Natural or Artificial? Habitat-Use by the Bull Shark,Carcharhinus leucas

Background

Despite accelerated global population declines due to targeted and illegal fishing pressure for many top-level shark species, the impacts of coastal habitat modification have been largely overlooked. We present the first direct comparison of the use of natural versus artificial habitats for the bull shark, Carcharhinus leucas, an IUCN ‘Near-threatened’ species - one of the few truly euryhaline sharks that utilises natural rivers and estuaries as nursery grounds before migrating offshore as adults. Understanding the value of alternate artificial coastal habitats to the lifecycle of the bull shark is crucial for determining the impact of coastal development on this threatened but potentially dangerous species.

Methodology/Findings

We used longline surveys and long-term passive acoustic tracking of neonate and juvenile bull sharks to determine the ontogenetic value of natural and artificial habitats to bull sharks associated with the Nerang River and adjoining canals on the Gold Coast, Australia. Long-term movements of tagged sharks suggested a preference for the natural river over artificial habitat (canals). Neonates and juveniles spent the majority of their time in the upper tidal reaches of the Nerang River and undertook excursions into adjoining canals. Larger bull sharks ranged further and frequented the canals closer to the river mouth.

Conclusions/Significance

Our work suggests with increased destruction of natural habitats, artificial coastal habitat may become increasingly important to large juvenile bull sharks with associated risk of attack on humans. In this system, neonate and juvenile bull sharks utilised the natural and artificial habitats, but the latter was not the preferred habitat of neonates. The upper reaches of tidal rivers, often under significant modification pressure, serve as nursery sites for neonates. Analogous studies are needed in similar systems elsewhere to assess the spatial and temporal generality of this research.     

Swimming against the flow—Environmental DNA can detect bull sharks (Carcharhinus leucas) across a dynamic deltaic interface

Human activities in coastal areas are accelerating ecosystem changes at an unprecedented pace, resulting in habitat loss, hydrological modifications, and predatory species declines. Understanding how these changes potentially cascade across marine and freshwater ecosystems requires knowing how mobile euryhaline species link these seemingly disparate systems. As upper trophic level predators, bull sharks (Carcharhinus leucas) play a crucial role in marine and freshwater ecosystem health. Telemetry studies in Mobile Bay, Alabama, suggest that bull sharks extensively use the northern portions of the bay, an estuarine–freshwater interface known as the Mobile‐Tensaw Delta. To assess whether bull sharks use freshwater habitats in this region, environmental DNA surveys were conducted during the dry summer and wet winter seasons in 2018. In each season, 5 × 1 L water samples were collected at each of 21 sites: five sites in Mobile Bay, six sites in the Mobile‐Tensaw Delta, and ten sites throughout the Mobile‐Tombigbee and Tensaw‐Alabama Rivers. Water samples were vacuum‐filtered, DNA extractions were performed on the particulate, and DNA extracts were analyzed with Droplet Digital™ Polymerase Chain Reaction using species‐specific primers and an internal probe to amplify a 237‐base pair fragment of the mitochondrial NADH dehydrogenase subunit 2 gene in bull sharks. One water sample collected during the summer in the Alabama River met the criteria for a positive detection, thereby confirming the presence of bull shark DNA. While preliminary, this finding suggests that bull sharks use less‐urbanized, riverine habitats up to 120 km upriver during Alabama''s dry summer season.     

