Fishing profile and commercial landings of shark and batoids in a global elasmobranchs conservation hotspot.

This paper describes the fishing profile and the temporal variation in the commercial landings of elasmobranchs in a global hotspot for their conservation and investigates the variables that influenced the landings. Census data on commercial catches were obtained between April 2008 and October 2010 from nine landing sites in Bragança (Pará, northern Brazil). Five vessel types, four fishing gears, and eight fishing techniques engaged with elasmobranch capture were identified. A total of 2,357 landings were recorded, with a total production of 354 t. The highest yields were recorded in 2009, with sharks being harvested mostly by small and medium-sized vessels, and batoids, by small vessels and canoes. Drifting nets and longlines played a prominent role in elasmobranch fisheries. The results show that the landings were influenced by days at sea, which is common in tropical fisheries. The elasmobranch data series is discontinuous as statistics are absent for most fishing sites albeit imperative for proper management, as well as relevant for decision-makers focusing on their conservation.

Can denticle morphology help identify southeastern Australian elasmobranchs?

Elasmobranchs are covered in scale-like structures called dermal denticles, comprising dentine and enameloid. These structures vary across the body of an individual and between species, and are frequently shed and preserved in marine sediments. With a good understanding of denticle morphology, current and historical elasmobranch diversity and abundance might be assessed from sediment samples. Here, replicate samples of denticles from the bodies of several known (deceased) shark species were collected and characterized for morphology before being assigned morphotypes. These data were used to expand the established literature describing denticles and to investigate intra- and interspecific variability, with the aim of increasing the viability of using sediment samples to assess elasmobranch diversity and abundance. Denticle morphology was influenced more by life-history traits than by species, where demersal species were largely characterized by generalized function and defense denticles, whereas pelagic and benthopelagic species were characterized by drag-reduction denticles. Almost all species possessed abrasion strength or defense denticles on the snout, precluding their utility for separating species. In a separate manipulative experiment, samples of denticles were collected from sediments in two aquaria with known elasmobranchs to determine their utility for reliably separating species. Visual examination of denticles, morphometric measurements, scaled photographs, and reference collections allowed for some precise identification, but not always to the species level. Ongoing work to develop denticle reference collections could help to identify past and present families and, in some cases, species.

The fossil record of extant elasmobranchs.

Sharks and their relatives (Elasmobranchii) are highly threatened with extinction due to various anthropogenic pressures. The abundant fossil record of fossil taxa has allowed the tracing of the evolutionary history of modern elasmobranchs to at least 250 MYA; nonetheless, exactly how far back the fossil record of living taxa goes has never been collectively surveyed. In this study, the authors assess the representation and extent of the fossil record of elasmobranchs currently living in our oceans by collecting their oldest records and quantifying first appearance dates at different taxonomic levels (i.e., orders, families, genera and species), ecological traits (e.g., body size, habitat and feeding mechanism) and extinction risks (i.e., threatened, not threatened and data deficient). The results of this study confirm the robust representation of higher taxonomic ranks, with all orders, most of the families and over half of the extant genera having a fossil record. Further, they reveal that 10% of the current global species diversity is represented in the geological past. Sharks are better represented and extend deeper in time than rays and skates. While the fossil record of extant genera (e.g., the six gill sharks, Hexanchus) goes as far back as c. 190 MYA, the fossil record of extant species (e.g., the sand shark, Carcharias taurus Rafinesque 1810) extends c. 66 MYA. Although no significant differences were found in the extent of the fossil record between ecological traits, it was found that the currently threatened species have a significantly older fossil record than the not threatened species. This study demonstrate that the fossil record of extant elasmobranchs extends deep into the geologic time, especially in the case of threatened sharks. As such, the elasmobranch geological history has great potential to advance the understanding of how species currently facing extinction have responded to different stressors in the past, thereby providing a deep-time perspective to conservation.

Evolutionary developmental genetics of teeth and odontodes in jawed vertebrates: a perspective from the study of elasmobranchs.

