Dorsal fin in the white shark, Carcharodon carcharias: a dynamic stabilizer for fast swimming

Transverse sections of the skin in the dorsal fin of the white shark, Carcharodon carcharias, tiger shark, Galeocerdo cuvier, and spotted raggedtooth shark, Carcharias taurus, show large numbers of dermal fiber bundles, which extend from the body into the fin. The bundles are tightly grouped together in staggered formation (not arranged in a straight line or in rows). This arrangement of dermal fibers gives tensile strength without impeding fiber movement. Tangential sections indicate that the fibers in all three species are strained and lie at angles in excess of 60 degrees . Of the three species investigated the dermal fibers in C. carcharias are the most densely concentrated and extend furthest distally along the dorsal fin. The overall results indicate that the dorsal fin of C. carcharias functions as a dynamic stabilizer and that the dermal fibers are crucial to this role. The fibers work like riggings that stabilize a ship's mast. During fast swimming, when the problems of yaw and roll are greatest, hydrostatic pressure within the shark increases and the fibers around the body, including in the dorsal fin, become taut, thereby stiffening the fin. During slow swimming and feeding the hydrostatic pressure is reduced, the fibers are slackened, and the muscles are able to exert greater bending forces on the fin via the radials and ceratotrichia. In C. carcharias there is a trade-off for greater stiffness of the dorsal fin against flexibility.     

A streamlined, bi-organelle, multiplex PCR approach to species identification: Application to global conservation and trade monitoring of the great white shark, Carcharodon carcharias

The great white shark, Carcharodoncarcharias, is the most widely protectedelasmobranch in the world, and is classified asVulnerable by the IUCN and listed on AppendixIII of CITES. Monitoring of trade in whiteshark products and enforcement of harvest andtrade prohibitions is problematic, however, inlarge part due to difficulties in identifyingmarketed shark parts (e.g., dried fins, meatand processed carcasses) to species level. Toaddress these conservation and managementproblems, we have developed a rapid, moleculardiagnostic assay based on species-specific PCRprimer design for accurate identification ofwhite shark body parts, including dried fins. The assay is novel in several respects: Itemploys a multiplex PCR assay utilizing bothnuclear (ribosomal internal transcribed spacer2) and mitochondrial (cytochrome b) locisimultaneously to achieve a highly robustmeasure of diagnostic accuracy; it is verysensitive, detecting the presence of whiteshark DNA in a mixture of genomic DNAs from upto ten different commercially fished sharkspecies pooled together in a single PCR tube;and it successfully identifies white shark DNAfrom globally distributed animals. Inaddition to its utility for white shark trademonitoring and conservation applications, thishighly streamlined, bi-organelle, multiplex PCRassay may prove useful as a general model forthe design of genetic assays aimed at detectingbody parts from other protected and threatenedspecies.     

Caudal fin in the white shark, Carcharodon carcharias (Lamnidae): a dynamic propeller for fast, efficient swimming

