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Concept: Shark

333

Biofluorescence has recently been found to be widespread in marine fishes, including sharks. Catsharks, such as the Swell Shark (Cephaloscyllium ventriosum) from the eastern Pacific and the Chain Catshark (Scyliorhinus retifer) from the western Atlantic, are known to exhibit bright green fluorescence. We examined the spectral sensitivity and visual characteristics of these reclusive sharks, while also considering the fluorescent properties of their skin. Spectral absorbance of the photoreceptor cells in these sharks revealed the presence of a single visual pigment in each species. Cephaloscyllium ventriosum exhibited a maximum absorbance of 484 ± 3 nm and an absorbance range at half maximum (λ1/2max) of 440-540 nm, whereas for S. retifer maximum absorbance was 488 ± 3 nm with the same absorbance range. Using the photoreceptor properties derived here, a “shark eye” camera was designed and developed that yielded contrast information on areas where fluorescence is anatomically distributed on the shark, as seen from other sharks' eyes of these two species. Phylogenetic investigations indicate that biofluorescence has evolved at least three times in cartilaginous fishes. The repeated evolution of biofluorescence in elasmobranchs, coupled with a visual adaptation to detect it; and evidence that biofluorescence creates greater luminosity contrast with the surrounding background, highlights the potential importance of biofluorescence in elasmobranch behavior and biology.

Concepts: Photoreceptor cell, Great white shark, Shark, Chondrichthyes, Scyliorhinidae, Catshark, Cephaloscyllium, Swellshark

206

Parthenogenesis is a natural form of asexual reproduction in which embryos develop in the absence of fertilisation. Most commonly found in plants and invertebrate organisms, an increasing number of vertebrate species have recently been reported employing this reproductive strategy. Here we use DNA genotyping to report the first demonstration of an intra-individual switch from sexual to parthenogenetic reproduction in a shark species, the zebra shark Stegostoma fasciatum. A co-housed, sexually produced daughter zebra shark also commenced parthenogenetic reproduction at the onset of maturity without any prior mating. The demonstration of parthenogenesis in these two conspecific individuals with different sexual histories provides further support that elasmobranch fishes may flexibly adapt their reproductive strategy to environmental circumstances.

Concepts: Reproduction, Organism, Life, Vertebrate, Asexual reproduction, Sexual reproduction, Shark, Parthenogenesis

188

Comparative genomic and/or transcriptomic analyses involving elasmobranchs remain limited, with genome level comparisons of the elasmobranch immune system to that of higher vertebrates, non-existent. This paper reports a comparative RNA-seq analysis of heart tissue from seven species, including four elasmobranchs and three teleosts, focusing on immunity, but concomitantly seeking to identify genetic similarities shared by the two lamnid sharks and the single billfish in our study, which could be linked to convergent evolution of regional endothermy.

Concepts: Immune system, Gene, Genetics, Organism, Innate immune system, Humoral immunity, Immunity, Shark

174

Modern selachians and their supposed sister group (hybodont sharks) have a long and successful evolutionary history. Yet, although selachian remains are considered relatively common in the fossil record in comparison with other marine vertebrates, little is known about the quality of their fossil record. Similarly, only a few works based on specific time intervals have attempted to identify major events that marked the evolutionary history of this group.

Concepts: Evolution, Phylogenetics, Batoidea, Fossil, Paleontology, Dinosaur, Shark, Chondrichthyes

167

While shark-based tourism is a rapidly growing global industry, there is ongoing controversy about the effects of provisioning on the target species. This study investigated the effect of feeding on whale sharks (Rhincodon typus) at a provisioning site in Oslob, Cebu, in terms of arrival time, avoidance and feeding behaviour using photo-identification and focal follows. Additionally, compliance to the code of conduct in place was monitored to assess tourism pressure on the whale sharks. Newly identified sharks gradually arrived earlier to the provisioning site after their initial sighting, indicating that the animals learn to associate the site with food rewards. Whale sharks with a long resighting history showed anticipatory behaviour and were recorded at the site on average 5 min after the arrival of feeder boats. Results from a generalised linear mixed model indicated that animals with a longer resighting history were less likely to show avoidance behaviour to touches or boat contact. Similarly, sequential data on feeding behaviour was modelled using a generalised estimating equations approach, which suggested that experienced whale sharks were more likely to display vertical feeding behaviour. It was proposed that the continuous source of food provides a strong incentive for the modification of behaviours, i.e., learning, through conditioning. Whale sharks are large opportunistic filter feeders in a mainly oligotrophic environment, where the ability to use novel food sources by modifying their behaviour could be of great advantage. Non-compliance to the code of conduct in terms of minimum distance to the shark (2 m) increased from 79% in 2012 to 97% in 2014, suggesting a high tourism pressure on the whale sharks in Oslob. The long-term effects of the observed behavioural modifications along with the high tourism pressure remain unknown. However, management plans are traditionally based on the precautionary principle, which aims to take preventive actions even if data on cause and effect are still inconclusive. Hence, an improved enforcement of the code of conduct coupled with a reduction in the conditioning of the whale sharks through provisioning were proposed to minimise the impacts on whale sharks in Oslob.

