Teeth of Embryos in Lamniform Sharks (Chondrichthyes: Elasmobranchii)

The dentitions of lamniform sharks possess a unique heterodonty, the lamnoid tooth pattern. However, in embryos, there are 'embryonic' and 'adult' dentitions. The teeth in the embryonic dentition are peg-like and appear to be attached to the jaw in an acrodont fashion. The adult dentition is characterized by the presence of replacement tooth series with the lamnoid tooth pattern. The embryonic–adult transition in dentitions appears at around 30–60cm TL. Tooth replacement generally begins before birth in embryos with adult dentitions. The adult dentition becomes functional just before or after parturition. An embryo of one species (Lamna nasus) shows a tooth directly on the symphysis of the upper jaws, marking the first record of a medial tooth for the order Lamniformes.     

A review of the sensory biology of chimaeroid fishes (Chondrichthyes; Holocephali)

The chimaeroid fishes (Chondrichthyes: Holocephali) are a small, ancient and poorly studied group of cartilaginous fishes that have puzzled and intrigued taxonomists, ichthyologists and evolutionary biologists for over 100 years. Like their close relatives, the elasmobranchs (sharks, skates and rays), chimaeroids possess an extensive battery of sense organs that allow them to detect information about the external environment in order to find mates, locate food and preferred habitats and avoid predators. In recent years the sensory systems of elasmobranchs have received an up-swell of attention from biologists, which has resulted in a greater understanding of the sensory capabilities and behaviour of these fishes. However, very little recent work has been done on the chimaeroids. The aim of this review is to provide a survey of the existing literature on the major senses (vision, smell, taste, mechanoreception, hearing and electroreception) in chimaeroids, in order to stimulate and identify areas for future research. In chimaeroids information on sensory systems is largely restricted to one or two species (with the exception of some aspects of the visual system) and for some sensory systems essentially nothing is known. Most studies are anatomical in nature and so there is a demand for a greater degree of neurophysiological and behavioural assessment of sensory capability in these fishes. The majority of chimaeroids occupy deep-sea habitats and are becoming increasingly threatened by the expansion of deep-sea fisheries, so an understanding of the sensory biology and behaviour of chimaeroids may be important for the protection and management of these fascinating fishes.     

The fine structure of spermatozoa of Hydrolagus colliei (Chondrichthyes, Holocephali)

The ultrastructure of spermatozoa in Hydrolagus colliei is described. Basic similarities of structure to the sperm of the related elasmobranch fish are noted. The most significant features of sperm structure in Hydrolagus that differ from those of elasmobranch fish occur in the tail. The axoneme is eccentrically located and forms a double helix with a single longitudinal column. A second longitudinal column is reduced to a short remnant at the base of the tail. Microtubules within the axoneme are also helically disposed, a feature that is consistent with the rotating motion of the sperm. Abundant glycogen reserves are stored along the length of the tail.     

Chimaera opalescens n. sp., a new chimaeroid (Chondrichthyes: Holocephali) from the north-eastern Atlantic Ocean

A new species of chimaeroid, Chimaera opalescens n. sp., was described from 31 type specimens caught in the north-eastern Atlantic Ocean at 950-1400 m depth. This species differed from all its congeners by the combination of the following characters: body uniformly pale brown to bronze, iridescent on fresh specimens; unpaired fins brown to purple, uniformly coloured or with pale or whitish edges; iris black; claspers tripartite and divided for a third of their length, not extending beyond the pelvic fins in adults; dorsal spine equal to or shorter than first dorsal fin; ventral caudal lobe equal to or deeper than dorsal caudal lobe. Comparison of DNA sequences of the CO1 gene with those of related species supported C. opalescens n. sp. as a distinct species. Specimens of C. opalescens n. sp. were previously misidentified as Chimaera monstrosa, a species sharing a similar geographical distribution. Chimaera opalescens n. sp. lives in deeper water and is a larger-bodied species than C. monstrosa. The two species were newly exploited by deep-sea fisheries and confused under a single landing name.     

