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Bone is a defining characteristic of the vertebrate skeleton, and while chondrichthyans (sharks, skates, and other cartilaginous fishes) are vertebrates, they are hypothesized to have ...lost the ability to make bone during their evolution. Multiple descriptions of a bone-like tissue in neural arches of vertebrae in various shark species (selachians), however, challenge this hypothesis. Here, we extend this argument by analyzing vertebrae of two members of the batoids (the little skate Leucoraja erinacea and Eaton’s skate Bathyraja eatonii), the sister group to selachians within elasmobranchs. Micro-CT images showed a bone-like mineralization pattern in neural arches of each skate species, and histological analyses confirmed that this bone-like tissue surrounded a cartilage core, exactly as described in sharks. Another mineralization pattern identified in skate vertebrae was distinct from the polygonal tesseral and areolar patterns that classically are associated with the chondrichthyan endoskeleton. Many regions of the vertebrae, including the neural spine and transverse processes, showed this perichondral mineralization pattern, termed here trabecular tesseral. Other than the cartilage core of the neural arch, all mineralized tissues in skate vertebrae had flattened cells surrounded by matrix with bone-like histology. Analyses of quantitative microstructural parameters revealed that, compared to rat vertebrae, the bone-like mineralization pattern in the neural arches of skate vertebrae was more similar to compact bone than trabecular bone. In contrast, the thickness of the trabecular tesseral pattern was more similar to trabecular bone than compact bone of rat vertebrae. In conclusion, a bone-like tissue in neural arches of skate vertebrae appears to be a novel elasmobranch synapomorphy. We propose that the trabecular tesseral mineralization pattern in the skate might have deep homology to the mineralization pattern utilized in trabecular bone.
Mineralization patterns of skeletal tissues have not been investigated thoroughly in all vertebrate clades. Despite their designation as ‘cartilaginous fish’, chondrichthyans clearly evolved from ancestral vertebrates that made bone. The consensus that chondrichthyans lost the ability to make bone during their evolution, however, is challenged by reports of bone and bone-like tissues in the neural arches of vertebrae in extant sharks (selachians). Here, we provide evidence from micro-CT imaging and histological analyses to support our hypothesis that a bone-like tissue is present in the neural arches of batoids (the sister group to selachians within elasmobranchs). These results argue strongly that the neural arch bone-like tissue is a previously unknown synapomorphy of elasmobranchs. In addition to the bone-like mineralization pattern identified in the neural arches, micro-CT images also showed a novel mineralization pattern which we described as trabecular tesseral. Quantitative microstructural features shared between trabecular tesseral pattern and trabecular bone (from homologous rat vertebrae) suggest that both patterns might derive from an ancestral gene network driving trabecular mineralization (i.e., deep homology).
Overexploitation of marine communities can lead to modifications in the structure of the food web and can force organisms like elasmobranchs to change their feeding habits. To evaluate the impact ...that fisheries have on food webs and on the interactions between species, it is necessary to describe and quantify the diet of the species involved and follow it through time. This study compares the diet of five skate species using the data obtained from the by‐catch of the Argentine hake (Merluccius hubbsi) fishery in north and central Patagonia, Argentina. Diet composition was assessed by analysing the digestive tract contents and trophic overlapping between species of the genus Bathyraja: Bathyraja albomaculata, Bathyraja brachyurops, Bathyraja macloviana, Bathyraja magellanica and Bathyraja multispinis. A total of 184 stomachs were analysed. The diets of B. albomaculata and B. macloviana mainly comprised annelids, whereas that of B. brachyurops primarily comprised fish, including hake heads discarded by the fishery. The diets of B. magellanica and B. multispinis were largely based on crustaceans. Despite the morphological similarities and their shared preference for benthic habitats, no complete diet overlaps were found between the different species. These results suggest that these skate species have undergone a process of diet specialisation. This is a common feeding strategy that occurs to successfully eliminate competition when resources are limited, which corresponds to the conditions found in an environment being affected by the pressures of overfishing.
A new species of Potamotrygon is described from the Rio Negro drainage, Amazonas, Brazil. In spite of being cited or pictured several times in the scientific and aquarium fish literature since the ...19th Century, it had been misidentified and still lacked a scientific name. Potamotrygon wallacei, n. sp., is diagnosed by the following characters: dorsal surface of disc light brown, with black irregularly-shaped vermiculate markings forming an amphora- or Ω-shaped figure on mid-disc, delimiting light brown reniform areas at disc center, and with subcircular light brown ocellate markings on disc margins; small body size (smallest known Potamotrygon species; largest examined specimen measured 310 mm DW); dorsal spines on tail usually rather low, without broad bases, in one to rarely three irregular rows, but extending posteriorly only to tail mid-length and not to caudal stings, with altogether relatively few spines; denticles on posterior mid-disc and tail base Y-shaped, with a central, anterior, bulbous cusp and usually two posterior pairs of smaller, rounded cusps; and single (anterior) angular cartilage. The new species is similar to P. orbignyi and other "reticulated" species in having a single (anterior) angular cartilage and in the color pattern of the tail, but is easily distinguished based on its size, dorsal tail spine arrangement, and specific details of color pattern.
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.
