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Background The evolutionary processes that shape patterns of species richness in marine ecosystems are complex and may differ between organismal groups. There has been considerable interest in understanding the evolutionary processes that led to marine species richness being concentrated in specific geographical locations. In this study we focus on the evolutionary history of a group of small-to-medium sized sharks known as carpet sharks. While a few carpet shark species are widespread, the majority of carpet shark species richness is contained within a biodiversity hotspot at the boundary of the Indian and Pacific oceans. We address the significance of this biodiversity hotspot in carpet shark evolution and speciation by leveraging a rich fossil record and molecular phylogenetics to examine the prehistoric distribution of carpet sharks. Results We find that carpet sharks species richness was greatest in shallow seas connected to the Atlantic Ocean during the Late Cretaceous, but that there was a subsequent loss of biodiversity in Atlantic waters. Fossil evidence from sites in close geographic proximity to the current center of carpet shark diversity are generally restricted to younger geologic strata. Conclusions From this data we conclude that (1) center of carpet shark biodiversity has shifted during the last 100 million years, (2) carpet sharks have repeatedly dispersed to nascent habitat (including to their current center of diversity), and (3) the current center of carpet shark biodiversity conserves lineages that have been extirpated from this prehistoric range and is a source of new carpet shark species. Our findings provide insights into the roles of marine biodiversity hotspots for higher-tropic level predators and the methods applied here can be used for additional studies of shark evolution.

Karyotyping, traditionally performed using cytogenetic techniques, is indispensable for validating genome assemblies whose sequence lengths can be scaled up to chromosome sizes using modern methods. Karyotype reports of chondrichthyans are scarce because of the difficulty in cell culture. Here, we focused on carpet shark species and the culture conditions for fibroblasts and lymphocytes. The utility of the cultured cells enabled the high-fidelity characterization of their karyotypes, namely 2n = 102 for the whale shark (Rhincodon typus) and zebra shark (Stegostoma fasciatum), and 2n = 106 for the brownbanded bamboo shark (Chiloscyllium punctatum) and whitespotted bamboo shark (C. plagiosum). We identified heteromorphic XX/XY sex chromosomes for the two latter species and demonstrated the first-ever fluorescence in situ hybridization of shark chromosomes prepared from cultured cells. Our protocols are applicable to diverse chondrichthyan species and will deepen the understanding of early vertebrate evolution at the molecular level., Shark cells are difficult to culture in the laboratory as the high osmolality physiological conditions do not reflect those of commonly cultured mammalian or teleost fish cell lines. Uno et al. present a new method of culturing shark fibroblasts and lymphocytes, and characterized the karyotypes of four carpet shark species, enabling whole genome analysis of these species.

Karyotyping is indispensable for validating genome assemblies whose sequence lengths can be scaled up to chromosome sizes using modern methods and is traditionally performed using cytogenetic techniques. Karyotype reports of chondrichthyans are scarce, mainly because of their unique osmoregulatory mechanism, which hinders cell culture. Here, we focused on carpet shark species and the culture conditions for fibroblasts and lymphocytes. Using this method, we performed high-fidelity characterization of their karyotypes, namely 2n = 102 for the whale shark (Rhincodon typus) and zebra shark (Stegostoma fasciatum), and 2n = 106 for the brownbanded bamboo shark (Chiloscyllium punctatum) and whitespotted bamboo shark (C. plagiosum). We identified heteromorphic XX/XY sex chromosomes for the two latter species and demonstrated the first-ever fluorescence in situ hybridization of shark chromosomes prepared from cultured cells. Our technical solution is applicable to diverse chondrichthyan species and will deepen the understanding of early vertebrate evolution at the molecular level.