Trophic ecology of juvenile bull sharks (Carcharhinus leucas) in the Coyote estuary,Costa Rica

Bull shark (Carcharhinus leucas) is a near-threatened elasmobranch species capable of moving between the fresh and salty waters of tropical and subtropical coastal areas, for which we still lack important ecological information. During their first years of life, bull sharks use estuarine systems as nursery areas, making them highly susceptible to environmental and anthropogenic pressures. We studied the trophic ecology of juveniles found in the Coyote estuary, a potential nursery area in Costa Rica, to understand the potential impact of further bull shark declines and gain knowledge that could aid in their conservation. We analysed the trophic ecology of juvenile bull sharks [81–103 cm total length (TL)] in the Coyote estuary, Costa Rica, using stable isotopes of δ¹⁵N and δ¹³C. Since one problem using this technique in juveniles is the confounding effect of the maternal signature, we sampled different tissues (muscle and plasma), verified the status of the shark's umbilical scar and identified the size at which the isotope signature is a result of the animal's current diet. The isotopic values of the muscle tissue reflected the maternal isotopic signature. In contrast, plasma values reflected the diet of juvenile bull sharks >95 cm TL and with a closed umbilical scar. Juvenile bull sharks fed primarily on teleost fishes of the order Anguilliformes and Siluriformes, and have a high trophic position (≥4.0) in the Coyote estuary. Our findings suggest that this estuary is an important feeding site for juvenile bull sharks of the Pacific of Costa Rica. Thus, the protection of essential habitats such as the Coyote estuary will benefit not only bull shark conservation, but also the conservation of an array of fish species that also use this habitat as a rookery, many of which are of commercial interest.     

The shark nursery of Bulls Bay,South Carolina,with a review of the shark nurseries of the southeastern coast of the United States

Synopsis Shark nurseries, or nursery areas, are geographically discrete parts of a species range where the gravid females of most species of coastal sharks deliver their young or deposit their eggs, and where their young spend their first weeks, months, or years. These areas are usually located in shallow, energy rich coastal areas where the young find abundant food and have little predation by larger sharks. Nurseries are characterized by the presence of both gravid females and free swimming neonates. Neonates are young bearing fresh, unhealed umbilical scars in the case of placental species, or those at or near the birth size in aplacental species. Bulls Bay, South Carolina, is a nursery for the blacknose, spinner, finetooth, blacktip, sandbar, dusky, Atlantic sharpnose, scalloped hammerhead, and smooth dogfish sharks. The lemon shark has its nursery in shallow waters of south Florida and the Bahamas. The bull shark has its nursery in the lagoons of the east coast of central Florida.     

Residency,Habitat Use and Sexual Segregation of White Sharks,Carcharodon carcharias in False Bay,South Africa

White sharks (Carcharodon carcharias) are threatened apex predators and identification of their critical habitats and how these are used are essential to ensuring improved local and ultimately global white shark protection. In this study we investigated habitat use by white sharks in False Bay, South Africa, using acoustic telemetry. 56 sharks (39 female, 17 male), ranging in size from 1.7–5 m TL, were tagged with acoustic transmitters and monitored on an array of 30 receivers for 975 days. To investigate the effects of season, sex and size on habitat use we used a generalized linear mixed effects model. Tagged sharks were detected in the Bay in all months and across all years, but their use of the Bay varied significantly with the season and the sex of the shark. In autumn and winter males and females aggregated around the Cape fur seal colony at Seal Island, where they fed predominantly on young of the year seals. In spring and summer there was marked sexual segregation, with females frequenting the Inshore areas and males seldom being detected. The shift from the Island in autumn and winter to the Inshore region in spring and summer by females mirrors the seasonal peak in abundance of juvenile seals and of migratory teleost and elasmobranch species respectively. This study provides the first evidence of sexual segregation at a fine spatial scale and demonstrates that sexual segregation in white sharks is not restricted to adults, but is apparent for juveniles and sub-adults too. Overall, the results confirm False Bay as a critical area for white shark conservation as both sexes, across a range of sizes, frequent the Bay on an annual basis. The finding that female sharks aggregate in the Inshore regions when recreational use peaks highlights the need for ongoing shark-human conflict mitigation strategies.     

