From rivers to ocean basins: The role of ocean barriers and philopatry in the genetic structuring of a cosmopolitan coastal predator.

The Bull Shark (Carcharhinus leucas) faces varying levels of exploitation around the world due to its coastal distribution. Information regarding population connectivity is crucial to evaluate its conservation status and local fishing impacts. In this study, we sampled 922 putative Bull Sharks from 19 locations in the first global assessment of population structure of this cosmopolitan species. Using a recently developed DNA-capture approach (DArTcap), samples were genotyped for 3400 nuclear markers. Additionally, full mitochondrial genomes of 384 Indo-Pacific samples were sequenced. Reproductive isolation was found between and across ocean basins (eastern Pacific, western Atlantic, eastern Atlantic, Indo-West Pacific) with distinct island populations in Japan and Fiji. Bull Sharks appear to maintain gene flow using shallow coastal waters as dispersal corridors, whereas large oceanic distances and historical land-bridges act as barriers. Females tend to return to the same area for reproduction, making them more susceptible to local threats and an important focus for management actions. Given these behaviors, the exploitation of Bull Sharks from insular populations, such as Japan and Fiji, may instigate local decline that cannot readily be replenished by immigration, which can in turn affect ecosystem dynamics and functions. These data also supported the development of a genetic panel to ascertain the population of origin, which will be useful in monitoring the trade of fisheries products and assessing population-level impacts of this harvest.

Inter- and intra-specific trophic interactions of coastal delphinids off the eastern coast of South Africa inferred from stable isotope analysis.

Dietary tracers, such as bulk stable carbon (δ13C) and nitrogen (δ15N) isotopes, can be used to investigate the trophic interactions of marine predators, which is useful to assess their ecological roles within communities. These tracers have also been used to elucidate population structure and substructure, which is critical for the better identification of management units for these species affected by a range of threats, particularly bycatch in fishing gears. Off eastern South Africa, large populations of Indo-Pacific bottlenose (Tursiops aduncus) and common dolphins (Delphinus delphis) co-occur and are thought to follow the pulses of shoaling sardines (Sardinops sagax) heading north-east in the austral winter. Here we used δ13C and δ15N to investigate the trophic interactions and define ecological units of these two species along a ≈800 km stretch of the east coast of South Africa, from Algoa Bay to the coast of KwaZulu-Natal. Common and bottlenose dolphin dietary niche overlapped by 39.7% overall in our study area, with the highest overlap occurring off the Wild Coast (40.7% at Hluleka). Both stable isotopes were significantly enriched in bottlenose dolphins sampled in the western part of our study area (i.e., Algoa Bay and Amathole) compared to eastern animals (i.e., from Hluleka, Pondoland, and KZN). In areas where genetic information is not available or is insufficient, food web tracers (such as stable isotopes) can be used to group individuals based on trophic ecology, which can provide ecological units for management of populations. The distinct isotope signatures found here for bottlenose dolphins can, therefore, be used as management units for conservation efforts in the future.

The extinct shark Otodus megalodon was a transoceanic superpredator: Inferences from 3D modeling.

Although shark teeth are abundant in the fossil record, their bodies are rarely preserved. Thus, our understanding of the anatomy of the extinct Otodus megalodon remains rudimentary. We used an exceptionally well-preserved fossil to create the first three-dimensional model of the body of this giant shark and used it to infer its movement and feeding ecology. We estimate that an adult O. megalodon could cruise at faster absolute speeds than any shark species today and fully consume prey the size of modern apex predators. A dietary preference for large prey potentially enabled O. megalodon to minimize competition and provided a constant source of energy to fuel prolonged migrations without further feeding. Together, our results suggest that O. megalodon played an important ecological role as a transoceanic superpredator. Hence, its extinction likely had large impacts on global nutrient transfer and trophic food webs.

Determining the appropriate pretreatment procedures and the utility of liver tissue for bulk stable isotope (δ13 C and δ15 N) studies in sharks.

