Grow or go? Energetic constraints on shark pup dispersal from pupping areas.

Many sharks and other marine taxa use natal areas to maximize survival of young, meaning such areas are often attributed conservation value. The use of natal areas is often linked to predator avoidance or food resources. However, energetic constraints that may influence dispersal of young and their use of natal areas are poorly understood. We combined swim-tunnel respirometry, calorimetry, lipid class analysis and a bioenergetics model to investigate how energy demands influence dispersal of young in a globally distributed shark. The school shark (a.k.a. soupfin, tope), Galeorhinus galeus, is Critically Endangered due to overfishing and is one of many sharks that use protected natal areas in Australia. Energy storage in neonate pups was limited by small livers, low overall lipid content and low levels of energy storage lipids (e.g. triacylglycerols) relative to adults, with energy stores sufficient to sustain routine demands for 1.3-4 days (mean ± SD: 2.4 ± 0.8 days). High levels of growth-associated structural lipids (e.g. phospholipids) and high energetic cost of growth suggested large investment in growth during residency in natal areas. Rapid growth (~40% in length) between birth in summer and dispersal in late autumn-winter likely increased survival by reducing predation and improving foraging ability. Delaying dispersal may allow prioritization of growth and may also provide energy savings through improved swimming efficiency and cooler ambient temperatures (daily ration was predicted to fall by around a third in winter). Neonate school sharks are therefore ill-equipped for large-scale dispersal and neonates recorded in the northwest of their Australian distribution are likely born locally, not at known south-eastern pupping areas. This suggests the existence of previously unrecorded school shark pupping areas. Integrated bioenergetic approaches as applied here may help to understand dispersal from natal areas in other taxa, such as teleost fishes, elasmobranchs and invertebrates.

The importance of sample size in marine megafauna tagging studies.

Telemetry is a key, widely used tool to understand marine megafauna distribution, habitat use, behavior, and physiology; however, a critical question remains: "How many animals should be tracked to acquire meaningful data sets?" This question has wide-ranging implications including considerations of statistical power, animal ethics, logistics, and cost. While power analyses can inform sample sizes needed for statistical significance, they require some initial data inputs that are often unavailable. To inform the planning of telemetry and biologging studies of marine megafauna where few or no data are available or where resources are limited, we reviewed the types of information that have been obtained in previously published studies using different sample sizes. We considered sample sizes from one to >100 individuals and synthesized empirical findings, detailing the information that can be gathered with increasing sample sizes. We complement this review with simulations, using real data, to show the impact of sample size when trying to address various research questions in movement ecology of marine megafauna. We also highlight the value of collaborative, synthetic studies to enhance sample sizes and broaden the range, scale, and scope of questions that can be answered.

Convergence of marine megafauna movement patterns in coastal and open oceans.

The extent of increasing anthropogenic impacts on large marine vertebrates partly depends on the animals' movement patterns. Effective conservation requires identification of the key drivers of movement including intrinsic properties and extrinsic constraints associated with the dynamic nature of the environments the animals inhabit. However, the relative importance of intrinsic versus extrinsic factors remains elusive. We analyze a global dataset of ∼2.8 million locations from >2,600 tracked individuals across 50 marine vertebrates evolutionarily separated by millions of years and using different locomotion modes (fly, swim, walk/paddle). Strikingly, movement patterns show a remarkable convergence, being strongly conserved across species and independent of body length and mass, despite these traits ranging over 10 orders of magnitude among the species studied. This represents a fundamental difference between marine and terrestrial vertebrates not previously identified, likely linked to the reduced costs of locomotion in water. Movement patterns were primarily explained by the interaction between species-specific traits and the habitat(s) they move through, resulting in complex movement patterns when moving close to coasts compared with more predictable patterns when moving in open oceans. This distinct difference may be associated with greater complexity within coastal microhabitats, highlighting a critical role of preferred habitat in shaping marine vertebrate global movements. Efforts to develop understanding of the characteristics of vertebrate movement should consider the habitat(s) through which they move to identify how movement patterns will alter with forecasted severe ocean changes, such as reduced Arctic sea ice cover, sea level rise, and declining oxygen content.

Distinguishing globally-driven changes from regional- and local-scale impacts: The case for long-term and broad-scale studies of recovery from pollution.

