Dermal Denticles Research Articles

Revision of Eocene electric rays (Torpediniformes, Batomorphii) from the Bolca Konservat-Lagerstätte, Italy, reveals the first fossil embryo in situ in marine batoids and provides new insights into the origin of trophic novelties in coral reef fishes

The Eocene electric ray †Titanonarke Carvalho, 2010 from the Bolca Konservat-Lagerstätte, north-eastern Italy, is redescribed in detail based upon new material from recent excavations. This taxon exhibits a combination of features (large voids between the pectoral and the axial skeleton filled in life by electric organs, anteriorly directed fan-shaped antorbital cartilages, lack of dermal denticles, long prepelvic processes, and rounded basibranchial copula with a small caudal tab) that clearly supports its assignment to the order Torpediniformes. The analysis of new material also demonstrates that the previous apparent absence of typical narcinoid characters used to diagnose †Titanonarke was the result of taphonomic biases. †Titanonarke shares at least three synapomorphies (presence of a rostral fontanelle, low number of ribs, and rostral cartilage connected to the antorbital cartilage through lateral appendices) with the extant genera Benthobatis, Diplobatis, Discopyge and Narcine, with which it forms a clade (family Narcinidae) recognized herein as unquestionably monophyletic. Moreover, based upon a single specimen of †Titanonarke that exhibits a unique combination of morphometric and meristic features, a new species of Eocene numbfish, †T. megapterygia sp. nov., is recognized. The presence of several specimens representing different ontogenetic stages of at least two species of numbfishes suggests a close association of this taxon with shallow-water habitats corresponding to coral reefs as hypothesized for the Monte Postale palaeoenvironment. The occurrence of a fossilized marine batoid embryo is reported here for the first time. Moreover, the analysis of the gut contents suggests that the dietary adaptations of †Titanonarke can be related, at least in part, to an opportunistic strategy in the context of abundant larger foraminifera in the Monte Postale palaeobiotope, suggesting that this kind of feeding mode, known to occur in present-day reefs, already was realized 50 million years ago.http://zoobank.org/urn:lsid:zoobank.org:pub:EDD6E170-CA64-4FFB-8DD1-AED2D61D5504

Squalus bassi sp. nov., a new long-snouted spurdog (Chondrichthyes: Squaliformes: Squalidae) from the Agulhas Bank.

The long-snouted African spurdog Squalus bassi sp. nov. is described based on material collected from the outer shelf and upper continental slope off South Africa and Mozambique. Squalus bassi shares with S. mitsukurii, S. montalbani, S. chloroculus, S. grahami, S. griffini, S. edmundsi, S. quasimodo and S. lobularis a large snout with prenarial length greater than distance between nostrils and upper labial furrows, dermal denticles tricuspidate and rhomboid and elevated number of vertebrae. Squalus bassi can be distinguished from all its congeners by a combination of body and fin colouration, external morphometrics, vertebral counts and shape of dermal denticles. Similar long-snouted congeners from the Indo-Pacific region, including S. montalbani, S. edmundsi and S. lalannei are compared in detail with the new species. This new species has been misidentified as the Japanese S. mitsukurii and the Mediterranean S. blainvillei due to the lack of comparative morphological analyses. The validity of the nominal species S. mitsukurii in the south-eastern Atlantic Ocean and western Indian Ocean is also clarified herein, indicating it has a more restricted geographical distribution in the North Pacific Ocean.

Systematic revision of the Potamotrygon scobina Garman, 1913 species-complex (Chondrichthyes: Myliobatiformes: Potamotrygonidae), with the description of three new freshwater stingray species from Brazil and comments on their distribution and biogeography