Stable-isotope analysis (SIA) provides a valuable tool to address complex questions pertaining to elasmobranch ecology. Liver, a metabolically active, high turnover tissue (~166 days for 95% turnover), has the potential to reveal novel insights into recent feeding/movement behaviours of this diverse group. To date, limited work has used this tissue, but ecological application of SIA in liver requires consideration of tissue preparation techniques given the potential for high concentrations of urea and lipid that could bias δ13 C and δ15 N values (i.e., result in artificially lower δ13 C and δ15 N values). Here we investigated the effectiveness of (a) deionized water washing (WW) for urea removal from liver tissue and (b) chloroform-methanol for extraction of lipids from this lipid rich tissue. We then (a) established C:N thresholds for deriving ecologically relevant liver isotopic values given complications of removing all lipid and (b) undertook a preliminary comparison of δ13 C values between tissue pairs (muscle and liver) to test if observed isotopic differences correlated with known movement behaviour. Tests were conducted on four large shark species: the dusky (DUS, Carcharhinus obscurus), sand tiger (RAG, Carcharias taurus), scalloped hammerhead (SCA, Sphyrna lewini) and white shark (GRE, Carcharodon carcharias). There was no significant difference in δ15 N values between lipid-extracted (LE) liver and lipid-extracted/water washed (WW) treatments, however, WW resulted in significant increases in %N, δ13 C and %C. Following lipid extraction (repeated three times), some samples were still biased by lipids. Our species-specific "C:N thresholds" provide a method to derive ecologically viable isotope data given the complexities of this lipid rich tissue (C:N thresholds of 4.0, 3.6, 4.7 and 3.9 for DUS, RAG, SCA and GRE liverLEWW tissue, respectively). The preliminary comparison of C:N threshold corrected liver and muscle δ13 C values corresponded with movement/habitat behaviours for each shark; minor differences in δ13 C values were observed for known regional movements of DUS and RAG (δ13 CDiffs = 0.24 ± 0.99‰ and 0.57 ± 0.38‰, respectively), while SCA and GRE showed greater differences (1.24 ± 0.63‰ and 1.08 ± 0.71‰, respectively) correlated to large-scale movements between temperate/tropical and pelagic/coastal environments. These data provide an approach for the successful application of liver δ13 C and δ15 N values to examine elasmobranch ecology.

Are you really what you eat? Stomach content analysis and stable isotope ratios do not uniformly estimate dietary niche characteristics in three marine predators.

Calculation of dietary niche characteristics using stable isotopes has become a popular approach to understand the functional role of taxa across food webs. An underlying assumption of this approach is that stable isotopes accurately reflect the dietary breadth of a species over a temporal duration defined by tissue-specific isotopic turnover rates. In theory, dietary niche estimates derived from fast turnover rate tissues (e.g., blood plasma and liver) may augment stomach content-derived estimates more agreeably than slower turnover rate tissues (e.g., muscle or fin). We tested this hypothesis by comparing commonly used dietary niche estimates derived from stomach contents (nicheSCA: Levins', Shannon-Wiener's, and Smith's), with those estimated using stable isotopes [nicheSIA: standard ellipse area (SEA), convex hull total area (TA), theta (θ), and ellipse eccentricity (E)] of liver and muscle tissue. Model species were three large-bodied sharks: white (Carcharodon carcharias), dusky (Carcharhinus obscurus), and scalloped hammerhead (Sphyrna lewini). Within-technique comparisons for nicheSCA and nicheSIA metrics (i.e., SEA vs. TA) were often correlated; however, we did not observe any statistically significant correlations between nicheSCA and liver/muscle tissue nicheSIA (i.e., Levins' vs. SEA). We conclude that nicheSCA and nicheSIA do not provide comparable estimates of dietary niche, at least for the three predator species examined. This fundamental discrepancy highlights technique-specific limitations to estimating organismal dietary niche and identifies a need for the use of clearly defined niche metrics, i.e., the standardized use and reporting of the term isotopic niche as proposed by Newsome et al. (Front Ecol Environ 5:429-436, 2007). Finally, further investigation into the factors underpinning nicheSIA is required to better contextualize this popular ecological metric when compared to nicheSCA.

