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The white shark (Carcharodon carcharias) is a wide-ranging apex predator in the northeastern Pacific (NEP). Electronic tagging has demonstrated that white sharks exhibit a regular migratory pattern, occurring at coastal sites during the late summer, autumn and early winter and moving offshore to oceanic habitats during the remainder of the year, although the purpose of these migrations remains unclear. The purpose of this study was to use stable isotope analysis (SIA) to provide insight into the trophic ecology and migratory behaviors of white sharks in the NEP. Between 2006 and 2009, 53 white sharks were biopsied in central California to obtain dermal and muscle tissues, which were analyzed for stable isotope values of carbon (δ(13)C) and nitrogen (δ(15)N). We developed a mixing model that directly incorporates movement data and tissue incorporation (turnover) rates to better estimate the relative importance of different focal areas to white shark diet and elucidate their migratory behavior. Mixing model results for muscle showed a relatively equal dietary contribution from coastal and offshore regions, indicating that white sharks forage in both areas. However, model results indicated that sharks foraged at a higher relative rate in coastal habitats. There was a negative relationship between shark length and muscle δ(13)C and δ(15)N values, which may indicate ontogenetic changes in habitat use related to onset of maturity. The isotopic composition of dermal tissue was consistent with a more rapid incorporation rate than muscle and may represent more recent foraging. Low offshore consumption rates suggest that it is unlikely that foraging is the primary purpose of the offshore migrations. These results demonstrate how SIA can provide insight into the trophic ecology and migratory behavior of marine predators, especially when coupled with electronic tagging data.

Elucidating how mobile ocean predators utilize the pelagic environment is vital to understanding the dynamics of oceanic species and ecosystems. Pop-up archival transmitting (PAT) tags have emerged as an important tool to describe animal migrations in oceanic environments where direct observation is not feasible. Available PAT tag data, however, are for the most part limited to geographic position, swimming depth and environmental temperature, making effective behavioral observation challenging. However, novel analysis approaches have the potential to extend the interpretive power of these limited observations. Here we developed an approach based on clustering analysis of PAT daily time-at-depth histogram records to distinguish behavioral modes in white sharks (Carcharodon carcharias). We found four dominant and distinctive behavioral clusters matching previously described behavioral patterns, including two distinctive offshore diving modes. Once validated, we mapped behavior mode occurrence in space and time. Our results demonstrate spatial, temporal and sex-based structure in the diving behavior of white sharks in the northeastern Pacific previously unrecognized including behavioral and migratory patterns resembling those of species with lek mating systems. We discuss our findings, in combination with available life history and environmental data, and propose specific testable hypotheses to distinguish between mating and foraging in northeastern Pacific white sharks that can provide a framework for future work. Our methodology can be applied to similar datasets from other species to further define behaviors during unobservable phases.

Satellite telemetry studies of 20 adult and sub-adult white sharks (360-530 cm estimated total length (TL)) in the eastern North Pacific during 1999-2005 revealed long distance seasonal migrations from the coast of California to an offshore focal area 2,500 km west of the Baja Peninsula, as well as the Hawaii Islands. Three tags were recovered allowing detailed behavioral analyses, including one shark's migration cycle from the coast to the offshore focal area and back. While near pinniped rookeries in autumn and winter, sharks avoided the surface and used water to 50 m depth, consistent with a silhouette-based hunting strategy. Offshore migrations were initiated during November-March and followed periods of decreasing pinniped abundance. Migrations were highly directed, taking 23 +/- 5 days to reach the offshore focal area along similar paths among sharks and years, defining a migration corridor. Sharks exhibited a broad depth distribution (0-644 m) in the offshore focal area, and remained there for up to 167 days during spring and summer, though primary productivity and fishery data suggest that forage resources are scarcer there than in other regions of the eastern North Pacific. Archival data from one shark revealed intensive oscillatory movements while in the offshore focal area, a behavior that may be related to foraging or mating. Sharks traveling to Hawaii remained near the islands up to 122 days, potentially feeding on pelagic fishes and marine mammals that concentrate around the islands. [PUBLICATION ABSTRACT]