Potamotrygon scobina Garman, 1913, described from Cametá, rio Tocantins, lower Amazon basin, is taxonomically reviewed through an extensive morphological study of internal and external features. Specimens identified as P. scobina from different locations of the Amazon basin were also studied. The identity of P. scobina, its morphology, and distribution are redefined. In addition, three new species closely related to P. scobina are described from Brazil: Potamotrygon adamastor, sp. nov., from rio Uraricoera, rio Branco system, upper Amazon basin; Potamotrygon amazona, sp. nov., from rio Juruá, upper Amazon basin; and Potamotrygon garmani, sp. nov., from the mid to upper rio Tocantins. These three new species do not occur sympatrically with P. scobina, as far as known. Together with P. scobina and P. limai Fontenelle, Silva & Carvalho, 2014, the new species have three angular cartilages of different sizes, a condition not seen in any other potamotrygonin (which have either one or two angulars). Diagnostic characters in combination are primarily morphometric proportions, especially tail length and width, dermal denticles and tail thorns (their relative size, number, morphology, and distribution), tooth size and number of rows, and color pattern. Printed copy of this book is available from US$24.6 plus postage, Click here to order.

A new chondrichthyan fauna from the Late Jurassic of the Swiss Jura (Kimmeridgian) dominated by hybodonts, chimaeroids and guitarfishes

AbstractThe fossil record of chondrichthyans (sharks, rays and chimaeroids) principally consists of isolated teeth, spines and dermal denticles, their cartilaginous skeleton being rarely preserved. Several Late Jurassic chondrichthyan assemblages have been studied in Europe based on large bulk samples, mainly in England, France, Germany and Spain. The first study of this kind in Switzerland is based on controlled excavations in Kimmeridgian deposits related to the construction of the A16 motorway in the Swiss Jura (Porrentruy, NW Switzerland). This study is based on more than 2000 isolated chondrichthyan remains (teeth, dental plates, spines and dermal denticles) and adds to our knowledge of the chondrichthyan distribution at a regional scale in Europe. We describe and identify this new fauna, define a new species of hybodont with crushing‐type dentition (Asteracanthus udulfensis sp. nov.) and report for the first time the carcharhiniform Corysodon cirinensis in Switzerland. By the Late Jurassic, modern neoselachian sharks had overtaken hybodonts in European marine realms, the latter being gradually confined to brackish or freshwater environments. However, while the associated fauna of the Porrentruy platform indicates marine conditions, neoselachian sharks are surprisingly rare. The chondrichthyan assemblage is largely dominated by hybodonts, guitarfishes (rays) and chimaeroids that are all known to be euryhaline. This unexpected chondrichthyan faunal composition questions the presence of fresh to brackish water in the vicinity of the platform, and the occurrence of salinity fluctuations within a general context marine. This could explain the scarcity of neoselachian sharks and the extended success of hybodonts in the Porrentruy area as late as the Late Jurassic.

The skin microbiome of the common thresher shark (Alopias vulpinus) has low taxonomic and gene function β-diversity.

The health of sharks, like all organisms, is linked to their microbiome. At the skin interface, sharks have dermal denticles that protrude above the mucus, which may affect the types of microbes that occur here. We characterized the microbiome from the skin of the common thresher shark (Alopias vulpinus) to investigate the structure and composition of the skin microbiome. On average 618812 (80.9% ± S.D. 0.44%) reads per metagenomic library contained open reading frames; of those, between 7.6% and 12.8% matched known protein sequences. Genera distinguishing the A. vulpinus microbiome from the water column included, Pseudoalteromonas (12.8% ± 4.7 of sequences), Erythrobacter (5. 3% ± 0.5) and Idiomarina (4.2% ± 1.2) and distinguishing gene pathways included, cobalt, zinc and cadmium resistance (2.2% ± 0.1); iron acquisition (1.2% ± 0.1) and ton/tol transport (1.3% ± 0.08). Taxonomic community overlap (100 - dissimilarity index) was greater in the skin microbiome (77.6), relative to the water column microbiome (70.6) and a reference host-associated microbiome (algae: 71.5). We conclude the A. vulpinus skin microbiome is influenced by filtering processes, including biochemical and biophysical components of the shark skin and result in a structured microbiome.