Examination of seven specimens of an undescribed species of bamboo shark (Orectolobiformes: Hemiscylliidae), currently referred to as Chiloscyllium sp. 1, from Sri Lanka yielded three new species of 'tetraphyllidean' cestodes, one each in the genera Carpobothrium Shipley et Hornell, 1906, Spiniloculus Southwell, 1925, and Yorkeria Southwell, 1927. Carpobothrium marjorieae Caira, Otto et Jensen sp. n. differs from its three valid congeners in total length, number of testes, and size of the apical sucker of its anterior bothridial flap. Like Carpobothrium eleanorae Koontz et Caira, 2016 it possesses spherical eggs with highly elongate bipolar filaments, but its eggs are conspicuously larger than those of the latter species. Spiniloculus akshayi Caira, Otto et Jensen sp. n. can be distinguished from its five congeners in total length and number of proglottids. Yorkeria sachiniae Caira, Otto et Jensen sp. n. differs from its 16 congeners in number of proglottids and testes, total length, terminal proglottid, cirrus sac, and cephalic peduncle length, hook size, genital pore position, and bothridial shape. The eggs of Y. sachiniae Caira, Otto et Jensen sp. n. are spindle-shaped and bear a single long, polar filament. All three species were found to exhibit microthrix patterns on their scolices like those of the majority of their congeners that have been examined to date with scanning electron microscopy. Microtriches on the distal surfaces of the bothridial pouches of a species of Carpobothrium were characterised for the first time; C. marjorieae Caira, Otto et Jensen sp. n. was found to possess a modified form of gladiate spinithrix on this surface. Among the seven members of the genus, this is the first species of Chiloscyllium Muller et Henle that has been found to host representatives of all three of these 'tetraphyllidean' genera. However, unlike most of its congeners, a representative of the non-hooked genus Scyphophyllidium Woodland, 1927 was not found infecting Chiloscyllium sp. 1.

[Figure][1] A grey carpet shark (top) and an epaulette shark (bottom). Photo credit: Jules Devaux. When humans suffer a stroke or heart attack, the supply of oxygen to the brain is interrupted, with disastrous consequences. ‘This is because mitochondria, famously referred to as the “

The carpet shark Cephaloscyllium isabellum is abundant and widespread around the New Zealand coastal shelf from the tidal zone to about 500 m. Their overall depth distribution deepens with decreasing latitude, indicating a distribution moderated by water temperature. Cephaloscyllium isabellum is a generalist benthic carnivore and scavenger that uses both ambush and active searching hunting strategies. It has a broad diet (mainly fish, crustaceans, cephalopods, gastropods, and holothurians) and an apparent ability to manipulate prey to remove the shells of gastropods and pagurids before ingesting these animals. Length at maturity is about 60 cm for male and 76 cm for female C. isabellum. They may have an egg-laying peak around summer but probably reproduce throughout the year; the egg laying process is described. Females attain a larger size (c. 92 cm) than males (c. 80 cm). Growth and mortality rates are unknown, and constitute the main knowledge gap for C. isabellum and for Cephaloscyllium species in general.

Exposure to anoxia leads to rapid ATP depletion, alters metabolic pathways and exacerbates succinate accumulation. Upon re-oxygenation, the preferential oxidation of accumulated succinate most often impairs mitochondrial function. Few species can survive prolonged periods of hypoxia and anoxia at tropical temperatures and those that do may rely on mitochondria plasticity in response to disruptions to oxygen availability. Two carpet sharks, the epaulette shark (Hemiscyllium ocellatum; ES) and the grey carpet shark (Chiloscyllium punctatum; GCS) display different adaptive responses to prolonged anoxia: while the ES enters energy conserving metabolic depression, the GCS temporarily elevates its haematocrit prolonging oxygen delivery. High-resolution respirometry was used to investigate mitochondrial function in the cerebellum, a highly metabolically active organ that is oxygen sensitive and vulnerable to injury after anoxia/re-oxygenation (AR). Succinate was titrated into cerebellar preparations in vitro, with or without pre-exposure to AR, then the activity of mitochondrial complexes was examined. Like most vertebrates, GCS mitochondria significantly increased succinate oxidation rates, with impaired complex I function post-AR. In contrast, ES mitochondria inhibited succinate oxidation rates and both complex I and II capacities were conserved, resulting in preservation of oxidative phosphorylation capacity post-AR. Divergent mitochondrial plasticity elicited by elevated succinate post A/R parallels the inherently divergent physiological adaptations of these animals to prolonged anoxia, namely the absence (GCS) and presence of metabolic depression (ES). Since anoxia tolerance in these species also occurs at temperatures close to that of humans, examining their mitochondrial responses to AR could provide insights for novel interventions in clinical settings.

