Scholarly Work - Ecology

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    Diverse habitats shape the movement ecology of a top marine predator, the white shark Carcharodon carcharias
    (Wiley, 2024-04) Jewell, Oliver J. D.; Chappel, Taylor K.; Jorgensen, Salvador J.; Kanive, Paul; Moxely, Jerry H.; Tweedley, James R.; Anderson, Scot; Block, Barbara A.; Gleiss, Adrian C.
    An animal's movement is influenced by a plethora of internal and external factors, leading to individual- and habitat-specific movement characteristics. This plasticity is thought to allow individuals to exploit diverse environments efficiently. We tested whether the movement characteristics of white sharks Carcharodon carcharias differ across ontogeny and among habitats along the coast of Central California. In doing so, we elucidate how changes in internal state (physiological changes coinciding with body size) and external environments (differing seascapes and/or diel phases) shape the movement of this globally distributed predator. Twenty-one white sharks, from small juveniles to large adults, were equipped with motion-sensitive biologging tags at four contrasting seascapes: two islands, a headland, and an inshore cove. From multisensor biologging data, 20 metrics characterizing movement (i.e., depth use, vertical velocities, activity, turning rates, and bursting events) were derived and subjected to multivariate analyses. Movement characteristics were most different across seascapes, followed by ontogeny and diel phase. Juvenile sharks, which were only encountered at the cove, displayed the most distinct movement characteristics. Sharks at this seascape remained close to the shore traveling over smaller areas, shallower depth ranges, and with lower levels of tail beat frequencies, when corrected for size, than sub-adult and adult sharks tagged elsewhere. Distinct tortuous daytime versus linear nighttime horizontal movements were recorded from sharks at island seascapes but not from those at the headland or inshore cove. At the offshore islands, the linear nighttime swimming patterns coincided with repeated dives to and from deeper water. The availability of prey and access to deeper water are likely drivers of the differences in movement characteristics described, with varying demographics of pinniped prey found at the subadult and adult aggregation areas and juvenile sharks being piscivorous and their habitat neither adjacent to pinniped haul out areas nor deeper water. This study demonstrates plasticity in the movements of a top predator, which adapts its routine to suit the habitat it forages within.
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    Connectivity between white shark populations off Central California, USA and Guadalupe Island, Mexico
    (Frontiers Media SA, 2023-07) Kanive, Paul E.; Rotella, Jay J.; Chapple, Taylor K.; Anderson, Scot D.; Hoyos-Padilla, Mauricio; Klimley, Abbott Peter; Galván-Magaña, Felipe; Andrzejaczek, Samantha; Block, Barbara A.; Jorgensen, Salvador J.
    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.
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    Estimates of regional annual abundance and population growth rates of white sharks off central California
    (Elsevier BV, 2021-05) Kanive, Paul E.; Rotella, Jay J.; Chapple, Taylor K.; Anderson, Scot D.; White, Timothy D.; Block, Barbara A.; Jorgensen, Salvador J.
    Determining population trends is critical for evaluating management actions and prioritizing species protections. In this study, we used empirical data to produce an estimate of the population trend for sub-adult and adult white sharks in central California. We used the unique dorsal fin morphology to build a mark-recapture data set in a modified Jolly-Seber model (POPAN formulation) to estimate annual abundance and then investigate population growth rates using parametric bootstrapping methods for sub-adult and adult sharks (males and females). For all demographic groups combined, we found equivocal evidence for a positive regional population growth (λ = 1.07 (95% CI = 0.91 to 1.23)). However, sex- and size-specific population growth rate estimates provided some evidence of population increases for reproductively mature males (λ = 1.06 (95% CI = 0.99 to 1.13)) and females (λ = 1.06 (95% CI = 0.95 to 1.17)). For sub-adult male and female white sharks, point estimates of λ were positive but uncertainty prevents strong inference (λ = 1.07 (95% CI = 0.85 to 1.29)) and (λ = 1.08 (95% CI = 0.88 to 1.28)), respectively. Our findings of a potential increase in reproductive-aged white sharks in central California may be a result of regional fluxes in density or attributed in part to current protection efforts and subsequent increase in abundance of pinnipeds as well as reduced gill-net fisheries mortality of juveniles. A trend estimate for the entire northeastern Pacific will require obtaining similar data across known aggregation areas along the west coast of North America.
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    Size-specific apparent survival rate estimates of white sharks using mark–recapture models
    (Canadian Science Publishing, 2019-02) Kanive, Paul E.; Rotella, Jay J.; Jorgensen, Salvador J.; Chapple, Taylor K.; HInes, James E.; Anderson, Scot D.; Block, Barbara A.
