First records of the Pacific sleeper shark Somniosus cf. pacificus in the western tropical Pacific.

Technological advances have enabled the expansion of ocean exploration to include the deep ocean, providing new species observations. Here, the authors present two new observations, captured by deep-sea cameras, of the sleeper shark Somniosus cf. pacificus from the Solomon Islands and Palau. This presents the first observation of S. cf. pacificus in the western Pacific tropics and extends its range about 2000 nautical miles south. The observations presented here provide much-needed information on the range of this species which can help guide future management and conservation actions.

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.

Age and Growth of Elasmobranchs and Applications to Fisheries Management and Conservation in the Northeast Pacific Ocean.

In addition to being an academic endeavour, the practical purpose of conducting age and growth studies on fishes is to provide biological data to stock assessment scientists and fisheries managers so they may better understand population demographics and manage exploitation rates. Age and size data are used to build growth models, which are a critical component of stock assessments. Though age determination of elasmobranchs in the northeast Pacific Ocean (NEP) began in the 1930s, the field has evolved substantially in recent years, allowing scientists to incorporate age data into assessments for more species than ever before. Owing to the highly diverse biology of this group of fishes, each species has its own set of challenges with regard to age determination. Age determination methods typically rely on semicalcified hard structures that form regular growth patterns; however, the structure selected and preparation method used is often species specific. New staining techniques have improved the ability to assess age and improve ageing precision for some species, and advances in microchemical methods have allowed for independent means of estimating age and validating age determination accuracy. Here we describe current age determination methods for NEP elasmobranchs. While the library of available techniques is increasing, there are still some NEP species for which reliable ageing methods have yet to be defined; we discuss these challenges and potential avenues of future research. Finally, we conclude by describing how age estimates are used in growth models and subsequently in stock assessments of selected NEP elasmobranchs.

The secret of the mermaid's purse: phylogenetic affinities within the Rajidae and the evolution of a novel reproductive strategy in skates.

The systematics of the skates in the family Rajidae have been contentious for over 250years, with most studies inferring relationships among geographically clustered species, and non-overlapping taxa and data sets. Rajid skates are oviparous, and lay egg capsules with a single embryo. However, two species exhibit a derived form of egg laying, with multiple embryos per egg capsule. We provide a molecular assessment of the phylogenetic relationships of skates in the family Rajidae based on three mitochondrial genes. The resulting topology supports monophyly the family. However the genusRajais polyphyletic, and several species assemblages need to be revised. We propose a new assemblage, the Rostrajini, which organizesrajid species into three well-supported tribal lineages for the first time. Further, these data provide an independent assessment of monophyly for the two species exhibiting multiple embryos per egg capsule, supporting their status as the unique genusBeringraja. In addition, we find that among the different size classes of egg capsules, ranging from 1 to 8 embryos per capsule in this genus, there is variation in frequency and survivorship. InBeringraja binoculata, the strategy of having two embryos per egg capsule occurs most frequently and with the highest fitness.

Overfishing and climate change elevate extinction risk of endemic sharks and rays in the southwest Indian Ocean hotspot.

Here, we summarise the extinction risk of the sharks and rays endemic to coastal, shelf, and slope waters of the southwest Indian Ocean and adjacent waters (SWIO+, Namibia to Kenya, including SWIO islands). This region is a hotspot of endemic and evolutionarily distinct sharks and rays. Nearly one-fifth (n = 13 of 70, 18.6%) of endemic sharks and rays are threatened, of these: one is Critically Endangered, five are Endangered, and seven are Vulnerable. A further seven (10.0%) are Near Threatened, 33 (47.1%) are Least Concern, and 17 (24.3%) are Data Deficient. While the primary threat is overfishing, there are the first signs that climate change is contributing to elevated extinction risk through habitat reduction and inshore distributional shifts. By backcasting their status, few endemic species were threatened in 1980, but this changed soon after the emergence of targeted shark and ray fisheries. South Africa has the highest national conservation responsibility, followed by Mozambique and Madagascar. Yet, while fisheries management and enforcement have improved in South Africa over recent decades, substantial improvements are urgently needed elsewhere. To avoid extinction and ensure robust populations of the region's endemic sharks and rays and maintain ecosystem functionality, there is an urgent need for the strict protection of Critically Endangered and Endangered species and sustainable management of Vulnerable, Near Threatened, and Least Concern species, underpinned by species-level data collection and reduction of incidental catch.

