Congruent trends in long-term zooplankton decline in the north-east Atlantic and basking shark (Cetorhinus maximus ) fishery catches off west Ireland

Trends in basking shark (Cetorhinus maximus) fishery catches off Achill Island, west Ireland between 1949 and 1975 were examined in relation to zooplankton (total copepod) abundance in four adjacent sea areas over a 27-year period. The numbers of basking sharks caught and copepod abundance showed downward trends and were positively correlated (r-value range, 0.44–0.74). A possible explanation for the downward trend in shark catches was that progressively fewer basking sharks occurred there between 1956 and 1975 because fewer copepods, their food resource, occurred near the surface off west Ireland over the same period. We suggest that the decline in basking sharks may have been due to a distributional shift of sharks to more productive areas, rather than a highly philopatric, localized stock that was over-exploited.

Quantifying the impact of environmental variables upon catch-per-unit-effort of the Blue Shark Prionace glauca in the Western English Channel

The effect of environmental variables on blue shark Prionace glauca catch per unit effort (CPUE) in a recreational fishery in the western English Channel, between June and September 1998–2011, was quantified using generalized additive models (GAMs). Sea surface temperature (SST) explained 1·4% of GAM deviance, and highest CPUE occurred at 16·7° C, reflecting the optimal thermal preferences of this species. Surface chlorophyll a concentration (CHL) significantly affected CPUE and caused 27·5% of GAM deviance. Additionally, increasing CHL led to rising CPUE, probably due to higher productivity supporting greater prey biomass. The density of shelf-sea tidal mixing fronts explained 5% of GAM deviance, but was non-significant, with increasing front density negatively affecting CPUE. Time-lagged frontal density significantly affected CPUE, however, causing 12·6% of the deviance in a second GAM and displayed a positive correlation. This outcome suggested a delay between the evolution of frontal features and the subsequent accumulation of productivity and attraction of higher trophic level predators, such as P. glauca.

Quantifying the impact of environmental variables upon catch-per-unit-effort of the Blue Shark Prionace glauca in the Western English Channel

The effect of environmental variables on blue shark Prionace glauca catch per unit effort (CPUE) in a recreational fishery in the western English Channel, between June and September 1998–2011, was quantified using generalized additive models (GAMs). Sea surface temperature (SST) explained 1·4% of GAM deviance, and highest CPUE occurred at 16·7° C, reflecting the optimal thermal preferences of this species. Surface chlorophyll a concentration (CHL) significantly affected CPUE and caused 27·5% of GAM deviance. Additionally, increasing CHL led to rising CPUE, probably due to higher productivity supporting greater prey biomass. The density of shelf-sea tidal mixing fronts explained 5% of GAM deviance, but was non-significant, with increasing front density negatively affecting CPUE. Time-lagged frontal density significantly affected CPUE, however, causing 12·6% of the deviance in a second GAM and displayed a positive correlation. This outcome suggested a delay between the evolution of frontal features and the subsequent accumulation of productivity and attraction of higher trophic level predators, such as P. glauca.

Shark personalities? Repeatability of social network traits in a widely distributed predatory fish. Behavioral Ecology and Sociobiology, 68, 1995-2003.

Interest in animal personalities has generated a burgeoning literature on repeatability in individual traits such as boldness or exploration through time or across different contexts. Yet, repeatability can be influenced by the interactive social strategies of individuals, for example, consistent inter-individual variation in aggression is well documented. Previous work has largely focused on the social aspects of repeatability in animal behaviour by testing individuals in dyadic pairings. Under natural conditions, individuals interact in a heterogeneous polyadic network. However, the extent to which there is repeatability of social traits at this higher order network level remains unknown. Here, we provide the first empirical evidence of consistent and repeatable animal social networks. Using a model species of shark, a taxonomic group in which repeatability in behaviour has yet to be described, we repeatedly quantified the social networks of ten independent shark groups across different habitats, testing repeatability in individual network position under changing environments. To understand better the mechanisms behind repeatable social behaviour, we also explored the coupling between individual preferences for specific group sizes and social network position. We quantify repeatability in sharks by demonstrating that despite changes in aggregation measured at the group level, the social network position of individuals is consistent across treatments. Group size preferences were found to influence the social network position of individuals in small groups but less so for larger groups suggesting network structure, and thus, repeatability was driven by social preference over aggregation tendency.

To Madagascar and back: long-distance, return migration across open ocean by a pregnant female bull shark Carcharhinus leucas

A large, pregnant, female bull shark Carcharhinus leucas was tracked migrating from Seychelles across open ocean to south-east Madagascar, c. 2000 km away, and back again. In Madagascar, the shark spent a prolonged period shallower than 5 m, consistent with entering estuarine habitat to pup, and upon return to Seychelles the shark was slender and no longer gravid. This represents an unprecedented return migration across the open ocean for a C. leucas and highlights the need for international collaboration to manage the regional C. leucas population sustainably.

