Shark Pharyngeal Muscles and Early Vertebrate Evolution

This study provides a new classification of shark pharyngeal muscles, leading to a re-interpretation of the evolutionary history of these muscles. It begins with Edgeworth's developmental classification but also incorporates phylogenetic and functional information. The two basic groups of pharyngeal muscles are (1) branchial (including deep interbranchials and superficial constrictors) and (2) spinal (including epaxial and hypobranchial muscles). Most of the branchial muscles are also present anteriorly in the mandibular and hyoid segments, indicating that these were once typical branchial segments. The extrabranchial cartilages and gill septa are used as landmarks for dividing the interbranchials from the superficial constrictors. These landmarks reveal that the most important expiratory muscles in the anterior pharynx (most parts of "constrictor hyoideus", interhyoideus, levator palatoquadrati, spiracularis, and intermandibularis) are actually enlarged interbranchial muscles that have taken on a surface location. "Shrinking" these enlarged interbranchials and returning them to a deep location produces a simple, metameric reconstruction of the pre-gnathostome pharynx that fits what is known about thelodonts and other fossil agnatha related to gnathostomes. Finally, the pharyngeal muscles of bony fishes are considered, and concluded to have evolved from a sharklike condition through an acanthodian-like intermediate. © 1997 The Royal Swedish Academy of Sciences. Published by Elsevier Science Ltd.     

Shark attacks in the rivers of southern Iran

Synopsis The literature on shark attacks in freshwaters of southern Iran is reviewed and 11 attacks with 3 fatalities recorded from local informants for the period 1953 to 1985. The species of shark responsible is probably the bull shark, Carcharhinus leucas, whose presence in the Tigris River is confirmed by preserved specimens in the British Museum (Natural History). The Iranian records represent a significant proportion (28%) of the documented cases world-wide for unprovoked, freshwater attacks.     

The Effects of Hypoxia on Three Sympatric Shark Species: Physiological and Behavioral Responses

Behavioral and physiological responses to hypoxia were examined in three sympatric species of sharks: bonnethead shark Sphyrna tiburo, blacknose shark, Carcharhinus acronotus, and Florida smoothhound shark, Mustelus norrisi, using closed system respirometry. Sharks were exposed to normoxic and three levels of hypoxic conditions. Under normoxic conditions (5.5–6.4mg l^–1), shark routine swimming speed averaged 25.5 and 31.0cm s^–1 for obligate ram-ventilating S. tiburo and C. acronotus respectively, and 25.0cm s^–1 for buccal-ventilating M. norrisi. Routine oxygen consumption averaged about 234.6 mg O₂kg^–1h^–1 for S. tiburo, 437.2mg O₂kg^–1h^–1 for C. acronotus, and 161.4mg O₂ kg^–1 h^–1 for M. norrisi. For ram-ventilating sharks, mouth gape averaged 1.0cm whereas M. norrisi gillbeats averaged 56.0 beats min^–1. Swimming speeds, mouth gape, and oxygen consumption rate of S. tiburo and C. acronotus increased to a maximum of 37–39cm s^–1, 2.5–3.0cm and 496 and 599mg O₂ kg^–1 h^–1 under hypoxic conditions (2.5–3.4mg l^–1), respectively. M. norrisi decreased swimming speeds to 16cm s^–1 and oxygen consumption rate remained similar. Results support the hypothesis that obligate ram-ventilating sharks respond to hypoxia by increasing swimming speed and mouth gape while buccal-ventilating smoothhound sharks reduce activity.     