The caudal peduncle and caudal fin of Carcharodon carcharias together form a dynamic locomotory structure. The caudal peduncle is a highly modified, dorsoventrally compressed and rigid structure that facilitates the oscillations of the caudal fin. Its stiffness appears to be principally achieved by a thick layer of adipose tissue ranging from 28-37% of its cross-sectional area, reinforced by cross-woven collagen fibers. Numerous overlying layers of collagen fibers of the stratum compactum, oriented in steep left- and right-handed helices (approximately 65 degrees to the shark's long axis), prevent bowstringing of the perimysial fibers, which lie just below the dermal layer. Perimysial fibers, muscles, and the notochord are restricted to the dorsal lobe of the caudal fin and comprise the bulk of its mass. Adipose tissue reinforces the leading edge of the dorsal lobe of the caudal fin and contributes to maintaining the ideal cross-sectional geometry required of an advanced hydrofoil. Most of the mass of the ventral lobe consists of the ceratotrichia or fin rays separated by thin partitions of connective tissue. Dermal fibers of the stratum compactum of the dorsal lobe occur in numerous distinct layers. The layers are more complex than in other sharks and appear to reflect a hierarchical development in C. carcharias. The fiber layer comprises a number of thick fiber bundles along the height of the layer and the layers get thicker deeper into the stratum compactum. Each of these layers alternates with a layer a single fiber-bundle deep, a formation thought to give stability to the stratum compactum and to enable freer movements of the fiber system. In tangential sections of the stratum compactum the fiber bundles in the dorsal lobe can be seen oriented with respect to the long axis of the shark at approximately 55-60 degrees in left- and right-handed helices. Because of the backward sweep of the dorsal lobe (approximately 55 degrees to the shark's long axis) the right-handed fibers also parallel the lobe's long axis. In the dorsal lobe, ceratotrichia are present only along the leading edge (embedded within connective tissue), apparently as reinforcement. Stratum compactum fiber bundles of the ventral lobe, viewed in transverse section, lack the well-ordered distinctive layers of the dorsal lobe, but rather occur as irregularly arranged masses of tightly compacted fiber bundles of various sizes. In tangential sections the fiber bundles are oriented at angles of approximately 60 degrees, generally in one direction, i.e., lacking the left- and right-handed helical pattern. Tensile load tests on the caudal fin indicate high passive resistance to bending by the skin. The shear modulus G showed that the skin's contribution to stiffness (average values from three specimens at radians 0.52 and 1.05) is 33.5% for the dorsal lobe and 41.8% for the ventral. The load tests also indicate greater bending stiffness of the ventral lobe compared to the dorsal. Overall, the anatomy and mechanics of the dorsal lobe of C. carcharias facilitate greater control of movement compared to the ventral lobe. The helical fiber architecture near the surface of the caudal fin is analogous to strengthening of a thin cylinder in engineering. High fiber angles along the span of the dorsal lobe are considered ideal for resisting the bending stresses that the lobe is subjected to during the locomotory beat cycle. They are also ideal for storing strain energy during bending of the lobe and consequently may be of value in facilitating the recovery stroke. The complex fiber architecture of the caudal fin and caudal peduncle of C. carcharias provides considerable potential for an elastic mechanism in the animal's swimming motions and consequently for energy conservation.     

Implications of life history uncertainty when evaluating status in the Northwest Atlantic population of white shark (Carcharodon carcharias)

To effectively protect at‐risk sharks, resource managers and conservation practitioners must have a good understanding of how fisheries removals contribute to changes in abundance and how regulatory restrictions may impact a population trajectory. This means they need to know the number of animals being removed from a population and whether a given number of removals will lead to population increases or declines. For white shark (Carcharodon carcharias), theoretical quantities like the intrinsic rate of population increase or rebound potential (ability to increase in size following decline) are difficult to conceptualize in terms of real‐world abundance changes, which limits our ability to answer practical management questions. To address this shortfall, we designed a simulation model to evaluate how our understanding of longevity and life history variability of white shark affects our understanding of population trends in the Northwest Atlantic. Then, we quantified the magnitude of removals that could have caused historical population declines, compared these to biologically based reference points, and explored the removal scenarios which would result in population increase. Our results suggest that removals on the order of 100s of juveniles per year could have resulted in population‐level declines in excess of 60% during the 1970s and 1980s. Conservation actions implemented since the 1990s would have needed to be nearly 100% effective at preventing fishing mortality in order for the population to double in abundance over the last 30 years. Total removals from all fleets needed to be exceptionally small to keep them below biological reference points for white shark in the Northwest Atlantic. The population's inherent vulnerability to fishing pressure reaffirms the need for restrictive national and international conservation measures, even under a situation of abundance increase.     

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     

Genetic profiling reveals illegal international trade in fins of the great white shark, <Emphasis Type="Italic">Carcharodon carcharias</Emphasis>

Great white sharks are protected by national legislation in several countries, making this species the most widely protected elasmobranch in the world. Although the market demand for shark fins in general has continued to grow, the value and extent of utilization of white shark fins in trade has been controversial. We combine law enforcement with genetic profiling to demonstrate that illegal trade in fins of this species is occurring in the contemporary international market. Furthermore, we document the presence of fins from very young white sharks in the trade, suggesting a multiple-use market (food to trophies) exists for fins of this species. The presence of small fins in the trade contradicts the view that white shark fins have market value only as large display trophies, and not as food. Our findings indicate that effective conservation of protected shark species will require international management regimes that include monitoring of the shark fishery and trade on a species-specific basis.     

