Age and Growth of Elasmobranchs and Applications to Fisheries Management and Conservation in the Northeast Pacific Ocean.

In addition to being an academic endeavour, the practical purpose of conducting age and growth studies on fishes is to provide biological data to stock assessment scientists and fisheries managers so they may better understand population demographics and manage exploitation rates. Age and size data are used to build growth models, which are a critical component of stock assessments. Though age determination of elasmobranchs in the northeast Pacific Ocean (NEP) began in the 1930s, the field has evolved substantially in recent years, allowing scientists to incorporate age data into assessments for more species than ever before. Owing to the highly diverse biology of this group of fishes, each species has its own set of challenges with regard to age determination. Age determination methods typically rely on semicalcified hard structures that form regular growth patterns; however, the structure selected and preparation method used is often species specific. New staining techniques have improved the ability to assess age and improve ageing precision for some species, and advances in microchemical methods have allowed for independent means of estimating age and validating age determination accuracy. Here we describe current age determination methods for NEP elasmobranchs. While the library of available techniques is increasing, there are still some NEP species for which reliable ageing methods have yet to be defined; we discuss these challenges and potential avenues of future research. Finally, we conclude by describing how age estimates are used in growth models and subsequently in stock assessments of selected NEP elasmobranchs.

Comparative analyses of animal-tracking data reveal ecological significance of endothermy in fishes.

Despite long evolutionary separations, several sharks and tunas share the ability to maintain slow-twitch, aerobic red muscle (RM) warmer than ambient water. Proximate causes of RM endothermy are well understood, but ultimate causes are unclear. Two advantages often proposed are thermal niche expansion and elevated cruising speeds. The thermal niche hypothesis is generally supported, because fishes with RM endothermy often exhibit greater tolerance to broad temperature ranges. In contrast, whether fishes with RM endothermy cruise faster, and achieve any ecological benefits from doing so, remains unclear. Here, we compiled data recorded by modern animal-tracking tools for a variety of free-swimming marine vertebrates. Using phylogenetically informed allometry, we show that both cruising speeds and maximum annual migration ranges of fishes with RM endothermy are 2-3 times greater than fishes without it, and comparable to nonfish endotherms (i.e., penguins and marine mammals). The estimated cost of transport of fishes with RM endothermy is twice that of fishes without it. We suggest that the high energetic cost of RM endothermy in fishes is offset by the benefit of elevated cruising speeds, which not only increase prey encounter rates, but also enable larger-scale annual migrations and potentially greater access to seasonally available resources.

Stable isotope analysis of vertebrae reveals ontogenetic changes in habitat in an endothermic pelagic shark.

Ontogenetic changes in habitat are driven by shifting life-history requirements and play an important role in population dynamics. However, large portions of the life history of many pelagic species are still poorly understood or unknown. We used a novel combination of stable isotope analysis of vertebral annuli, Bayesian mixing models, isoscapes and electronic tag data to reconstruct ontogenetic patterns of habitat and resource use in a pelagic apex predator, the salmon shark (Lamna ditropis). Results identified the North Pacific Transition Zone as the major nursery area for salmon sharks and revealed an ontogenetic shift around the age of maturity from oceanic to increased use of neritic habitats. The nursery habitat may reflect trade-offs between prey availability, predation pressure and thermal constraints on juvenile endothermic sharks. The ontogenetic shift in habitat coincided with a reduction of isotopic niche, possibly reflecting specialization upon particular prey or habitats. Using tagging data to inform Bayesian isotopic mixing models revealed that adult sharks primarily use neritic habitats of Alaska yet receive a trophic subsidy from oceanic habitats. Integrating the multiple methods used here provides a powerful approach to retrospectively study the ecology and life history of migratory species throughout their ontogeny.

A re-evaluation of the size of the white shark (Carcharodon carcharias) population off California, USA.