Meiotic chromatin diminution in a vertebrate,the holocephalan fish Hydrolagus collie (Chondrichthyes,Holocephali)

A histochemical, microdensitometric, and electron microscopic study of testes of the ratfish Hydrolagus colliei shows that an instance of the rare phenomenon of germ line chromatin diminution occurs in this vertebrate species. In primary spermatocytes at metaphase I a spherical mass of heterochromatin accumulates at one side of the metaphase plate. At anaphase I the heterochromatic mass is left in the equatorial cytoplasm and is passed into one of the two secondary spermatocytes formed during cytokinesis. As nuclear membranes are being restored, a double membrane envelope is also formed around the heterochromatic mass, which is then termed the ‘chromatin diminution body’ (CDB). At second meiotic division the CDB is included in the cytoplasm of one of the four spermatids and retained there, apparently unchanged, until mid-spermiogenesis. At that time the CDB becomes adherent to the spermatid plasma membrane and is pinched off from the spermatid by a process of apocrine exocytosis, taking a layer of spermatid plasma membrane along with it. Simultaneously this tri-membrane CDB is taken into the adjacent Sertoli cell by endocytosis, thereby acquiring a fourth membrane layer, a part of the Sertoli cell plasma membrane. The CDBs are subsequently phagocytized, possibly first fusing with dense, multilaminate bodies in the Sertoli cell cytoplasm. The CDB chromatin mass is strongly positive with the Feulgen method for DNA and the alkaline fast green method for histones. Microdensitometric analysis shows that the discarded chromatin amounts to about 10% of the diploid nuclear content and that it appears to be part of the normal diploid complement rather than DNA amplified during meiosis.     

The Sizes of Elephant Groups in Zoos: Implications for Elephant Welfare

This study examined the distribution of 495 Asian elephants (Elephas maximus) and 336 African elephants (Loxodonta africana) in 194 zoos, most of which were located in Europe (49.1%) and North America (32.6%). Cows outnumbered bulls 4 to 1 (Loxodonta) and 3 to 1 (Elephas). Groups contained 7 or fewer: mean, 4.28 (σ = 5.73). One fifth of elephants lived alone or with one conspecific. Forty-six elephants (5.5%) had no conspecific. Many zoos ignore minimum group sizes of regional zoo association guidelines. The American Zoo and Aquarium Association recommends that breeding facilities keep herds of 6 to 12 elephants. The British and Irish Association of Zoos and Aquariums recommends keeping together at least 4 cows over 2 years old. Over 69% Asian and 80% African cow groups—including those under 2 years—consisted of fewer than 4 individuals. Recently, Europe and North America have made progress with some zoos no longer keeping elephants and with others investing in improved facilities and forming larger herds. The welfare of individual elephants should outweigh all other considerations; zoos should urgently seek to integrate small groups into larger herds.     

Glial architecture of the ghost shark (Callorhinchus milii, Holocephali, Chondrichthyes) as revealed by different immunohistochemical markers

This article presents the first study on the glial architecture of a representative species of Holocephali, Callorhinchus milii (ghost shark). Holocephali are a small subclass of Chondrichthyes, with only a few extant genera, and those are considered to have a brain organization more similar to squalomorph sharks than to galeomorph sharks, skates, and rays. Three different astroglial markers--glial fibrillary acidic protein, S-100 protein, and glutamine synthetase (GS)--were investigated by immunohistochemical methods, applying both diaminobenzidine (DAB) and fluorescent techniques. They revealed similar glial structures, although most of them were detected by immunohistochemical reaction against GS and visualized by DAB. The predominant elements were radial ependymoglia spanning the area between the ventricular and meningeal surfaces, as in squalomorph sharks. Other similar features were the light appearance of myelinated neural tracts devoid of immunoreactivity, and the glial architecture of the reticular formation of the brain stem, cerebellum, and tectum, the latter with recognizable layers. The immunoreactivity of the vascular walls was similar; however, it is believed that different cell types form the blood-brain barrier in chimeras and in elasmobranchs. Some glial structures, however, resembled those of skates, rays, and galeomorph sharks. In C. milii astrocyte-like elements were observed in the telencephalon, using GS and S-100, although typical astrocyte-rich regions were not found. In some areas, especially the telencephalon, not only endfeet but also cell bodies were observed to be attached to the meningeal surface, with processes extending into the brain substance.     