Effective number of white shark (Carcharodon carcharias,Linnaeus) breeders is stable over four successive years in the population adjacent to eastern Australia and New Zealand

Population size is a central parameter for conservation; however, monitoring abundance is often problematic for threatened marine species. Despite substantial investment in research, many marine species remain data‐poor presenting barriers to the evaluation of conservation management outcomes and the modeling of future solutions. Such is the case for the white shark (Carcharodon carcharias), a highly mobile apex predator for whom recent and substantial population declines have been recorded in many globally distributed populations. Here, we estimate the effective number of breeders that successfully contribute offspring in one reproductive cycle (Nb) to provide a snapshot of recent reproductive effort in an east Australian–New Zealand population of white shark. Nb was estimated over four consecutive age cohorts (2010, 2011, 2012, and 2013) using two genetic estimators (linkage disequilibrium; LD and sibship assignment; SA) based on genetic data derived from two types of genetic markers (single nucleotide polymorphisms; SNPs and microsatellite loci). While estimates of Nb using different marker types produced comparable estimates, microsatellite loci were the least precise. The LD and SA estimates of Nb within cohorts using SNPs were comparable; for example, the 2013 age cohort Nb(SA) was 289 (95% CI 200–461) and Nb(LD) was 208.5 (95% CI 116.4–712.7). We show that over the time period studied, Nb was stable and ranged between 206.1 (SD ± 45.9) and 252.0 (SD ± 46.7) per year using a combined estimate of Nb(LD+SA) from SNP loci. In addition, a simulation approach showed that in this population the effective population size (Ne) per generation can be expected to be larger than Nb per reproductive cycle. This study demonstrates how breeding population size can be monitored over time to provide insight into the effectiveness of recovery and conservation measures for the white shark, where the methods described here may be applicable to other data‐poor species of conservation concern.     

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.     

Behavior of Cape fur seals (Arctocephalus pusillus pusillus) in response to spatial variation in white shark (Carcharodon carcharias) predation risk

Foraging and predation risk are often separated at rookeries of marine central place foragers, thus offering an opportunity to gain insight into how predator‐avoidance shapes the behavior of prey. Here we compare the behavior of Cape fur seals (Arctocephalus pusillus pusillus) at two island rookeries with and without white shark (Carcharodon carcharias) predations, and assess seal behavior in relation to marked spatiotemporal variation in risk at the high‐risk site (Seal Island, South Africa). Our results show that seal behavior at the two sites is comparatively similar in summer, when predation risk is low at both sites, but not in winter. Compared to seals at the “low‐risk” site, seals at Seal Island avoided deep‐water habitat around the island at high risk times and restricted their use of this habitat in favor of safe, shallow waters when engaging in social and thermoregulatory behaviors. Seals increased their frequency of jostling, porpoising, and diving when moving through the danger zone and seals in groups were safer than single individuals. Overall, our results suggest that seal behavior around the high‐risk site is strongly affected by predation risk, and show this rookery to be an excellent predator‐prey system at which to evaluate long‐standing ecological hypotheses.     

Behavior of Cape fur seals (Arctocephalus pusillus pusillus) in relation to temporal variation in predation risk by white sharks (Carcharodon carcharias) around a seal rookery in False Bay,South Africa

The marked differences in predation risk posed by white sharks (Carcarodon carcarias) at island rookeries of Cape fur seals (Arctocephalus pusillus pusillus) offer a quasi‐experimental design within a natural system for exploring how prey adjust their behavior in response to temporal variation in predation risk. Here we compare movement of juvenile and adult Cape fur seals at a high risk (Seal Island) and low risk (Egg Island) rookery. We further compare juveniles and adults at Seal Island in low and high risk seasons and at low and high risk times of day within those seasons. Adult fur seals at Seal Island avoided traversing the zone of high white shark predation risk during the high risk period (0700–0959) in the season of high risk (winter), but not during the low risk season (summer). By contrast, adult fur seals at Egg Island showed no temporal discretion in either season. Unlike juvenile fur seals at Egg Island, juveniles at Seal Island adjusted their temporal movement patterns to more closely mimic adult seal movement patterns. This suggests that exposure to predators is the primary driver of temporal adjustments to movement by prey species commuting from a central place.     

