The Last Frontier: Catch Records of White Sharks (Carcharodon carcharias) in the Northwest Pacific Ocean

White sharks are highly migratory apex predators, globally distributed in temperate, sub-tropical, and tropical waters. Knowledge of white shark biology and ecology has increased recently based on research at known aggregation sites in the Indian, Atlantic, and Northeast Pacific Oceans; however, few data are available for the Northwest Pacific Ocean. This study provides a meta-analysis of 240 observations of white sharks from the Northwest Pacific Ocean between 1951 and 2012. Records comprise reports of bycatch in commercial fisheries, media accounts, personal communications, and documentation of shark-human interactions from Russia (n = 8), Republic of Korea (22), Japan (129), China (32), Taiwan (45), Philippines (1) and Vietnam (3). Observations occurred in all months, excluding October-January in the north (Russia and Republic of Korea) and July-August in the south (China, Taiwan, Philippines, and Vietnam). Population trend analysis indicated that the relative abundance of white sharks in the region has remained relatively stable, but parameterization of a 75% increase in observer effort found evidence of a minor decline since 2002. Reliably measured sharks ranged from 126–602 cm total length (TL) and 16–2530 kg total weight. The largest shark in this study (602 cm TL) represents the largest measured shark on record worldwide. For all countries combined the sex ratio was non-significantly biased towards females (1∶1.1; n = 113). Of 60 females examined, 11 were confirmed pregnant ranging from the beginning stages of pregnancy (egg cases) to near term (140 cm TL embryos). On average, 6.0±2.2 embryos were found per litter (maximum of 10) and gestation period was estimated to be 20 months. These observations confirm that white sharks are present in the Northwest Pacific Ocean year-round. While acknowledging the difficulties of studying little known populations of a naturally low abundance species, these results highlight the need for dedicated research to inform regional conservation and management planning.     

The Occurrence of White Sharks, Carcharodon carcharias, Around the Balearic Islands (Western Mediterranean Sea)

The regular presence of the white shark, Carcharodon carcharias, off the Balearic Islands (western Mediterranean) is shown from 27 captures carried out with trap nets between the 1920s and 1970s and from eight attacks on cetaceans and marine turtles from the 1990s to the present. The geographic distribution, seasonality and population structure of the species in the area are analysed and discussed in relation to environmental conditions and to the proposed distribution of this species in the Mediterranean Sea.     

White Sharks (Carcharodon carcharias) Scavenging on Whales and Its Potential Role in Further Shaping the Ecology of an Apex Predator

Scavenging, a result of a temporary pulse of resources, occurs in virtually all ecosystems containing carnivores, and is an important energy transfer pathway that can impact ecosystem structure and function, and this ecological significance has largely been considered from a terrestrial standpoint; however, little is known about the role of scavenging in shaping the behavioral ecology of marine species, specifically apex predators. Here we present findings from multiple opportunistic observations of white sharks scavenging on whale carcasses in False Bay, South Africa. Observations of white sharks scavenging over successive days provided evidence of strategic and selective scavenging by this species. Moreover, extended daily observations permitted recordings of unique social, aggregative, and feeding behaviors. We further compare these data against observations of natural predation by sharks on seals in the study area. We discuss these data in relation to environmental conditions, shark social interactions, migration patterns, whale biology, and behaviorally-mediated trophic cascades. While the appearance of a whale carcass is largely a stochastic event, we propose that white shark scavenging on whales may represent an underestimated, yet significant component to the overall foraging ecology of this species, especially as individuals attain sexual maturity.     

Space Utilization and Swimming Depth of White Sharks, Carcharodon carcharias, at the South Farallon Islands, Central California

