Sheldon F. J. Dudley

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 (δ15N) provide a powerful tool to estimate TP but are limited by a pragmatic assumption that isotope discrimination is constant (change in δ15N between predator and prey, Δ15N = 3.4‰), resulting in an additive framework that omits known Δ15N variation. Through meta-analysis, we determine narrowing discrimination from an empirical linear relationship between experimental Δ15N and δ15N values of prey consumed. The resulting scaled Δ15N 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 Δ15N framework will greatly improve the accuracy of trophic estimates widely used in ecosystem-based management.

Population status of 14 shark species caught in the protective gillnets off KwaZulu–Natal beaches, South Africa, 1978–2003

Shark nets have been set off the beaches of KwaZulu–Natal, South Africa, since 1952 to reduce the risk of shark attack. The nets fish in fixed localities 400 m from shore and both directly affect local shark populations and act as fisheries-independent monitoring devices. Reliable catch information at the species level was available for the period 1978–2003. Trends in catch rate and size were used to assess the population status of 14 commonly caught shark species. In addition, a demographic modelling approach was used in conjunction with the catch information to assess the potential effect of the nets on populations. Catch rates of four species (Carcharhinus leucas, C. limbatus, Sphyrna lewini and S. mokarran) showed a significant decline, as did the mean or median length of three species (Carcharhinus amboinensis, C. limbatus and female Carcharodon carcharias). For three species that showed declining catch rates or length the potential effect of the shark nets was assessed to be low, suggesting that other sources of catch were responsible for the declining status. The potential effect of the shark nets was assessed to be high for two species (Carcharhinus obscurus and Carcharias taurus, neither of which showed declines in catch rate or length), because of very low intrinsic rates of population increase.

Maternal investment and size-specific reproductive output in carcharhinid sharks

1. Life-history theory predicts that organisms will provide an optimal level of parental investment for offspring survival balanced against the effects on their own survival and future reproductive potential. 2. Optimal resource allocation models also predict an increase in reproductive output with age as expected future reproductive effort decreases. To date, maternal investment in sharks has received limited attention. 3. We found that neonatal dusky sharks (Carcharhinus obscurus) are not independent from maternal resource allocation at the point of parturition but instead are provisioned with energy reserves in the form of an enlarged liver that constitutes approximately 20% of total body mass. 4. Analysis of long-term archived data sets showed that a large proportion of this enlarged liver is utilized during the first weeks or months of life suggesting that the reported weight loss of newborn sharks signifies a natural orientation process and is not necessarily related to prey abundance and/or indicative of high mortality rates. 5. Interrogation of near-term pup mass in two carcharhinids, the dusky and spinner shark (Carcharhinus brevipinna), further revealed an increase in reproductive output with maternal size, with evidence for a moderate decline in the largest mothers. 6. For the dusky shark, there was a trade-off between increasing litter size and near-term pup mass in support of optimal offspring size theory. 7. For both the dusky and spinner shark, there was a linear increase in near-term pup mass with month, which may indicate variable parturition strategies and/or that carcharhinids are able to adjust the length of the gestation period. 8. The identification of optimal size-specific reproductive output has direct implications for improving the reproductive potential of exploited shark populations and for structuring future management strategies.

Nursery grounds, movement patterns and growth rates of dusky sharks, Carcharhinus obscurus: a long-term tag and release study in South African waters

Knowledge of movement patterns of sharks in coastal waters is critical for the structuring of regional management plans. Through a long-term tag–recapture program, 9716 dusky sharks (Carcharhinus obscurus) were tagged and released along the east coast of South Africa. A total of 648 C. obscurus, principally small sharks ( 200 km that indicated a southerly migration between KZN and Eastern/Southern Cape (E/SC) between June and November. Seasonal northerly migrations were less well defined. The largest southerly and northerly movements were 1323 km and 1374 km, respectively. For sharks moving 1–100 km south from their tagging locality in KZN, an increase in displacement occurred between June and September identifying animals beginning their migration to the E/SC. With increasing displacement, there was also an increase in minimum swimming speed. Calculated growth rates of small sharks of 10.3–11.5 cm year–1 were in agreement with current literature values. Established tag–recapture programs provide an important tool in understanding the ecology of early life-stages of coastal shark species.