The complete mitochondrial genome of the grey bamboo shark, Chiloscyllium griseum (Orectolobiformes: Hemiscylliidae) was obtained, with a total of 16 755 base pairs (bp). The mitogenome encoded 13 protein-coding genes, two ribosomal RNAs, 22 transfer RNAs and a noncoding control region, a gene arrangement identical to the most common type found in vertebrates. All tRNA sequences were folded into typical cloverleaf secondary structure with the exception of tRNA-Ser2, in which the DHU arm stem was replaced with 12 unpaired nucleotides and formed a simple loop. In the control region, the putative termination-associated sequences (TAS) with hairpin-loop structure and the conserved sequence blocks (CSB) I-III were identified and considered to be associated with the replication and transcription of mtDNA. No repeat motifs were found in the mitogenome of C. griseum. On the basis of the mitogenomic data available in the Class Chondrichthyes from both Genbank and this study, and the phylogenetic analyses using maximum likelihood (ML) and Bayesian inference (BI) methods, the results strongly support the basal division between batoids and sharks and the monophyly of the Superorder Galeomorphii, and confirm the phylogenetic position of C. griseum in the Subclass Elasmobranchii. This study suggests that the mitogenomic data are more robust for reconstructing phylogenetic relationships than individual genes in the Elasmobranchii.

Re-examination of previously reported micro-vertebrate specimens collected from the upper portion of the Dakota Formation (Middle Cenomanian) in Russell County, Kansas, has identified two teeth of the extinct Carpet Shark, Orectoloboides sp. While the teeth of Orectoloboides have been described from Africa and Europe, the only previous report of the genus in North America is from the Cenomanian of Saskatchewan in Canada. The presence of Orectoloboides in Kansas represents a more than 2000 km southward extension of the range of this group into the central portion of the Western Interior Seaway.

We compared the hematological responses of wild and captive populations of two closely related sharks to a standardized anoxic challenge and during a 12 hr recovery period in normoxia: the epaulette shark (Hemiscyllium ocellatum, Bonnaterre, 1788) and the grey carpet shark (Chiloscyllium punctatum, Muller and Henle, 1838). Compared to normoxic controls, a significant increase in hematocrit (captive 22.3%; wild 35.9%) coupled with a decline in mean corpuscular hemoglobin concentration occurred in epaulette sharks indicating erythrocyte swelling in response to anoxia. However, the grey carpet shark had a significantly increased hematocrit (captive 27.2%; wild 29.3%), erythrocyte count (captive 37.6%; wild 46.3%) and hemoglobin concentration (captive 31.9%; wild 31.5%), suggesting a release of erythrocytes into the circulation and/or hemoconcentration in response to anoxia. Plasma glucose concentrations were maintained in both wild and captive epaulette sharks and in wild grey carpet sharks during anoxia but increased significantly after 2 hr of re-oxygenation (epaulette: captive 55.8%; wild 50.1%; grey carpet shark: wild 70.3%) and remained elevated for 12 hr. Captive grey carpet sharks had an immediate increase in plasma glucose concentrations after anoxia (96.4%), which was sustained for 12 hr of re-oxygenation. Lactate concentrations significantly increased in captive and wild animals of both species after anoxia, reaching a peak at 2 hr of re-oxygenation. Both species showed significant, yet divergent, hematological changes in response to anoxia and re-oxygenation, which may not only prolong their survival and assist in recovery but also reflect their respective ecophysiological adaptations to the extreme environments that they inhabit. J. Exp. Zool. 311A:422–438, 2009. © 2009 Wiley-Liss, Inc.