Distribution and Dynamic Habitat Use of Young Bull Sharks Carcharhinus leucas in a Highly Stratified Northern Gulf of Mexico Estuary

Understanding how animals alter habitat use in response to changing abiotic conditions is important for effective conservation management. For bull sharks (Carcharhinus leucas), habitat use has been widely examined in the eastern and western Gulf of Mexico; however, knowledge of their movements and the factors influencing them is lacking for populations in the more temperate north-central Gulf of Mexico. To examine how changes in hydrographic conditions affected the presence of young bull sharks in Mobile Bay, Alabama, thirty-five sharks were fitted with internal acoustic transmitters and monitored with an acoustic monitoring array consisting of thirty-three receivers between June 2009 and December 2010. Tagged sharks ranged in size from 60 to 114 cm fork length and were detected between the upper and lower portions of Mobile Bay. Despite a variety of freshwater sources associated with this highly productive estuary, sharks were most consistently detected at the largest input to the system – the Mobile and Tensaw Rivers. Our findings suggest a combination of hydrographic factors interact to influence the distribution of juvenile bull sharks in Mobile Bay. The factors affecting the probability of detecting at least one bull shark varied both temporally (2009 vs 2010) and spatially (upper vs lower bay). Electivity analysis demonstrated that bull sharks showed highest affinity for warm water (29–32°C), moderate salinities (10–11 psu) and normoxic waters (5–7 mg/l), although these patterns were not consistent between regions or across years. We suggest future studies coupling telemetry and hydrographic variables should, when possible, consider the interactions of multiple environmental parameters when defining the dynamic factors explaining the spatial distribution of coastal sharks.     

Spatial separation without territoriality in shark communities

Spatial separation within predator communities can arise via territoriality but also from competitive interactions among and within species. However, linking competitive interactions to predator distribution patterns is difficult and theoretical models predict different habitat selection patterns dependent on habitat quality and how competition manifests itself. While models generally consider competitors to be either equal in ability, or for one phenotype to have a fixed advantage over the other, few studies consider that an animal may only have a competitive advantage in specific habitats. We used  10 years of telemetry data, habitat surveys and behavioral experiments, to show spatial partitioning between and within two species of reef shark (grey reef Carcharhinus amblyrhinchos and blacktip reef sharks C. melanopterus) at an unfished Pacific atoll. Within a species, sharks remained within small ‘sub‐habitats’ with very few movements of individuals between sub‐habitats, which previous models have suggested could be caused by intra‐specific competition. Blacktip reef sharks were more broadly distributed across habitat types but a greater proportion used lagoon and backreef habitats, while grey reef sharks preferred forereef habitats. Grey reef sharks at a nearby atoll where blacktip reef sharks are absent, were distributed more broadly between habitat types than when both species were present. A series of individual‐based models predict that habitat separation would only arise if there are competitive interactions between species that are habitat‐specific, with grey reefs having a competitive advantage on the forereefs and blacktips in the lagoons and backreef. We provide compelling evidence that competition helps drive distribution patterns and spatial separation of a marine predator community, and highlight that competitive advantages may not be constant but rather dependent on habitats.     

Dramatic increase in sea otter mortality from white sharks in California

Although southern sea otters (Enhydra lutris nereis) are not considered prey for white sharks (Carcharodon carcharias), sharks do nonetheless bite sea otters. We analyzed spatial and temporal trends in shark bites on sea otters in California, assessing the frequency of shark bite wounds in 1,870 carcasses collected since 1985. The proportion of stranded sea otters having shark bites has increased sharply since 2003, and white shark bites now account for >50% of recovered carcasses. The trend was most pronounced in the southern part of the range, from Estero Bay to Point Conception, where shark bite frequency has increased eightfold. Seasonal trends were also evident: most shark‐bitten carcasses are recovered in late summer and fall; however, the period of elevated shark bite frequency has lengthened. The causes of these trends are unclear, but possible contributing factors include increased white shark abundance and/or changes in white shark behavior and distribution. In particular, the spatiotemporal patterns of shark‐bitten sea otters match increases in pinniped populations, and the increased availability of marine mammal prey for white sharks may have led to more sharks spending more time in nearshore waters utilized by both sea otters and pinnipeds.     