Marine ecosystems are subject to anthropogenic change at global, regional and local scales. Global drivers interact with regional- and local-scale impacts of both a chronic and acute nature. Natural fluctuations and those driven by climate change need to be understood to diagnose local- and regional-scale impacts, and to inform assessments of recovery. Three case studies are used to illustrate the need for long-term studies: (i) separation of the influence of fishing pressure from climate change on bottom fish in the English Channel; (ii) recovery of rocky shore assemblages from the Torrey Canyon oil spill in the southwest of England; (iii) interaction of climate change and chronic Tributyltin pollution affecting recovery of rocky shore populations following the Torrey Canyon oil spill. We emphasize that "baselines" or "reference states" are better viewed as envelopes that are dependent on the time window of observation. Recommendations are made for adaptive management in a rapidly changing world.

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.

Feeding requirements of white sharks may be higher than originally thought.

Quantifying the energy requirements of animals in nature is critical for understanding physiological, behavioural, and ecosystem ecology; however, for difficult-to-study species such as large sharks, prey intake rates are largely unknown. Here, we use metabolic rates derived from swimming speed estimates to suggest that feeding requirements of the world's largest predatory fish, the white shark (Carcharodon carcharias), are several times higher than previously proposed. Further, our estimates of feeding frequency identify a clear benefit in seasonal selection of pinniped colonies - a white shark foraging strategy seen across much of their range.

Inferred global connectivity of whale shark Rhincodon typus populations.

Ten years have passed since the last synopsis of whale shark Rhincodon typus biogeography. While a recent review of the species' biology and ecology summarized the vast data collected since then, it is clear that information on population geographic connectivity, migration and demography of R. typus is still limited and scattered. Understanding R. typus migratory behaviour is central to its conservation management considering the genetic evidence suggesting local aggregations are connected at the generational scale over entire ocean basins. By collating available data on sightings, tracked movements and distribution information, this review provides evidence for the hypothesis of broad-scale connectivity among populations, and generates a model describing how the world's R. typus are part of a single, global meta-population. Rhincodon typus occurrence timings and distribution patterns make possible a connection between several aggregation sites in the Indian Ocean. The present conceptual model and validating data lend support to the hypothesis that R. typus are able to move among the three largest ocean basins with a minimum total travelling time of around 2-4 years. The model provides a worldwide perspective of possible R. typus migration routes, and suggests a modified focus for additional research to test its predictions. The framework can be used to trim the hypotheses for R. typus movements and aggregation timings, thereby isolating possible mating and breeding areas that are currently unknown. This will assist endeavours to predict the longer-term response of the species to ocean warming and changing patterns of human-induced mortality.

Identifying reproductive events using archival tags: egg-laying behaviour of the small spotted catshark Scyliorhinus canicula.

The use of archival depth telemetry as a means of remotely assessing the reproductive rates of free-ranging fishes is explored. This is achieved by electronically tracking the vertical movements of individual female small spotted catsharks Scyliorhinus canicula in the natural environment, whilst simultaneously evaluating the temporal and vertical distributions of egg-laying in this species. Distinctive patterns of short-term (0·3-3·7 h), shallow-water activity are documented in the time-depth profiles of female S. canicula that occur at an appropriate depth (1·0-2·3 m) and periodicity (every 10-12 days) to represent egg-laying behaviour. Putative egg-laying behaviour was exhibited simultaneously by two individually tracked female S. canicula during late-spring and early-summer. The results highlight that, provided species behaviour is suitable and complementary methods such as previous data, laboratory experiments and field surveys can be used to validate the patterns observed, archival depth telemetry offers an unobtrusive means by which egg production and egg-laying behaviour of free-living fishes can be estimated. As precise information regarding life-history parameters is difficult to obtain for free-ranging fish species, this technique could be used to improve the parameterization of species demographic models that are relevant to the management of wild fish populations.

The effect of familiarity on aggregation and social behaviour in juvenile small spotted catsharks Scyliorhinus canicula.

This study was designed to address whether juvenile small spotted catsharks Scyliorhinus canicula aggregate and to determine whether potential aggregation is underpinned by social preferences for conspecifics. Using controlled and replicated experiments, the role of familiarity as a potential mechanism driving aggregation and social behaviour in this species was considered. Observed S. canicula association data compared to null model simulations of random distributions revealed differences in aggregation under different social contexts. Only familiar juvenile S. canicula aggregated more than would be expected from random distribution across their habitat. Familiarity increased the mean number of groups but did not significantly affect mean group size. Significant preference and avoidance behaviour across all groups were also observed. Furthermore, the strength of social attraction, quantified by the mean association index, was significantly higher in groups containing familiar individuals. Mixed familiar and unfamiliar treatments were also conducted to test for within- and between-group effects, finding high variation across replicates with some groups assorting by familiarity and others not. It is believed that this study is the first to examine experimentally the influence of conspecific familiarity on aggregation behaviour in sharks. These results not only imply a functional benefit to aggregation, but also suggest that persistent social affiliation is likely to influence dispersal following hatching in this small benthic elasmobranch.