Fishing for oil and meat drives irreversible defaunation of deepwater sharks and rays.

The deep ocean is the last natural biodiversity refuge from the reach of human activities. Deepwater sharks and rays are among the most sensitive marine vertebrates to overexploitation. One-third of threatened deepwater sharks are targeted, and half the species targeted for the international liver-oil trade are threatened with extinction. Steep population declines cannot be easily reversed owing to long generation lengths, low recovery potentials, and the near absence of management. Depth and spatial limits to fishing activity could improve conservation when implemented alongside catch regulations, bycatch mitigation, and international trade regulation. Deepwater sharks and rays require immediate trade and fishing regulations to prevent irreversible defaunation and promote recovery of this threatened megafauna group.

Lost and found: recovery of the holotype of the ocellated angelshark, Squatina tergocellatoides Chen, 1963 (Squatinidae), with comments on western Pacific squatinids.

The ocellated angelshark, Squatina tergocellatoides, Chen, 1963 is redescribed from the holotype, which was thought to be lost. Its recent recovery has allowed for a revised description, including new data, and comparison to other Western Pacific squatinids. Squatina tergocellatoides can be distinguished from its congeners by three pairs of prominent large black spots, each with a diameter greater than eye length; two on each pectoral fin at anterior and posterior angles and one on each side near the tail base; another three pairs of lesser defined spots, one large spot on base of each dorsal fin and one located laterally on each side of tail located below first dorsal fin. Ventral surface is uniformly white to cream coloured, and margins of pectoral fins and tail similar in colour to dorsal side. Pectoral fins with angular lateral apices and rounded posterior lobe, pelvic fin tips not reaching origin of first dorsal fin, strongly fringed nasal barbels, small inter-orbital space, head and mouth lengths, broad internarial width and pelvic fin base, a very small pelvic girdle width, and a caudal fin with triangular ventral lobe greater in length than dorsal lobe. Comments on additional specimens are provided, as well as observations on biogeography. A review of western Pacific squatinids is also provided.

Pristiophorus lanae sp. nov., a new sawshark species from the western North Pacific, with comments on the genus Pristiophorus Müller & Henle, 1837 (Chondrichthyes: Pristiophoridae).

A new species of sawshark, Pristiophorus lanae sp. nov., is described from off the Philippine Islands. The new species is the second member of the genus Pristiophorus described from the western North Pacific and can be separated from its closest geographic congener, P. japonicus, by having fewer rostral teeth in front of rostral barbels (17-26 versus 25-32), mouth at corners extending forward to below the rear margin of the eye versus extending below the rear one-third of eye margin, a greater mouth width at 6.9-7.8 times into pre-oral length (versus 5.8-6.9), eye length into head length (15.6-15.9 versus 9.8-13.2), mouth width into head length 9.0-10.0 versus 7.4-8.5 times, head width at nostrils 5.2-6.1 times into pre-orbital length versus 3.9-4.9 times, shorter prebarbel length (from snout tip to barbel) of 50.7-54.5% of preoral length versus 53.6-59.2%, a snout angle of 10.6-13.0 degrees versus 12.4 degrees - 14.6 degrees, and lateral trunk denticles with flat crowns that are imbricated versus erect crowns that are not imbricated. The number of monospondylous vertebrae is slightly lowe in P. lanae (43-48) versus P. japonicus (51-52). The genus is reviewed, with a revised key to its species presented.

Redescription of Hexanchus nakamurai Teng 1962, (Chondrichthyes: Hexanchiformes: Hexanchidae), with designation of a neotype.