An integrated risk assessment for climate change: analysing the vulnerability of sharks and rays on Australia's Great Barrier Reef

An Integrated Risk Assessment for Climate Change (IRACC) is developed and applied to assess the vulnerability of sharks and rays on Australia's Great Barrier Reef (GBR) to climate change. The IRACC merges a traditional climate change vulnerability framework with approaches from fisheries ecological risk assessments. This semi‐quantitative assessment accommodates uncertainty and can be applied at different spatial and temporal scales to identify exposure factors, at‐risk species and their key biological and ecological attributes, critical habitats a`nd ecological processes, and major knowledge gaps. Consequently, the IRACC can provide a foundation upon which to develop climate change response strategies. Here, we describe the assessment process, demonstrate its application to GBR shark and ray species, and explore the issues affecting their vulnerability to climate change. The assessment indicates that for the GBR, freshwater/estuarine and reef associated sharks and rays are most vulnerable to climate change, and that vulnerability is driven by case‐specific interactions of multiple factors and species attributes. Changes in temperature, freshwater input and ocean circulation will have the most widespread effects on these species. Although relatively few GBR sharks and rays were assessed as highly vulnerable, their vulnerability increases when synergies with other factors are considered. This is especially true for freshwater/estuarine and coastal/inshore sharks and rays. Reducing the impacts of climate change on the GBR's sharks and rays requires a range of approaches including mitigating climate change and addressing habitat degradation and sustainability issues. Species‐specific conservation actions may be required for higher risk species (e.g. the freshwater whipray, porcupine ray, speartooth shark and sawfishes) including reducing mortality, preserving coastal catchments and estuarine habitats, and addressing fisheries sustainability. The assessment identified many knowledge gaps concerning GBR habitats and processes, and highlights the need for improved understanding of the biology and ecology of the sharks and rays of the GBR.     

The shark assemblage at French Frigate Shoals atoll, Hawai'i: species composition, abundance and habitat use

Empirical data on the abundance and habitat preferences of coral reef top predators are needed to evaluate their ecological impacts and guide management decisions. We used longline surveys to quantify the shark assemblage at French Frigate Shoals (FFS) atoll from May to August 2009. Fishing effort consisted of 189 longline sets totaling 6,862 hook hours of soak time. A total of 221 sharks from 7 species were captured, among which Galapagos (Carcharhinus galapagensis, 36.2%), gray reef (Carcharhinus amblyrhynchos, 25.8%) and tiger (Galeocerdo cuvier, 20.4%) sharks were numerically dominant. A lack of blacktip reef sharks (Carcharhinus melanopterus) distinguished the FFS shark assemblage from those at many other atolls in the Indo-Pacific. Compared to prior underwater visual survey estimates, longline methods more accurately represented species abundance and composition for the majority of shark species. Sharks were significantly less abundant in the shallow lagoon than adjacent habitats. Recaptures of Galapagos sharks provided the first empirical estimate of population size for any Galapagos shark population. The overall recapture rate was 5.4%. Multiple closed population models were evaluated, with Chao M(h) ranking best in model performance and yielding a population estimate of 668 sharks with 95% confidence intervals ranging from 289-1720. Low shark abundance in the shallow lagoon habitats suggests removal of a small number of sharks from the immediate vicinity of lagoonal islets may reduce short-term predation on endangered monk seal (Monachus schauinslandi) pups, but considerable fishing effort would be required to catch even a small number of sharks. Additional data on long-term movements and habitat use of sharks at FFS are required to better assess the likely ecological impacts of shark culling.     

