks inside and outside of Namena, Fiji’s largest reserve (60.6 km²). Using stereo baited remote underwater video systems (stereo-BRUVs), we found that the abundance and biomass of sharks was approximately two and four times greater in shallow and deep locations, respectively, within the Namena reserve compared to adjacent fished areas. The greater abundance and biomass of reef sharks inside Namena is likely a result of greater prey availability rather than protection from fishing. This study demonstrates that marine reserves can benefit sharks.     

Observations on Abundance of Bluntnose Sixgill Sharks,Hexanchus griseus,in an Urban Waterway in Puget Sound, 2003-2005

The bluntnose sixgill shark, Hexanchus griseus, is a widely distributed but poorly understood large, apex predator. Anecdotal reports of diver-shark encounters in the late 1990’s and early 2000’s in the Pacific Northwest stimulated interest in the normally deep-dwelling shark and its presence in the shallow waters of Puget Sound. Analysis of underwater video documenting sharks at the Seattle Aquarium’s sixgill research site in Elliott Bay and mark-resight techniques were used to answer research questions about abundance and seasonality. Seasonal changes in relative abundance in Puget Sound from 2003–2005 are reported here. At the Seattle Aquarium study site, 45 sixgills were tagged with modified Floy visual marker tags, along with an estimated 197 observations of untagged sharks plus 31 returning tagged sharks, for a total of 273 sixgill observations recorded. A mark-resight statistical model based on analysis of underwater video estimated a range of abundance from a high of 98 sharks seen in July of 2004 to a low of 32 sharks seen in March of 2004. Both analyses found sixgills significantly more abundant in the summer months at the Seattle Aquarium’s research station.     

A review of shark agonistic displays: comparison of display features and implications for shark–human interactions

Agonistic displays in 23 species of sharks of six families are described and illustrated. These displays are reviewed in terms of ethological concepts and shark hydrodynamic models. Shark agonistic displays feature many common elements rendering them readily distinguishable from normal swimming and pseudodisplays caused by sharksucker irritation. Shark agonistic displays are most readily elicited by rapid, direct diver approach when food is absent and potential escape routes restricted. Such displays appear to be motivated by defence of self or the immediately surrounding space rather than defence of territory or resources. Costs and benefits of display versus attack in shark–shark and shark–diver contests are evaluated using payoff matrices and optimal strategies are identified. Shark–human interactions are modelled in terms of a system of nested critical approach distances. For divers faced with a displaying shark, responses which may decrease the likelihood of defensive attack are suggested. Recommendations for future work on shark agonistic behaviour are offered.     

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.     

Segregation and foraging ecology of whale sharks, Rhincodon typus, in the southwestern Gulf of California

Sharks segregate by sex and size, but few studies have attempted to explain such behaviors. To address this, we examined aggregations and the foraging ecology of whale sharks in Bahía de La Paz (BLP) with aerial and ship surveys and direct observation. Zooplankton abundance and composition, and hydrographic conditions were analyzed in relation to whale shark occurrence to explore underlying factors causing segregations. We observed large aggregations of juveniles (<9 m total length, TL) inshore, comprised by 60 % male individuals, and small aggregations of adults (>9 m TL) offshore, composed of 84 % females. Juvenile sharks were associated to turbid shallow waters in BLP, where they performed stationary and dynamic suction feeding on dense copepod swarms. Adults occurred in oceanic waters and fed by ram-filtering on diffuse patches of euphausiids, with no association to oceanographic conditions. Such segregation may be advantageous to juvenile R. typus utilizing shallow coastal waters to find abundant preferred prey needed for their fast growth rates. Our studies suggest that the main driving forces of whale shark segregation by sex and size in BLP may be diet preference for juveniles and habitat preference for adult sharks.     

Utility of mesohabitat features for determining habitat associations of subadult sharks in Georgia’s estuaries

We examined the affects of selected water quality variables on the presence of subadult sharks in six of nine Georgia estuaries. During 231 longline sets, we captured 415 individuals representing nine species. Atlantic sharpnose shark (Rhizoprionodon terranovae), bonnethead (Sphyrna tiburo), blacktip shark (Carcharhinus limbatus) and sandbar shark (C. plumbeus) comprised 96.1% of the catch. Canonical correlation analysis (CCA) was used to assess environmental influences on the assemblage of the four common species. Results of the CCA indicated Bonnethead Shark and Sandbar Shark were correlated with each other and with a subset of environmental variables. When the species occurred singly, depth was the defining environmental variable; whereas, when the two co-occurred, dissolved oxygen and salinity were the defining variables. Discriminant analyses (DA) were used to assess environmental influences on individual species. Results of the discriminant analyses supported the general CCA findings that the presence of bonnethead and sandbar shark were the only two species that correlated with environmental variables. In addition to depth and dissolved oxygen, turbidity influenced the presence of sandbar shark. The presence of bonnethead shark was influenced primarily by salinity and turbidity. Significant relationships existed for both the CCA and DA analyses; however, environmental variables accounted for <16% of the total variation in each. Compared to the environmental variables we measured, macrohabitat features (e.g., substrate type), prey availability, and susceptibility to predation may have stronger influences on the presence and distribution of subadult shark species among sites.     