Population structure, connectivity, and demographic history of an apex marine predator, the bull shark Carcharhinus leucas.

Knowledge of population structure, connectivity, and effective population size remains limited for many marine apex predators, including the bull shark Carcharhinus leucas. This large-bodied coastal shark is distributed worldwide in warm temperate and tropical waters, and uses estuaries and rivers as nurseries. As an apex predator, the bull shark likely plays a vital ecological role within marine food webs, but is at risk due to inshore habitat degradation and various fishing pressures. We investigated the bull shark's global population structure and demographic history by analyzing the genetic diversity of 370 individuals from 11 different locations using 25 microsatellite loci and three mitochondrial genes (CR, nd4, and cytb). Both types of markers revealed clustering between sharks from the Western Atlantic and those from the Western Pacific and the Western Indian Ocean, with no contemporary gene flow. Microsatellite data suggested low differentiation between the Western Indian Ocean and the Western Pacific, but substantial differentiation was found using mitochondrial DNA. Integrating information from both types of markers and using Bayesian computation with a random forest procedure (ABC-RF), this discordance was found to be due to a complete lack of contemporary gene flow. High genetic connectivity was found both within the Western Indian Ocean and within the Western Pacific. In conclusion, these results suggest important structuring of bull shark populations globally with important gene flow occurring along coastlines, highlighting the need for management and conservation plans on regional scales rather than oceanic basin scale.

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.

Weak population structure of the Spot-tail shark Carcharhinus sorrah and the Blacktip shark C. limbatus along the coasts of the Arabian Peninsula, Pakistan, and South Africa.

The increase in demand for shark meat and fins has placed shark populations worldwide under high fishing pressure. In the Arabian region, the spot-tail shark Carcharhinus sorrah and the Blacktip shark Carcharhinus limbatus are among the most exploited species. In this study, we investigated the population genetic structure of C. sorrah (n = 327) along the coasts of the Arabian Peninsula and of C. limbatus (n = 525) along the Arabian coasts, Pakistan, and KwaZulu-Natal, South Africa, using microsatellite markers (15 and 11 loci, respectively). Our findings support weak population structure in both species. Carcharhinus sorrah exhibited a fine structure, subdividing the area into three groups. The first group comprises all samples from Bahrain, the second from the UAE and Yemen, and the third from Oman. Similarly, C. limbatus exhibited population subdivision into three groups. The first group, comprising samples from Bahrain and Kuwait, was highly differentiated from the second and third groups, comprising samples from Oman, Pakistan, the UAE, and Yemen; and South Africa and the Saudi Arabian Red Sea, respectively. Population divisions were supported by pairwise F ST values and discriminant analysis of principal components (DAPC), but not by STRUCTURE. We suggest that the mostly low but significant pairwise F ST values in our study are suggestive of fine population structure, which is possibly attributable to behavioral traits such as residency in C. sorrah and site fidelity and philopatry in C. limbatus. However, for all samples obtained from the northern parts of the Gulf (Bahrain and/or Kuwait) in both species, the higher but significant pairwise F ST values could possibly be a result of founder effects during the Tethys Sea closure. Based on DAPC and F ST results, we suggest each population to be treated as independent management unit, as conservation concerns emerge.

Satellite tagging highlights the importance of productive Mozambican coastal waters to the ecology and conservation of whale sharks.