Survival and abundance of polar bears in Alaska’s Beaufort Sea, 2001–2016

The Arctic Ocean is undergoing rapid transformation toward a seasonally ice‐free ecosystem. As ice‐adapted apex predators, polar bears (Ursus maritimus) are challenged to cope with ongoing habitat degradation and changes in their prey base driven by food‐web response to climate warming. Knowledge of polar bear response to environmental change is necessary to understand ecosystem dynamics and inform conservation decisions. In the southern Beaufort Sea (SBS) of Alaska and western Canada, sea ice extent has declined since satellite observations began in 1979 and available evidence suggests that the carrying capacity of the SBS for polar bears has trended lower for nearly two decades. In this study, we investigated the population dynamics of polar bears in Alaska''s SBS from 2001 to 2016 using a multistate Cormack–Jolly–Seber mark–recapture model. States were defined as geographic regions, and we used location data from mark–recapture observations and satellite‐telemetered bears to model transitions between states and thereby explain heterogeneity in recapture probabilities. Our results corroborate prior findings that the SBS subpopulation experienced low survival from 2003 to 2006. Survival improved modestly from 2006 to 2008 and afterward rebounded to comparatively high levels for the remainder of the study, except in 2012. Abundance moved in concert with survival throughout the study period, declining substantially from 2003 and 2006 and afterward fluctuating with lower variation around an average of 565 bears (95% Bayesian credible interval [340, 920]) through 2015. Even though abundance was comparatively stable and without sustained trend from 2006 to 2015, polar bears in the Alaska SBS were less abundant over that period than at any time since passage of the U.S. Marine Mammal Protection Act. The potential for recovery is likely limited by the degree of habitat degradation the subpopulation has experienced, and future reductions in carrying capacity are expected given current projections for continued climate warming.     

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.     

Development and microstructure of tooth histotypes in the blue shark,Prionace glauca (Carcharhiniformes: Carcharhinidae) and the great white shark,Carcharodon carcharias (Lamniformes: Lamnidae)

Elasmobranchs exhibit two distinct arrangements of mineralized tissues in the teeth that are known as orthodont and osteodont histotypes. Traditionally, it has been said that orthodont teeth maintain a pulp cavity throughout tooth development whereas osteodont teeth are filled with osteodentine and lack a pulp cavity when fully developed. We used light microscopy, scanning electron microscopy, and high‐resolution micro‐computed tomography to compare the structure and development of elasmobranch teeth representing the two histotypes. As an example of the orthodont histotype, we studied teeth of the blue shark, Prionace glauca (Carcharhiniformes: Carcharhinidae). For the osteodont histotype, we studied teeth of the great white shark, Carcharodon carcharias (Lamniformes: Lamnidae). We document similarities and differences in tooth development and the microstructure of tissues in these two species and review the history of definitions and interpretations of elasmobranch tooth histotypes. We discuss a possible correlation between tooth histotype and tooth replacement and review the history of histotype differentiation in sharks. We find that contrary to a long held misconception, there is no orthodentine in the osteodont teeth of C. carcharias. J. Morphol. 276:797–817, 2015. © 2015 Wiley Periodicals, Inc.     

Population dynamics of little brown bats (Myotis lucifugus) at summer roosts: Apparent survival,fidelity, abundance,and the influence of winter conditions

1. White‐nose syndrome (WNS) has caused the death of millions of bats, but the impacts have been more difficult to identify in western North America. Understanding how WNS, or other threats, impacts western bats may require monitoring other roosts, such as maternity roosts and night roosts, where bats aggregate in large numbers.
2. Little brown bats (Myotis lucifugus) are experiencing some of the greatest declines from WNS. Estimating survival and understanding population dynamics can provide valuable data for assessing population declines and informing conservation efforts.
3. We conducted a 5‐year mark–recapture study of two M. lucifugus roosts in Colorado. We used the robust design model to estimate apparent survival, fidelity, and abundance to understand population dynamics, and environmental covariates to understand how summer and winter weather conditions impact adult female survival. We compared the fidelity and capture probability of M. lucifugus between colonies to understand how bats use such roosts.
4. Overwinter survival increased with the number of days with temperatures below freezing (β > 0.100, SE = 0.003) and decreased with the number of days with snow cover (β < −0.40, SE < 0.13). Adult female fidelity was higher at one maternity roost than the other. Overwinter and oversummer adult female survival was high (>0.90), and based on survival estimates and fungal‐swabbing results, we believe these populations have yet to experience WNS.
5. Recapture of M. lucifugus using antennas that continuously read passive integrated transponder tags allows rigorous estimation of bat population parameters that can elucidate trends in abundance and changes in survival. Monitoring populations at summer roosts can provide unique population ecology data that monitoring hibernacula alone may not. Because few adult males are captured at maternity colonies, and juvenile males have low fidelity, additional effort should focus on understanding male M. lucifugus population dynamics.