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.     

A Comparison of Spatial and Movement Patterns between Sympatric Predators: Bull Sharks (Carcharhinus leucas) and Atlantic Tarpon (Megalops atlanticus)

Background

Predators can impact ecosystems through trophic cascades such that differential patterns in habitat use can lead to spatiotemporal variation in top down forcing on community dynamics. Thus, improved understanding of predator movements is important for evaluating the potential ecosystem effects of their declines.

Methodology/Principal Findings

We satellite-tagged an apex predator (bull sharks, Carcharhinus leucas) and a sympatric mesopredator (Atlantic tarpon, Megalops atlanticus) in southern Florida waters to describe their habitat use, abundance and movement patterns. We asked four questions: (1) How do the seasonal abundance patterns of bull sharks and tarpon compare? (2) How do the movement patterns of bull sharks and tarpon compare, and what proportion of time do their respective primary ranges overlap? (3) Do tarpon movement patterns (e.g., straight versus convoluted paths) and/or their rates of movement (ROM) differ in areas of low versus high bull shark abundance? and (4) Can any general conclusions be reached concerning whether tarpon may mitigate risk of predation by sharks when they are in areas of high bull shark abundance?

Conclusions/Significance

Despite similarities in diet, bull sharks and tarpon showed little overlap in habitat use. Bull shark abundance was high year-round, but peaked in winter; while tarpon abundance and fishery catches were highest in late spring. However, presence of the largest sharks (>230 cm) coincided with peak tarpon abundance. When moving over deep open waters (areas of high shark abundance and high food availability) tarpon maintained relatively high ROM in directed lines until reaching shallow structurally-complex areas. At such locations, tarpon exhibited slow tortuous movements over relatively long time periods indicative of foraging. Tarpon periodically concentrated up rivers, where tracked bull sharks were absent. We propose that tarpon trade-off energetic costs of both food assimilation and osmoregulation to reduce predation risk by bull sharks.     

Seasonal variation in occurrence of the sevengill shark,Notorynchus cepedianus,in two inshore habitats of southern New Zealand

Despite being a large, relatively abundant predator, the distribution and seasonal occurrence of the broadnose sevengill shark, Notorynchus cepedianus, in New Zealand is poorly understood. During 71 sampling trips conducted from July 2013 to May 2015, sharks were attracted to coastal sampling sites in southern New Zealand at ōtākou/Otago Harbour and Te Whaka ā Te Wera/Paterson Inlet, Stewart Island, using chum. Using a logistic regression model, water temperature was identified as a key predictor of encountering sevengill sharks. In addition, location, cloud cover and sea state were also identified as influential predictors. At ōtākou, a clear seasonal pattern of sevengill shark sightings emerged. Sharks were detected on 86% of survey trips in summer, whilst no sharks were detected in winter or spring. At Te Whaka ā Te Wera, sharks were sighted throughout all seasons, but a decline in shark encounters occurred during winter. This study represents the first systematic data on seasonal habitat use by sevengill sharks in New Zealand.     

Quantifying Shark Distribution Patterns and Species-Habitat Associations: Implications of Marine Park Zoning

Quantifying shark distribution patterns and species-specific habitat associations in response to geographic and environmental drivers is critical to assessing risk of exposure to fishing, habitat degradation, and the effects of climate change. The present study examined shark distribution patterns, species-habitat associations, and marine reserve use with baited remote underwater video stations (BRUVS) along the entire Great Barrier Reef Marine Park (GBRMP) over a ten year period. Overall, 21 species of sharks from five families and two orders were recorded. Grey reef Carcharhinus amblyrhynchos, silvertip C. albimarginatus, tiger Galeocerdo cuvier, and sliteye Loxodon macrorhinus sharks were the most abundant species (>64% of shark abundances). Multivariate regression trees showed that hard coral cover produced the primary split separating shark assemblages. Four indicator species had consistently higher abundances and contributed to explaining most of the differences in shark assemblages: C. amblyrhynchos, C. albimarginatus, G. cuvier, and whitetip reef Triaenodon obesus sharks. Relative distance along the GBRMP had the greatest influence on shark occurrence and species richness, which increased at both ends of the sampling range (southern and northern sites) relative to intermediate latitudes. Hard coral cover and distance across the shelf were also important predictors of shark distribution. The relative abundance of sharks was significantly higher in non-fished sites, highlighting the conservation value and benefits of the GBRMP zoning. However, our results also showed that hard coral cover had a large effect on the abundance of reef-associated shark species, indicating that coral reef health may be important for the success of marine protected areas. Therefore, understanding shark distribution patterns, species-habitat associations, and the drivers responsible for those patterns is essential for developing sound management and conservation approaches.