The shark-beds of the Eyam Limestone Formation (Lower Carboniferous, Viséan) of Steeplehouse Quarry, Wirksworth, Derbyshire, UK

The Eyam Limestone Formation of Steeplehouse Quarry, Wirksworth, Derbyshire, UK yields a diverse assemblage of Lower Carboniferous vertebrate remains. The assemblage is dominated by dermal denticles of the enigmatic selachian Petrodus patelliformis M’Coy, 1848, but also contains teeth of petalodonts, hybodonts and neoselachians. Actinopterygian remains also occur. The assemblage has yielded the earliest Neoselachian, Cooleyella fordi (Duffin and Ward, 1983) and the earliest British lonchidiid, Reesodus wirksworthensis (Duffin 1985). The first occurrence of the enigmatic spiny shark Acanthorhachis (Listracanthidae) is reported from the Viséan, extending its range back some 10 million years. Associated invertebrate remains and sedimentological data indicates a thriving fore-reef environment, deposited in a low energy off-reef setting. The vertebrate remains are well preserved with little abrasion, indicating short transport distances. Conodont elements indicating a late Brigantian age (Early Carboniferous, Viséan) have unusual and extensive euhedral apatite overgrowths.

Bythaelurus vivaldii, a new deep-water catshark (Carcharhiniformes, Scyliorhinidae) from the northwestern Indian Ocean off Somalia.

A new very small deep-water catshark, Bythaelurus vivaldii, is described based on two female specimens caught off Somalia in the northwestern Indian Ocean during the German 'Valdivia' expedition in 1899. It is morphologically closest to the recently described B. bachi, which is the only other Bythaelurus species in the western Indian Ocean that shares a stout body of large specimens and the presence of oral papillae. It further resembles B. vivaldii in the broad mouth and broad posterior head, but differs in the presence of composite oral papillae and a higher diversity in dermal denticle morphology. Additionally, the new species differs from all congeners in the western Indian Ocean in a larger pre-second dorsal fin length, a longer head, a larger interdorsal space, a larger intergill length, a longer pectoral-fin posterior margin, a shorter caudal fin, an intermediate caudal fin preventral margin, and a larger internarial width. Furthermore, the second dorsal fin of the new species is smaller than in its congeners in the western Indian Ocean except for B. lutarius, which is easily distinguished by the slender body and virtual absence of oral papillae, as well as the aforementioned further characters. An updated key to all valid species of Bythaelurus is provided.

Developing an ancient epithelial appendage: FGF signalling regulates early tail denticle formation in sharks

BackgroundVertebrate epithelial appendages constitute a diverse group of organs that includes integumentary structures such as reptilian scales, avian feathers and mammalian hair. Recent studies have provided new evidence for the homology of integumentary organ development throughout amniotes, despite their disparate final morphologies. These structures develop from conserved molecular signalling centres, known as epithelial placodes. It is not yet certain whether this homology extends beyond the integumentary organs of amniotes, as there is a lack of knowledge regarding their development in basal vertebrates. As the ancient sister lineage of bony vertebrates, extant chondrichthyans are well suited to testing the phylogenetic depth of this homology. Elasmobranchs (sharks, skates and rays) possess hard, mineralised epithelial appendages called odontodes, which include teeth and dermal denticles (placoid scales). Odontodes constitute some of the oldest known vertebrate integumentary appendages, predating the origin of gnathostomes. Here, we used an emerging model shark (Scyliorhinus canicula) to test the hypothesis that denticles are homologous to other placode-derived amniote integumentary organs. To examine the conservation of putative gene regulatory network (GRN) member function, we undertook small molecule inhibition of fibroblast growth factor (FGF) signalling during caudal denticle formation.ResultsWe show that during early caudal denticle morphogenesis, the shark expresses homologues of conserved developmental gene families, known to comprise a core GRN for early placode morphogenesis in amniotes. This includes conserved expression of FGFs, sonic hedgehog (shh) and bone morphogenetic protein 4 (bmp4). Additionally, we reveal that denticle placodes possess columnar epithelial cells with a reduced rate of proliferation, a conserved characteristic of amniote skin appendage development. Small molecule inhibition of FGF signalling revealed placode development is FGF dependent, and inhibiting FGF activity resulted in downregulation of shh and bmp4 expression, consistent with the expectation from comparison to the amniote integumentary appendage GRN.ConclusionOverall, these findings suggest the core GRN for building vertebrate integumentary epithelial appendages has been highly conserved over 450 million years. This provides evidence for the continuous, historical homology of epithelial appendage placodes throughout jawed vertebrates, from sharks to mammals. Epithelial placodes constitute the shared foundation upon which diverse vertebrate integumentary organs have evolved.