Morphological scaling of body form in four shark species differing in ecology and life history

Body form can change across ontogeny, and can influence how animals of different sizes move and feed. Scaling data on live apex predatory sharks are rare and, therefore, we examined patterns of scaling in ontogenetic series of four sympatric shark species exhibiting a range of sizes, ecologies and life histories (tiger, bull, blacktip, and nurse shark). We evaluated 13 linear morphological variables and two areas (caudal and dorsal) that could influence both animal condition and locomotor performance. These measurements included dimensions of the dorsal, pectoral, and caudal fins, as well as several dimensions of body circumference, and of the head. For all four species, the body axis (eye‐to‐eye, lateral span, frontal span, proximal span) scaled close to isometry (expected slope of 1.0). The two largest sharks (tiger and bull sharks) also showed significant negative allometry for elements of the caudal fin. We found significant negative allometry in the lengths of the upper lobe of the caudal fin (caudal fin 1) and the overall height of the caudal fin (caudal fin 2) in tiger and bull sharks, with slopes ranging from about 0.60 to 0.73. Further, tiger sharks showed negative allometry in caudal fin area. These results suggest that in terms of overall body dimensions, small sharks are roughly geometrically similar to large sharks, at least within the species we examined. However, juvenile tiger (and to a lesser extent bull sharks) are notable in having proportionately larger caudal fins compared to adult sharks. As the caudal fin contributes to generating thrust during forward locomotion, this scaling implies differences among adult and juvenile sharks in locomotor ability. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 126–135.     

Defining distribution and habitat use of west‐central Florida’s coastal sharks through a research and education program

Identifying critical habitat for highly mobile species such as sharks is difficult, but essential for effective management and conservation. In regions where baseline data are lacking, non‐traditional data sources have the potential to increase observational capacity for species distribution and habitat studies. In this study, a research and education organization conducted a 5‐year (2013–2018) survey of shark populations in the coastal waters of west‐central Florida, an area where a diverse shark assemblage has been observed but no formal population analyses have been conducted. The objectives of this study were to use boosted regression tree (BRT) modeling to quantify environmental factors impacting the distribution of the shark assemblage, create species distribution maps from the model outputs, and identify spatially explicit hot spots of high shark abundance. A total of 1036 sharks were captured, encompassing eleven species. Abundance hot spots for four species and for immature sharks (collectively) were most often located in areas designated as “No Internal Combustion Engine” zones and seagrass bottom cover, suggesting these environments may be fostering more diverse and abundant populations. The BRT models were fitted for immature sharks and five species where n > 100: the nurse shark (Ginglymostoma cirratum), blacktip shark (Carcharhinus limbatus), blacknose shark (C. acronotus), Atlantic sharpnose shark (Rhizoprionodon terraenovae), and bonnethead (Sphyrna tiburo). Capture data were paired with environmental variables: depth (m), sea surface temperature (°C), surface, middle, and bottom salinity (psu), dissolved oxygen (mg/L), and bottom type (seagrass, artificial reef, or sand). Depth, temperature, and bottom type were most frequently identified as predictors with the greatest marginal effect on shark distribution, underscoring the importance of nearshore seagrass and barrier island habitats to the shark assemblage in this region. This approach demonstrates the potential contribution of unconventional science to effective management and conservation of coastal sharks.     

Habitat partitioning and vulnerability of sharks in the Great Barrier Reef Marine Park

Sharks present a critical conservation challenge, but little is known about their spatial distribution and vulnerability, particularly in complex seascapes such as Australia’s Great Barrier Reef Marine Park (GBRMP). We review (1) the distribution of shark species among the primary habitats of the GBRMP (coral reefs, inshore/shelf, pelagic and deep-water habitats) (2) the relative exploitation of each species by fisheries, and (3) how current catch rates interact with their vulnerability and trophic index. Excluding rays and chimaeras, we identify a total of 82 shark species in the GBRMP. We find that shark research in the GBRMP has yielded little quantitative information on most species. Reef sharks are largely site-fidelic, but can move large distances and some regularly use non-reef habitats. Inshore and shelf sharks use coastal habitats either exclusively or during specific times in their life cycle (e.g. as nurseries). Virtually nothing is known about the distribution and habitat use of the GBRMP’s pelagic and deep-water sharks. At least 46 species (53.5 %) are caught in one or more fisheries, but stock assessments are lacking for most. At least 17 of the sharks caught are considered highly vulnerable to exploitation. We argue that users of shark resources should be responsible for demonstrating that a fishery is sustainable before exploitation is allowed to commence or continue. This fundamental change in management principle will safeguard against stock collapses that have characterised many shark fisheries.     