The whale shark Rhincodon typus is an endangered, highly migratory species with a wide, albeit patchy, distribution through tropical oceans. Ten aerial survey flights along the southern Mozambican coast, conducted between 2004-2008, documented a relatively high density of whale sharks along a 200 km stretch of the Inhambane Province, with a pronounced hotspot adjacent to Praia do Tofo. To examine the residency and movement of whale sharks in coastal areas around Praia do Tofo, where they may be more susceptible to gill net entanglement, we tagged 15 juveniles with SPOT5 satellite tags and tracked them for 2-88 days (mean = 27 days) as they dispersed from this area. Sharks travelled between 10 and 2,737 km (mean = 738 km) at a mean horizontal speed of 28 ± 17.1 SD km day-1. While several individuals left shelf waters and travelled across international boundaries, most sharks stayed in Mozambican coastal waters over the tracking period. We tested for whale shark habitat preferences, using sea surface temperature, chlorophyll-a concentration and water depth as variables, by computing 100 random model tracks for each real shark based on their empirical movement characteristics. Whale sharks spent significantly more time in cooler, shallower water with higher chlorophyll-a concentrations than model sharks, suggesting that feeding in productive coastal waters is an important driver of their movements. To investigate what this coastal habitat choice means for their conservation in Mozambique, we mapped gill nets during two dedicated aerial surveys along the Inhambane coast and counted gill nets in 1,323 boat-based surveys near Praia do Tofo. Our results show that, while whale sharks are capable of long-distance oceanic movements, they can spend a disproportionate amount of time in specific areas, such as along the southern Mozambique coast. The increasing use of drifting gill nets in this coastal hotspot for whale sharks is likely to be a threat to regional populations of this iconic species.

Possible maternal offloading of metals in the plasma, uterine and capsule fluid of pregnant ragged-tooth sharks (Carcharias taurus) on the east coast of South Africa.

We studied the possible metal offloading onto the progeny of three pregnant female ragged-tooth sharks (Carcharias taurus) (C. taurus). The presences of five metals, i.e. aluminium (Al), arsenic (As), cadmium (Cd), lead (Pb) and selenium (Se) were validated by mass spectrometry in the maternal plasma as well as the intracapsular and uterine fluids (UF) in which embryos develop. Metals were ranked in a decreasing concentration as follows: Plasma: As > Al > Se > Pb > Cd; ICF: As > Se > Al > Cd > Pb and UF: As > Se > Al > Cd > Pb. As was present in the highest concentration in all three sharks. Al, Pb and Cd were found to be the highest within the plasma, while concentrations of Se were similar in all three fluids. These results indicate that C. taurus embryos are exposed to metals during early development, but the impact of this exposure remains unknown. To the best of our knowledge, this is the first investigation to confirm the presence of metals in the fluids that surround the developing C. taurus embryos, a species that is already listed as vulnerable.

Diet and trophic ecology of the tiger shark (Galeocerdo cuvier) from South African waters.

Knowledge of the diet and trophic ecology of apex predators is key for the implementation of effective ecosystem as well as species-based management initiatives. Using a combination of stomach content data and stable isotope analysis (δ15N and δ13C) the current study provides information on size-based and sex-specific variations in diet, trophic position (TP) and foraging habitat of tiger sharks (Galeocerdo cuvier) caught in the KwaZulu-Natal Sharks Board bather protection program. This study presents the longest time-series and most detailed analysis of stomach content data for G. cuvier worldwide. Prey identified from 628 non-empty stomachs revealed a size-based shift in diet. Reptiles, birds, mysticetes, and large shark species increased in dietary importance with G. cuvier size, concomitant with a decrease in smaller prey such as batoids and teleosts. Seasonal and decadal shifts in diet driven primarily by changes in the importance of elasmobranchs and mammal (cetacean) prey were recorded for medium sized (150-220 cm) G. cuvier. Both stomach content and stable isotope analysis indicated that G. cuvier is a generalist feeder at the population level. Size-based δ13C profiles indicated a movement to offshore foraging habitats by larger G. cuvier. Calculated TP varied by method ranging from 4.0 to 5.0 (TPSCA for stomach contents) and from 3.6 to 4.5 (TPscaled and TPadditive for δ15N). Large (> 220 cm) G. cuvier did not feed at discrete trophic levels, but rather throughout the food web. These data provide key information on the ecological role of G. cuvier to improve the accuracy of regional food web modelling. This will enable a better understanding of the ecological impacts related to changes in the abundance of this predator.

Profiling and Spatial Variation Analysis of Persistent Organic Pollutants in South African Delphinids.