This paper presents information on the movements of white sharks, Carcharodon carcharias, at the South Farallon Islands (SFI), central California. Acoustic telemetry techniques provided preliminary data on the diurnal space utilization, movement patterns and swimming depths of four white sharks, ranging from approximately 3.7 to 4.9m in length. Sharks swam within about 10m of the bottom to depths of approximately 30m, but in deeper water they tended to stray more from the bottom. Activity spaces for time periods tracked ranged from 1.84 to 9.15km². Indications are that an inverse relationship exists between length and activity space. During the time tracked, larger individuals swam within particular areas around the islands whereas smaller individuals did not restrict their movements in the same manner. Values of a site attachment index were inversely related to length for all sharks tracked. The site attachment indices, apparent inverse relationship between total length and activity space and observations on telemetered and other known individuals support a hypothesis that larger sharks possess site fidelity in their search for prey at SFI, within and between years. With the high frequency of predation by white sharks on juvenile northern elephant seals at SFI in the fall, the majority of the sharks' movements are probably related to their search for these pinniped prey. These data provide preliminary evidence that white sharks at SFI may search for prey by swimming in a particular area over a number of days or weeks, traversing the area in a manner which maximizes coverage, and swimming close to the bottom or at a distance far enough from the surface to remain cryptic from prey.     

The Influence of Environmental Variables on the Presence of White Sharks,Carcharodon carcharias at Two Popular Cape Town Bathing Beaches: A Generalized Additive Mixed Model

Shark attacks on humans are high profile events which can significantly influence policies related to the coastal zone. A shark warning system in South Africa, Shark Spotters, recorded 378 white shark (Carcharodon carcharias) sightings at two popular beaches, Fish Hoek and Muizenberg, during 3690 six-hour long spotting shifts, during the months September to May 2006 to 2011. The probabilities of shark sightings were related to environmental variables using Binomial Generalized Additive Mixed Models (GAMMs). Sea surface temperature was significant, with the probability of shark sightings increasing rapidly as SST exceeded 14°C and approached a maximum at 18°C, whereafter it remains high. An 8 times (Muizenberg) and 5 times (Fish Hoek) greater likelihood of sighting a shark was predicted at 18°C than at 14°C. Lunar phase was also significant with a prediction of 1.5 times (Muizenberg) and 4 times (Fish Hoek) greater likelihood of a shark sighting at new moon than at full moon. At Fish Hoek, the probability of sighting a shark was 1.6 times higher during the afternoon shift compared to the morning shift, but no diel effect was found at Muizenberg. A significant increase in the number of shark sightings was identified over the last three years, highlighting the need for ongoing research into shark attack mitigation. These patterns will be incorporated into shark awareness and bather safety campaigns in Cape Town.     

Insights into Young of the Year White Shark, Carcharodon carcharias, Behavior in the Southern California Bight

A young of the year female white shark, Carcharodon carcharias, was tagged with a pop-up satellite archival tag off Southern California in early June of 2000. The tag was recovered after 28 days, and records of temperature, depth and light intensity were extracted. Depth and temperature records indicate a number of interesting behaviors, including a strong diurnal pattern. At night the shark remained in the top 50 m, often making shallow repetitive vertical excursions. Most dives below the mixed layer were observed during the day, 91% of which occurred from 05:00 to 21:00 h, with depths extending to 240 m. Many of the dives exhibited secondary vertical movements that were consistent with the shark swimming at the bottom (at depths from 9 to 165 m) where it was most likely foraging. The white shark experienced dramatic and rapid changes in temperature, and demonstrated a considerable tolerance for cold waters. Temperatures ranged from 9°C to 22°C, and although 89% of the total time was spent in waters 16–22°C, on some days the small shark spent as much as 32% of the time in 12°C waters. The deep dives into cold waters separate the white sharks from mako sharks, which share the California Bight nursery ground but appear to remain primarily in the mixed layer and thermocline. Movement information (derived from light-based geolocation, bottom depths and sea surface temperatures) indicated that the white shark spent the 28 days in the Southern California Bight, possibly moving as far south as San Diego, California. While the abundance and diversity of prey, warm water and separation from adults make this region an ideal nursery ground, the potential for interaction with the local fisheries should be examined.     