To Madagascar and back: long-distance, return migration across open ocean by a pregnant female bull shark Carcharhinus leucas

A large, pregnant, female bull shark Carcharhinus leucas was tracked migrating from Seychelles across open ocean to south-east Madagascar, c. 2000 km away, and back again. In Madagascar, the shark spent a prolonged period shallower than 5 m, consistent with entering estuarine habitat to pup, and upon return to Seychelles the shark was slender and no longer gravid. This represents an unprecedented return migration across the open ocean for a C. leucas and highlights the need for international collaboration to manage the regional C. leucas population sustainably.

DNA from historical and trophy samples provides insights into white shark population origins and genetic diversity

Characterizing genetic variation by retrospective genotyping of trophy or historical artifacts from endangered species is an important conservation tool. Loss of genetic diversity in top predators such as the white shark Carcharodon carcharias remains an issue, exacerbated in this species by declining, sometimes isolated philopatric populations. We successfully sequenced mitochondrial DNA (mtDNA) D-loop from osteodentine of contemporary South African white shark teeth (from 3 jaws), and from 34 to 129 yr old dried cartilage and skin samples from 1 Pacific Ocean and 5 Mediterranean sharks. Osteodentine-derived sequences from South African fish matched those derived from an individual’s finclips, but were generally of poorer quality than those from skin and cartilage of historical samples. Three haplotypes were identified from historical Mediterranean samples (n = 5); 2 individuals had unique sequences and 3 shared the contemporary Mediterranean haplotype. Placement of previously undescribed mtDNA haplotypes from historical material within both the Mediterranean and Pacific clades fits with the accepted intra-specific phylogeny derived from contemporary material, verifying our approaches. The utility of our methodology is in its provision of additional genetic resources from osteodentine (for species lacking tooth pulp) and cartilage of rare and endangered species held in often uncurated, contemporary and historical dry collections. Such material can usefully supplement estimates of connectivity, population history, and stock viability. We confirm the depauperate haplotype diversity of historical Mediterranean sharks, consistent with founding by a small number of Pacific colonizers. The consequent lack of diversity suggests serious challenges for the maintenance of this top predator and the Mediterranean ecosystem.

DNA from historical and trophy samples provides insights into white shark population origins and genetic diversity

Characterizing genetic variation by retrospective genotyping of trophy or historical artifacts from endangered species is an important conservation tool. Loss of genetic diversity in top predators such as the white shark Carcharodon carcharias remains an issue, exacerbated in this species by declining, sometimes isolated philopatric populations. We successfully sequenced mitochondrial DNA (mtDNA) D-loop from osteodentine of contemporary South African white shark teeth (from 3 jaws), and from 34 to 129 yr old dried cartilage and skin samples from 1 Pacific Ocean and 5 Mediterranean sharks. Osteodentine-derived sequences from South African fish matched those derived from an individual’s finclips, but were generally of poorer quality than those from skin and cartilage of historical samples. Three haplotypes were identified from historical Mediterranean samples (n = 5); 2 individuals had unique sequences and 3 shared the contemporary Mediterranean haplotype. Placement of previously undescribed mtDNA haplotypes from historical material within both the Mediterranean and Pacific clades fits with the accepted intra-specific phylogeny derived from contemporary material, verifying our approaches. The utility of our methodology is in its provision of additional genetic resources from osteodentine (for species lacking tooth pulp) and cartilage of rare and endangered species held in often uncurated, contemporary and historical dry collections. Such material can usefully supplement estimates of connectivity, population history, and stock viability. We confirm the depauperate haplotype diversity of historical Mediterranean sharks, consistent with founding by a small number of Pacific colonizers. The consequent lack of diversity suggests serious challenges for the maintenance of this top predator and the Mediterranean ecosystem.

Encounter success of free-ranging marine predator movements across a dynamic prey landscape

Movements of wide-ranging top predators can now be studied effectively using satellite and archival telemetry. However, the motivations underlying movements remain difficult to determine because trajectories are seldom related to key biological gradients, such as changing prey distributions. Here, we use a dynamic prey landscape of zooplankton biomass in the north-east Atlantic Ocean to examine active habitat selection in the plankton-feeding basking shark Cetorhinus maximus. The relative success of shark searches across this landscape was examined by comparing prey biomass encountered by sharks with encounters by random-walk simulations of ‘model’ sharks. Movements of transmitter-tagged sharks monitored for 964 days (16754km estimated minimum distance) were concentrated on the European continental shelf in areas characterized by high seasonal productivity and complex prey distributions. We show movements by adult and sub-adult sharks yielded consistently higher prey encounter rates than 90% of random-walk simulations. Behavioural patterns were consistent with basking sharks using search tactics structured across multiple scales to exploit the richest prey areas available in preferred habitats. Simple behavioural rules based on learned responses to previously encountered prey distributions may explain the high performances. This study highlights how dynamic prey landscapes enable active habitat selection in large predators to be investigated from a trophic perspective, an approach that may inform conservation by identifying critical habitat of vulnerable species.