Assessing wave energy effects on biodiversity: the wave hub experience.

Marine renewable energy installations harnessing energy from wind, wave and tidal resources are likely to become a large part of the future energy mix worldwide. The potential to gather energy from waves has recently seen increasing interest, with pilot developments in several nations. Although technology to harness wave energy lags behind that of wind and tidal generation, it has the potential to contribute significantly to energy production. As wave energy technology matures and becomes more widespread, it is likely to result in further transformation of our coastal seas. Such changes are accompanied by uncertainty regarding their impacts on biodiversity. To date, impacts have not been assessed, as wave energy converters have yet to be fully developed. Therefore, there is a pressing need to build a framework of understanding regarding the potential impacts of these technologies, underpinned by methodologies that are transferable and scalable across sites to facilitate formal meta-analysis. We first review the potential positive and negative effects of wave energy generation, and then, with specific reference to our work at the Wave Hub (a wave energy test site in southwest England, UK), we set out the methodological approaches needed to assess possible effects of wave energy on biodiversity. We highlight the need for national and international research clusters to accelerate the implementation of wave energy, within a coherent understanding of potential effects-both positive and negative.

Oceans apart? Short-term movements and behaviour of adult bull sharks Carcharhinus leucas in Atlantic and Pacific Oceans determined from pop-off satellite archival tagging.

Adult bull sharks Carcharhinus leucas were monitored with electronic tags to investigate horizontal and vertical movements in the Atlantic and Pacific Oceans. In both locations, C. leucas showed some fidelity to specific coastal areas with only limited horizontal movements away from the tagging sites after tag attachment. Fish tagged in the Bahamas were detected mostly in the upper 20 m of the water column in water 25-26° C, whereas C. leucas tagged in Fiji spent most of their time below 20 m in water usually >26° C. The results highlight the importance of coastal inshore habitats for this species.

Deep-diving behaviour of a whale shark Rhincodon typus during long-distance movement in the western Indian Ocean.

A whale shark Rhincodon typus satellite tagged off the coast of Mozambique showed a highly directional movement across the Mozambique Channel and around the southern tip of Madagascar, a minimum distance of 1200 km in 87 days. Dives to depths well into the mesopelagic and bathypelagic zones (1286 m maximum depth) were recorded in a bathymetrically non-constraining habitat. The water temperature range recorded during the fish's movement was 3.4-29.9 degrees C.

Shifts in a fish's resource holding power during a contact paired interaction: the influence of a copper-contaminated diet in rainbow trout.

The influence of sublethal chronic dietary copper (Cu) exposure on the dominant-subordinate relationship between pairs of juvenile rainbow trout (Oncorhynchus mykiss) was examined. Fish were fed either a normal (11 mg Cu kg(-1) food) or Cu-contaminated (721 mg Cu kg(-1) food) diet for 8 wk. Paired interactions were observed--control versus control, Cu-exposed versus Cu-exposed, and control versus Cu-exposed fish--using a computer-aided video tracking system to measure duration of interactions, total distance moved, and the number of encounters during each contest. In concurrence with game theory, each interaction became escalated with a lesser size disparity between contestants. However, in Cu-exposed versus Cu-exposed fish interactions, the dominant-subordinate relationship was decided sooner and with less aggression than a control versus control fish interaction with fish of a similar relative body mass disparity. During control versus Cu-exposed fish interactions, control fish would normally dominate interactions (12 out of 16 bouts) unless the Cu-exposed fish had a 15% body mass advantage. Muscle glycogen and lactate levels after each contest reflected the duration of bouts and winners of the contests, irrespective of Cu exposure. We conclude that Cu-contaminated fish are less able to compete and have lower resource holding power than controls and will withdraw from a contest at a lower level of interaction, unless a size advantage in the Cu-exposed fish increases the probability of winning.

Shark skin: a function in feeding.

