Current and future considerations for shark conservation in the Northeast and Eastern Central Pacific Ocean.

Sharks are iconic and ecologically important predators found in every ocean. Because of their ecological role as predators, some considered apex predators, and concern over the stability of their populations due to direct and indirect overfishing, there has been an increasing amount of work focussed on shark conservation, and other elasmobranchs such as skates and rays, around the world. Here we discuss many aspects of current shark science and conservation and the path to the future of shark conservation in the Northeastern and Eastern Central Pacific. We explore their roles in ecosystems as keystone species; the conservation measures and laws in place at the international, national, regional and local level; the conservation status of sharks and rays in the region, fisheries for sharks in the Northcentral Pacific specifically those that target juveniles and the implications to shark conservation; a conservation success story: the recovery of Great White Sharks in the Northeast Pacific; public perceptions of sharks and the roles zoos and aquariums play in shark conservation; and the path to the future of shark conservation that requires bold partnerships, local stakeholders and innovative measures.

Functional genetic diversity in an exploited marine species and its relevance to fisheries management.

The timing of reproduction influences key evolutionary and ecological processes in wild populations. Variation in reproductive timing may be an especially important evolutionary driver in the marine environment, where the high mobility of many species and few physical barriers to migration provide limited opportunities for spatial divergence to arise. Using genomic data collected from spawning aggregations of Pacific herring (Clupea pallasii) across 1600 km of coastline, we show that reproductive timing drives population structure in these pelagic fish. Within a specific spawning season, we observed isolation by distance, indicating that gene flow is also geographically limited over our study area. These results emphasize the importance of considering both seasonal and spatial variation in spawning when delineating management units for herring. On several chromosomes, we detected linkage disequilibrium extending over multiple Mb, suggesting the presence of chromosomal rearrangements. Spawning phenology was highly correlated with polymorphisms in several genes, in particular SYNE2, which influences the development of retinal photoreceptors in vertebrates. SYNE2 is probably within a chromosomal rearrangement in Pacific herring and is also associated with spawn timing in Atlantic herring (Clupea harengus). The observed genetic diversity probably underlies resource waves provided by spawning herring. Given the ecological, economic and cultural significance of herring, our results support that conserving intraspecific genetic diversity is important for maintaining current and future ecosystem processes.

Conclusions: Do we eat them or watch them, or both? Challenges for conservation of sharks in Mexico and the NEP.

Mexico is a country that makes heavy use of the shark populations that inhabit the southern portion of the Northeast Pacific Ocean (NEP). Shark meat has become an essential food source in this country, while shark fins are used to supply traditional Asian markets. In addition to consumptive utilization, charismatic shark species support an ecotourism industry that has gained significance in several tourist resorts across the country. In this concluding chapter, we recap the contents of chapters included in volumes 83 and 85 in the Advances in Marine Biology series. The chapters in these volumes address biodiversity, conservation genetics, trophic ecology, migratory movements, fisheries, and shark ecotourism, allowing us to understand the state of knowledge relevant to human: shark interactions in the Mexican Pacific. We discuss the challenges for the sustainable use and conservation of sharks in the southern NEP and highlight the need for a more holistic management approach that includes economic and social factors. To meet these challenges, we recommend updating the Mexican National Plan of Action for Sharks published in, 2004, such that it may continue serving as a roadmap for the conservation and management of sharks in the southern NEP during the years to come.

Introduction: The sharks of Pacific Mexico and their conservation: Why should we care?

Sharks exhibit a broad range of body forms, occupy diverse ecological niches, and are found in every ocean on earth. They have roles as both predator and prey and have an evolutionary history far longer than that of human beings. As long as humans have interacted with the ocean they have interacted with sharks and, as a result, sharks have featured in the mythology, history, and fisheries of diverse cultures around the world. Only in the last 100 years or less, however, have humans begun to study and fully realize the susceptibility of these animals to intensive fishing pressure, ecological degradation, and other anthropogenic factors. This volume highlights the biodiversity and biological attributes of, and conservation efforts targeted at, populations of sharks that reside in the Northeast Pacific Ocean (NEP) bordering the west coast of Mexico. Together with a pair of companion volumes published in 2017 that focused on the NEP adjacent to the United States and Canada, this volume provide readers a detailed backdrop of shark biology and conservation in the Mexican waters of the Northeast Pacific against which to consider their own actions, and those of resource managers, academics, and educators, as they relate to the long-term conservation of sharks and their relatives.