Seasonal movements and behaviour of basking sharks from archival tagging: No evidence of winter hibernation

Habitat selection processes in highly migratory animals such as sharks and whales are important to understand because they influence patterns of distribution, availability and therefore catch rates. However, spatial strategies remain poorly understood over seasonal scales in most species, including, most notably, the plankton-feeding basking shark Cetorhinus maximus. It was proposed nearly 50 yr ago that this globally distributed species migrates from coastal summer-feeding areas of the northeast Atlantic to hibernate during winter in deep water on the bottom of continental-shelf slopes. This view has perpetuated in the literature even though the 'hibernation theory' has not been tested directly. We have now tracked basking sharks for the first time over seasonal scales (1.7 to 6.5 mo) using 'pop-up' satellite archival transmitters. We show that they do not hibernate during winter but instead undertake extensive horizontal (up to 3400 km) and vertical (> 750 m depth) movements to utilise productive continental-shelf and shelf-edge habitats during summer, autumn and winter. They travel long distances (390 to 460 km) to locate temporally discrete productivity 'hotspots' at shelf-break fronts, but at no time were prolonged movements into open-ocean regions away from shelf waters observed. Basking sharks have a very broad vertical diving range and can dive beyond the known range of planktivorous whales. Our study suggests this species can exploit shelf and slope-associated zooplankton communities in mesopelagic (200 to 1000 m) as well as epipelagic habitat (0 to 200 m).

Basking sharks and oceanographic fronts: quantifying associations in the north‐east Atlantic

Understanding the mechanisms linking oceanographic processes and marine vertebrate habitat use is critical to effective management of populations of conservation concern. The basking shark Cetorhinus maximus has been shown to associate with oceanographic fronts – physical interfaces at the transitions between water masses – to exploit foraging opportunities resulting from aggregation of zooplankton. However, the scale, significance and variability of these observed associations have not yet been established. Here, we quantify the influence of mesoscale (10s – 100s km) frontal activity on habitat use over timescales of weeks to months. We use animal-mounted archival tracking with composite front mapping via Earth Observation (EO) remote sensing to provide an oceanographic context to individual shark movements. We investigate levels of association with fronts occurring over two spatio-temporal scales, (i) broad-scale seasonally persistent frontal zones and (ii) contemporaneous mesoscale thermal and chl-a fronts. Using random walk simulations and logistic regression within an iterative generalised linear mixed modelling (GLMM) framework, we find that seasonal front frequency is a significant predictor of shark presence. Temporally-matched oceanographic metrics also indicate that sharks demonstrate a preference for productive regions, and associate with contemporaneous thermal and chl-a fronts more frequently than could be expected at random. Moreover, we highlight the importance of cross-frontal temperature change and persistence, which appear to interact to affect the degree of prey aggregation along thermal fronts. These insights have clear implications for understanding the preferred habitats of basking sharks in the context of anthropogenic threat management and marine spatial planning in the northeast Atlantic.

Basking sharks and oceanographic fronts: quantifying associations in the north-east Atlantic

Understanding the mechanisms linking oceanographic processes and marine vertebrate habitat use is critical to effective management of populations of conservation concern. The basking shark Cetorhinus maximus has been shown to associate with oceanographic fronts – physical interfaces at the transitions between water masses – to exploit foraging opportunities resulting from aggregation of zooplankton. However, the scale, significance and variability of these observed associations have not yet been established. Here, we quantify the influence of mesoscale (10s – 100s km) frontal activity on habitat use over timescales of weeks to months. We use animal-mounted archival tracking with composite front mapping via Earth Observation (EO) remote sensing to provide an oceanographic context to individual shark movements. We investigate levels of association with fronts occurring over two spatio-temporal scales, (i) broad-scale seasonally persistent frontal zones and (ii) contemporaneous mesoscale thermal and chl-a fronts. Using random walk simulations and logistic regression within an iterative generalised linear mixed modelling (GLMM) framework, we find that seasonal front frequency is a significant predictor of shark presence. Temporally-matched oceanographic metrics also indicate that sharks demonstrate a preference for productive regions, and associate with contemporaneous thermal and chl-a fronts more frequently than could be expected at random. Moreover, we highlight the importance of cross-frontal temperature change and persistence, which appear to interact to affect the degree of prey aggregation along thermal fronts. These insights have clear implications for understanding the preferred habitats of basking sharks in the context of anthropogenic threat management and marine spatial planning in the northeast Atlantic.