    Abstract: For species that exist at low abundance or are otherwise difficult to study, it is challenging to estimate vital rates such as survival and fecundity and common to assume that survival rates are constant across ages and sexes. Population assessments based on overly simplistic vital rates can lead to erroneous conclusions. We estimated sex- and length-based annual apparent survival rates for white sharks (Carcharodon carcharias). We found evidence that annual apparent survival differed over ontogeny in a system with competitive foraging aggregations, from 0.63 (standard error (SE) = 0.08) for newly recruiting subadults to 0.95 (SE = 0.02) for the largest sharks. Our results reveal a potential challenge to ontogenetic recruitment in a long-lived, highly mobile top marine predator, as survival rates for subadult white sharks may be lower than previously assumed. Alternatively, younger and competitively inferior individuals may be forced to permanently emigrate from primary foraging sites. This study provides new methodology for estimating apparent survival as a function of diverse covariates by capture–recapture study, including when sex assignment is uncertain. Résumé : Pour les espèces qui existent en faible abondance ou dont l’étude présente par ailleurs un défi, il est difficile d’estimer des indices vitaux comme la survie et la fécondité et il est couramment présumé que les taux de survie ne varient pas selon le sexe et le groupe d’âge. Des évaluations de populations reposant sur des indices vitaux trop simplistes peuvent mener à des conclusions erronées. Nous avons estimé les taux de survie annuels apparents en fonction du sexe et de la longueur pour de grands requins blancs (Carcharodon carcharias). Nous avons relevé des preuves de variation du taux de survie annuel apparent au fil de l’ontogénie dans un système caractérisé par des regroupements concurrents d’individus en quête de nourriture, ce taux allant de 0,63 (l’écart-type (ÉT) = 0,08) pour les individus subadultes récemment recrutés à 0,95 (ÉT = 0,02) pour les requins les plus grands. Nos résultats révèlent une difficulté potentielle en ce qui concerne le recrutement ontogénique chez un prédateur marin de niveau trophique supérieur très mobile et longévif, puisque les taux de survie de grands requins blancs subadultes pourraient être plus faibles que présumés auparavant. Une autre explication est que les individus plus jeunes ou moins concurrentiels pourraient être forcés d’émigrer des meilleurs sites d’approvisionnement de manière permanente. L’étude présente une nou-velle méthodologie pour estimer la survie apparente en fonction de différentes variables reliées, par une approche de capture–recapture, y compris pour les cas où l’affectation du sexe est incertaine. [Traduit par la Rédaction]
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    Estimates of regional annual abundance and population growth rates of white sharks off central California
    (2021-05) Kanive, Paul E.; Rotella, Jay J.; Chapple, Taylor K.; Anderson, Scot D.; White, Timothy D.; Block, Barbara A.; Jorgensen, Salvador J.
    Determining population trends is critical for evaluating management actions and prioritizing species protections. In this study, we used empirical data to produce an estimate of the population trend for sub-adult and adult white sharks in central California. We used the unique dorsal fin morphology to build a mark-recapture data set in a modified Jolly-Seber model (POPAN formulation) to estimate annual abundance and then investigate population growth rates using parametric bootstrapping methods for sub-adult and adult sharks (males and females). For all demographic groups combined, we found equivocal evidence for a positive regional population growth (λ = 1.07 (95% CI = 0.91 to 1.23)). However, sex- and size-specific population growth rate estimates provided some evidence of population increases for reproductively mature males (λ = 1.06 (95% CI = 0.99 to 1.13)) and females (λ = 1.06 (95% CI = 0.95 to 1.17)). For sub-adult male and female white sharks, point estimates of λ were positive but uncertainty prevents strong inference (λ = 1.07 (95% CI = 0.85 to 1.29)) and (λ = 1.08 (95% CI = 0.88 to 1.28)), respectively. Our findings of a potential increase in reproductive-aged white sharks in central California may be a result of regional fluxes in density or attributed in part to current protection efforts and subsequent increase in abundance of pinnipeds as well as reduced gill-net fisheries mortality of juveniles. A trend estimate for the entire northeastern Pacific will require obtaining similar data across known aggregation areas along the west coast of North America.
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    Killer whales redistribute white shark foraging pressure on seals
    (2019-04) Jorgensen, Salvador J.; Anderson, Scot; Ferretti, Francesco; Tietz, James R.; Chapple, Taylor; Kanive, Paul; Bradley, Russell W.; Moxley, Jerry H.; Block, Barbara A.
    Predatory behavior and top-down effects in marine ecosystems are well-described, however, intraguild interactions among co-occurring marine top predators remain less understood, but can have far reaching ecological implications. Killer whales and white sharks are prominent upper trophic level predators with highly-overlapping niches, yet their ecological interactions and subsequent effects have remained obscure. Using long-term electronic tagging and survey data we reveal rare and cryptic interactions between these predators at a shared foraging site, Southeast Farallon Island (SEFI). In multiple instances, brief visits from killer whales displaced white sharks from SEFI, disrupting shark feeding behavior for extended periods at this aggregation site. As a result, annual predations of pinnipeds by white sharks at SEFI were negatively correlated with close encounters with killer whales. Tagged white sharks relocated to other aggregation sites, creating detectable increases in white shark density at Ano Nuevo Island. This work highlights the importance of risk effects and intraguild relationships among top ocean predators and the value of long-term data sets revealing these consequential, albeit infrequent, ecological interactions.
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    Estimating apparent survival of sub-adult and adult white sharks (Carcharodon carcharias) in central California using mark-recapture methods
    (2015-04) Kanive, Paul Edward, Jr.; Rotella, Jay J.; Jorgensen, Salvador J.; Chapple, Taylor K.; Anderson, Scot D.; Klimley, A. Peter; Block, Barbara A.
    Quantifying life history parameters of marine top predators is challenging, as observations are difficult and uncertainty in sex assignment can confound the determination of sex specific parameters. 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 6 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.
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