Shark depredation: future directions in research and management.

Shark depredation is a complex social-ecological issue that affects a range of fisheries worldwide. Increasing concern about the impacts of shark depredation, and how it intersects with the broader context of fisheries management, has driven recent research in this area, especially in Australia and the United States. This review synthesises these recent advances and provides strategic guidance for researchers aiming to characterise the occurrence of depredation, identify the shark species responsible, and test deterrent and management approaches to reduce its impacts. Specifically, the review covers the application of social science approaches, as well as advances in video camera and genetic methods for identifying depredating species. The practicalities and considerations for testing magnetic, electrical, and acoustic deterrent devices are discussed in light of recent research. Key concepts for the management of shark depredation are reviewed, with recommendations made to guide future research and policy development. Specific management responses to address shark depredation are lacking, and this review emphasizes that a "silver bullet" approach for mitigating depredation does not yet exist. Rather, future efforts to manage shark depredation must rely on a diverse range of integrated approaches involving those in the fishery (fishers, scientists and fishery managers), social scientists, educators, and other stakeholders.

Diurnal activity patterns and habitat use of juvenile Pastinachus ater in a coral reef flat environment.

Stingrays are thought to play important ecological roles in coral reef ecosystems. However, little is known about juvenile stingray movement patterns and habitat use in coral reefs. This study used active acoustic telemetry to determine fine-scale diel movement patterns and habitat use of juvenile cowtail stingrays (Pastinachus ater) in a coral reef flat environment. Seven cowtail stingrays (4 males and 3 females) were manually tracked between April and December 2016. Each individual was tracked over 2 days, generating a total of 14 active tracks ranging from 4.91 to 9 h. Specimens moved at an average speed of 2.44 m min-1 ± 0.87 SE, with minimum distances travelled ranging from 546 to 1446 m. Tracking data showed that juvenile cowtail stingrays move in response to tidal cycles, moving faster and in straighter pathways during incoming and outgoing tides. Juvenile cowtail stingrays also showed a strong affinity to sand flat areas and mangrove edge areas. These areas provide food resources and potential refuges for juvenile rays to avoid predators. Coral reef flats were identified as secondary refuge for juveniles during the lowest tides. Future research is necessary to fully unveil the major drivers of juvenile cowtail stingray seasonal and ontogenetic movement patterns and habitat use within coral reef flat environments. This information is important to establish a full understanding of juvenile cowtail stingray ecology, but could also improve management and conservation policies.

The importance of sample size in marine megafauna tagging studies.

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

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

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

Crossing lines: a multidisciplinary framework for assessing connectivity of hammerhead sharks across jurisdictional boundaries.

Conservation and management of migratory species can be complex and challenging. International agreements such as the Convention on Migratory Species (CMS) provide policy frameworks, but assessments and management can be hampered by lack of data and tractable mechanisms to integrate disparate datasets. An assessment of scalloped (Sphyrna lewini) and great (Sphyrna mokarran) hammerhead population structure and connectivity across northern Australia, Indonesia and Papua New Guinea (PNG) was conducted to inform management responses to CMS and Convention on International Trade in Endangered Species listings of these species. An Integrated Assessment Framework (IAF) was devised to systematically incorporate data across jurisdictions and create a regional synopsis, and amalgamated a suite of data from the Australasian region. Scalloped hammerhead populations are segregated by sex and size, with Australian populations dominated by juveniles and small adult males, while Indonesian and PNG populations included large adult females. The IAF process introduced genetic and tagging data to produce conceptual models of stock structure and movement. Several hypotheses were produced to explain stock structure and movement patterns, but more data are needed to identify the most likely hypothesis. This study demonstrates a process for assessing migratory species connectivity and highlights priority areas for hammerhead management and research.

Diet-tissue discrimination factors and turnover of carbon and nitrogen stable isotopes in tissues of an adult predatory coral reef fish, Plectropomus leopardus.