A Comparison of Spatial and Movement Patterns between Sympatric Predators: Bull Sharks (Carcharhinus leucas) and Atlantic Tarpon (Megalops atlanticus)

Background

Predators can impact ecosystems through trophic cascades such that differential patterns in habitat use can lead to spatiotemporal variation in top down forcing on community dynamics. Thus, improved understanding of predator movements is important for evaluating the potential ecosystem effects of their declines.

Methodology/Principal Findings

We satellite-tagged an apex predator (bull sharks, Carcharhinus leucas) and a sympatric mesopredator (Atlantic tarpon, Megalops atlanticus) in southern Florida waters to describe their habitat use, abundance and movement patterns. We asked four questions: (1) How do the seasonal abundance patterns of bull sharks and tarpon compare? (2) How do the movement patterns of bull sharks and tarpon compare, and what proportion of time do their respective primary ranges overlap? (3) Do tarpon movement patterns (e.g., straight versus convoluted paths) and/or their rates of movement (ROM) differ in areas of low versus high bull shark abundance? and (4) Can any general conclusions be reached concerning whether tarpon may mitigate risk of predation by sharks when they are in areas of high bull shark abundance?

Conclusions/Significance

Despite similarities in diet, bull sharks and tarpon showed little overlap in habitat use. Bull shark abundance was high year-round, but peaked in winter; while tarpon abundance and fishery catches were highest in late spring. However, presence of the largest sharks (>230 cm) coincided with peak tarpon abundance. When moving over deep open waters (areas of high shark abundance and high food availability) tarpon maintained relatively high ROM in directed lines until reaching shallow structurally-complex areas. At such locations, tarpon exhibited slow tortuous movements over relatively long time periods indicative of foraging. Tarpon periodically concentrated up rivers, where tracked bull sharks were absent. We propose that tarpon trade-off energetic costs of both food assimilation and osmoregulation to reduce predation risk by bull sharks.     

Evidence of Partial Migration in a Large Coastal Predator: Opportunistic Foraging and Reproduction as Key Drivers?

Understanding animal movement decisions that involve migration is critical for evaluating population connectivity, and thus persistence. Recent work on sharks has shown that often only a portion of the adult population will undertake migrations, while the rest may be resident in an area for long periods. Defining the extent to which adult sharks use specific habitats and their migratory behaviour is essential for assessing their risk of exposure to threats such as fishing and habitat degradation. The present study used acoustic telemetry to examine residency patterns and migratory behaviour of adult bull sharks (Carcharhinus leucas) along the East coast of Australia. Fifty-six VR2W acoustic receivers were used to monitor the movements of 33 bull sharks in the central Great Barrier Reef (GBR). Both males and females were detected year-round, but their abundance and residency peaked between September and December across years (2012–2014). High individual variability in reef use patterns was apparent, with some individuals leaving the array for long periods, whereas others (36%) exhibited medium (0.20–0.40) or high residency (> 0.50). A large portion of the population (51%) undertook migrations of up to 1,400 km to other coral reefs and/or inshore coastal habitats in Queensland and New South Wales. Most of these individuals (76%) were mature females, and the timing of migrations coincided with the austral summer (Dec-Feb). All migrating individuals (except one) returned to the central GBR, highlighting its importance as a potential foraging ground. Our findings suggest that adult bull sharks appear to be highly dependent on coral reef resources and provide evidence of partial migration, where only a portion of the female population undertook seasonal migrations potentially to give birth. Given that estuarine habitats face constant anthropogenic pressures, understanding partial migration and habitat connectivity of large coastal predators should be a priority for their management.     