Hexanchus nakamurai Teng, 1962 is one of two valid nominal species in the genus Hexanchus; the other being H. griseus. The taxonomic history of H. nakamurai is somewhat convoluted due to questions about the validity of whether it constituted a publication or an unpublished dissertation. The issue appeared to have been resolved once it was determined that the Teng's original description met the criteria under the International Commission on Zoological Nomenclature code (Article 8a) of a publication. However, recent molecular studies have indicated that the western North Atlantic H. nakamurai (= ? H. vitulus) may be distinct from western Indian Ocean H. nakamurai. Compounding the issue is the loss of the holotype of H. nakamurai from Taiwan. A neotype is herein designated and the species redescribed based on the neotype and eight additional Taiwanese specimens.

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.

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.

Revision of the genus Centrophorus (Squaliformes: Centrophoridae): Part 1--Redescription of Centrophorus granulosus (Bloch & Schneider), a senior synonym of C. acus Garman and C. niaukang Teng.

The genus Centrophorus is one of the most taxonomically complex and confusing elasmobranch groups. A revision of this group is currently underway and this first paper sets an important foundation in this process by redescribing the type species of the genus--Centrophorus granulosus. This taxon name has been previously applied to two different morphotypes: a large species > 1.5 m TL and a smaller species -1 m TL. Centrophorus acus and C. niaukang are the most commonly used names applied to the larger morphotype. The original description of C. granulosus was based on a large specimen of -1.5 m TL, but subsequent redescriptions were based on either of the large or small morphotypes. Centrophorus granulosus is herein redescribed as a large species and a neotype is designated. Centrophorus acus and C. niaukang are found to be junior synonyms of C. granulosus. Centrophorus granulosus is distinguishable from its congeners by its large size, dermal denticle shape, colouration and a number of morphological and biological characteristics. Ontogenetic changes in morphology, dentition and denticle shape for this species are described in detail.

An annotated checklist of the chondrichthyans of Taiwan.

An annotated checklist of chondrichthyan fishes (sharks, batoids, and chimaeras) occurring in Taiwanese waters is presented. The checklist is the result of a biodiversity workshop held in Mach 2012 as well as on-going systematic revisions by the authors. The chondrichthyan fauna of Taiwan is one of the richest in the world with the number of species totaling 181, comprising 52 families and 98 genera. It includes 31 families, 64 genera, and 119 species of sharks, 19 families, 31 genera, and 58 species of batoids, and 2 families, 3 genera, and 4 species of chimaeras. The most species-rich families are the Carcharhinidae with 22 species followed by the Scyliorhinidae with 17. The most species-rich batoid families are the Dasyatidae with 11 species and and the Rajidae with 10. Verified voucher material is provided for each species where available and potential taxonomic issues are high-lighted when applicable. This represents the first detailed, evidence-based checklist of chondrichthyans from Taiwanese waters in over 40 years.

A taxonomic revision of Northeast Pacific softnose skates (Rajiformes: Arhynchobatidae: Bathyraja Ishiyama).

Softnose skates (Rajiformes: Arhynchobatidae: Bathyraja Ishiyama) are the most diverse skate genus, with 54 species, and are readily distinguishable from other genera in the family by their poorly calcified, flexible rostral cartilages. Six Bathyraja species are considered valid in the eastern North Pacific, including: B. abyssicola, B. aleutica, B. kincaidii, microtrachys, B. spinosissima, and B. trachura. Similar to other skate genera, eastern North Pacific Bathyraja lack a robust species-specific identification, which leads to issues with setting catch limits and creating management plans. This study identifies and formally redescribes the eastern North Pacific softnose skate species based on morphometric and meristic measurements and includes an Alaskan species, Bering Skate, B. interrupta due to its close morphological relationship to B. kincaidii. A lectotype for B. interrupta is designated. Multivariate tests determined that significant differences existed between the study species. Parsimonious phylogenetic trees showed that B kincaidii represents the basal condition, with B. abyssicola and B. aleutica being the most derived species in the study. The formerly synonymized species B. interrupta and B. kincaidii were shown to be separate, as were the previously synonymized species B. microtrachys and B. trachura. Improved Bathyraja species identification will hopefully assist fisheries managers in developing conservation policies easing the impacts of deep-sea fishing expansion.