     

Genetic population structure and demography of an apex predator,the tiger shark Galeocerdo cuvier

Population genetics has been increasingly applied to study large sharks over the last decade. Whilst large shark species are often difficult to study with direct methods, improved knowledge is needed for both population management and conservation, especially for species vulnerable to anthropogenic and climatic impacts. The tiger shark, Galeocerdo cuvier, is an apex predator known to play important direct and indirect roles in tropical and subtropical marine ecosystems. While the global and Indo‐West Pacific population genetic structure of this species has recently been investigated, questions remain over population structure and demographic history within the western Indian (WIO) and within the western Pacific Oceans (WPO). To address the knowledge gap in tiger shark regional population structures, the genetic diversity of 286 individuals sampled in seven localities was investigated using 27 microsatellite loci and three mitochondrial genes (CR, COI, and cytb). A weak genetic differentiation was observed between the WIO and the WPO, suggesting high genetic connectivity. This result agrees with previous studies and highlights the importance of the pelagic behavior of this species to ensure gene flow. Using approximate Bayesian computation to couple information from both nuclear and mitochondrial markers, evidence of a recent bottleneck in the Holocene (2,000–3,000 years ago) was found, which is the most probable cause for the low genetic diversity observed. A contemporary effective population size as low as 111 [43,369] was estimated during the bottleneck. Together, these results indicate low genetic diversity that may reflect a vulnerable population sensitive to regional pressures. Conservation measures are thus needed to protect a species that is classified as Near Threatened.     

A global synthesis of survival estimates for microbats

Accurate survival estimates are needed to construct robust population models, which are a powerful tool for understanding and predicting the fates of species under scenarios of environmental change. Microbats make up 17% of the global mammalian fauna, yet the processes that drive differences in demographics between species are poorly understood. We collected survival estimates for 44 microbat species from the literature and constructed a model to determine the effects of reproductive, feeding and demographic traits on survival. Our trait-based model indicated that bat species which produce more young per year exhibit lower apparent annual survival, as do males and juveniles compared with females and adults, respectively. Using 8 years of monitoring data for two Australian species, we demonstrate how knowledge about the effect of traits on survival can be incorporated into Bayesian survival analyses. This approach can be applied to any group and is not restricted to bats or even mammals. The incorporation of informative priors based on traits can allow for more timely construction of population models to support management decisions and actions.     

A population estimate of Heaviside's dolphins, Cephalorhynchus heavisidii, at the southern end of their range

Heaviside's dolphins, Cephalorhynchus heavisidii , are endemic to southwestern Africa, where they are exposed to unknown levels of anthropogenic threats, including inshore set netting. Using photo-ID data collected over 3 yr on the west coast of South Africa, we calculated Chapman's-modified Petersen estimates of the number of distinctive individuals at three spatial scales. Sample sizes were small and recapture rates low resulting in high variance. Total population abundance was extrapolated from the proportion of well-marked animals in the population (14%–17%) with between-year estimates adjusted for mortality using data from Commerson's dolphin. The total population size was calculated as 527 animals (CV = 0.35, CI 272–1,020) in the 1999 study area (20 km of coastline, within season), 3,429 animals (CV = 0.36, CI 1,721–6,828) in the central study area (150 km of coastline, 3 yr), and 6,345 animals (CV = 0.26, CI 3,573–11,267) in the full study area (390 km of coastline, 2 yr). Dolphins fitted with satellite transmitters varied in their use of the inshore photographic study area from 39.5% to 94.7% of transmission days (38–51 total). Given the known or suspected biases in the data, these abundance estimates are likely to be biased downward.