Comparative genomics approach to detecting split-coding regions in a low-coverage genome: lessons from the chimaera Callorhinchus milii (Holocephali, Chondrichthyes)

Recent development of deep sequencing technologies has facilitated de novo genome sequencing projects, now conducted even by individual laboratories. However, this will yield more and more genome sequences that are not well assembled, and will hinder thorough annotation when no closely related reference genome is available. One of the challenging issues is the identification of protein-coding sequences split into multiple unassembled genomic segments, which can confound orthology assignment and various laboratory experiments requiring the identification of individual genes. In this study, using the genome of a cartilaginous fish, Callorhinchus milii, as test case, we performed gene prediction using a model specifically trained for this genome. We implemented an algorithm, designated ESPRIT, to identify possible linkages between multiple protein-coding portions derived from a single genomic locus split into multiple unassembled genomic segments. We developed a validation framework based on an artificially fragmented human genome, improvements between early and recent mouse genome assemblies, comparison with experimentally validated sequences from GenBank, and phylogenetic analyses. Our strategy provided insights into practical solutions for efficient annotation of only partially sequenced (low-coverage) genomes. To our knowledge, our study is the first formulation of a method to link unassembled genomic segments based on proteomes of relatively distantly related species as references.     

The role of a Sertoli cell actin-myosin system in sperm bundle formation in the ratfish, Hydrolagus colliei (Chondrichthyes, Holocephali)

Sertoli cells in the ratfish entirely surround a clone of spermatids to form a spermatocyst. As spermiogenesis proceeds within the cyst cavity, the acrosome areas become apposed to the Sertoli cell plasma membrane lining the spermatocyst. The spermatids elongate and are gathered into an increasingly compact bundle oriented with acrosomal tips directed toward the Sertoli cell base. As all acrosome areas move closer together, Sertoli cell microfilaments oriented parallel to the long spermatid axis appear and increase in concentration. Actin and myosin were demonstrated in the microfilament area with fluorescent antibodies and NBD-Phallacidin. Simultaneously, endocytosis of Sertoli cell membrane between spermatid attachment sites removes the intervening membrane and allows the latter sites to approach each other. Sertoli cell endocytosis is spatially and temporally related to a unique projection at the basal rim of each acrosome. During midspermiogenesis, structured intercellular material appears between the Sertoli cell and the acrosomal region of each spermatid. Its periodicity is closely related to periodic arrangement of Sertoli cell actin and material within the spermatids. These attachment sites move together upon endocytosis, gathering a clone of spermatids into a closely packed bundle.     

Development and Evolution of Dentition Pattern and Tooth Order in the Skates And Rays (Batoidea; Chondrichthyes)

Shark and ray (elasmobranch) dentitions are well known for their multiple generations of teeth, with isolated teeth being common in the fossil record. However, how the diverse dentitions characteristic of elasmobranchs form is still poorly understood. Data on the development and maintenance of the dental patterning in this major vertebrate group will allow comparisons to other morphologically diverse taxa, including the bony fishes, in order to identify shared pattern characters for the vertebrate dentition as a whole. Data is especially lacking from the Batoidea (skates and rays), hence our objective is to compile data on embryonic and adult batoid tooth development contributing to ordering of the dentition, from cleared and stained specimens and micro-CT scans, with 3D rendered models. We selected species (adult and embryonic) spanning phylogenetically significant batoid clades, such that our observations may raise questions about relationships within the batoids, particularly with respect to current molecular-based analyses. We include developmental data from embryos of recent model organisms Leucoraja erinacea and Raja clavata to evaluate the earliest establishment of the dentition. Characters of the batoid dentition investigated include alternate addition of teeth as offset successional tooth rows (versus single separate files), presence of a symphyseal initiator region (symphyseal tooth present, or absent, but with two parasymphyseal teeth) and a restriction to tooth addition along each jaw reducing the number of tooth families, relative to addition of successor teeth within each family. Our ultimate aim is to understand the shared characters of the batoids, and whether or not these dental characters are shared more broadly within elasmobranchs, by comparing these to dentitions in shark outgroups. These developmental morphological analyses will provide a solid basis to better understand dental evolution in these important vertebrate groups as well as the general plesiomorphic vertebrate dental condition.     