Population assignment and local adaptation along an isolation-by-distance gradient in Pacific cod (Gadus macrocephalus).

The discernment of populations as management units is a fundamental prerequisite for sustainable exploitation of species. A lack of clear stock boundaries complicates not only the identification of spatial management units, but also the assessment of mixed fisheries by population assignment and mixed stock analysis. Many marine species, such as Pacific cod, are characterized by isolation by distance, showing significant differentiation but no clear stock boundaries. Here, we used restriction-site-associated DNA (RAD) sequencing to investigate population structure and assess power to genetically assign Pacific cod to putative populations of origin. Samples were collected across the species range in the eastern Pacific Ocean, from the Salish Sea to the Aleutian Islands. A total of 6,425 putative biallelic single nucleotide polymorphisms were identified from 276 individuals. We found a strong isolation-by-distance signal along coastlines that mirrored previous microsatellite results and pronounced genetic differentiation between coastal samples and those from the inland waters of the Salish Sea, with no evidence for hybridization between these two populations. Individual assignment success based on two methods was high overall (≥84%) but decreased from south to north. Assignment to geographic location of origin also was successful, with average distance between capture and assignment location of 220 km. Outlier analyses identified more loci potentially under selection along the coast than between Salish Sea and coastal samples, suggesting more diverse adaptation to latitudinal environmental factors than inshore vs. offshore environments. Our results confirm previous observations of sharp genetic differentiation of the Salish Sea population and isolation by distance along the coast, but also highlight the feasibility of using modern genomic techniques to inform stock boundaries and fisheries management in a low FST marine species.

Conclusions: The Future of Shark Management and Conservation in the Northeast Pacific Ocean.

Human interactions with sharks in the Northeast Pacific Ocean (NEP) have occurred for millennia but were largely limited to nearshore encounters as target and nontarget catch in fisheries. The arrival of Spanish explorers in the mid-1500s, followed by subsequent waves of explorers and colonizers from Europe and Russia, did little to change this relationship, until the mid-1800s. As technological advances conferred the ability to exploit marine fish further offshore and in deeper water, substantial fisheries developed and many of these encountered, and sometimes directly targeted, sharks. As these fisheries rose and fell with market demands and fluctuations in the abundance of target species, the collective consciousness of the nations fishing this region came to realize that adequate management plans with clear policy guidance rooted in conservation were crucial to sustaining both biodiversity and abundance of marine resources. With explicitly defined management regions governed by scientifically informed bodies that consider both societal and ecological needs, systems have been in place to manage and conserve marine species, including sharks, for over four decades now in the NEP. While policy evolution has largely limited directed fishing pressure as a threat for most shark species, bycatch is still a concern. Additionally, habitat degradation and destruction, ocean acidification, and global climate change are anticipated to fundamentally alter the ecosystems sharks are an integral part of in coming decades and centuries. Adequate conservation and management of sharks in the NEP, and around the world, moving into this period of uncertainty will rely upon comprehensive, integrated management of the ecosystem rooted in international coordination and cooperation. Far from being an unattainable goal, steps are being made each day to 'move the needle' in this direction-for the benefit of all.

Introduction to Northeast Pacific Shark Biology, Research, and Conservation, Part B.

Sharks are iconic, sometimes apex, predators found in every ocean. Because of their ecological role as predators and concern over the stability of their populations, there has been an increasing amount of work focused on shark conservation around the world in recent decades. The populations of sharks that reside in the Northeast Pacific (NEP) Ocean bordering the west coast of the United States reside in one of the most economically and ecologically important oceanic regions in the world. Volume 78 of Advances in Marine Biology (AMB) is a companion to Volume 77, which focused primarily on NEP shark biodiversity, organismal biology, and ecology. Volume 78 highlights fisheries and the conservation implications of fisheries management; shark population modelling and the conservation impacts of these models given that many life history metrics of NEP sharks necessary to accurately run these models are still unknown; the value of captive sharks to the biology, outreach, and conservation of NEP sharks; and the conservation value of citizen science and shark ecotourism. Together these volumes encapsulate the current state of the knowledge for sharks in the NEP and lay the foundation for protecting, managing, and learning from these species in the face evolving natural conditions and societal opinions.

Introduction to Northeast Pacific Shark Biology, Ecology, and Conservation.