Comparative analyses of the complete mitochondrial genomes of two southern African endemic guitarfish, Acroteriobatus annulatus and A. blochii.

Shark-like rays (order Rhinopristiformes) are among the most threatened cartilaginous fish globally. Despite this, unresolved taxonomic issues still exist within the group. To date, no studies have used complete mitochondrial genomes to assess the phylogenetic placement of Acroteriobatus within the non-monophyletic family Rhinobatidae. The current study reports the first complete mitochondrial genomes for Acroteriobatus annulatus and A. blochii. Similar to other rhinopristiforms, the complete sequences of A. annulatus (16,773 bp) and A. blochii (16,771 bp) were circular molecules with gene organisations identical to that of the typical vertebrate mitogenome. The A + T content was higher than the G + C content, with a bias towards A and C nucleotides observed in all complete mitogenomes. The stem-and-loop secondary structures of the putative origin of light-strand replication were found to have highly conserved synthesis and stem regions, with all substitutions and indels restricted to the loop structure. The ratios of non-synonymous to synonymous substitution rates indicated that purifying selection has been the dominant driver of evolution in rhinopristiform mitogenomes. Phylogenetic reconstructions placed Acroteriobatus as a sister-group to Rhinobatos, confirming its affiliation with the family Rhinobatidae. However, based on its apparent polyphyly with the aforementioned genera, the familial assignment of Pseudobatos is not fully resolved and requires further investigation.

Revision of the genus Centrophorus (Squaliformes: Centrophoridae): Part 3Redescription of Centrophorus uyato (Rafinesque) with a discussion of its complicated nomenclatural history.

Centrophorus uyato (Rafinesque, 1810) has a complicated nomenclatural history which has led to multiple scientific names being ascribed to this species. In the Mediterranean Sea, and elsewhere in its range, this species was previously referred to as C. granulosus (Bloch Schneider, 1801). The first paper in this revision series clarified that C. granulosus refers to a much larger species of gulper shark which attains at least 1.7 m length and is absent from the Mediterranean Sea. Further complicating the nomenclature of this species is the fact that the original description clearly refers to a Squalus species, and not a Centrophorus species. In this third part of the revision of the genus Centrophorus, this problematic species is redescribed. A detailed synonymy is provided and it is confirmed that C. machiquensis, C. bragancae and C. zeehaani are junior synonyms of this species. To preserve nomenclatural stability within the genus, the name Centrophorus uyato is retained for this species with a neotype from close to the original type locality off Italy being designated. Intraspecific variation within Centrophorus uyato is also discussed, particularly relating to denticle morphology and body morphology.

An annotated checklist of the chondrichthyans of South Africa.

An annotated checklist of chondrichthyan fishes (sharks, batoids, and chimaeras) occurring in South African waters is presented. The checklist is the result of decades of research and on-going systematic revisions of the regional fauna. The chondrichthyan fauna of South Africa is one of the richest in the world with 191 species, comprising 50 families and 103 genera. It consists of 30 families, 64 genera, and 111 species of sharks; 17 families, 36 genera, and 72 species of batoids; and, 3 families, 5 genera, and 8 species of chimaeras. The most species-rich shark families are the whaler sharks Carcharhinidae with 20 species followed by the deepwater catsharks Pentanchidae with 13 species. The most species-rich batoid families are the hardnose stakes Rajidae with at least 21 species followed by the stingrays Dasyatidae with 13 species. This monograph represents the first detailed annotated checklist of chondrichthyans from South Africa in over 30 years.

Overfishing drives over one-third of all sharks and rays toward a global extinction crisis.