Resolving the ambiguities in the identification of two smooth-hound sharks (Mustelus mustelus and Mustelus punctulatus) using genetics and morphology

Elasmobranch species are among the most threatened marine fish, and sound biological data for many of them are lacking. In this context, the smooth-hounds (Mustelus spp., Triakidae, Carcharhiniformes) represent an intriguing genus, being characterized by unclear and sometimes contrasting diagnostic traits. Here, we present new insights into the identification of two closely related species, M. mustelus and M. punctulatus. The use of three different molecular markers on a dataset of 588 specimens highlighted a genetic distinction between the two species. However, whereas microsatellites provided unambiguous results in all the samples, the other two markers were not able to assign a fraction of the individuals (6.6% for cytochrome c oxidase subunit 1, 14.4% for Internal Transcribed Spacer 2), suggesting the occurrence of heteroplasmy and introgression due to past hybridisation events. The comparison between morphological traits, previously suggested as diagnostic for species identification, and genetic identification, enabled validation of the most reliable and practical morphological traits for species identification. The shape of the dermal denticles represents the most reliable trait, but its use during fieldwork may be impractical. The black spots on body sides were present almost exclusively in M. punctulatus, allowing its identification; however, their absence is not diagnostic for M. mustelus. The distance of the nostrils and the shape of the mouth are useful to distinguish the two species during fieldwork. Our results will allow an accurate identification of the two species leading to the collection of reliable data on their distribution, life history traits, exploitation and status of conservation.

Two new species of short-snouted dogfish sharks of the genus Squalus Linnaeus, 1758, from southern Africa (Chondrichthyes: Squaliformes: Squalidae)

Species composition of the genus Squalus from the eastern Atlantic and western Indian oceans is still poorly known, with three to four species often recognized: S. acanthias, S. megalops, S. blainvillei, and S. mitsukurii. A fifth species, S. acutipinnis, was recently redescribed as valid in the region, although morphological variation was noted among its type specimens that indicated the presence of two different forms. The second form is described herein as Squalus margaretsmithae sp. nov. and distinguished from S. acutipinnis based on external morphological characters. Squalus mahia sp. nov. is also described from southern Africa based mostly on material collected from Madagascar. Squalus mahia sp. nov. stands out from other Squalus species by having a conspicuously slender body, elongate and thin dorsal fin spines, narrow caudal fin, and other external proportions. These two species are often misidentified with the Australian S. megalops because they share a short snout, white postventral caudal margins, lanceolate and unicuspid dermal denticles, and a low number of vertebrae. Comparative analyses are provided here between Squalus margaretsmithae sp. nov., Squalus mahia sp. nov., and morphologically similar species that also share these features, including S. brevirostris, S. crassispinus, S. acutipinnis, S. altipinnis, and S. hemipinnis. A taxonomic evaluation of the nominal species S. megalops and its occurrence in the region is also discussed.

Microscopic aspects of the nictitating membrane in Carcharhinidae and Sphyrnidae sharks: a preliminary study

The nictitating membrane is an anatomic structure exclusively exhibited by Carcharhiniformes, the largest order among sharks. Here we present a detailed description of morphological characteristics of the nictitating membrane through light microscopy (LM) and scanning electron microscopy (SEM) in the following shark species: Carcharhinus limbatus, Galeocerdo cuvier, Prionace glauca, Rhizoprionodon lalandii, R. porosus, Sphyrna lewini and S. zygaena. Differences in the microscopic aspects of dermal denticles from the species studied were observed. P. glauca, a pelagic shark, showed a well-developed protection apparatus when compared with other pelagic species, while coastal sharks showed even higher structural complexity. In the blue shark the denticles are enameled, presenting an extensive pulp cavity and a base inserted in a connective tissue. Moreover, the species exhibits the higher number of ridges (up to nine) of varied size and shape and the muscular tissue is inserted in the ventral region of the connective tissue. Dermal denticles from C. limbatus, R. lalandii, R. porosus, S. zygaena and G. cuvier exhibit up to five ridges with hexagonal ornamentations in the crown. In S. lewini and S. zygaena, the denticles are rounded shaped and glandular cells are present. The patterns observed in the present study suggest a high level of specialization and evolutionary conservation shaped by the function of the structure. In addition, we hypothesize that the morphological simplification observed in the membrane when compared to the dermal denticles from the skin, is an evolutionary trait that evolved to improve the dynamic and biomechanics of this highly mobile structure allowing this way, a rapid and efficient protection against abrasion, mainly during predation events.