Predation by White Sharks Carcharodon carcharias (Chondrichthyes: Lamnidae) Upon Chelonians, with New Records from the Mediterranean Sea and a First Record of the Ocean Sunfish Mola mola (Osteichthyes: Molidae) as Stomach Contents

The occurrence of marine turtles in the diet of white sharks, Carcharodon carcharias, is reviewed worldwide. Four records of chelonians eaten by white sharks in the Mediterranean Sea are described, which on the basis of carapace remnants confirmed both the loggerhead Caretta caretta and green turtle Chelonia mydas to be preyed upon in those waters. The condition of these remains indicates that large white sharks can ingest turtles essentially intact. As well as falling prey to white sharks, we suspect that some interactions involve turtles being grab-released in a non-predatory mannner and their survivability from such low-intensity bites or other mouthings may be quite high. The white shark may be the chief marine predator of adult chelonians in the Mediterranean Sea, albeit the impact of this predation upon turtle populations is nominal compared to other sources of mortality. Further, we give an account describing an adult ocean sunfish, Mola mola, in the stomach of a white shark taken in Italian waters.     

Regulation of body temperature in the white shark, Carcharodon carcharias

Stomach temperatures of three white sharks, Carcharodoncarcharias, (one reported previously and two new individuals) were intermittently recorded by acoustic telemetry at the South Farallon Islands, central California. Temperature profiles of the water column were obtained concurrently. Stomach temperatures were elevated over ambient water temperatures by as much as 14.3 °C. Stomach temperatures varied within a narrow range while ambient water temperature fluctuated over a much larger range, showing that this species regulates its body temperature. These data, in combination with previous work on the physiology and anatomy of white sharks, indicate that the white shark is endothermic. It appears that the heat retention system in lamnid sharks has allowed them to inhabit cold water and remain active predators of swift and agile prey. Accepted: 17 February 1997     

Effects of Smart Position Only (SPOT) tag deployment on white sharks Carcharodon carcharias in South Africa

We present 15 individual cases of sub-adult white sharks that were SPOT tagged in South Africa from 2003-2004 and have been re-sighted as recently as 2011. Our observations suggest SPOT tags can cause permanent cosmetic and structural damage to white shark dorsal fins depending on the duration of tag attachment. SPOT tags that detached within 12-24 months did not cause long term damage to the dorsal fin other than pigmentation scarring. Within 12 months of deployment, tag fouling can occur. After 24 months of deployment permanent damage to the dorsal fin occurred. A shark survived this prolonged attachment and there seems little compromise on the animal's long term survival and resultant body growth. This is the first investigation detailing the long term effects of SPOT deployment on the dorsal fin of white sharks.     

Fine scale movements and activity areas of white sharks (Carcharodon carcharias) in Mossel Bay, South Africa

Previous work on white sharks indicate the species show seasonally limited movement patters, at certain aggregation sites small areas may play vital roles in the life history of a large amount of the population. Acoustic telemetry was used to estimate habitat use of white sharks, Carcharodon carcharias, while aggregating at Mossel Bay, South Africa. Total range of all shark tracks combined accumulated 782 h and covered an area of 93.5 km² however, within this range, sharks were found to highly utilise a core habitat (50 % Kernel, K50) of just 1.05 km² over a reef system adjacent to a river mouth. Individual tracks revealed additional core habitats, some of which were previously undocumented and one adjacent to a commercial harbor. Much was found to be dependent on the size of the shark, with larger sharks (>400 cm) occupying smaller activity areas than subadult (300–399 cm) and juvenile (<300 cm) conspecifics, while Index of Reuse (IOR) and Index of Shared Space (IOSS) were both found to increase with shark size. Such results provide evidence that larger white sharks are more selective in habitat use, which indicates they have greater experience within aggregation sites. Furthermore, the focused nature of foraging means spatially restricted management strategies would offer a powerful tool to aid enforcement of current protective legislation for the white shark in similar environments of limited resources and capacity.     

Using stable isotope analysis to understand the migration and trophic ecology of northeastern Pacific white sharks (Carcharodon carcharias)