Sharks are iconic, sometimes apex, predators found in every ocean and, as a result, they have featured prominently in the mythology, history, and fisheries of diverse human cultures around the world. Because of their regional significance to fisheries and ecological role as predators, and as a result of concern over long-term stability of their populations, there has been an increasing amount of work focused on shark conservation in recent decades. This volume highlights the biodiversity and biological attributes of, and conservation efforts targeted at, populations of sharks that reside in the Northeast Pacific Ocean bordering the west coast of the United States and Canada, one of the most economically and ecologically important oceanic regions in the world. A companion volume addresses details of fisheries and ecotourism in the same region, as well as delving into the relationship between captive husbandry of sharks and education/outreach efforts aimed at fostering a conservation mindset in the public at large. Together, these volumes provide readers a detailed backdrop against which to consider their own actions, and those of resource managers, academics, and educators, as they relate to the long-term conservation of sharks and their relatives.

Population assessment using multivariate time-series analysis: A case study of rockfishes in Puget Sound.

Estimating a population's growth rate and year-to-year variance is a key component of population viability analysis (PVA). However, standard PVA methods require time series of counts obtained using consistent survey methods over many years. In addition, it can be difficult to separate observation and process variance, which is critical for PVA. Time-series analysis performed with multivariate autoregressive state-space (MARSS) models is a flexible statistical framework that allows one to address many of these limitations. MARSS models allow one to combine surveys with different gears and across different sites for estimation of PVA parameters, and to implement replication, which reduces the variance-separation problem and maximizes informational input for mean trend estimation. Even data that are fragmented with unknown error levels can be accommodated. We present a practical case study that illustrates MARSS analysis steps: data choice, model set-up, model selection, and parameter estimation. Our case study is an analysis of the long-term trends of rockfish in Puget Sound, Washington, based on citizen science scuba surveys, a fishery-independent trawl survey, and recreational fishery surveys affected by bag-limit reductions. The best-supported models indicated that the recreational and trawl surveys tracked different, temporally independent assemblages that declined at similar rates (an average of -3.8% to -3.9% per year). The scuba survey tracked a separate increasing and temporally independent assemblage (an average of 4.1% per year). Three rockfishes (bocaccio, canary, and yelloweye) are listed in Puget Sound under the US Endangered Species Act (ESA). These species are associated with deep water, which the recreational and trawl surveys sample better than the scuba survey. All three ESA-listed rockfishes declined as a proportion of recreational catch between the 1970s and 2010s, suggesting that they experienced similar or more severe reductions in abundance than the 3.8-3.9% per year declines that were estimated for rockfish populations sampled by the recreational and trawl surveys.

Feeding Behavior of Subadult Sixgill Sharks (Hexanchus griseus) at a Bait Station.

This is the first in-situ study of feeding behaviors exhibited by bluntnose sixgill sharks. Bait was placed beneath the Seattle Aquarium pier situated on the waterfront in Elliott Bay, Puget Sound, Washington at 20m of water depth. Cameras and lights were placed around the bait box to record sixgill shark presence and behavior while feeding. Analysis of feeding behavior revealed that sixgills utilize a bite comparable to many other elasmobranchs and aquatic vertebrates, have the ability to protrude their upper jaw, change their feeding behavior based on the situation, and employ sawing and lateral tearing during manipulation. The versatility of their feeding mechanism and the ability of sixgills to change their capture and food manipulation behaviors may have contributed to the species' worldwide distribution and evolutionary success.

Preliminary Observations of Population Genetics and Relatedness of the Broadnose Sevengill Shark, Notorynchus cepedianus, in Two Northeast Pacific Estuaries.

The broadnose sevengill shark, Notorynchus cepedianus, a common coastal species in the eastern North Pacific, was sampled during routine capture and tagging operations conducted from 2005-2012. One hundred and thirty three biopsy samples were taken during these research operations in Willapa Bay, Washington and in San Francisco Bay, California. Genotypic data from seven polymorphic microsatellites (derived from the related sixgill shark, Hexanchus griseus) were used to describe N. cepedianus genetic diversity, population structure and relatedness. Diversity within N. cepedianus was found to be low to moderate with an average observed heterozygosity of 0.41, expected heterozygosity of 0.53, and an average of 5.1 alleles per microsatellite locus. There was no evidence of a recent population bottleneck based on genetic data. Analyses of genetic differences between the two sampled estuaries suggest two distinct populations with some genetic mixing of sharks sampled during 2005-2006. Relatedness within sampled populations was high, with percent relatedness among sharks caught in the same area indicating 42.30% first-order relative relationships (full or half siblings). Estuary-specific familial relationships suggest that management of N. cepedianus on the U.S. West Coast should incorporate stock-specific management goals to conserve this ecologically important predator.