Seasonal aggregations commonly occur in the marine environment where typically wide-ranging organisms come together to exploit temporary resources or find conspecifics for mating events. The zebra shark Stegostoma fasciatum is a demersal carpet shark that aggregates over the austral summer months in the coastal waters of southeast Queensland, Australia. This study employed photo-identification and mark-recapture methods over a 3 yr period (2003 to 2006) to investigate the population size and structure of this aggregation. In total 327 individual zebra sharks were identified from 570 photographs. Numbered dart-tags on 15 zebra sharks were used to confirm that pigmentation patterns were unique and persistent in wild zebra sharks for up to 810 d. Pollock’s robust design resulted in an annual population estimate of 458 individuals (95% CI = 298–618). The mean number of zebra sharks observed on a single day was 8 (± 8 SE) and the maximum number of zebra sharks seen on a single day was 34. In total, 27% of the sharks were sighted in more than one summer aggregation period and males had greater re-capture probabilities than females. The aggregation consisted exclusively of large (>1800 mm total length) adults with an overall female sex bias of 3.8:1 though sex-ratios varied temporally. Predictable visitation of large, presumably mature individuals to the site raises conservation concerns if aggregations of similar size and structure occur in regions where zebra sharks are fished.

[Extract] Temperature can impact embryonic development in nearly all vertebrates. This may be particularly evident in ectotherms—especially embryos developing in eggs outside of the female. Until hatching, the eggs can be restricted to their local thermal environment. Temperature can affect development rates, and skeletal abnormalities and abnormal colouration and patterns (e.g., snakes; Vinegar 1974) can result from elevated temperatures. While perhaps not directly life-threatening, temperature-mediated changes in colouration and pattern development may impact biological fitness because patterns are often required for camouflage, to attract mates, and/or for deterring predators.

We examined gill morphology and ventilatory function in the carpet shark family Parascylliidae using 14 preserved specimens of Parascyllium ferrugineum, P. variolatum, P. collare and Cirrhoscyllium japonicum, and two live specimens of P. ferrugineum and P. variolatum. Morphological examinations revealed eight morphological characteristics related to the fifth gill, based on comparisons with other elasmobranchs, viz. large fifth gill slit without gill filaments, anatomical modifications in the fourth ceratobranchial cartilage and coraco-branchialis muscle, and the hypaxialis muscle associated with the fifth gill arch. Ventilation examinations using dyed seawater and prey items showed different water flows through the gill slits for respiration and prey-capture actions. For respiration, water sucked into the mouth was expelled equally through the first to fourth gill slits via a "double-pump" action, there being no involvement of the fifth gill slit. In prey-capture, however, water sucked into the mouth was discharged only via the widely opened fifth gill slit. This form of water flow is similar to that in other benthic suction-feeding sharks (e.g., Chiloscyllium plagiosum), except for the active water discharge by wide expansion and contraction of the fifth parabranchial cavity. The latter is dependent upon the morphological modifications of the fourth and fifth gill arches, derived phylogenetically as a mechanistic suction specialization in Parascylliidae.

The nominal Eucrossorhinus microcuspidatus and Cretorectolobus olsoni from the Campanian Judith River Formation of Montana, correspond to teeth from juvenile and adult individuals of one single species of orectoloboid shark. The type series of the latter is heterogeneous and includes, besides the holotype, a tooth belonging to a different orectoloboid of a new generic type named Cederstroemia. This genus is unique among Orectolobiformes by comprising sharks with dental adaptations towards a strict cutting dentition. Two new species of Cederstroemia are described: C. triangulata from the Judith River Formation of Montana and C. nilsi from the early Campanian of the Kristianstad Basin, Skåne, Sweden. An emended diagnosis is provided for Cretorectolobus, clarifying the dental characteristics of its members.

The epaulette shark (Hemiscyllium ocellatum) and the grey carpet shark (Chiloscyllium punctatum) are commonly found in periodically hypoxic environments. The ecophysiological time available for these animals to safely exploit these niches during different seasonal temperatures was examined. The time to loss of righting reflex (T LRR) was examined in response to an open ended anoxic challenge at three seasonal temperatures (23, 25 and 27°C). Ventilation rates were measured in an open ended anoxic challenge at 23°C and during 1.5 h of anoxia followed by 2 h of re-oxygenation at 23 and 25°C. The mean T LRR of epaulette and grey carpet sharks was inversely proportional to temperature. The T LRR was similar between species at 23°C; however, grey carpet sharks had significantly reduced T LRR at higher temperatures. During the standardised anoxic challenge, epaulette sharks entered into ventilatory depression significantly earlier at 25°C. During re-oxygenation, epaulette sharks exposed to anoxia at 23°C had no significant increase in ventilation rates. However, after anoxic challenge and re-oxygenation at 25°C, epaulette sharks showed a significant increase in ventilation rates during re-oxygenation. Grey carpet sharks displayed no evidence of ventilatory depression during anoxia. However, during re-oxygenation, grey carpet sharks had significantly elevated ventilation rates above pre-experimental levels and control animals. These data demonstrate that the anoxia tolerance times of both species were temperature dependent, with a significant reduction in the T LRR occurring at higher temperatures. Epaulette sharks had a significantly greater T LRR at higher temperatures than grey carpet sharks, which did not enter into a ventilatory depression.