A Review of Elasmobranch Reproductive Behavior with a Case Study on the Nurse Shark, Ginglymostoma Cirratum

Elasmobranch reproductive behavior has been inferred from freshly caught specimens, laboratory examinations of reproductive structures and function, or determined from direct observations of captive or free swimming wild animals. Several general behaviors have been described including seasonal sexual segregation, courtship and copulation. Courtship behavior was inferred for many species from the presence of scars and tooth cuts on the female's body, and noted in more detail from underwater observations. Copulation has been directly observed in captive settings for several species of elasmobranchs in large aquaria, and in the wild for three species of urolophids and for Triaenodon obesus and Ginglymostoma cirratum. A detailed case history of nurse shark reproductive behavior is presented that may be used as a template for future work on shark reproductive behavior of other species. Our studies, using diver identifiable tags and in situ behavioral observations, provide unprecedented information on social structure and mating behavior in this species. Since 1993, 115 G. cirratum, 45 adults and 70 juveniles have been tagged in the Dry Tortugas, Florida. Observations show that adult males visit the study site every year with three males dominant. Individual adult females visit the study area to mate in alternate years. Polygyny and polyandry are common. Future research on reproductive behavior of elasmobranchs should address questions on male access to females, sexual selection and dominance hierarchies.     

Age and Growth of the Whiskery Shark, Furgaleus macki, from Southwestern Australia

Age and growth of the whiskery shark, Furgaleus macki, from southwestern Australia were examined using vertebral ageing and tag-recapture data. The readability of bands on the vertebral centra varied markedly between individuals. Four readers were used to make band counts, with the most experienced reader having the lowest index of average percent error and the highest level of agreement with final counts. Marginal increment analysis indicated that opaque bands form in January. With parturition occurring from August to October, size data suggests that the first band is probably formed 15–17 months after birth. The age at maturity was estimated to be 4.5 years for males, and 6.5 years for females. The oldest male was 10.5 years, and oldest female was 11.5 years. Von Bertalanffy growth parameters for males were L =121.5cm fork length, K=0.423 year^–1, t ₀=–0.472 years, were L =120.7cm fork length, K=0.369 year^–1, t ₀=–0.544 years for females, and were L =118.1cm fork length, K=0.420 year^–1, t ₀=–0.491 years for combined sexes. Data from a tag recapture study were analysed using a maximum likelihood method to verify the estimates of growth parameters from vertebral ageing. Von Bertalanffy growth parameters from the tag recapture study were L =128.2cm fork length, K=0.288 year^–1, t ₀=–0.654 years. The two methods of estimating growth parameters produced similar results, with rapid growth until approximately 5 years of age, after which there was little increase in length.     

Surfactants as Chemical Shark Repellents: Past, Present, and Future

The development of Shark Chaser by the U.S. Navy during World War II was the first serious effort to develop a chemical shark repellent. In the decade following the war reports of Shark Chaser ineffectiveness led the Office of Naval Research to search for a more efficacious shark repellent. After years without success, ONR eventually canceled the use of Shark Chaser and abandoned the search for a chemical shark repellent. In the early 1970s, interest in chemical shark repellents was renewed by the discovery of pardaxin, a natural shark repellent secreted by the Red Sea Moses sole, Pardachirus marmoratus. The surfactant-like nature of pardaxin led investigators to test the potential of various surfactants as repellents. Subsequent studies indicated that the shark repellent efficacy of the effective alkyl sulfate surfactants was due to their hydrophobic nature. Here we report tests conducted on juvenile swell sharks, Cephaloscyllium ventriosum, to determine if the noxious quality of alkyl sulfates is affected by surfactant hydrophobicity [carbon chain length and ethylene oxide (EO) groups] and counterions. Our results indicate that the aversive response of sharks to alkyl sulfate surfactants increases with carbon chain length from octyl to dodecyl, decreases with the addition of EO groups and is not affected by counterions. This study confirms that dodecyl sulfate is the most effective surfactant shark repellent, but it does not meet the Navy's potency requirement for a nondirectional surrounding-cloud type repellent of 100 parts per billion (0.1ugml^–1). Thus, dodecyl sulfate is only practical as a directional repellent such as in a squirt application. Future research should test the action of alkyl sulfates on cell membranes, the potential of other biotoxic agents, and semiochemicals in the search for an effective chemical shark repellent.     