The continuous disposal of persistent organic pollutants (POPs) in South Africa (SA) warrants concern about their detrimental effects on humans and wildlife. We surveyed six dolphin species (n = 90) incidentally captured in shark net installations or stranded off the SA east and south coast from 2005 to 2009 to study the POP exposure. Sousa plumbea, an inshore and estuarine species, was found to be the most contaminated by total POPs (21 100 ng g(-1) lw) of all the dolphins off SA, followed by Tursiops aduncus (19 800 ng g(-1) lw), Lagenodelphis hosei (13 600 ng g(-1) lw), and Delphinus capensis (5500 ng g(-1) lw), whereas POP levels in the offshore or pelagic delphinids were much lower. In all delphinids, dominant pollutants were dichlorodiphenyltrichloroethanes (DDTs), which represented more than 60% of the total concentration of total POPs, followed by polychlorinated biphenyls (PCBs, 30%). Concentrations of DDTs in S. plumbea and T. aduncus off SA were among the highest levels reported in delphinids globally. Approximately half of the adult T. aduncus had PCB concentrations above the effect threshold for impairment of immune functions. The concentrations of Mirex and Dieldrin in SA delphinids were higher than those found in species from other regions of the Southern Hemisphere.

Global versus local causes and health implications of high mercury concentrations in sharks from the east coast of South Africa.

Conservation concern regarding the overharvest of global shark populations for meat and fin consumption largely surrounds documented deleterious ecosystem effects, but may be further supported by improved knowledge of possibly high levels in their edible tissues (particularly meat) of the neurotoxin, methylmercury (CH3Hg). For many regions, however, little data exist on shark tissue Hg concentrations, and reasons for Hg variation within and among species or across regions are poorly understood. We quantified total Hg (THg) in 17 shark species (total n=283) from the east coast of South Africa, a top Hg emitter globally. Concentrations varied from means of around 0.1 mg kg(-1) dry weight (dw) THg in hardnose smoothhound (Mustelus mosis) and whale (Rhincodon typus) sharks to means of over 10 mg kg(-1) dw in shortfin mako (Isurus oxyrinchus), scalloped hammerhead (Sphyrna lewini), white (Carcharodon carcharias) and ragged-tooth (Carcharias taurus) sharks. These sharks had higher THg levels than conspecifics sampled from coastal waters of the North Atlantic and North, mid-, and South Pacific, and although sampling year and shark size may play a confounding role, this result suggests the potential importance of elevated local emissions. Values of THg showed strong, species-specific correlations with length, and nearly half the remaining variation was explained by trophic position (using nitrogen stable isotopes, δ(15)N), whereas measures of foraging habitat (using carbon stable isotopes, δ(13)C) were not significant. Mercury concentrations were above the regulatory guidelines for fish health effects and safe human consumption for 88% and 70% of species, respectively, suggesting on-going cause for concern for shark health, and human consumers of shark meat.

Comparative organochlorine accumulation in two ecologically similar shark species (Carcharodon carcharias and Carcharhinus obscurus) with divergent uptake based on different life history.

Trophic position and body mass are traits commonly used to predict organochlorine burdens. Sharks, however, have a variety of feeding and life history strategies and metabolize lipid uniquely. Because of this diversity, and the lipid-association of organochlorines, the dynamics of organochlorine accumulation in sharks may be predicted ineffectively by stable isotope-derived trophic position and body mass, as is typical for other taxa. The present study compared ontogenetic organochlorine profiles in the dusky shark (Carcharhinus obscurus) and white shark (Carcharodon carcharias), which differ in metabolic thermoregulation and trophic position throughout their ontogeny. Although greater organochlorine concentrations were observed in the larger bodied and higher trophic position white shark (e.g., p,p'-dichlorodiphenyldichloroethylene: 20.2 ± 2.7 ng/g vs 9.3 ± 2.2 ng/g in the dusky shark), slopes of growth-dilution corrected concentrations with age were equal to those of the dusky shark. Similar ontogenetic trophic position increases in both species, less frequent white shark seal predation than previously assumed, or inaccurate species-specific growth parameters are possible explanations. Inshore habitat use (indicated by δ(13)C values) and mass were important predictors in white and dusky sharks, respectively, of both overall compound profiles and select organochlorine concentrations. The present study clarified understanding of trophic position and body mass as reliable predictors of interspecific organochlorine accumulation in sharks, whereas regional endothermy and diet shifting were shown to have less impact on overall rates of accumulation.