RATIONALE: Stable isotope ratios (δ(13)C and δ(15)N values) provide a unique perspective into the ecology of animals because the isotope ratio values of consumers reflect the values in food. Despite the value of stable isotopes in ecological studies, the lack of species-specific experimentally derived diet-tissue discrimination factors (DTDFs) and turnover rates limits their application at a broad scale. Furthermore, most aquatic feeding experiments use temperate, fast-growing fish species and few have considered medium- to large-sized adults with low growth rates from tropical ecosystems. METHODS: A controlled-diet stable isotope feeding trial was conducted over a 196-day period for the adult predatory reef fish leopard coralgrouper (Plectropomus leopardus). This study calculated δ(13)C and δ(15)N DTDFs and turnover rates in five tissues (liver, plasma, red blood cells (RBC), fin, and muscle) using a continuous flow isotope ratio mass spectrometer equipped with an elemental analyzer. In addition, the effect of chemical lipid extraction (LE) on stable isotope values was examined for each tissue. RESULTS: Turnover was mainly influenced by metabolism (as opposed to growth) with LE δ(15)N half-life values lowest in fin (37 days) and plasma (66 days), and highest in RBC (88 days) and muscle (126 days). The diet-tissue discrimination factors for δ(15)N values in all tissues (Δ(15)N: -0.15 to 1.84‰) were typically lower than commonly reported literature values. Lipid extraction altered both δ(15) N and δ(13)C values compared with untreated samples; however, for the δ(15)N values, the differences were small (mean δ(15)N(LE-Bulk) <0.46‰ in all tissues). CONCLUSIONS: This study informs future interpretation of stable isotope data for medium- to large-sized fish and demonstrates that DTDFs developed for temperate fish species, particularly for δ(15)N values, may not apply to tropical species. Sampling of muscle and/or RBC is recommended for a relatively long-term representation of feeding habits, while plasma and/or fin should be used for a more recent indication of diet.

Regional movement patterns of a small-bodied shark revealed by stable-isotope analysis.

This study used stable-isotope analysis to define the nearshore regional residency and movements of the small-bodied Australian sharpnose shark Rhizoprionodon taylori. Plasma and muscle δ(13) C and δ(15) N of R. taylori were collected from across five embayments and compared with values of seagrass and plankton from each bay. Linear distances between adjacent bays ranged from 30 to 150 km. There was a positive geographic correlation between R. taylori tissue and environmental δ(13) C values. Populations with the highest tissue δ(15) N were collected from bays that had the highest environmental δ(15) N values. These results suggest that R. taylori did not forage more than 100 km away from their capture location within 6 months to 1 year. The successful application of isotope analysis to define R. taylori movement demonstrates that this technique may be used in addition to traditional methods to study the movement of sharks, even within similar habitats across regionally small spatial scales (<100 km).

Age and growth parameters of shark-like batoids.

Estimates of life-history parameters were made for shark-like batoids of conservation concern Rhynchobatus spp. (Rhynchobatus australiae, Rhynchobatus laevis and Rhynchobatus palpebratus) and Glaucostegus typus using vertebral ageing. The sigmoid growth functions, Gompertz and logistic, best described the growth of Rhynchobatus spp. and G. typus, providing the best statistical fit and most biologically appropriate parameters. The two-parameter logistic was the preferred model for Rhynchobatus spp. with growth parameter estimates (both sexes combined) L(∞) = 2045 mm stretch total length, LST and k = 0·41 year⁻¹. The same model was also preferred for G. typus with growth parameter estimates (both sexes combined) L∞ = 2770 mm LST and k = 0·30 year⁻¹. Annual growth-band deposition could not be excluded in Rhynchobatus spp. using mark-recaptured individuals. Although morphologically similar G. typus and Rhynchobatus spp. have differing life histories, with G. typus longer lived, slower growing and attaining a larger maximum size.

Population structure and residency patterns of the blacktip reef shark Carcharhinus melanopterus in turbid coastal environments.

This study examined the characteristics of a blacktip reef shark Carcharhinus melanopterus population in turbid coastal habitats through a multi-year fishery-independent sampling and tag-recapture programme. Results revealed a highly structured population comprised almost entirely of juveniles and adult females with individuals between 850 and 1050 mm total length effectively absent. Mature males were also rarely encountered with adult sex ratio highly biased towards females (female:male = 7:1). Mating scars were observed on adult females between December and April, and parturition was observed from December to March. Regression analysis showed that catch rates were significantly higher during the summer wet season between November and May. Recapture data suggested a highly resident population with a recapture rate of 21% and a mean recapture distance of 0·8 km. In addition, 33% of recaptured animals were captured multiple times, indicating long-term residency. Most recaptures were, however, of adults with few juveniles recaptured. Widespread sampling at the study site and in adjacent areas suggested that the population was highly localized to a specific bay. The bimodal and sex-segregated population structure observed here differs from previous reports for this species, and in combination with reproductive observations, suggests population structuring to facilitate reproductive and recruitment success. These data also highlight the potential ecosystem functions performed by coastal habitats in sustaining C. melanopterus populations.

Blood lactate loads of redthroat emperor Lethrinus miniatus associated with angling stress and exhaustive exercise.