The white shark (Carcharodon carcharias) is a wide-ranging apex predator in the northeastern Pacific (NEP). Electronic tagging has demonstrated that white sharks exhibit a regular migratory pattern, occurring at coastal sites during the late summer, autumn and early winter and moving offshore to oceanic habitats during the remainder of the year, although the purpose of these migrations remains unclear. The purpose of this study was to use stable isotope analysis (SIA) to provide insight into the trophic ecology and migratory behaviors of white sharks in the NEP. Between 2006 and 2009, 53 white sharks were biopsied in central California to obtain dermal and muscle tissues, which were analyzed for stable isotope values of carbon (δ(13)C) and nitrogen (δ(15)N). We developed a mixing model that directly incorporates movement data and tissue incorporation (turnover) rates to better estimate the relative importance of different focal areas to white shark diet and elucidate their migratory behavior. Mixing model results for muscle showed a relatively equal dietary contribution from coastal and offshore regions, indicating that white sharks forage in both areas. However, model results indicated that sharks foraged at a higher relative rate in coastal habitats. There was a negative relationship between shark length and muscle δ(13)C and δ(15)N values, which may indicate ontogenetic changes in habitat use related to onset of maturity. The isotopic composition of dermal tissue was consistent with a more rapid incorporation rate than muscle and may represent more recent foraging. Low offshore consumption rates suggest that it is unlikely that foraging is the primary purpose of the offshore migrations. These results demonstrate how SIA can provide insight into the trophic ecology and migratory behavior of marine predators, especially when coupled with electronic tagging data.     

Travelling light: white sharks (Carcharodon carcharias) rely on body lipid stores to power ocean-basin scale migration

Many species undertake long-distance annual migrations between foraging and reproductive areas. Such migrants depend on the efficient packaging, storage and utilization of energy to succeed. A diverse assemblage of organisms accomplishes this through the use of lipid reserves; yet, it remains unclear whether the migrations of elasmobranchs, which include the largest gill breathers on Earth, depend on such a mechanism. We examine depth records from pop-up satellite archival tags to discern changes in buoyancy as a proxy for energy storage in Eastern Pacific white sharks, and assess whether lipid depletion fuels long-distance (approx. 4000 km) migrations. We develop new algorithms to assess body condition, buoyancy and drift rate during drift dives and validate the techniques using a captive white shark. In the wild, we document a consistent increase in drift rate over the course of all migrations, indicating a decrease in buoyancy caused by the depletion of lipid reserves. These results comprise, to our knowledge, the first assessment of energy storage and budgeting in migrating sharks. The methods provide a basis for further insights into using electronic tags to reveal the energetic strategies of a wide range of elasmobranchs.     

On the geometry and mechanics of tooth position in the white shark,Carcharodon carcharias

The teeth of captured specimens, of prepared museum specimens, and of high-speed videotape images of the white shark, Carcharodon carcharias, were compared with respect to (1) deviation of each tooth from the animal's midline and (2) the crown angle of the functional teeth along the jaw margin. Tooth position was measured either directly using a meter stick apparatus or derived from tracings of the video footage. Tooth positions were not statistically unique in any region of the upper or lower jaw but demonstrated less variability in crown angle within 30° of the midline (71.48° ± 10°). Videotape analysis of feeding sharks indicated an 8.7° increase in crown angle of the centermost teeth during bites where the jaws were closed through an angle of 20–35° and a 15.7° reduction in this same parameter during jaw adduction through 35° or more. Such changes in tooth orientation (relative to the rear of the buccal cavity) are ascribed to flexure of the cartilaginous jaws and cranium by the cranial musculature and possibly also to sliding of the tooth bed over the jaw. Outward rotation of the teeth and jaw rami describes a plucking action during feeding or prey sampling, while larger bites rotate the frontmost teeth inward towards the gullet. Functionally, this may make the teeth more effective at grasping small prey items or gouging chunks from larger prey. However, testing of the load required to remove teeth showed no significant increase in tensile resistance with reduced crown angle. © 1995 Wiley-Liss, Inc.     

A first estimate of white shark, Carcharodon carcharias, abundance off Central California

The decline of sharks in the global oceans underscores the need for careful assessment and monitoring of remaining populations. The northeastern Pacific is the home range for a genetically distinct clade of white sharks (Carcharodon carcharias). Little is known about the conservation status of this demographically isolated population, concentrated seasonally at two discrete aggregation sites: Central California (CCA) and Guadalupe Island, Mexico. We used photo-identification of dorsal fins in a sequential Bayesian mark-recapture algorithm to estimate white shark abundance off CCA. We collected 321 photographs identifying 130 unique individuals, and estimated the abundance off CCA to be 219 mature and sub-adult individuals ((130, 275) 95% credible intervals), substantially smaller than populations of other large marine predators. Our methods can be readily expanded to estimate shark population abundance at other locations, and over time, to monitor the status, population trends and protection needs of these globally distributed predators.     