Most extant vertebrates display a high variety of tooth and tooth-like organs (odontodes) that vary in shape, position over the body and nature of composing tissues. The development of these structures is known to involve similar genetic cascades and teeth and odontodes are believed to share a common evolutionary history. Gene expression patterns have previously been compared between mammalian and teleost tooth development but we highlight how the comparative framework was not always properly defined to deal with different tooth types or tooth developmental stages. Larger-scale comparative analyses also included cartilaginous fishes: sharks display oral teeth and dermal scales for which the gene expression during development started to be investigated in the small-spotted catshark Scyliorhinus canicula during the past decade. We report several descriptive approaches to analyse the embryonic tooth and caudal scale gene expressions in S. canicula. We compare these expressions wih the ones reported in mouse molars and teleost oral and pharyngeal teeth and highlight contributions and biases that arise from these interspecific comparisons. We finally discuss the evolutionary processes that can explain the observed intra and interspecific similarities and divergences in the genetic cascades involved in tooth and odontode development in jawed vertebrates.

The early elasmobranch Phoebodus: phylogenetic relationships, ecomorphology and a new time-scale for shark evolution.

Anatomical knowledge of early chondrichthyans and estimates of their phylogeny are improving, but many taxa are still known only from microremains. The nearly cosmopolitan and regionally abundant Devonian genus Phoebodus has long been known solely from isolated teeth and fin spines. Here, we report the first skeletal remains of Phoebodus from the Famennian (Late Devonian) of the Maïder region of Morocco, revealing an anguilliform body, specialized braincase, hyoid arch, elongate jaws and rostrum, complementing its characteristic dentition and ctenacanth fin spines preceding both dorsal fins. Several of these features corroborate a likely close relationship with the Carboniferous species Thrinacodus gracia, and phylogenetic analysis places both taxa securely as members of the elasmobranch stem lineage. Identified as such, phoebodont teeth provide a plausible marker for range extension of the elasmobranchs into the Middle Devonian, thus providing a new minimum date for the origin of the chondrichthyan crown-group. Among pre-Carboniferous jawed vertebrates, the anguilliform body shape of Phoebodus is unprecedented, and its specialized anatomy is, in several respects, most easily compared with the modern frilled shark Chlamydoselachus. These results add greatly to the morphological, and by implication ecological, disparity of the earliest elasmobranchs.

Genetic and historical evidence of common sawsharks Pristiophorus cirratus in the waters of southern Queensland.

In 2011, a male pristiophorid was caught by a prawn trawler north east of Cape Moreton, Queensland, Australia. Molecular analyses confirmed the specimen to be the common sawshark Pristiophorus cirratus. Historical catch data indicate the occurrence of the species in the region but this is the first verified record of P. cirratus occurring in the waters of southern Queensland. Together, these records extend the recognised northern limit of P. cirratus by c. 500 km, which suggests that further investigation of its distribution is warranted.

Evolutionary Plasticity in Detoxification Gene Modules: The Preservation and Loss of the Pregnane X Receptor in Chondrichthyes Lineages.

To appraise how evolutionary processes, such as gene duplication and loss, influence an organism's xenobiotic sensitivity is a critical question in toxicology. Of particular importance are gene families involved in the mediation of detoxification responses, such as members of the nuclear receptor subfamily 1 group I (NR1I), the pregnane X receptor (PXR), and the constitutive androstane receptor (CAR). While documented in multiple vertebrate genomes, PXR and CAR display an intriguing gene distribution. PXR is absent in birds and reptiles, while CAR shows a tetrapod-specific occurrence. More elusive is the presence of PXR and CAR gene orthologs in early branching and ecologically-important Chondrichthyes (chimaeras, sharks and rays). Therefore, we investigated various genome projects and use them to provide the first identification and functional characterization of a Chondrichthyan PXR from the chimaera elephant shark (Callorhinchus milii, Holocephali). Additionally, we substantiate the targeted PXR gene loss in Elasmobranchii (sharks and rays). Compared to other vertebrate groups, the chimaera PXR ortholog displays a diverse expression pattern (skin and gills) and a unique activation profile by classical xenobiotic ligands. Our findings provide insights into the molecular landscape of detoxification mechanisms and suggest lineage-specific adaptations in response to xenobiotics in gnathostome evolution.

Mineral homeostasis and regulation of mineralization processes in the skeletons of sharks, rays and relatives (Elasmobranchii).