Baseline, post-angling and maximum attainable blood lactate concentrations were measured for the fishery species redthroat emperor Lethrinus miniatus to gain insight into the condition of fish released following c. 30 s angling and <45 s air exposure. Mean ± S.D. baseline blood lactate was 1·5 ± 0·6 mmol l⁻¹, which increased and plateaued around 6 mmol l⁻¹ at 15-30 min post-angling. These values were significantly lower than those obtained from fish maximally exhausted with a prolonged chase and air exposure protocol following capture (10·9 ± 1·8 mmol l⁻¹), suggesting that L. miniatus is not maximally exhausted during standard angling practices.

Habitat use and spatial segregation of adult spottail sharks Carcharhinus sorrah in tropical nearshore waters.

An array of acoustic receivers deployed in Cleveland Bay, north Queensland, Australia, passively tracked 20 adult spottail sharks Carcharhinus sorrah over 2 years (2009-2010) to define patterns in movement and habitat use. Individuals were present in the study site for long periods, ranging from 8 to 408 days (mean = 185). Size and location of home ranges did not vary over time. A high level of segregation occurred among C. sorrah, with individuals using different types of habitat and showing strong attachment to specific regions. The depth of habitat individuals used varied between sexes. Males tended to use a narrow range of habitat depths within the study site (2·8-6·0 m), whereas females used shallower habitats (1·4-6·2 m) and displayed a seasonal shift in the depth of habitat used. Mean monthly habitat depth used varied by as much as 2 m for females, with individuals using shallower habitats during the winter months. Long-term presence and consistent home ranges suggest that Cleveland Bay provides important habitat for C. sorrah. By defining patterns in the use of nearshore habitats for C. sorrah, this study improves the understanding of the movement and habitat use of smaller-bodied coastal sharks and may help provide guidance for the management of their populations.

Habitat use and movement patterns of bull sharks Carcharhinus leucas determined using pop-up satellite archival tags.

Habitat use, movement and residency of bull sharks Carcharhinus leucas were determined using satellite pop-up archival transmitting (PAT) tags throughout coastal areas in the U.S., Gulf of Mexico and waters off the south-east U.S. From 2005 to 2007, 18 fish (mean size = 164 cm fork length, L(F)) were tagged over all seasons. Fish retained tags for up to 85 days (median = 30 days). Based on geolocation data from initial tagging location to pop-off location, C. leucas generally travelled c. 5-6 km day(-1) and travelled an average of 143.6 km. Overall, mean proportions of time at depth revealed C. leucas spent the majority of their time in waters <20 m. They exhibited significant differences among depths but were not found at a particular depth regardless of diurnal period. Most fish occupied temperatures c. 32 degrees C with individuals found mostly between 26 and 33 degrees C. Geolocation data for C. leucas were generally poor and varied considerably but tracks for two individuals revealed long distance movements. One fish travelled from the south-east coast of the U.S. to coastal Texas near Galveston while another moved up the east coast of the U.S. to South Carolina. Data on C. leucas movements indicated that they are found primarily in shallower waters and tend to remain in the same location over long periods. While some individuals made large-scale movements over open ocean areas, the results emphasize the importance of the coastal zone for this species as potential essential habitat, particularly in areas of high freshwater inflow.

Science or slaughter: need for lethal sampling of sharks.

General consensus among scientists, commercial interests, and the public regarding the status of shark populations is leading to an increasing need for the scientific community to provide information to help guide effective management and conservation actions. Experience from other marine vertebrate taxa suggests that public, political, and media pressures will play an increasingly important part in setting research, management, and conservation priorities. We examined the potential implications of nonscientific influences on shark research. In particular, we considered whether lethal research sampling of sharks is justified. Although lethal sampling comes at a cost to a population, especially for threatened species, the conservation benefits from well-designed studies provide essential data that cannot be collected currently in any other way. Methods that enable nonlethal collection of life-history data on sharks are being developed (e.g., use of blood samples to detect maturity), but in the near future they will not provide widespread or significant benefits. Development of these techniques needs to continue, as does the way in which scientists coordinate their use of material collected during lethal sampling. For almost half of the known shark species there are insufficient data to determine their population status; thus, there is an ongoing need for further collection of scientific data to ensure all shark populations have a future. Shark populations will benefit most when decisions about the use of lethal sampling are made on the basis of scientific evidence that is free from individual, political, public, and media pressures.

Microsatellite and mitochondrial DNA analyses of the genetic structure of blacktip shark (Carcharhinus limbatus) nurseries in the northwestern Atlantic, Gulf of Mexico, and Caribbean Sea.