White sharks are highly migratory and segregate by sex, age and size. Unlike marine mammals, they neither surface to breathe nor frequent haul-out sites, hindering generation of abundance data required to estimate population size. A recent tag-recapture study used photographic identifications of white sharks at two aggregation sites to estimate abundance in "central California" at 219 mature and sub-adult individuals. They concluded this represented approximately one-half of the total abundance of mature and sub-adult sharks in the entire eastern North Pacific Ocean (ENP). This low estimate generated great concern within the conservation community, prompting petitions for governmental endangered species designations. We critically examine that study and find violations of model assumptions that, when considered in total, lead to population underestimates. We also use a Bayesian mixture model to demonstrate that the inclusion of transient sharks, characteristic of white shark aggregation sites, would substantially increase abundance estimates for the adults and sub-adults in the surveyed sub-population. Using a dataset obtained from the same sampling locations and widely accepted demographic methodology, our analysis indicates a minimum all-life stages population size of >2000 individuals in the California subpopulation is required to account for the number and size range of individual sharks observed at the two sampled sites. Even accounting for methodological and conceptual biases, an extrapolation of these data to estimate the white shark population size throughout the ENP is inappropriate. The true ENP white shark population size is likely several-fold greater as both our study and the original published estimate exclude non-aggregating sharks and those that independently aggregate at other important ENP sites. Accurately estimating the central California and ENP white shark population size requires methodologies that account for biases introduced by sampling a limited number of sites and that account for all life history stages across the species' range of habitats.

White shark offshore habitat: a behavioral and environmental characterization of the eastern Pacific shared offshore foraging area.

BACKGROUND: Although much is known about the behavior of white sharks in coastal regions, very little is known about their vertical movements offshore in the eastern Pacific where they spend up to five months. We provide the first detailed description of the offshore habitat use of white sharks in the eastern North Pacific. METHODOLOGY/PRINCIPAL FINDINGS: This study uses 2-min data from four recovered pop-up satellite archival tags deployed at Guadalupe Island (2002 and 2005). Deployments ranged from 5.4 to 8.2 months. Two predominant vertical patterns were described. The first was a bimodal vertical pattern with time spent at the surface and at depth, which was observed while traveling. The second was a repetitive oscillatory diving mode displayed by sharks in the Shared Offshore Foraging Area (SOFA). For all four datasets the average maximum daily dive depths ranged from 442.5 to 492.8 m and were typically associated with dissolved oxygen concentrations of above 1.7 ml L(-1). Although infrequent, occasional dives to near 1000 m with a minimum temperature of 3.9 degrees C and a minimum O(2) level of 0.3 ml L(-1) were observed. CONCLUSIONS/SIGNIFICANCE: Recovered pop-up satellite tags from Guadalupe Island white sharks advance our understanding of the vertical habitat use of white sharks while offshore. The bimodal vertical pattern during traveling is most likely related to geolocation. The oscillatory dive pattern is likely associated with foraging. While feeding is not documented, foraging is likely occurring in association with the deep scattering layer. Diving depths were not limited by temperature but were constrained by O(2) levels below approximately 1.5 ml L(-1). While oxygen may limit the extent of sharks' vertical movements, it will also impact prey distribution. Consequently, the shallow oxygen minimum zone in the SOFA may act to concentrate prey, thus enhancing foraging opportunities in these oligotrophic waters.

Satellite tagging and cardiac physiology reveal niche expansion in salmon sharks.

Shark populations are declining globally, yet the movements and habitats of most species are unknown. We used a satellite tag attached to the dorsal fin to track salmon sharks (Lamna ditropis) for up to 3.2 years. Here we show that salmon sharks have a subarctic-to-subtropical niche, ranging from 2 degrees to 24 degrees C, and they spend winter periods in waters as cold as 2 degrees to 8 degrees C. Functional assays and protein gels reveal that the expression of excitation-contraction coupling proteins is enhanced in salmon shark hearts, which may underlie the shark's ability to maintain heart function at cold temperatures and their niche expansion into subarctic seas.