In the tails of resting Cephaloscyllium, left and right radial muscles below the spine contract synchronously in a caudal-rostral sequence. This pulls the skin above the origin of the muscles inward and compresses the underlying caudal sinus. This series of connected valved chambers is part of the secondary venous system; it communicates, near the base of the tail, with the caudal vein. Evidence from surface mechanograms and electromyograms of the muscles and of records of blood pressure in the cutaneous veins, caudal sinus, and caudal vein of intact fish and of isolated perfused tails is presented. It indicates that blood is aspirated into the caudal sinus past inlet valves in veins from the trunk and tail and is expelled into the caudal vein. It is concluded that the caudal heart of Cephaloscyllium, powered by unmodified, metamerically repeated radial muscles is of a less specialized grade of construction than that of hagfish and teleosts.

This study utilizes retrograde axonal transport of cobaltous-lysine, and conventional silver and Golgi staining techniques to study the abducens motor nucleus innervating the external rectus muscle of the carpet shark. The nucleus consists of 300-400 motoneurons located immediately ventrolateral to the medial longitudinal fasciculus (MLF), distributed over about 1.25 mm in a rostrocaudal direction at the level of exit of the VI nerve. The axons of the motoneurons form seven or eight discrete ventrally directed fascicles which, having exited from the brainstem, group together to form the abducens (VI) nerve. The motoneurons are on average about 16 micron in diameter, are bipolar, and their dendrites have a transverse orientation. Typically one set of dendrites penetrates the MLF and the other set extends ventrally into the reticular formation.

1. Carpet Shark haemoglobin shows co-operative equilibrium CO binding at pH 7.0 (n = 1.8). 2. Flash photolysis shows that the T form of the protein is stabilised at low pH and in the presence of Inositol Hexaphosphate (I.H.P.). 3. At high pH the R form of the protein is stabilised and exhibits kinetic heterogeneity, assigned to different chain reactivities. 4. Stopped flow and flash photolysis under various conditions allows estimates to be made of the association and dissociation rates of CO binding for the two forms of the protein. 5. The functionality of the protein, with regard to CO binding, can be described by a modified two state model.

The central neuronal pathways and connections required to process mechanosensory lateral line information probably arose, concomitantly with the peripheral receptors, in the first vertebrates. The cartilaginous fishes (Chondrichthyes) share the mechanosensory component of the lateral line system with all other anamniotes, although there are vast differences in the central nervous systems of those vertebrates that possess lateral line sensory capabilities. Even among the elasmobranchs, there are different levels of brain organization and profound variations within each level (Northcutt 1978). It is important to identify mechanosensory lateral line centers and their fiber relations with other octavolateralis centers among those elasmobranchs that possess different patterns of brain organization, in order to establish homologies and to better understand the evolution of mechanoreception in chondrichthyans. However, our knowledge of the central mechanosensory lateral line pathways is limited to only a few elasmobranchs, e.g., the batoids (thornback guitarfish Platyrhinoidis triseriata, clearnose skate Raja eglanteria, and the little skate Raja erinacea), the squalomorph sharks (spiny dogfish Squalus acanthias), and the galeomorph sharks (carpet shark Cephaloscyllium isabella). Central lateral line pathways have been studied mostly in skates that are considered advanced elasmobranchs in terms of brain organization (Northcutt 1978), but some information is available in the spiny dogfish and carpet shark which, in contrast to skates, are more primitive in terms of brain organization. The purpose of this chapter is to delineate some of the central mechanosensory lateral line centers and their connections at each level of the neuraxis and to establish homologies in these few species of cartilaginous fishes.