Sharks, rays and other elasmobranch fishes are characterized by a skeletal type that is unique among living vertebrates, comprised predominantly of an unmineralized cartilage, covered by a thin outer layer of sub-millimeter, mineralized tiles called tesserae. The mineralized portion of the skeleton appears to grow only by apposition, adding material at the edges of each tessera; maintenance of non-mineralized joints between tesserae is therefore vital, with precise control of mineral deposition and inhibition at the many thousands of growth fronts in the skeleton. Yet, we have only scattered evidence as to how the elasmobranchs mineralize and grow their skeletons. In this review, we take an "environment to skeleton" approach, drawing together research from a vast range of perspectives to track calcium and phosphate from the typical elasmobranch habitats into and through the body, to their deposition at tesseral growth fronts. In the process, we discuss the available evidence for skeletal resorption capability, mineral homeostasis hormones, and nucleation inhibition mechanisms. We also outline relevant theories in crystal nucleation and typical errors in measurements of serum calcium and phosphate in the study of vertebrate biology. We assemble research that suggests consensus in some concepts in elasmobranch skeletal development, but also highlight the very large gaps in our knowledge, particularly in regards to endocrine functional networks and biomineralization mechanisms. In this way, we lay out frameworks for future directions in the study of elasmobranch skeletal biology with stronger and more comparative links to research in other disciplines and into other taxa.

Biodiversity, Life History, and Conservation of Northeastern Pacific Chondrichthyans.

The sharks, batoids, and chimaeras, collectively the class Chondrichthyes, are one of the most successful groups of fishes, with over 1250 species globally. Recent taxonomic revisions have increased their diversity by about 20% over the past 17 years (2000-2016). The Northeast Pacific Ocean is one of the top 20 most diverse regions/countries on the globe with 77 chondrichthyan species, a number less than a quarter that of the most species-rich area (Australia) but that has increased by 10% since 2000 to include three new species (two skates and a chimaera). In this chapter we discuss the species richness of chondrichthyans occurring in the Northeast Pacific Ocean, characterize their life histories, briefly review several fisheries, and summarize the conservation status of those chondrichthyans occurring in the region. Detailed descriptions and evaluations of fisheries can be found in Chapter 7 of AMB Volume 78.

Early development of rostrum saw-teeth in a fossil ray tests classical theories of the evolution of vertebrate dentitions.

In classical theory, teeth of vertebrate dentitions evolved from co-option of external skin denticles into the oral cavity. This hypothesis predicts that ordered tooth arrangement and regulated replacement in the oral dentition were also derived from skin denticles. The fossil batoid ray Schizorhiza stromeri (Chondrichthyes; Cretaceous) provides a test of this theory. Schizorhiza preserves an extended cartilaginous rostrum with closely spaced, alternating saw-teeth, different from sawfish and sawsharks today. Multiple replacement teeth reveal unique new data from micro-CT scanning, showing how the 'cone-in-cone' series of ordered saw-teeth sets arrange themselves developmentally, to become enclosed by the roots of pre-existing saw-teeth. At the rostrum tip, newly developing saw-teeth are present, as mineralized crown tips within a vascular, cartilaginous furrow; these reorient via two 90° rotations then relocate laterally between previously formed roots. Saw-tooth replacement slows mid-rostrum where fewer saw-teeth are regenerated. These exceptional developmental data reveal regulated order for serial self-renewal, maintaining the saw edge with ever-increasing saw-tooth size. This mimics tooth replacement in chondrichthyans, but differs in the crown reorientation and their enclosure directly between roots of predecessor saw-teeth. Schizorhiza saw-tooth development is decoupled from the jaw teeth and their replacement, dependent on a dental lamina. This highly specialized rostral saw, derived from diversification of skin denticles, is distinct from the dentition and demonstrates the potential developmental plasticity of skin denticles.

Systematics and morphology of Potamotrygon orbignyi (Castelnau, 1855) and allied forms (Chondrichthyes: Myliobatiformes: Potamotrygonidae).