In viviparous (live-bearing) animals, embryos face an embryo-specific defecation issue: faecal elimination in utero can cause fatal contamination of the embryonic environment. Our data from the ...viviparous red stingray (Hemitrygon akajei) reveals how viviparous elasmobranchs circumvent this issue. The exit of the embryonic intestine is maintained closed until close to birth, which allows the accumulation of faeces in the embryonic body. Faecal accumulation abilities are increased by (1) the large intestine size (represents about 400-600% of an adult intestine, proportionally), and (2) the modification in the intestinal inner wall structure, specialized to increase water uptake from the faecal matter. According to the literature, faecal accumulation may occur in embryos of the lamniform white shark as well. The reproductive biology of myliobatiform stingrays and lamniform sharks is characterized by the onset of oral feeding before birth (i.e. drinking of uterine milk and eating of sibling eggs, respectively), which is expected to result in the production of large amounts of faeces during gestation. The strong ability of faecal accumulation in these lineages is therefore likely an adaptation to their unique embryonic nutrition mechanism.
The application of a photographic identification methodology using the unique ventral surface markings (natural spot patterns) of an observed population in southern Mozambique enabled many aspects of ...the reproductive ecology of reef manta rays Manta alfredi to be examined. The region encompassing the study site was identified as a mating ground for M. alfredi based on observations of mating events and fresh mating scars on females. The distribution of these pectoral fin scars was highly biased and indicated a strong lateralized behavioural trait, with 99% of these scars occurring only on the left pectoral fin. No other elasmobranch has been reported to display behavioural lateralization. The study region also acts as a birthing ground, with individuals typically giving birth in the austral summer period after a gestation of c. 1 year. Reproductive periodicity in M. alfredi was most commonly biennial, but a few individuals were pregnant in consecutive years, confirming an annual ovulatory cycle. The production of a single pup appears to be the normal situation, although observations in the wild as well as during opportunistic dissections of individuals killed by fisheries revealed that two pups are conceived on occasion. Many aspects of the study have contributed to the limited baseline data currently available for this species and have highlighted the potential need for more conservative conservation strategies.
Claspers of stingrays (Myliobatiformes) are poorly documented in comparison to claspers of skates, with our knowledge restricted mainly to external morphological characters and skeletal components ...included in descriptions of new species; more encompassing morphological comparative analyses are lacking. Concerning potamotrygonins, clasper morphology has been described for a handful of species, but without elucidating their variation and systematic potential. The present article analyzed clasper structures in all genera of the subfamily Potamotrygoninae (Potamotrygon, Paratrygon, Plesiotrygon, and Heliotrygon), which were compared to the clasper of Styracura and some dasyatid genera. Potamotrygon shows some morphometric variation among the species analyzed. Anatomically, we found variation mainly in the dorsal terminal 1 and accessory terminal 1 cartilages, which are considered diagnostic among potamotrygonin genera; external morphological structures did not present significant differences among potamotrygonins.
External clasper morphology of Heliotrygon rosai (MZUSP 104996).
Atlantoraja platana is an oviparous skate endemic to the south‐west Atlantic Ocean, and is one of the skate species most exploited by local industrial bottom trawl fisheries. Oviparous elasmobranchs ...encapsulate their eggs in complex egg cases produced by the oviductal gland (OG). This organ is exclusively present in these fishes and comprises four distinct zones: club, baffle, papillary and terminal. The relative size and structural complexity of these zones correlate with mode of reproduction. Glycans are known to play major roles in reproduction so their distribution in each zone of the OG could explain the functional multiplicity of the gland in skates, but this topic has not been previously investigated. In this study, morphological, histochemical and lectin‐histochemical analysis revealed various novel aspects of A. platana's OG. The club, papillary and terminal zones positively stained for periodic acid Schiff's reagent (PAS) and Alcian Blue (AB), indicating the presence of neutral and acid mucopolysaccharides. However, the buffle zone was negative for PAS and AB stains, but was positive for all the lectins used. Each zone of the OG had a characteristic pattern of glycan expression. Finally, we confirmed the presence of sperm but not sperm storage. This is the first lectin‐histochemical study of the OG in chondrichtyan fish and it has proven to be an important tool to understand some of the mechanisms of fertility and reproductive success in economic important species such as A. platana.
Encheliophis chardewalli was described from a single cleared and stained specimen. Twelve years later, additional specimens were found in the lagoon of Moorea (French Polynesia) in association with ...their host, the sea cucumber Actinopyga mauritiana. These fish were used to consolidate the species diagnosis, to validate species status and to record sound production. This species is remarkable because of its ability to penetrate inside the cloaca of sea cucumbers having anal teeth and the fact this species is largely unknown despite it lives in lagoons in 1m depth. Encheliophis chardewalli produced three sound types: long regular calls made of trains of numerous pulses, short irregular calls characterized by a constant lowering of its pulse period and short regular call (or knock) made of 3 to 6 pulses. Comparison with other sympatric Carapini supports a large and distinct repertoire. Morphological characteristics could be the result of reduced body size allowing to penetrate inside a new host, thus avoiding competition and conflict with other larger sympatric Carapini species.
This study highlights three different types of calls in a cryptic species and provides morphological information related to its way of life. The fish paedomorphic form could be due to requirement related to host distinctive features.
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