An Annotated List of Cartilaginous Fishes (Chondrichthyes: Elasmobranchii,Holocephali) of the Coastal Waters of Sakhalin Island and the Adjacent Southern Part of the Sea of Okhotsk

An annotated list of cartilaginous fish (Chondrichthyes: Elasmobranchii, Holocephali) is given for the first time in the 200-year history of studying the ichthyofauna of Sakhalin Island and adjacent waters of the southern part of the Sea of Okhotsk (including the coast of Hokkaido Island) and the northern Sea of Japan. The list includes 43 species in two classes, eight orders, 16 families, and 25 genera. Information on nature conservation status, English and Latin names, depths of habitat, and distribution within the coastal waters of Sakhalin are presented. For a number of species caught off the coast of Sakhalin and in the adjacent waters, information is provided on collection specimens confirming their presence in the region under study. For a number of species of the Rajiformes order (Arctoraja parmifera, A. smirnovi, A. simoterus), the modern ranges and taxonomic status are being refined in the light of new data. The taxonomic status of the so-called “disputed” taxa is discussed as well as the validity of the species considered in the Bathyraja matsubarai complex. Based on the study of the collections, Arctoraja simoterus, previously unknown in the waters of Russia, as well as Myliobatis tobijei, caught in the Bering Sea, has been discovered, which significantly expands the range of this species to the north.     

On the South Atlantic distribution of Callorhinchus callorhynchus (Holocephali: Callorhynchidae)

The four species of the family Callorhynchidae are restricted to the southern hemisphere ( Didier 1995 ) and have the most littoral-related distribution among holocephalans.     

Assessment of the dorsal fin spine for chimaeroid (Holocephali: Chimaeriformes) age estimation

Previous attempts to age chimaeroids have not rigorously tested assumptions of dorsal fin spine growth dynamics. Here, novel imaging and data-analysis techniques revealed that the dorsal fin spine of the spotted ratfish Hydrolagus colliei is an unreliable structure for age estimation. Variation among individuals in the relationship between spine width and distance from the spine tip indicated that the technique of transverse sectioning may impart imprecision and bias to age estimates. The number of growth-band pairs observed by light microscopy in the inner dentine layer was not a good predictor of body size. Mineral density gradients, indicative of growth zones, were absent in the dorsal fin spine of H. colliei , decreasing the likelihood that the bands observed by light microscopy represent a record of growth with consistent periodicity. These results indicate that the hypothesis of aseasonal growth remains plausible and it should not be assumed that chimaeroid age is quantifiable by standard techniques.     

Zoogeography and relationships of Australasian skates (Chondrichthyes: Rajidae)

Aim To investigate distributional patterns and derivation of skates in the Australasian realm. Location Australasia. Methods Genus‐group skate taxa were defined for this region for the first time and new systematic information, as well as bathymetric and geographical data, used to identify distribution patterns. Results The extant skate fauna of Australasia (Australia, New Zealand, New Caledonia and adjacent subAntarctic dependencies) is highly diverse and endemic with sixty‐two species from twelve currently recognized, nominal genus‐group taxa. These include the hardnose skate (rajin) groups Anacanthobatis, Amblyraja, Dipturus, Okamejei, Rajella and Leucoraja, and softnose skate (arhynchobatin) genera Arhynchobatis, Bathyraja, Insentiraja, Irolita, Pavoraja and Notoraja. Additional new and currently unrecognized nominal taxa of both specific and supraspecific ranks also occur in the region. The subfamily Arhynchobatinae is particularly speciose in Australasia, and the New Zealand/New Caledonian fauna is dominated by undescribed supraspecific taxa and species. The Australian fauna, although well represented by arhynchobatins, is dominated by Dipturus‐like skates and shows little overlap in species composition with the fauna of New Zealand and New Caledonia. Similarly, these faunas exhibit no overlap with the polar faunas of the Australian subAntarctic dependencies (Heard and Macdonald Islands) to the south. Skates appear to be absent from the Macquarie Ridge at the southern margin of the New Zealand Plateau. Their absence off New Guinea probably reflects inadequate sampling and the subsequent poor knowledge of that region's deepwater fish fauna. Main conclusions Skates appear to have existed in the eastern, Australasian sector of Gondwana before fragmentation in the late Cretaceous. The extant fauna appears to be derived from elements of Gondwanan origin, dispersal from the eastern and western Tethys Sea, and intraregional vicariance speciation.     