The Neotropical freshwater stingray Potamotrygom orbignyi (Castelnau, 1855), and other similar "reticulated" species occurring in northern South American basins, were submitted to a thorough taxonomic analysis based on an extensive external and internal morphological study. The identity of P. orbignyi and the taxonomic status of the related nominal species Potamotrygon dumerilii (Castelnau, 1855), Potamotrygon reticulata (Günther, 1880), and Potamotrygon humerosa Garman, 1913, are defined. Taxonomic and morphological analyses revealed that P. reticulata and P. dumerilii fall within the range of variation found in P. orbignyi and were consequently treated as junior synonyms, corroborating previous works. The extensive variation in coloration observed in P. orbignyi could not be divided into consistent morphotypes; P. orbignyi is therefore a widespread species in the upper, mid and lower Amazonas basin, the Orinoco drainage, and in rivers of Suriname and the Guianas. Additionally, P. humerosa and Potamotrygon marinae Deynat, 2007 were found to present characters that support their validity, and are redescribed based on newly collected material. Potamotrygon humerosa occurs predominantly in the mid and lower Amazonas River and in lower reaches of many of its affluents, whereas P. marinae is known only from French Guiana and Suriname. Characters that proved valuable as diagnostic indicators, either in combination or as derived features, are primarily from coloration, dermal denticles and spines (morphology, development and distribution), meristic features (e.g. numbers of tooth rows, vertebrae and mesopterygial radials), morphometric proportions (e.g. snout length, tail width at base and length), and size at sexual maturity.

Holocephalan embryos provide evidence for gill arch appendage reduction and opercular evolution in cartilaginous fishes.

Chondrichthyans possess endoskeletal appendages called branchial rays that extend laterally from their hyoid and gill-bearing (branchial) arches. Branchial ray outgrowth, like tetrapod limb outgrowth, is maintained by Sonic hedgehog (Shh) signaling. In limbs, distal endoskeletal elements fail to form in the absence of normal Shh signaling, whereas shortened duration of Shh expression correlates with distal endoskeletal reduction in naturally variable populations. Chondrichthyans also exhibit natural variation with respect to branchial ray distribution--elasmobranchs (sharks and batoids) possess a series of ray-supported septa on their hyoid and gill arches, whereas holocephalans (chimaeras) possess a single hyoid arch ray-supported operculum. Here we show that the elongate hyoid rays of the holocephalan Callorhinchus milii grow in association with sustained Shh expression within an opercular epithelial fold, whereas Shh is only transiently expressed in the gill arches. Coincident with this transient Shh expression, branchial ray outgrowth is initiated in C. milii but is not maintained, yielding previously unrecognized vestigial gill arch branchial rays. This is in contrast to the condition seen in sharks, where sustained Shh expression corresponds to the presence of fully formed branchial rays on the hyoid and gill arches. Considered in light of current hypotheses of chondrichthyan phylogeny, our data suggest that the holocephalan operculum evolved in concert with gill arch appendage reduction by attenuation of Shh-mediated branchial ray outgrowth, and that chondrichthyan branchial rays and tetrapod limbs exhibit parallel developmental mechanisms of evolutionary reduction.

Not all fish are equal: functional biodiversity of cartilaginous fishes (Elasmobranchii and Holocephali) in Chile.

A review of the primary literature on the cartilaginous fishes (sharks, skates, rays and chimaeras), together with new information suggests that 106 species occur in Chilean waters, comprising 58 sharks, 30 skates, 13 rays and five chimaeras. The presence of 93 species was confirmed, although 30 species were encountered rarely, through validated catch records and sightings made in artisanal and commercial fisheries and on specific research cruises. Overall, only 63 species appear to have a range distribution that normally includes Chilean waters. Actual reliable records of occurrence are lacking for 13 species. Chile has a cartilaginous fish fauna that is relatively impoverished compared with the global species inventory, but conservative compared with countries in South America with warm-temperate waters. The region of highest species richness occurs in the mid-Chilean latitudes of c. 30-40° S. This region represents a transition zone with a mix of species related to both the warm-temperate Peruvian province to the north and cold-temperate Magellan province to the south. This study provides clarification of species occurrence and the functional biodiversity of Chile's cartilaginous fish fauna.

Complementary redescription of Anacanthobatis ori (Wallace, 1967) and its assignment to Indobatis n. g. (Elasmobranchii, Anacanthobatidae), with comments on other legskates.