Phylogeography and conservation of the bull shark (Carcharhinus leucas) inferred from mitochondrial and microsatellite DNA

The bull shark (Carcharhinus leucas) is a widely distributed, large coastal shark species known to travel long distances. These characteristics, coupled with the species?? long life span and late age of maturity, would lead one to predict significant global genetic exchange among bull shark populations. By contrast, data show localized depletion in some areas of large coastal shark fisheries, indicating some geographic isolation may exist. We examined genetic variation in the control region of mitochondrial DNA and at five nuclear microsatellite loci in bull sharks sampled from the western Atlantic to investigate the degree of population subdivision. The average per sample haplotype and nucleotide diversity in the mtDNA (0.51 ± 0.26 and 0.12% ± 0.12, respectively) and expected heterozygosity (0.84) in the microsatellite loci contrast sharply in having lower and higher values (respectively) relative to many other shark species. Significant structure exists between the Brazilian and all northern populations at the mtDNA control region (pairwise ??_ST > 0.8, P < 0.001), but not at the nuclear microsatellite loci. Adjacent northern populations show weak to no genetic differentiation for both markers. These results are congruent with restricted maternal gene flow between populations caused by female site fidelity to nursery areas. We estimate the current effective population size to be around 160,000 and 221,000 individuals for the southern and northern Atlantic populations, respectively. The philopatric habits and the relatively low levels of mtDNA genetic diversity observed in bull sharks must be considered in the conservation of this species. Our results indicate that effective bull shark management strategies will require local, regional, and international attention and cooperation.     

Spatial and temporal analysis of juvenile blacktip reef shark (Carcharhinus melanopterus) demographies identifies critical habitats

Reef shark species have undergone sharp declines in recent decades, as they inhabit coastal areas, making them an easy target in fisheries (i.e., sharks are exploited globally for their fins, meat, and liver oil) and exposing them to other threats (e.g., being part of by-catch, pollution, and climate change). Reef sharks play a critical role in coral reef ecosystems, where they control populations of smaller predators and herbivorous fishes either directly via predation or indirectly via behavior, thus protecting biodiversity and preventing potential overgrazing of corals. The urgent need to conserve reef shark populations necessitates a multifaceted approach to policy at local, federal, and global levels. However, monitoring programmes to evaluate the efficiency of such policies are lacking due to the difficulty in repeatedly sampling free-ranging, wild shark populations. Over nine consecutive years, we monitored juveniles of the blacktip reef shark (Carcharhinus melanopterus) population around Moorea, French Polynesia, and within the largest shark sanctuary globally, to date. We investigated the roles of spatial (i.e., sampling sites) and temporal variables (i.e., sampling year, season, and month), water temperature, and interspecific competition on shark density across 10 coastal nursery areas. Juvenile C. melanopterus density was found to be stable over 9 years, which may highlight the effectiveness of local and likely federal policies. Two of the 10 nursery areas exhibited higher juvenile shark densities over time, which may have been related to changes in female reproductive behavior or changes in habitat type and resources. Water temperatures did not affect juvenile shark density over time as extreme temperatures proven lethal (i.e., 33°C) in juvenile C. melanopterus might have been tempered by daily variation. The proven efficiency of time-series datasets for reef sharks to identify critical habitats (having the highest juvenile shark densities over time) should be extended to other populations to significantly contribute to the conservation of reef shark species.     