Modulation in the feeding prey capture of the ant-lion, Myrmeleon crudelis.

Ant-lions are pit-building larvae (Neuroptera: Myrmeleontidae), which possess relatively large mandibles used for catching and consuming prey. Few studies involving terrestrial arthropod larva have investigated prey capture behavior and kinematics and no study has shown modulation of strike kinematics. We examined feeding kinematics of the ant-lion, Myrmeleon crudelis, using high-speed video to investigate whether larvae modulate strike behavior based on prey location relative to the mandible. Based on seven capture events from five M. crudelis, the strike took 17.60 ± 2.92 msec and was characterized by near-simultaneous contact of both mandibles with the prey. Modulation of the angular velocity of the mandibles based on prey location was clearly demonstrated. M. crudelis larvae attempted to simultaneously contact prey with both mandibles by increasing mean angular velocity of the far mandible (65 ± 21 rad sec(-1) ) compared with the near mandible (35 ± 14 rad sec(-1) ). Furthermore, kinematic results showed a significant difference for mean angular velocity between the two mandibles (P<0.005). Given the lengthy strike duration compared with other fast-striking arthropods, these data suggest that there is a tradeoff between the ability to modulate strike behavior for accurate simultaneous mandible contact and the overall velocity of the strike. The ability to modulate prey capture behavior may increase dietary breadth and capture success rate in these predatory larvae by allowing responsive adjustment to small-scale variations in prey size, presentation, and escape response.

Functional morphology of the feeding apparatus, feeding constraints, and suction performance in the nurse shark Ginglymostoma cirratum.

The nurse shark, Ginglymostoma cirratum, is an obligate suction feeder that preys on benthic invertebrates and fish. Its cranial morphology exhibits a suite of structural and functional modifications that facilitate this mode of prey capture. During suction-feeding, subambient pressure is generated by the ventral expansion of the hyoid apparatus and the floor of its buccopharyngeal cavity. As in suction-feeding bony fishes, the nurse shark exhibits expansive, compressive, and recovery kinematic phases that produce posterior-directed water flow through the buccopharyngeal cavity. However, there is generally neither a preparatory phase nor cranial elevation. Suction is generated by the rapid depression of the buccopharyngeal floor by the coracoarcualis, coracohyoideus, and coracobranchiales muscles. Because the hyoid arch of G. cirratum is loosely connected to the mandible, contraction of the rectus cervicis muscle group can greatly depress the floor of the buccopharyngeal cavity below the depressed mandible, resulting in large volumetric expansion. Suction pressures in the nurse shark vary greatly, but include the greatest subambient pressures reported for an aquatic-feeding vertebrate. Maximum suction pressure does not appear to be related to shark size, but is correlated with the rate of buccopharyngeal expansion. As in suction-feeding bony fishes, suction in the nurse shark is only effective within approximately 3 cm in front of the mouth. The foraging behavior of this shark is most likely constrained to ambushing or stalking due to the exponential decay of effective suction in front of the mouth. Prey capture may be facilitated by foraging within reef confines and close to the substrate, which can enhance the effective suction distance, or by foraging at night when it can more closely approach prey.

Relative importance of growth and behaviour to elasmobranch suction-feeding performance over early ontogeny.

Development of the ability to capture prey is crucial to predator survival. Trends in food-capture performance over early ontogeny were quantified for leopard sharks Triakis semifasciata and whitespotted bamboosharks Chiloscyllium plagiosum by measuring suction pressure and flow in front of the mouth during feeding. At any size, C. plagiosum produce greater subambient pressure and ingest more rounded water parcels. Maximum subambient pressure scaled with negative allometry in T. semifasciata and was accompanied by an increase in the time to reach maximum gape. Despite a similar trend in buccal expansion timing, maximum pressure in C. plagiosum scaled with isometry and was accompanied by an earlier onset of hyoid depression and a positive allometric increase in buccal reserve volume. Growth was the primary factor responsible for developmental trends in both species, with size-independent behavioural changes contributing little to overall performance variability. Ontogenetic dietary shifts are predicted for both species as a consequence of size-dependent changes in performance. Chiloscyllium plagiosum becomes anatomically and behaviourally canalized towards suction feeding, limiting the effective range of prey capture and possibly necessitating stalking. Triakis semifasciata, by contrast, retains the flexibility to employ both ram and suction and therefore captures more elusive prey with age.