Anacanthobatis ori is one of the least known species of the family Anacanthobatidae with only four juvenile specimens reported. The species remained assigned to the genus Anacanthobatis sensu lato due to the lack of an adult male as external and skeletal clasper characters are the essential diagnostic features for the differentiation of genera and subgenera within the family Anacanthobatidae. Since an adult male of A. ori became available, along with an adult female and six further juveniles, the authors reinvestigated the species and present its so far unknown diagnostic characters of clasper morphology and skeleton and scapulocoracoid. The clasper turned out to be the most complex one of all known anacanthobatids as the external components flag, slit, pseudosiphon-like cavity, pecten, and two sentinas are not known from any other anacanthobatid species. Furthermore, a dorsal terminal 1 cartilage is present but displaced proximally of the terminal clasper skeleton, the outer edge of dorsal terminal 2 is deeply serrated, the ventral terminal has a very long, curved, strap-like process, and the proximal part of accessory terminal 1 is embedded in the cavity of the baseball-glove-like head of accessory terminal 2. Due to the strong differences in external and internal clasper characters to all other known anacanthobatid species, A. ori is placed in its own, newly erected genus, Indobatis. 

Evolutionary trends in the elasmobranch neurocranium.

The neurocranium (braincase) is one of the defining vertebrate characters. Housing the brain and other key sensory organs, articulating with the jaws and contributing to the shape of the anteriormost portion of the body, the braincase is undoubtedly of great functional importance. Through studying relationships between braincase shape and ecology we can gain an improved understanding of form-function relationships in extant and fossil taxa. Elasmobranchii (sharks and rays) represent an important case study of vertebrate braincase diversity as their neurocranium is simplified and somewhat decoupled from other components of the cranium relative to other vertebrates. Little is known about the associations between ecology and braincase shape in this clade. In this study we report patterns of mosaic cranial evolution in Elasmobranchii that differ significantly from those present in other clades. The degree of evolutionary modularity also differs between Selachii and Batoidea. In both cases innovation in the jaw suspension appears to have driven shifts in patterns of integration and modularity, subsequently facilitating ecological diversification. Our results confirm the importance of water depth and biogeography as drivers of elasmobranch cranial diversity and indicate that skeletal articulation between the neurocranium and jaws represents a major constraint upon the evolution of braincase shape in vertebrates.

Seasonality and relative abundance within an elasmobranch assemblage near a major biogeographic divide.

Nearshore waters are utilized by elasmobranchs in various ways, including foraging, reproduction, and migration. Multiple elasmobranch species have been previously documented in the nearshore waters of North Carolina, USA, which has a biogeographic break at Cape Hatteras on the Atlantic coast. However, comprehensive understanding of the elasmobranch community in this region is still lacking. Monthly year-round trawling conducted along two ocean transects (near Cape Lookout and Masonboro Inlet in 5 to 18 m depth) in Onslow Bay, North Carolina provided the opportunity to examine the dynamics and seasonal patterns of this community using a multivariate approach, including permutational multivariate analysis of variance and nonparametric BIO-ENV analysis. From November 2004 to April 2008, 21,149 elasmobranchs comprised of 20 species were caught, dominated by spiny dogfish (Squalus acanthias) and clearnose skate (Rostroraja eglanteria). All species exhibited seasonal variation in abundance, but several key species contributed the most to seasonal differences in species composition within each transect. Spiny dogfish was most abundant in the winter at both locations, comprised mainly of mature females. Although clearnose skate was caught in all seasons, the species was most abundant during the spring and fall. Atlantic sharpnose (Rhizoprionodon terraenovae) was one of the most abundant species in the summer, and two distinct size cohorts were documented. Temperature appeared to be the main abiotic factor driving the community assemblage. The extensive year-round sampling provided the ability to better understand the dramatic seasonal variation in species composition and provides new information on the relative abundance of several understudied elasmobranch species that may be of significant ecological importance. Our results underscore the importance of inner continental shelf waters as important elasmobranch habitat and provide baseline data to examine for future shifts in timing and community structure at the northern portion of the biogeographic break at Cape Hatteras.

Planonasus parini n. g. and n. sp., a new genus and species of false cat sharks (Carchariniformes, Pseudotriakidae) from the deep northwestern Indian Ocean off Socotra Islands.