The effect of habitat on modern shark diversification

Sharks occupy marine habitats ranging from shallow, inshore environments to pelagic, and deepwaters, and thus provide a model system for testing how gross habitat differences have shaped vertebrate macroevolution. Palaeontological studies have shown that onshore lineages diversify more quickly than offshore taxa. Among onshore habitats, coral reef‐association has been shown to increase speciation rates in several groups of fishes and invertebrates. In this study, we investigated whether speciation rates are habitat dependent by generating the first comprehensive molecular timescale for shark divergence. Using phylogenetic comparative methods, we rejected the hypothesis that shelf (i.e. onshore) lineages have higher speciation rates compared to those occupying deepwater and oceanic (i.e. offshore) habitats. Our results, however, support the hypothesis of increased speciation rates in coral reef‐associated lineages within the Carcharhinidae. Our new timetree suggests that the two major shark lineages leading to the extant shark diversity began diversifying mostly after the end‐Permian mass extinction: the squalimorphs into deepwater and the galeomorphs into shelf habitats. We suggest that the breakdown of the onshore–offshore speciation rate pattern in sharks is mediated by success in deepwater environments through ecological partitioning, and in some cases, the evolution of morphological novelty.     

Comparative Feeding Ecology of Bull Sharks (Carcharhinus leucas) in the Coastal Waters of the Southwest Indian Ocean Inferred from Stable Isotope Analysis

As apex predators, sharks play an important role shaping their respective marine communities through predation and associated risk effects. Understanding the predatory dynamics of sharks within communities is, therefore, necessary to establish effective ecologically based conservation strategies. We employed non-lethal sampling methods to investigate the feeding ecology of bull sharks (Carcharhinus leucas) using stable isotope analysis within a subtropical marine community in the southwest Indian Ocean. The main objectives of this study were to investigate and compare the predatory role that sub-adult and adult bull sharks play within a top predatory teleost fish community. Bull sharks had significantly broader niche widths compared to top predatory teleost assemblages with a wide and relatively enriched range of δ¹³C values relative to the local marine community. This suggests that bull sharks forage from a more diverse range of δ¹³C sources over a wider geographical range than the predatory teleost community. Adult bull sharks appeared to exhibit a shift towards consistently higher trophic level prey from an expanded foraging range compared to sub-adults, possibly due to increased mobility linked with size. Although predatory teleost fish are also capable of substantial migrations, bull sharks may have the ability to exploit a more diverse range of habitats and appeared to prey on a wider diversity of larger prey. This suggests that bull sharks play an important predatory role within their respective marine communities and adult sharks in particular may shape and link ecological processes of a variety of marine communities over a broad range.     

Feeding of the megamouth shark (Pisces: Lamniformes: Megachasmidae) predicted by its hyoid arch: A biomechanical approach

Studies of the megamouth shark, one of three planktivorous sharks, can provide information about their evolutionary history. Megamouth shark feeding has never been observed in life animals, but two alternative hypotheses on biomechanics suggest either feeding, i.e., ram feeding or suction feeding. In this study, the second moment of area of the ceratohyal cartilages, which is an indicator of the flexural stiffness of the cartilages, is calculated for 21 species of ram‐ and suction‐feeding sharks using computed tomography. The results indicate that suction‐feeding sharks have ceratohyal cartilages with a larger second moment of area than ram‐feeding sharks. The result also indicates that the ram–suction index, which is an indicator of relative contribution of ram and suction behavior, is also correlated with the second moment of area of the ceratohyal. Considering that large bending stresses are expected to be applied to the ceratohyal cartilage during suction, the larger second moment of area of the ceratohyal of suction‐feeding sharks can be interpreted as an adaptation for suction feeding. Based on the small second moment of area of the ceratohyal cartilage of the megamouth shark, the feeding mode of the megamouth shark is considered to be ram feeding, similar to the planktivorous basking shark. From these results, an evolutionary scenario of feeding mechanics of three species of planktivorous sharks can be suggested. In this scenario, the planktivorous whale shark evolved ram feeding from a benthic suction‐feeding ancestor. Ram feeding in the planktivorous megamouth shark and the basking shark evolved from ram feeding swimming‐type ancestors and that both developed their unique filtering system to capture small‐sized prey. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.