The Biology of the Nurse Shark,Ginglymostoma cirratum,Off the Florida East Coast and the Bahama Islands

The nurse shark is an extremely abundant shallow water species in Florida and the Caribbean, yet its biology is poorly known. Moreover, there is a great deal of misinformation about it in the literature. The maximum size and weight attained by the nurse shark have often been exaggerated. None of the specimens measured in this study exceeded 265cm TL and 114.5kg, and none of the specimens actually measured by other researchers exceeded 280cm. Females reach maturity at a length of 223–231cm, or at 86% of their maximum size. Males reach maturity between 214 and 214.6cm in length or at about 83% of their maximum size. Mating primarily occurs from mid-June to early July. The embryos are enclosed in sturdy egg capsules for the first 12–14 weeks of gestation. In a gravid female, the embryos are at different stages of development during the first four months of gestation. Embryos are lecithotrophic and there is no evidence of any supplemental mode of embryonic nourishment. Embryos measure 28–30.5cm at birth. The gestation period is estimated at about five to six months. Brood sizes are large, ranging from 21 to 50 young, with a median of 34 young. The reproductive cycle of the nurse shark consists of a five to six-month gestation period and a two-year ovarian cycle. Thus, the reproductive cycle is biennial and a female produces a brood every two years. The nurse shark is an opportunistic predator that consumes a wide range of small fishes, primarily grunts (Haemulidae).     

Electrical Stimulation of the Shark Brain

Early studies using various means to electrically activate portionsof the brain in immobilized elasmobranchs are critically reviewed.The results of brain stimulation experiments using modern techniquesare then considered in light of this background material. Preliminaryexperiments on anesthetized-immobilized sharpnose and bonnetheadsharks are given as an outline of the types of responses thatcan be studied using acute preparations of pelagic sharks. Theresponses evoked can be grouped into the following categories:arousal and swimming; body, fin and eye movements suggestiveof circling and loss of equilibrium; possible components ofagonistic and reproductive behavior; head, mouth and respiratorymovements and possible color changes. Chronic studies on free-swimmingsharpnose sharks are discussed with respect to potential areasfor future study. Responses suggestive of portions of agonisticand feeding behavior were evoked but no attempt was made tolocalize the areas stimulated. The behavioral effects of electricalstimulation of 85 electrodes in free-swimming juvenile nursesharks are presented in some detail. The anatomical distributionof sites from which the following responses have been elicitedare plotted on representative sections of the brain: arousalescape;coughing; head shaking; barbel movement; eye retraction; bitingor mouthing food; snapping up gravel; slow dragging of the barbelsacross the substrate; circling; rolling about the long axis;continuous swimming and interrupted swimming. Several possiblefunctional-anatomical systems are suggested to account for someof these results.     

Aspects of Reproductive Biology of the Scalloped Hammerhead Shark, Sphyrna Lewini, off Northeastern Brazil

Ninety four scalloped hammerhead sharks, Sphyrna lewini (53 females and 41 males) ranging in size from 121 to 321cm total length (TL), were collected from surface gillnetters operating off northeastern Brazil and throughout the southwestern equatorial Atlantic Ocean between January and December 1996. A common regression for TL and eviscerated weight (EW) was calculated as, logEW = –11.786 + 2.889 logTL. Females and males were categorised into reproductive stages (4 and 2, respectively) according to morphological changes in their gonads. Size at sexual maturity for females was estimated to be 240cm, while males appeared to mature at between 180 and 200cm. Gravid females had between 2 and 21 embryos or pups, varying in TL from 3 to 38cm. There was no relationship between maternal length and size of litter. Copulation and parturition appear to occur outside the sampled area and possibly closer to the coast. With the exception of slightly lower uterine and ovarian fecundities, the results support the few existing data on the reproductive cycle of S. lewini in other areas.     