Concepts: Psychology, Maldives, Shark, Megafauna, Basking shark, Whale shark, Filter feeder, Megamouth shark

166

The hunting strategies of pelagic thresher sharks (Alopias pelagicus) were investigated at Pescador Island in the Philippines. It has long been suspected that thresher sharks hunt with their scythe-like tails but the kinematics associated with the behaviour in the wild are poorly understood. From 61 observations recorded by handheld underwater video camera between June and October 2010, 25 thresher shark shunting events were analysed. Thresher sharks employed tail-slaps to debilitate sardines at all times of day. Hunting events comprised preparation, strike, wind-down recovery and prey item collection phases, which occurred sequentially. Preparation phases were significantly longer than the others, presumably to enable a shark to windup a tail-slap. Tail-slaps were initiated by an adduction of the pectoral fins, a manoeuvre that changed a thresher shark’s pitch promoting its posterior region to lift rapidly, and stall its approach. Tail-slaps occurred with such force that they may have caused dissolved gas to diffuse out of the water column forming bubbles. Thresher sharks were able to consume more than one sardine at a time, suggesting that tail-slapping is an effective foraging strategy for hunting schooling prey. Pelagic thresher sharks appear to pursue sardines opportunistically by day and night, which may make them vulnerable to fisheries. Alopiids possess specialist pectoral and caudal fins that are likely to have evolved, at least in part, for tail-slapping. The evidence is now clear; thresher sharks really do hunt with their tails.

Concepts: Hunting, Commercial fish, Shark, Thresher shark, Common thresher, Pelagic thresher, Bigeye thresher, Alopiidae

159

Five striking and prey capture events of two goblin sharks were videotaped at sea for the first time, showing their extraordinary biting process. The goblin sharks swung their lower jaw downward and backward to attain a huge gape and then rapidly protruded the jaws forward a considerable distance. The jaws were projected at a maximum velocity of 3.1 m/s to 8.6-9.4% of the total length of the shark, which is by far the fastest and greatest jaw protrusion among sharks. While the jaws were being retracted, the mouth opened and closed again, which was considered a novel feeding event for sharks. Phylogenetic evidence suggested that their feeding behavior has evolved as an adaptation to food-poor deep-sea environments, possibly as a trade-off for the loss of strong swimming ability.

Concepts: Great white shark, Ocean, Jaw, Shark, Lamniformes, Goblin shark, Mitsukurinidae, Scapanorhynchus

152

Data on the occurrence of whale sharks, Rhincodon typus, in the Arabian Gulf and Gulf of Oman were collected by dedicated boat surveys and via a public-sightings scheme during the period from 2011 to 2014. A total of 422 individual whale sharks were photo-identified from the Arabian Gulf and the northern Gulf of Oman during that period. The majority of sharks (81%, n = 341) were encountered at the Al Shaheen area of Qatar, 90 km off the coast, with the Musandam region of Oman a secondary area of interest. At Al Shaheen, there were significantly more male sharks (n = 171) than females (n = 78; X2 = 17.52, P < 0.05). Mean estimated total length (TL) for sharks was 6.90 m ± 1.24 (median = 7 m; n = 296). Males (7.25 m ± 1.34; median = 8 m, n = 171) were larger than females (6.44 m ±1.09; median = 7 m, n = 78; Mann-Whitney U test, p < 0.01). Of the male sharks assessed for maturity 63% were mature (n = 81), with 50% attaining maturity by 7.29 m and 100% by 9.00 m. Two female sharks of >9 m individuals were visually assessed as pregnant. Connectivity among sharks sighted in Qatari, Omani and UAE waters was confirmed by individual spot pattern matches. A total of 13 identified sharks were re-sighted at locations other than that at which they were first sighted, including movements into and out of the Arabian Gulf through the Strait of Hormuz. Maximum likelihood techniques were used to model an estimated combined population for the Arabian Gulf and Gulf of Oman of 2837 sharks ± 1243.91 S.E. (95% C.I. 1720-6295). The Al Shaheen aggregation is thus the first site described as being dominated by mature males while the free-swimming pregnant females are the first reported from the Indian Ocean.

Concepts: Arabian Peninsula, United Arab Emirates, Qatar, Persian Gulf, Shark, Gulf of Oman, Oman, Strait of Hormuz

143

Conservation and management efforts for white sharks (Carcharodon carcharias) remain hampered by a lack of basic demographic information including age and growth rates. Sharks are typically aged by counting growth bands sequentially deposited in their vertebrae, but the assumption of annual deposition of these band pairs requires testing. We compared radiocarbon (Δ(14)C) values in vertebrae from four female and four male white sharks from the northwestern Atlantic Ocean (NWA) with reference chronologies documenting the marine uptake of (14)C produced by atmospheric testing of thermonuclear devices to generate the first radiocarbon age estimates for adult white sharks. Age estimates were up to 40 years old for the largest female (fork length [FL]: 526 cm) and 73 years old for the largest male (FL: 493 cm). Our results dramatically extend the maximum age and longevity of white sharks compared to earlier studies, hint at possible sexual dimorphism in growth rates, and raise concerns that white shark populations are considerably more sensitive to human-induced mortality than previously thought.

Concepts: Demography, Fish, Gerontology, Great white shark, Shark, Lamniformes, Megalodon, Lamnidae

136

The Greenland shark (Somniosus microcephalus), an iconic species of the Arctic Seas, grows slowly and reaches >500 centimeters (cm) in total length, suggesting a life span well beyond those of other vertebrates. Radiocarbon dating of eye lens nuclei from 28 female Greenland sharks (81 to 502 cm in total length) revealed a life span of at least 272 years. Only the smallest sharks (220 cm or less) showed signs of the radiocarbon bomb pulse, a time marker of the early 1960s. The age ranges of prebomb sharks (reported as midpoint and extent of the 95.4% probability range) revealed the age at sexual maturity to be at least 156 ± 22 years, and the largest animal (502 cm) to be 392 ± 120 years old. Our results show that the Greenland shark is the longest-lived vertebrate known, and they raise concerns about species conservation.

Concepts: Species, Arctic Ocean, Arctic, Shark, Greenland shark, Somniosus, Pacific sleeper shark, Hákarl