A new genus and species of the carcharhiniform family Pseudotriakidae is described based on three specimens caught near the Socotra Islands in the northwestern Indian Ocean. The first specimen and holotype of Planonasus parini g. n. and sp. n. was caught during cruise 17 of RV 'Vityaz' in 1988/89 along the deep western Indian Ocean. Two further specimens of the new genus and species were caught somewhat later by commercial trawlers close to the locality of the holotype. The new genus differs from the two other pseudotriakid genera Gollum and Pseudotriakis by the presence of oral papillae, the absence of nicitating eyelids, a longer head, an intermediate prenarial snout length, an intermediate number of tooth rows per jaw, a first dorsal fin of intermediate height and length and with a white free rear tip, a caudal peduncle of inter-mediate length, and fewer vertebrae.

Introduction to the systematics and biodiversity of sharks, rays, and chimaeras (Chondrichthyes) of Taiwan.

All 13 orders of chondrichthyan fishes occur in Taiwanese waters, representing 52 chondrichthyan families (31 shark, 19 batoid, 2 chimaeroid) and 98 genera (64 shark, 31 batoid, 3 chimaeroid). A total of 119 shark, 58 batoid, and 4 chimaera species may occur in the waters surrounding Taiwan, pending taxonomic resolution of some groups. Of the 34 nominally described species from Taiwan, 17 are currently considered valid. The majority of named species occurred during two peak periods in Taiwanese chondrichthyan research; the first between 1959-63, when 13 nominal species were described, of which 7 remain valid today, and a second peak period between 2003-13 when 9 nominal species were described, of which 6 remain valid. The overall species diversity of Taiwan's chondrichthyan fauna is comparable to that of other adjacent marine zoogeographic hotspots, e.g. Japan (126 shark, 75 batoid, 11 chimaeroid species) and the Philippines (81 shark, 46 batoid, 2 chimaeroid species). The Carcharhiniformes, Squaliformes, Myliobatiformes, and Rajiformes are the most dominant orders in terms of abundance and species-richness within this region. Each of these groups may increase in relative diversity with improved taxonomic resolution resulting from the incorporation of molecular tools and renewed morphological studies. Improved identification of Taiwan's chondrichthyan fauna will aid in developing better conservation and management practices.

A new species of wedgefish, Rhynchobatus immaculatus (Chondrichthyes, Rhynchobatidae), from Taiwan.

A new species of wedgefish, Rhynchobatus immaculatus sp. nov., is described from a small collection of specimens obtained from fish markets in northern Taiwan. It is probably a medium-sized species (probably attaining ca. 1.5 m TL) because the largest known specimen, an immature male (ca. I m TL), has prolongated dorsal and caudal fins typical of adult wedgefishes. Rhynchobatus immaculatus is unique within the family in having a very high vertebral count (within the range of 165-170 total free centra) and in lacking a dark pectoral marking. Other Rhynchobatus species occurring in Taiwanese seas appear to attain a larger adult size, possess a dark pectoral marking at least in young, and have lower vertebral counts (fewer than 161 total fee centra). Rhynchobatus yentinesis, which was described from a specimen taken nearby at Wenzhou, China, has not yet been attributed to a currently recognised species. However, based on the illustration of the holotype, which reveals a broad-snouted species with a dark pectoral spot, it is closest to either R. palpebratus or R. springeri.

Records of new and rare elasmobranchs from Taiwan.

Five new records of elasmobranchs collected from eastern Taiwan fish markets, Da-xi and Cheng-gong, are presented. Samples were caught by deepsea longliners and bottom trawlers which operate in northeastern waters off Taiwan between 2004 and 2012. These five new species records include the smalltooth sandtiger, Odontaspis ferox (Risso, 1810) (Lamniformes: Odontaspididae), salamander shark, Parmaturus pilosus Garman, 1906 (Carcharhiniformes: Scyliorhinidae), leadhued skate Notoraja tobitukai (Hiyama, 1940) (Rajiformes: Arhynchobatidae), giant skate Dipturus gigas (Ishiyama, 1958) (Rajiformes: Rajidae), and the pelagic stingray Pteroplatytrygon violacea (Bonaparte, 1832) (Myliobatiformes: Dasyatidae), Diagnostic characteristics for each species are given and a key to the genera Parmaturus and Dipturus from Taiwan is presented.