Dermal denticles are unique tooth-like structures embedded in the skin of sharks and rays that protect them from predators and ectoparasites, reduce mechanical abrasion and possibly minimize swimming-induced drag. Here, we show that juvenile lesser spotted dogfish (Scyliorhinus canicula) also use this body armour to anchor food items near their tail so that bite-sized pieces can be torn away by rapid jaw and head movements. This scale-rasp behaviour is novel among fishes and suggests a new role for skin in the feeding ecology of sharks. Scale rasping may be important ecologically because it could function to increase the dietary breadth and growth potential of juveniles.

Timing of squid migration reflects North Atlantic climate variability.

The environmental and biotic conditions affecting fisheries for cephalopods are only partially understood. A problem central to this is how climate change may influence population movements by altering the availability of thermal resources. In this study we investigate the links between climate and sea-temperature changes and squid arrival time off southwestern England over a 20-year period. We show that veined squid (Loligo forbesi) migrate eastward in the English Channel earlier when water in the preceding months is warmer, and that higher temperatures and early arrival correspond with warm (positive) phases of the North Atlantic oscillation (NAO). The timing of squid peak abundance advanced by 120-150 days in the warmest years ('early' years) compared with the coldest ('late' years). Furthermore, sea-bottom temperature was closely linked to the extent of squid movement. Temperature increases over the five months prior to and during the month of peak squid abundance did not differ between early and late years, indicating squid responded to temperature changes independently of time of year. We conclude that the temporal variation in peak abundance of squid seen off Plymouth represents temperature-dependent movement, which is in turn mediated by climatic changes associated with the NAO. Such climate-mediated movement may be a widespread characteristic of cephalopod populations worldwide, and may have implications for future fisheries management because global warming may alter both the timing and location of peak population abundance.

Pre-operative ultrasonography guiding minimal, selective surgical approach in primary hyperparathyroidism.

BACKGROUND: The traditional surgical approach for the treatment of primary hyperparathyroidism (HPT) has been routine bilateral surgical exploration. Our aim was to evaluate pre-operative ultrasonography (U/S) in altering our practice to selective unilateral neck exploration. PATIENTS AND METHODS: A retrospective study was conducted involving 53 patients who had parathyroidectomy over a 5-year period (1989-1994), 41 of whom with the diagnosis of HPT had pre-operative neck U/S. RESULTS: Thirty-eight patients had a single adenoma, one had parathyroid cancer, and two had multiple adenomas. Thirty-six of 41 (88%) patients were true positives, two (5%) false positives, two (5%) false negatives and one (2%) was true negative. Based on the surgeon's personal preference and U/S results, 21/41 (51%) of patients had bilateral and 20/41 (49%) had unilateral neck exploration. All the 41 patients had resolution of the hypercalcemia postoperatively. Eighteen of 38 (47%) patients with a single adenoma had bilateral neck exploration and only in two patients was this necessary. CONCLUSIONS: Patients undergoing parathyroid surgery for HPT should have preoperative U/S and, if a single enlarged parathyroid gland is found, unilateral U/S guided neck exploration is safe and economical. In all the other patients, bilateral exploration is the preferred approach.

Annual social behaviour of basking sharks associated with coastal front areas.

Comparatively little is known about reproductive behaviour in wild sharks as it has proved extremely difficult to study, especially in large pelagic sharks. Here we describe annual courtship-like behaviour in the second-largest fish species, the basking shark (Cetorhinus maximus), from 25 separate episodes observed and tracked during a five-year study period (1995-1999) off south-west England. Social behaviours observed between paired, or three or four, sharks were consistent with courtship behaviours seen in other shark species, namely nose-to-tail following, close following, close flank approach, parallel and echelon swimming. Mature individuals between 5 and 8 m total body length (L(T)) exhibited these behaviours whereas smaller sharks (3-4 m L(T)) did not. Lead individuals were identified as female on a number of occasions and interactions were prolonged; the longest continuous observation of socializing was 1.8 h, although intermittent track data indicates bouts may last for up to 5-6 h. Locations of courtship-like behaviour events were not distributed randomly and were significantly associated with thermal fronts. Our results indicate that putative courtship behaviour occurs between May and July along oceanographic fronts, probably as a consequence of individuals aggregating to forage in rich prey patches before initiating courtship. Thus, locating the richest prey patches along fronts may be important for basking sharks to find mates as well as food in the pelagic ecosystem. As courtship-like behaviours occur annually off south-west England we speculate that this region may represent an annual breeding area for this protected species, but mating itself probably takes place at depth as it was not seen at the surface.