Oldest record of the Great White Shark (Lamnidae,Carcharodon; Miocene) in the Southern Atlantic

Lamnid teeth close to but more plesiomorph than Carcharodon carcharias were collected in the late Miocene beds of the Paraná Formation in the central eastern Argentina. We propose, as other authors, that some species formerly assigned to Isurus or to Cosmopolitodus should be included in Carcharodon. Some workers suggested that C. carcharias originated by phyletic evolution in the Pacific basin. The teeth from Paraná could pertain to a new species of Carcharodon already known from Perú. In contrast with the Pacific basin, Carcharodon sp. was sympatric with the wide toothed species Carcharodon plicatilis in central Argentina. This is the oldest record of Carcharodon in the southern Atlantic.     

Vertebral Bomb Radiocarbon Suggests Extreme Longevity in White Sharks

Conservation and management efforts for white sharks (Carcharodon carcharias) remain hampered by a lack of basic demographic information including age and growth rates. Sharks are typically aged by counting growth bands sequentially deposited in their vertebrae, but the assumption of annual deposition of these band pairs requires testing. We compared radiocarbon (Δ¹⁴C) values in vertebrae from four female and four male white sharks from the northwestern Atlantic Ocean (NWA) with reference chronologies documenting the marine uptake of ¹⁴C produced by atmospheric testing of thermonuclear devices to generate the first radiocarbon age estimates for adult white sharks. Age estimates were up to 40 years old for the largest female (fork length [FL]: 526 cm) and 73 years old for the largest male (FL: 493 cm). Our results dramatically extend the maximum age and longevity of white sharks compared to earlier studies, hint at possible sexual dimorphism in growth rates, and raise concerns that white shark populations are considerably more sensitive to human-induced mortality than previously thought.     

Plant-Soil-Contaminant Specificity Affects Phytoremediation of Organic Contaminants

The objective of this study was the evaluation of seven forage and conservation crop species for phytoremediation of trinitrotoluene (TNT) and pyrene-contaminated soils. TNT and pyrene were added to soil at 100 mg kg^-1. Crop species screening studies were conducted in a greenhouse and growth chambers on two soil types with different organic matter contents. Under high soil organic matter conditions, adsorption or covalent binding to the soil organic matter appeared to be a dominant force of removal limiting TNT and pyrene availability. In both soil types, pyrene dissipation could not be attributed to the presence of plants. However, in soils with lower organic matter content, all of the plant species treatments showed a significantly higher degree of TNT transformation compared with the unplanted control. Statistically significant differences in TNT transformation were observed among crop species grown in the low OM soil. Reed canary grass (Phalaris arundinacea L.) and switchgrass (Panicum virgatum L.) were the most effective species in enhancing TNT transformation. Our data indicated that use of plants was effective for phytoremediation of TNT-contaminated low OM content soils, but did not have any significant effect on pyrene dissipation. Based on these observations, it appears that plant-soil-contaminant interactions are very specific, and this specificity determines the effectiveness of phytoremediation schemes.     

Benefits of Vegetation for Soils with Organic Contaminants

Referee: Dr. Paul E. Olson, Colorado State University, Department of Biology, E414 Anatomy/Zoology Building, Fort Collins, CO 80523-1878 Plants have many beneficial effects on contaminated soils, including direct metabolism of some organic compounds, stimulation of microbial activity in the root zone, extraction of water, reduction of infiltration, improved aeration of the soil and stabilization against wind and water erosion. Some of these benefits have been explored and exploited in the newly named field of phytoremediation, whereas others remain largely in the domain of agronomy and ecology. This review attempts an assessment of the major environmental impacts of plants in general, recently termed phytotechnology, while including some case studies where particular species have been shown to have impacts on certain contaminants, more specifically termed phytoremediation. There is growing evidence that plants can have significant benefits in phytoremediating a range of complex organic molecules, including pesticides, solvents, explosives, and industrial byproducts.