Laser photogrammetry improves size and demographic estimates for whale sharks.

Whale sharks Rhincodon typus are globally threatened, but a lack of biological and demographic information hampers an accurate assessment of their vulnerability to further decline or capacity to recover. We used laser photogrammetry at two aggregation sites to obtain more accurate size estimates of free-swimming whale sharks compared to visual estimates, allowing improved estimates of biological parameters. Individual whale sharks ranged from 432-917 cm total length (TL) (mean ± SD = 673 ± 118.8 cm, N = 122) in southern Mozambique and from 420-990 cm TL (mean ± SD = 641 ± 133 cm, N = 46) in Tanzania. By combining measurements of stranded individuals with photogrammetry measurements of free-swimming sharks, we calculated length at 50% maturity for males in Mozambique at 916 cm TL. Repeat measurements of individual whale sharks measured over periods from 347-1,068 days yielded implausible growth rates, suggesting that the growth increment over this period was not large enough to be detected using laser photogrammetry, and that the method is best applied to estimating growth rates over longer (decadal) time periods. The sex ratio of both populations was biased towards males (74% in Mozambique, 89% in Tanzania), the majority of which were immature (98% in Mozambique, 94% in Tanzania). The population structure for these two aggregations was similar to most other documented whale shark aggregations around the world. Information on small (<400 cm) whale sharks, mature individuals, and females in this region is lacking, but necessary to inform conservation initiatives for this globally threatened species.

Comparison of total lipids and fatty acids from liver, heart and abdominal muscle of scalloped (Sphyrna lewini) and smooth (Sphyrna zygaena) hammerhead sharks.

Liver, heart and abdominal muscle samples from scalloped (Sphyrna lewini) and smooth (Sphyrna zygaena) hammerhead sharks were analysed to characterise their lipid and fatty acid profiles. Samples were compared both between and within species, but there were no significant differences in total lipids for either comparison, although much greater total amounts were found in the liver samples. Within the individual fatty acids, the only significant differences were greater amounts of 22:6n-3, total n-3 polyunsaturates and total polyunsaturates in smooth, when compared to scalloped, hammerhead liver. This may reflect the more wide spread distribution of this species into cooler waters. Within both species the liver levels of the same fatty acid fractions decreased from spring to summer, which may correlate with changes in fatty acid profile to adapt to any differences in amount or species of prey consumed, or other considerations, eg. buoyancy, however there was no data to clarify this.

Comparison of pinniped and cetacean prey tissue lipids with lipids of their elasmobranch predator.

BACKGROUND: The great white shark is known to include pinnipeds and cetaceans in its diet. Both groups of marine mammals deposit thick blubber layers around their bodies. Elasmobranchs do not produce adipose tissue, but rather store lipid in their livers, thus a great white predating on a marine mammal will deposit the lipids in its liver until required. MATERIALS AND METHODS: Samples from great white liver and muscle, Cape fur seal, Indian Ocean bottlenose dolphin and common dolphin liver, muscle and blubber were analyzed for their lipid and fatty acid profiles. RESULTS: The great white liver and marine mammal blubber samples showed a considerable degree of homogeneity, but there were significant differences when comparing between the muscle samples. Blubber from all three marine mammal species was calculated to provide greater than 95% of lipid intake for the great white shark from the tissues analyzed. CONCLUSION: Sampling of prey blubber may give a good indication of the lipids provided to the shark predator.

Effect of sample preparation techniques for carbon and nitrogen stable isotope analysis of hydroxyapatite structures in the form of elasmobranch vertebral centra.