Quantifying life history parameters of marine top predators such as sharks is challenging, as observations are difficult. However, these parameters are critical for accurate population assessments and understanding of population dynamics. Using mark recapture observations at white shark foraging aggregation sites, we tested for differences in survival between sexes and estimated apparent survival for sub-adult and adult white sharks in neritic waters off central California. We used six years of mark-recapture data and a model that accounted for imperfect detection and imperfect sex assignment. Empirical information based on direct observations suggests that there are no sex-specific or temporal differences in survival during the study period and that survival was estimated to be 0.90; SE = 0.04. Additionally, after animals whose sex was unknown throughout the study period were probabilistically assigned to sex, the ratio in this sample is estimated to be 2.1 males for every female observed. This estimated ratio is lower than the observed ratio of 3:1. We demonstrate that the estimated capture probability for males was roughly twice as high as that for females (0.41, SE = 0.06 and 0.19, SE = 0.07 respectively). Together these results suggest (1) that the sex ratio is uneven but not as skewed as uncorrected observation data would suggest and (2) that unequal mortality in older age classes are not the cause of the observed sex bias but more likely results from disparate mortality earlier in life or differences in behavior. Future research is needed to explore the potential causes of the observed sex bias.

Background Motion detecting archival data loggers such as accelerometers have become increasingly important in animal biotelemetry and offer unique insights into animal behavior, energetics, and kinematics. However, challenges remain for successful deployment and interpretation of data from captive and wild animals. Accelerometer sensors require being packaged in an archival tag that has a firm attachment in a fixed (known) orientation to accurately measure the relevant motion of the animal. This requirement can lead to handling stress and attachment techniques that can affect the tagged animal’s natural behavior and welfare, and lead to behavioral artifacts in the data. Accelerometer data also require careful interpretation to correctly identify behavioral events of interest such as foraging. For endothermic species, changes in stomach temperature can produce temperature signatures indicative of foraging events. In this paper, we present a novel method for recording foraging events in free-swimming white sharks. Methods We used a combination of accelerometer loggers and pop-up archival transmitting (PAT) tags (MK10, Wildlife Computers) to examine the feeding and kinematics of white sharks ( Carcharodon carcharias ) in the wild. We validated feeding results using a captive juvenile white shark where controlled feeding experiments could be conducted in an aquarium setting at the Monterey Bay Aquarium. We fed data logger instrument packages to eight free-swimming white sharks. Deployment durations ended naturally when the package was regurgitated and ranged from 2 to 12 days. While inside the stomach, the orientation of the data logger package was arbitrary and resulted in slow shifting over time, a challenge for normal analysis routines. We present one of these datasets to illustrate a novel methodology for calibrating accelerometer orientation, and evaluate the utility of resulting data. Results We obtained accurate accelerometer measurements including surge, heave, and sway from data loggers with shifting orientation through data post-processing. We measured consistent dips in stomach temperature followed by a steady prolonged heating during controlled feeding events in a captive white shark. Similar thermal signatures identified in wild white shark records confirmed feeding events while acceleration data characterized the associated prey capture behavior. Conclusions We provided proof of concept for a novel and non-invasive technique for accelerometer data logger deployment extending the possibilities for their use in bio-logging. The placement of data-logging tags in the stomachs of endothermic white sharks produced high quality tri-axial acceleration data in addition to stomach temperature data capable of detecting feeding events. The technique has the potential for distinguishing between attempted and successful foraging events. This approach to accelerometer coordinate correction could be applied in other systems where logger orientation is unknown or changes after deployment.

Mark-recapture techniques can be used to estimate white shark ( Carcharodon carcharias ) population abundance. These frameworks are based on assumptions that marks are conserved and animals are present at the sampling location over the entire duration of the study. Though these assumptions have been validated across short-time scales for white sharks, long-term studies of population trends are dependent on these assumptions being valid across longer periods. We use 22 years of photographic data from aggregation sites in central California to support the use of dorsal fin morphology as long-term individual identifiers. We identified five individuals over 16–22 years, which support the use of dorsal fins as long-time individual identifiers, illustrate strong yearly site fidelity to coastal aggregation sites across extended time periods (decades), and provide the first empirical validation of white shark longevity >22 years. These findings support the use of fin morphology in mark-recapture frameworks for white sharks.