Biodiversity and systematics of skates (Chondrichthyes: Rajiformes: Rajoidei)

Skates (Rajiformes: Rajoidei) are a highly diverse fish group, comprising more valid species than any other group of cartilaginous fishes. The high degree of endemism exhibited by the skates is somewhat enigmatic given their relatively conserved body morphology and apparent restrictive habitat, e.g. soft bottom substrates. Skates are primarily marine benthic dwellers found from the intertidal down to depths in excess of 3,000 m. They are most diverse at higher latitudes and in deepwater, but are replaced in shallower, warm temperate to tropical waters by stingrays (Myliobatodei). The number of valid skate species has increased exponentially, with more species having been described since 1950 (n = 126) than had been described in the previous 200 years (n = 119). Much of the renaissance in skate systematics has largely been through the efforts of a few individuals who through author–coauthor collaboration have accounted for 78 of the 131 species described since 1948 and for nine of 13 genera named since 1950. Furthermore, detailed regional surveys and accounts of skate biodiversity have also contributed to a better understanding of the diversity of the skates. A checklist of the living valid skate species is presented.     

The cardiac ultrastructure of Chimaera monstrosa L. (Elasmobranchii: Holocephali)

Summary The ultrastructure of the heart in Chimaera monstrosa L. is described. The endocardial and the epicardial cells are similar in the three cardiac regions. Myocardial cells show small variations.The myofibre, 4–6 m thick, contains one or a few myofibrils. Each myosin filament is surrounded by six actin filaments. The sarcomere banding pattern includes the Z-, A-, I-, M-, N-, and H-band. End-to-end attachments between myofibres are composed of alternating desmosomes and fasciae adhaerentes. Desmosomes and nexuses occur between longitudinally oriented cell surfaces. The sarcoplasmic reticulum is poorly developed but well defined. Peripheral coupling-like structures are common, T-tubules are absent. Membrane bound dense bodies occur in all regions. Areas with ribosomes and single myosin filaments are often seen.The epicardial cells have a regular hexagonal surface and are much thicker than the endocardial cells. Numerous short and a few longer cytoplasmic extensions face the pericardial cavity.The fiat endocardial cells contain a large nucleus and small amounts of cytoplasm.     

Body plan convergence in the evolution of skates and rays (Chondrichthyes: Batoidea)

Skates, rays and allies (Batoidea) comprise more than half of the species diversity and much of the morphological disparity among chondrichthyan fishes, the sister group to all other jawed vertebrates. While batoids are morphologically well characterized and have an excellent fossil record, there is currently no consensus on the interrelationships of family-level taxa. Here we construct a resolved, robust and time-calibrated batoid phylogeny using mitochondrial genomes, nuclear genes, and fossils, sampling densely across taxa. Data partitioning schemes, biases in the sequence data, and the relative informativeness of each fossil are explored. The molecular phylogeny is largely congruent with morphology crownward in the tree, but the branching orders of major batoid groups are mostly novel. Body plan convergence appears to be widespread in batoids. A depressed, rounded pectoral disk supported to the snout tip by fin radials, common to skates and stingrays, is indicated to have been derived independently by each group, while the long, spiny rostrum of sawfishes similarly appears to be convergent with that of sawsharks, which are not batoids. The major extant batoid lineages are inferred to have arisen relatively rapidly from the Late Triassic into the Jurassic, with long stems followed by subsequent radiations in each group around the Cretaceous/Tertiary boundary. The fossil record indicates that batoids were affected with disproportionate severity by the end-Cretaceous extinction event.