The scale and drivers of marine biodiversity loss are being revealed by the International Union for Conservation of Nature (IUCN) Red List assessment process. We present the first global reassessment of 1,199 species in Class Chondrichthyes-sharks, rays, and chimeras. The first global assessment (in 2014) concluded that one-quarter (24%) of species were threatened. Now, 391 (32.6%) species are threatened with extinction. When this percentage of threat is applied to Data Deficient species, more than one-third (37.5%) of chondrichthyans are estimated to be threatened, with much of this change resulting from new information. Three species are Critically Endangered (Possibly Extinct), representing possibly the first global marine fish extinctions due to overfishing. Consequently, the chondrichthyan extinction rate is potentially 25 extinctions per million species years, comparable to that of terrestrial vertebrates. Overfishing is the universal threat affecting all 391 threatened species and is the sole threat for 67.3% of species and interacts with three other threats for the remaining third: loss and degradation of habitat (31.2% of threatened species), climate change (10.2%), and pollution (6.9%). Species are disproportionately threatened in tropical and subtropical coastal waters. Science-based limits on fishing, effective marine protected areas, and approaches that reduce or eliminate fishing mortality are urgently needed to minimize mortality of threatened species and ensure sustainable catch and trade of others. Immediate action is essential to prevent further extinctions and protect the potential for food security and ecosystem functions provided by this iconic lineage of predators.

Leucoraja elaineae sp. nov., a new rough skate (Rajiformes: Rajidae) from the Western Indian Ocean.

A new species of the genus Leucoraja is described from off Kenya in the Western Indian Ocean (WIO). The new species was collected during a survey of the R/V Fridtiof Nansen. Leucoraja elaineae sp. n. is the third species in the genus found to occur in the WIO, and the only species in the genus Leucoraja known exclusively from this region. Its closest geographic congener L. wallacei, occurring from Namibia to southern Mozambique, can be separated by a combination of morphological and meristic characters including an elongated, pronounced snout (L. elaineae sp. n.) vs a short, blunt snout (L. wallacei), higher predorsal vertebral (107 vs 64-74) and pectoral fin radial counts (77 vs 61-64), a dorsal surface pattern consisting of a single ocelli at the midbase of each pectoral fin, paired small white spots, and no banding or crossbars on its tail vs numerous rosettes and whorls, no prominent ocelli at the pectoral fin bases, and 3-4 tail bands. The only other WIO Leucoraja species in the genus, L. compagnoi is known only from South Africa and has a short, blunt snout, a lower upper tooth count (38 vs 57 in L. elaineae sp. n.), a plain dorsal disc color with no markings and distinct bands on its tail. All other members of the genus occur in the North Atlantic, except one species that occurs in the eastern Central Atlantic and one in the eastern Indian Ocean.

Redescription and taxonomic status of Dipturus chilensis (Guichenot, 1848), and description of Dipturus lamillai sp. nov. (Rajiformes: Rajidae), a new species of long-snout skate from the Falkland Islands.

Recent molecular evidence has called into question the identity of skates collected in the waters off the Falkland Islands previously identified as Zearaja chilensis. NADH2 sequence data indicate that these specimens are not conspecific with those currently referred to as Z. chilensis from Chile and, in fact, represent a novel cryptic species. This study aimed to investigate this hypothesis based on morphological comparisons of specimens from the coasts of both western and eastern South America. In total, 50 specimens from Chile and 41 specimens from the Falkland Islands were collected and examined; morphometric data were generated for a subset of specimens from both areas. NADH2 sequence data were generated for a total of 19 specimens from both areas, as well as specimens of Z. nasuta from New Zealand, D. pullopunctatus from South Africa, D. oxyrinchus from the Azores, Okamejei hollandi, and O. cairae from Borneo, and O. kenojei from Japan. Based on morphological and molecular analyses, Zearaja is synonymized with Dipturus and species assigned to the former genus are transferred to the latter genus. A neotype is designated for D. chilensis and this species is redescribed. Dipturus lamillai sp. nov. is described based on specimens from the Falkland Islands. Comparison of our NADH2 data with data for mitochondrial genomes generated from tissue samples taken from two specimens originally identified as Z. chilensis, indicate that, while the sample from Chile came from a specimen of D. chilensis, that from the skate steak obtained from a restaurant in Korea actually came from a specimen of D. lamillai sp. nov. This emphasizes the importance of confirming both the provenance and identity of specimens from which sequence data are generated and submitted to GenBank if misidentifications are to be avoided.