Abstract We investigated the genetic structure of blacktip shark (Carcharhinus limbatus) continental nurseries in the northwestern Atlantic Ocean, Gulf of Mexico, and Caribbean Sea using mitochondrial DNA control region sequences and eight nuclear microsatellite loci scored in neonate and young-of-the-year sharks. Significant structure was detected with both markers among nine nurseries (mitochondrial PhiST = 0.350, P < 0.001; nuclear PhiST = 0.007, P < 0.001) and sharks from the northwestern Atlantic, eastern Gulf of Mexico, western Gulf of Mexico, northern Yucatan, and Belize possessed significantly different mitochondrial DNA haplotype frequencies. Microsatellite differentiation was limited to comparisons involving northern Yucatan and Belize sharks with nuclear genetic homogeneity throughout the eastern Gulf of Mexico, western Gulf of Mexico, and northwestern Atlantic. Differences in the magnitude of maternal vs. biparental genetic differentiation support female philopatry to northwestern Atlantic, Gulf of Mexico, and Caribbean Sea natal nursery regions with higher levels of male-mediated gene flow. Philopatry has produced multiple reproductive stocks of this commercially important shark species throughout the range of this study.

Plasma steroid hormone profiles and reproductive biology of the epaulette shark, Hemiscyllium ocellatum.

Examination of the reproductive biology of the oviparous epaulette shark, Hemiscyllium ocellatum, was conducted on a wild population. Male sharks were found to reach maturity at between 55-60 cm total length (TL) and female sharks mature around 55 cm TL. Blood samples collected from mature male and female sharks were analyzed for sex steroid hormones to examine seasonal hormone patterns. Plasma samples were analyzed via radioimmunoassay techniques with female samples measured for estradiol, progesterone, and androgen concentrations, and male samples measured for androgen concentrations. Male androgen concentrations showed a single broad peak from July to October with maximum hormone concentrations (60 ng/ml) occurring in August. Male androgen concentrations were lowest in December-February (<20 ng/ml), and appeared to correlate with reproductive activity and water temperature. Female androgen concentrations were an order of magnitude lower than those for males and showed peaks in June (6 ng/ml) and December (8 ng/ml). Estradiol concentrations in females peaked during the months of September-November (0.5 ng/ml) coinciding with the egg laying period. Progesterone concentrations ranged up to 0.5 ng/ml prior to the mating season. Observations of ova size and egg production showed eggs develop in pairs and ova are ovulated at a size of 25-27 mm. Females lay eggs from August to January. Males were observed with swollen claspers from July through December, with the highest amount of sperm storage in the epididymis occurring between August through November. Our observations indicate that epaulette sharks in the waters near Heron Island mate from July through December. J. Exp. Zool. 284:586-594, 1999.

Doppler ultrasonographic evidence of intrapartum brain-sparing effect in fetuses with low oxygen saturation according to pulse oximetry.

OBJECTIVE: This study was undertaken to verify by means of Doppler ultrasonography and simultaneous fetal pulse oximetry the redistribution of fetal blood flow in favor of the brain during intrapartum hypoxemia. STUDY DESIGN: During labor 11 term fetuses with abnormal heart rate patterns and arterial oxygen saturation <30% and 14 control term fetuses with normal oxygen saturation were simultaneously monitored by pulse oximetry and Doppler ultrasonography. The results were compared with the Student t test. RESULTS: The blood flow velocity in the middle cerebral artery was significantly higher in the presence of reduced oxygen saturation, implying lower pulsatility and resistance indices (P <.001). The reduction of blood flow in the umbilical artery was not significant (P =.61). CONCLUSION: Simultaneous intrapartum pulse oximetry and Doppler ultrasonography proved that reduced arterial oxygen saturation (<30%) is associated with profound changes in fetal hemodynamics and could be tolerated for only a limited period, which should be the subject of further studies.

The occurrence, distribution and pathology associated with gnathiid isopod larvae infecting the epaulette shark, Hemiscyllium ocellatum.

Gnathiid isopod praniza larvae were found to infect the epaulette shark Hemiscyllium ocellatum. All sharks carried larvae on their external body surface, with the preferred attachment site in both sexes around the cloaca (P<0.05). The claspers were the second site of preference in male sharks. Within the buccal and branchial cavities, about 16% of larvae were attached to the roof and floor of the mouth and 84% attached to the gills. A significant positive correlation existed between larval number and fish size. Histological examination showed that larval attachment in the buccal cavity elicited variable responses, the most severe being a loss of epithelium and compression of underlying tissue. No host cellular response or tissue proliferation was observed. Praniza attached preferentially to the efferent side of gill filaments (relative to blood flow), and caused loss of epithelium, compression of tissue, and a small amount of connective tissue proliferation. Attachment to the gill septum or to the afferent side of the gill filament caused lamellar disruption, a cellular inflammatory response, and connective tissue proliferation. Scanning electron microscopy showed little obvious praniza-induced gill damage, other than localised tissue distortion to form "pockets" around larvae attached between filaments. The results suggest that praniza larvae do not cause sufficient tissue damage to adversely affect the health of this shark species.