Some effects of shark nets in the Natal nearshore environment

SynopsisProtective gillnets (shark nets) have been successful in reducing the frequency of shark attacks on the coast of Natal, South Africa. This is achieved primarily through a reduction in numbers of large sharks. The nets also take a by catch of dolphins, sea turtles, batoids and teleosts. Concern has been expressed over the direct effect of mortalities on the various stocks and also the broader, indirect effects on the inshore system. Catch rates of most shark species declined initially but have shown no trend since the mid-1970s. A first estimate of the reduction in total shark numbers is provided, and factors such as stock identity and net avoidance are discussed. Turtle and teleost stocks do not appear to be threatened by net mortalities but marine mammalogists are investigating the sustainability of catches of two dolphin species. Certain batoids may have declined despite a high release rate, but more data are needed. A published contention that shark netting has resulted in a proliferation of small sharks through reduced predation is re-examined and considered to be exaggerated. Reduced predation on dolphins, as a result of shark netting, is estimated.

Reducing the environmental impact of shark-control programs: a case study from KwaZulu-Natal, South Africa

Large-scale shark-control programs at popular beaches in New South Wales and Queensland, Australia, and KwaZulu-Natal (KZN), South Africa, provide protection against shark attack. Although these programs have enhanced bathing safety, reducing the environmental impacts of decades of fishing for large sharks and the associated by-catch remains a challenge. Over the past three decades, there have been several interventions to reduce such impact in the KZN program.Thefirstwasthereleaseofalllivesharks,includingthosespeciesknowntoberesponsibleforfatalsharkattacks. Measures to reduce catches of sharks associated with the winter influx of shoals of sardines, Sardinops sagax, have been increasingly successful. In addition, extensive removal of nets has resulted in a major reduction in effort. Collectively, these initiatives reduced mortalities of sharks by 64%. Baited lines, termed drumlines, were introduced at 18 beaches, where they replaced some of the nets. The former had a far lower by-catch of rays, turtles and cetaceans and significantly lower catches of certain shark species. Replacement of some nets with drumlines is planned for the remaining beaches. Onlytwoattacks,bothnon-fatal,haveoccurredatprotectedbeachesinKZNoverthepastthreedecades,indicatingthatthe program has maintained its public safety mandate while it has succeeded in reducing its impact on the environment.

Large sharks and plastic debris in KwaZulu-Natal, South Africa

In total, 28 687 large sharks were caught between 1978 and 2000 in the nets that protect users of the popular swimming beaches of KwaZulu-Natal, South Africa, against shark attack. Over this 23-year period, 53 sharks (0.18% of the catch) were found with polypropylene strapping bands around the body. Less than 1% of the individuals from each of eight species were entangled in this manner. The dusky shark Carcharhinus obscurus was the most frequently entangled species, with 27 individuals (0.47% of the species catch). There was an increase in the incidence of entangled C. obscurus with time. Those examined in the laboratory were significantly underweight. Although entanglement may ultimately result in death, the low incidence recorded in this study is unlikely to affect the populations of sharks concerned. A total of 60 sharks (0.38% of those with recorded stomach contents) had ingested plastic debris. The most common items were packets or sheets. There was no increase in the ingestion of plastics with time. The highest frequency of occurrence was in the tiger shark Galeocerdo cuvier, with 38 individuals (7.5% of tiger sharks examined).

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.

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.

Revisiting acoustic deterrence devices: Long-term bycatch data from South Africa’s bather protection nets

The KwaZulu-Natal (KZN) Sharks Board in Durban, South Africa, has been managing bather protection nets off KZN’s most popular beaches since 1952 to mitigate shark-human interaction. The nets catch potentially dangerous sharks, but also take a bycatch of other marine megafauna, including the Indo-Pacific bottlenose dolphin Tursiops aduncus. A total of 1169 T. aduncus were caught between 1980 and 2015 (annual mean 32.5 ± 13.4). Most were young and immature. To mitigate cetacean bycatch, acoustic devices (pingers, whale alarms, and biosonar reflectors) were installed at selected beaches for some years between 1993 and 2015. On nets with pingers, most dolphins were caught close to active pingers, with 44 out of 72 animals (61%) caught within 50 m and 25 (35%) within 10 m. Bycatch varied inter-annually and between neighboring beaches both with and without pingers and there was no clear indication that pingers reduced bycatch. As this is an operational bather protection program, no controlled experiments on pin...