RATIONALE: Bulk stable isotope analysis (SIA) provides an important tool for the study of animal ecology. Elasmobranch vertebral centra can be serially sampled to obtain an isotopic history of an individual over ontogeny. The measured total δ(13)C value, however, may be misinterpreted due to the inclusion of the (13)C-rich inorganic portion. Hydrochloric acid (HCl) is commonly used to remove the inorganic portion of hydroxyapatite structures before undertaking SIA, but more recently ethylenediaminetetraacetic acid (EDTA) has been recommended for elasmobranch vertebrae. These acid treatments may introduce uncertainty on measured δ(13)C and δ(15)N values above instrument precision and the effect of small sample size remains untested for elasmobranch vertebrae. METHODS: Using a non-dilution program on an isotope ratio mass spectrometer the minimum sample weight of vertebrae required to obtain accurate isotopic values was determined for three shark species: white (Carcharodon carcharias), tiger (Galeocerdo cuvier), and sand tiger (Carcharias taurus). To examine if acid treatment completely removes the inorganic component of the vertebrae or whether the technique introduces its own uncertainty on measured δ(13)C and δ(15)N values, vertebrae samples were analyzed untreated and following EDTA treatment. RESULTS: The minimum sample weight required for accurate stable isotope values and the percentage sample yield following EDTA treatment varied within and among species. After EDTA treatment, white shark vertebrae were all enriched in (13)C and depleted in (15) N, tiger shark vertebrae showed both enrichment and depletion of (13)C and (15)N, and sand tiger shark vertebrae were all depleted in (13)C and (15)N. CONCLUSIONS: EDTA treatment of elasmobranch vertebrae produces unpredictable effects (i.e. non-linear and non-correctable) among species in both the percentage sample yield and the measured δ(13)C and δ(15)N values. Prior to initiating a large-scale study, we strongly recommend investigating (i) the minimum weight of vertebral material required to obtain consistent isotopic values and (ii) the effects of EDTA treatment, specific to the study species and the isotope ratio mass spectrometer employed.

Rescaling the trophic structure of marine food webs.

Measures of trophic position (TP) are critical for understanding food web interactions and human-mediated ecosystem disturbance. Nitrogen stable isotopes (δ(15) N) provide a powerful tool to estimate TP but are limited by a pragmatic assumption that isotope discrimination is constant (change in δ(15) N between predator and prey, Δ(15) N = 3.4‰), resulting in an additive framework that omits known Δ(15) N variation. Through meta-analysis, we determine narrowing discrimination from an empirical linear relationship between experimental Δ(15) N and δ(15) N values of prey consumed. The resulting scaled Δ(15) N framework estimated reliable TPs of zooplanktivores to tertiary piscivores congruent with known feeding relationships that radically alters the conventional structure of marine food webs. Apex predator TP estimates were markedly higher than currently assumed by whole-ecosystem models, indicating perceived food webs have been truncated and species-interactions over simplified. The scaled Δ(15) N framework will greatly improve the accuracy of trophic estimates widely used in ecosystem-based management.

A comparison of the heart and muscle total lipid and fatty acid profiles of nine large shark species from the east coast of South Africa.

We have assessed the fatty acid profiles of the hearts and different muscle tissues from nine large shark species (Carcharhinus limbatus (blacktip), Carcharhinus obscurus (dusky), Carcharhinus brevipinna (spinner), Carcharhinus leucas (Zambezi/bull), Galeocerdo cuvier (tiger), Sphyrna lewini (scalloped hammerhead), Sphyrna zygaena (smooth hammerhead), Carcharodon carcharias (great white) and Carcharias taurus (raggedtooth/grey nurse/sand tiger)) found off the east coast of South Africa. While there was generally little variation between the species, all species showed profiles rich in both n6 and n3 polyunsaturated fatty acids compared to terrestrial commercial meats that have low n3. Thus, utilizing skeletal muscle tissues from sharks caught as part of the bycatch when fishing for teleosts would avoid unnecessary wastage of a potentially valuable resource, with all the possible health benefits of high quality protein combined with balanced polyunsaturates, although contamination with high levels of metabolic wastes, such as urea, may be a negative consideration.