Basking Shark (Lamniformes: Cetorhinidae) from the Lower Oligocene of the Caucasus

Based on an almost complete impression from the base of the Pshekha Horizon (layer 2, Planorbella Beds, zone NP 21) of the North Caucasus (lower early Oligocene), we describe the basking shark Caucasochasma zherikhini gen. et sp. nov., differing from other members of family Cetorhinidae by higher number of vertebrae, weakly developed lower lobe of caudal fin, and details of the structure of gill rakers. Based on the structure of the body, it is not excluded that the described taxon was associated with the bottom to a greater degree than other species of the family (Keasius parvus and C. maximus), for which the structure of the body is known. The accumulations of plankton at the bottom could be a possible food resource for Caucasochasma.     

Shark lymphocytes: Primitive antigen reactive cells

Shark lymphocytes have been characterized by the presence or absence of surface immunoglobulin (SIg). Thus, SIg⁺ cells are B lymphocytes, and SIg⁻ cells are presumed to include the shark T cell subset, as well as other minor subsets of lymphocytes. Few functional studies have been performed to characterize the nature of either lymphocyte population. To date, there is no information concerning the shark T cell receptor. The majority of adult mammalian T cell receptors are composed of α and β chain heterodimers while a minority use γ and δ chains. The discussion presented here explores the evidence that the majority of shark lymphocytes are analogous to mammalian T and B lymphocytes that appear during early fetal development. The hypotheses considered suggest that shark T cells are similar to γδ T cell receptor (TCR)-bearing mammalian T lymphocytes, and that shark B cells are the primitive equivalent of neonatal and newborn primary B cells.     

Archival Tagging Of School Shark, Galeorhinus Galeus, in Australia: Initial Results

Archival tags were used to study the movement and depth behaviour of school sharks, Galeorhinus galeus, in southern australia. Thirty fish were tagged in late 1997, and to date there have been nine recaptures (30% recapture rate). Periods at liberty varied from 8 days to 18 months. The sharks spent about 80% of their time on the continental shelf, and appeared to swim close to the bottom during the day. At night they often ascended for periods of up to several hours, except at times around the full moon. When in deep water, the sharks typically descended at dawn to depths of up to 600m, before ascending at dusk. It was not possible to use the light data from the tags to estimate position when the sharks were in deep water, because they were often at depths beyond the sensitivity of the tag. In shallower water, longitude was estimated from the light data but latitude was estimated from the maximum daily depth, assuming the fish were on the bottom. The timing of the dives in deepwater appeared sufficiently regular to offer the prospect of using it to estimate longitude. We propose future research using archival tags on this species should address questions about female reproductive migrations, pelagic behaviour and vertical movements.     

DNA Barcoding of Shark Meats Identify Species Composition and CITES-Listed Species from the Markets in Taiwan

Background

An increasing awareness of the vulnerability of sharks to exploitation by shark finning has contributed to a growing concern about an unsustainable shark fishery. Taiwan’s fleet has the 4th largest shark catch in the world, accounting for almost 6% of the global figures. Revealing the diversity of sharks consumed by Taiwanese is important in designing conservation plans. However, fins make up less than 5% of the total body weight of a shark, and their bodies are sold as filets in the market, making it difficult or impossible to identify species using morphological traits.

Methods

In the present study, we adopted a DNA barcoding technique using a 391-bp fragment of the mitochondrial cytochrome oxidase I (COI) gene to examine the diversity of shark filets and fins collected from markets and restaurants island-wide in Taiwan.

Results

Amongst the 548 tissue samples collected and sequenced, 20 major clusters were apparent by phylogenetic analyses, each of them containing individuals belonging to the same species (most with more than 95% bootstrap values), corresponding to 20 species of sharks. Additionally, Alopias pelagicus, Carcharhinus falciformis, Isurus oxyrinchus, and Prionace glauca consisted of 80% of the samples we collected, indicating that these species might be heavily consumed in Taiwan. Approximately 5% of the tissue samples used in this study were identified as species listed in CITES Appendix II, including two species of Sphyrna, C. longimanus and Carcharodon carcharias.

Conclusion

DNA barcoding provides an alternative method for understanding shark species composition when species-specific data is unavailable. Considering the global population decline, stock assessments of Appendix II species and highly consumed species are needed to accomplish the ultimate goal of shark conservation.     