The prediction of fetal acidosis by means of intrapartum fetal pulse oximetry.

OBJECTIVES: The study's objectives were to verify a threshold value for fetal arterial oxygen saturation as the critical boundary for fetal compromise during labor and to investigate a method of predicting acidosis caused by hypoxemia. STUDY DESIGN: In a multicenter study involving 3 German obstetric centers, a total of 400 deliveries were monitored by fetal pulse oximetry (Nellcor-Puritan-Bennett Model N-400 Oxygen Saturation Monitor and FS-14 Sensor; Nellcor, Inc, Pleasanton, Calif). The durations of low (60%) fetal arterial oxygen saturations during the measurement were compared between neonates with a pH <7.15 versus >/=7.15 and a base excess <-12 mmol/L versus >-12 mmol/L in the umbilical artery post partum and in neonates with an Apgar score <7 versus >/=7 by Mann-Whitney U test. In 121 of the pulse oximetry measurements the durations of low, medium, and high fetal arterial oxygen saturations were measured from one fetal scalp blood sampling to the next and correlated with the change of scalp blood pH between samplings. Multiple regression analysis was performed to estimate the expected change of pH between 2 fetal scalp blood samplings, and receiver operating characteristic analysis was done to define a minimum duration of low fetal arterial oxygen saturation values to exclude or predict a significant decline of pH. RESULTS: Neonates with a 1-minute Apgar score <7 differed from those with 1-minute Apgar score >/=7 significantly in the duration of low fetal arterial oxygen saturation but not in the durations of medium and high fetal arterial oxygen saturations. The duration of low fetal arterial oxygen saturation had been significantly longer in children with pH <7.15 or base excess <-12 mmol/L in the umbilical artery compared with those with a pH >/=7.15 or base excess >/=-12 mmol/L. The duration of high fetal arterial oxygen saturation was significantly shorter for children with a pH <7.15 or base excess <12 mmol/L than for those with a pH >/=7.15 or base excess >/=12 mmol/L. There was no difference in the groups with respect to the duration of medium fetal arterial oxygen saturation values. The duration of low fetal arterial oxygen saturation proved to be the best predictor of a decline of scalp pH between 2 fetal scalp blood samples. The pH declined significantly with a longer duration of low fetal arterial oxygen saturation (0.02 per 10 minutes). No decrease of pH by more than 0.05 was observed unless fetal arterial oxygen saturation had remained at /=10 minutes. CONCLUSION: An arterial oxygen saturation of 30% was confirmed as the critical boundary for fetal compromise during labor. The development of acidosis seems to be predictable by the duration of hypoxemia, as indicated by fetal arterial oxygen saturation

Double dissociation of function within the hippocampus: a comparison of dorsal, ventral, and complete hippocampal cytotoxic lesions.

Rats with complete cytotoxic hippocampal lesions exhibited spatial memory impairments in both the water maze and elevated T maze. They were hyperactive in photocell cages; swam faster in the water maze; and were less efficient on a nonspatial, differential reinforcement of low rates (DRL) task. Performance on both spatial tasks was also impaired by selective dorsal but not ventral lesions; swim speed was increased by ventral but not dorsal lesions. Both partial lesions caused a comparable reduction in DRL efficiency, although these effects were smaller than those of complete lesions. Neither partial lesion induced hyperactivity when rats were tested in photocell cages, although both complete and ventral lesion groups showed increased activity after footshock in other studies (Richmond et al., 1999). These results demonstrate possible functional dissociations along the septotemporal axis of the hippocampus.

Sheina orri (Myodocopa: Cypridinidae), an ostracod parasitic on the gills of the epaulette shark, Hemiscyllium ocellatum (Elasmobranchii: Hemiscyllidae).

The cypridinid ostracod, Sheina orri, was found on the gills of healthy epaulette sharks, Hemiscyllium ocellatum, collected from the Great Barrier Reef, Australia. Seventeen of the 28 fish examined had ostracods attached to their gills. Detailed investigation of the gills and ostracods using light microscopy and scanning electron microscopy revealed that ostracods anchor themselves to the gill tissues using their mandibular and maxillular claws. These claws appear to be adapted for this purpose and the process of attachment causes some damage to the host tissues. The observation that ostracods were often located in distinct pockets, formed by local distortion of shark respiratory lamellae, strongly suggests that they had been attached to the gills for considerable time.