Marine animals often move beyond national borders and exclusive economic zones resulting in a need for trans-boundary management spanning multiple national jurisdictions. Highly migratory fish vulnerable to over-exploitation require protections at international level, as exploitation practices can be disparate between adjacent countries and marine jurisdictions. In this study we collaboratively conducted an analysis of white shark connectivity between two main aggregation regions with independent population assessment and legal protection programs; one off central California, USA and one off Guadalupe Island, Mexico. We acoustically tagged 326 sub-adult and adult white sharks in central California (n=210) and in Guadalupe Island (n=116) with acoustic transmitters between 2008-2019. Of the 326 tagged white sharks, 30 (9.20%) individuals were detected at both regions during the study period. We used a Bayesian implementation of logistic regression with a binomial distribution to estimate the effect of sex, maturity, and tag location to the response variable of probability of moving from one region to the other. While nearly one in ten individuals in our sample were detected in both regions over the study period, the annual rate of trans-regional movement was low (probability of movement = 0.015 yr -1 , 95% credible interval = 0.002, 0.061). Sub-adults were more likely than adults to move between regions and sharks were more likely to move from Guadalupe Island to central California, however, sex, and year were not important factors influencing movement. This first estimation of demographic-specific trans-regional movement connecting US and Mexico aggregations with high seasonal site fidelity represents an important step to future international management and assessment of the northeastern Pacific white shark population as a whole.

Over the last two decades, satellite tagging of adult and sub-adult white sharks Carcharodon carcharias off the west coast of North America has revealed a predictable onshore-offshore migratory cycle. Our current understanding of the vertical movements exhibited by white sharks while in their coastal foraging phase in the California Current, however, remains limited. Here, we used recovered datasets from 31 archival satellite tags to quantify vertical habitat use. Tags were deployed on individuals between 2000 and 2018 and recorded depth and temperature data at continuous 1–120 s intervals before being recovered up to a year after deployments. Four satellite-tagged individuals were concurrently tagged with acoustic tags, providing precise location data when detected by acoustic receivers that allowed us to explore how reported vertical habitat use varied spatially. While in the coastal shelf waters, white sharks moved at a mean depth ± SD of 14.3 ± 4.0 m and occupied significantly deeper depths during the day than the night. High individual, temporal and spatial variation was evident in vertical movements, while consistent diel and lunar effects emphasized the importance of light-level driving vertical behavior around hunting sites. The vertical movement behaviors reported here provide knowledge of how white sharks may directly and indirectly interact with their mammalian prey in a dynamic three-dimensional system during their capital foraging phase. Temporal patterns in vertical behavior, for instance, indicated that surface waters during early morning hours are the riskiest place for prey. Combining these novel findings with higher-resolution biologging techniques in future studies will allow us to further contextualize fine-scale vertical movement behaviors of white sharks and examine the specific foraging events that could not yet be isolated in the tagging data.

This paper presents information on the movements of white sharks, Carcharodon carcharias, at the South Farallon Islands (SFI), central California. Acoustic telemetry techniques provided preliminary data on the diurnal space utilization, movement patterns and swimming depths of four white sharks, ranging from approximately 3.7 to 4.9 m in length. Sharks swam within about 10 m of the bottom to depths of approximately 30 m, but in deeper water they tended to stray more from the bottom. Activity spaces for time periods tracked ranged from 1.84 to 9.15 km super(2). Indications are that an inverse relationship exists between length and activity space. During the time tracked, larger individuals swam within particular areas around the islands whereas smaller individuals did not restrict their movements in the same manner. Values of a site attachment index were inversely related to length for all sharks tracked. The site attachment indices, apparent inverse relationship between total length and activity space and observations on telemetered and other known individuals support a hypothesis that larger sharks possess site fidelity in their search for prey at SFI, within and between years. With the high frequency of predation by white sharks on juvenile northern elephant seals at SFI in the fall, the majority of the sharks' movements are probably related to their search for these pinniped prey. These data provide preliminary evidence that white sharks at SFI may search for prey by swimming in a particular area over a number of days or weeks, traversing the area in a manner which maximizes coverage, and swimming close to the bottom or at a distance far enough from the surface to remain cryptic from prey.