Preliminary Study of Vertebral Growth Rings in the Whale Shark, Rhincodon typus, from the East Coast of South Africa

Growth rings (GR) in vertebral centra of 15 whale sharks, Rhincodon typus, four female (418–750cm precaudal length), 10 male (422–770cm), and one of unknown sex (688cm), were examined using x-radiography. GR counts were made from scanned images and count precision was determined using the average percentage error index (4.19%) and the index of precision D (3.31%). In females, counts ranged from 19 GR (418cm) to 27 GR (750cm); in males from 20 GR (670cm) to 31 GR (770cm). Three mature males had 20 GR (670cm), 24 GR (744cm) and 27 GR (755cm). A female with 22 GR (445cm) was adolescent. There was a linear relationship between centrum dorsal diameter and body length, and back-calculated body lengths at number of GR are presented. A linear relationship between body length and number of GR prevented the calculation of von Bertalanffy parameters from either observed or back-calculated values.     

Shark detection and classification with machine learning

Suitable shark conservation depends on well-informed population assessments. Direct methods such as scientific surveys and fisheries monitoring are adequate for defining population statuses, but species-specific indices of abundance and distribution coming from these sources are rare for most shark species. We can rapidly fill these information gaps by boosting media-based remote monitoring efforts with machine learning and automation.We created a database of 53,345 shark images covering 219 species of sharks, and packaged object-detection and image classification models into a Shark Detector bundle. The Shark Detector recognizes and classifies sharks from videos and images using transfer learning and convolutional neural networks (CNNs). We applied these models to common data-generation approaches of sharks: collecting occurrence records from photographs taken by the public or citizen scientists, processing baited remote camera footage and online videos, and data-mining Instagram. We examined the accuracy of each model and tested genus and species prediction correctness as a result of training data quantity.The Shark Detector can classify 47 species pertaining to 26 genera. It sorted heterogeneous datasets of images sourced from Instagram with 91% accuracy and classified species with 70% accuracy. It located sharks in baited remote footage and YouTube videos with 89% accuracy, and classified located subjects to the species level with 69% accuracy. All data-generation methods were processed without manual interaction.As media-based remote monitoring appears to dominate methods for observing sharks in nature, we developed an open-source Shark Detector to facilitate common identification applications. Prediction accuracy of the software pipeline increases as more images are added to the training dataset. We provide public access to the software on our GitHub page.     

On the Field Study of Shark Behavior

The difficulties of observational study of active, wide-rangingsharks in the relatively concealing natural underwater environmentare discussed for both baited and non-baited situations. Thepossible technique of habituating sharks to the diver's presenceis considered in view of the logistical and safety problemsimposed by the physical limitations of unaided, unprotectedhuman divers. It is concluded that previous sharkobservationmethods are inadequate, and that supplementary study techniquesmust be developed and emphasized. These include remote monitoringby ultrasonic telemetry, telemetry-aided direct observation,and the use of miniature submersibles designed specificallyfor shark observation.     

Shark skin: a function in feeding

Dermal denticles are unique tooth-like structures embedded in the skin of sharks and rays that protect them from predators and ectoparasites, reduce mechanical abrasion and possibly minimize swimming-induced drag. Here, we show that juvenile lesser spotted dogfish (Scyliorhinus canicula) also use this body armour to anchor food items near their tail so that bite-sized pieces can be torn away by rapid jaw and head movements. This scale-rasp behaviour is novel among fishes and suggests a new role for skin in the feeding ecology of sharks. Scale rasping may be important ecologically because it could function to increase the dietary breadth and growth potential of juveniles.     

Shark fishing in fin soup

Shark DNA extracted from products used in trade(e.g. soup and dried fins) was amplified usingPCR. A strategy is described that permits theidentification of amplified material to species(for mtDNA Cytb and NADH2 loci) and thespecies-specific amplification of basking sharkDNA from mixed templates (for the Cytb locus). These methods will be useful for forensicapplications to govern trade in these heavilyexploited species.