Etmopterus samadiae n. sp., a new lanternshark (Squaliformes: Etmopteridae) from Papua New Guinea.

A new species of lanternshark, Etmopterus samadiae (Squaliformes: Etmopteridae), is described from off northern Papua New Guinea, in the western Central Pacific Ocean. The new species resembles other members of the "Etmopterus lucifer" clade in having linear rows of dermal denticles and most closely resembles E. brachyurus from the western North Pacific. The new species occurs along insular slopes between 340 and 785 m depth. The new species can be distinguished from other members of the E. lucifer clade by a combination of characteristics, including length of anterior flank branch markings being slightly shorter than its posterior branch, a longer caudal base marking, and irregular and variable number of black, horizontal, dash-like marks on sides of body. Molecular analysis based on the NADH2 marker further supports the distinction of E. samadiae from other members of the E. lucifer clade.

Dermal denticles as a tool to reconstruct shark communities

Dermal denticles as a tool to reconstruct shark communities

Etmopterus lailae sp. nov., a new lanternshark (Squaliformes: Etmopteridae) from the Northwestern Hawaiian Islands.

A new species of lanternshark, Etmopterus lailae (Squaliformes: Etmopteridae), is described from the Northwestern Hawaiian Islands, in the central North Pacific Ocean. The new species resembles other members of the "Etmopterus lucifer" clade in having linear rows of dermal denticles, and most closely resembles E. lucifer from Japan. The new species occurs along insular slopes around seamounts at depths between 314-384 m. It can be distinguished from other members of the E. lucifer clade by a combination of characteristics, including a longer anterior flank marking branch, arrangement of dermal denticles on the ventral snout surface and body, flank and caudal markings, and meristic counts including number of spiral valve turns, and precaudal vertebrate. A key to species of the Etmopterus lucifer-clade is included.

Methods for isolation and quantification of microfossil fish teeth and elasmobranch dermal denticles (ichthyoliths) from marine sediments

Methods for isolation and quantification of microfossil fish teeth and elasmobranch dermal denticles (ichthyoliths) from marine sediments

The oldest record of gnathostome fossils from Greece: Chondrichthyes from the Lopingian of Hydra Island

The Permian deposits of Hydra Island, Greece, have been known for over a century and host some of the best-studied and most diverse invertebrate assemblages of the ancient Paleotethys Ocean. However, until now, no Paleozoic fossils of jawed vertebrates had been reported from Greece. Recent fieldwork on Hydra Island brought to light rare cartilaginous fish remains, including a tooth belonging to an unknown hybodontiform shark, as well as an unidentifiable dermal denticle of an euselachian shark. Despite similarities with iconic Paleozoic and Mesozoic durophagous euselachians, the Hydriot tooth likely corresponds to a new species, but is provisionally left in open nomenclature until more material becomes available. The new chondrichthyan fossils from Hydra Island correspond to one of the few Lopingian (late Permian) occurrences known from the Paleotethys. Moreover, they constitute the oldest record of jawed-vertebrate fossils from Greece, predating younger occurrences by more than 50 million years.

Bythaelurus bachi n. sp., a new deep-water catshark (Carcharhiniformes,Scyliorhinidae) from the southwestern Indian Ocean, with a review of Bythaelurus species and a key to their identification.

A new deep-water catshark, Bythaelurus bachi, is described based on 44 specimens caught on the southern Madagascar Ridge in the southwestern Indian Ocean. The new species is the only stout-bodied Bythaelurus with oral papillae in the region and is distinguished from all congeners by the plain beige to light gray-brown coloration, high diversity in dermal denticle morphology, and presence of composite oral papillae. Despite resemblance in body shape, Bythaelurus bachi n. sp. is distinguished from its closest congener, B. naylori Ebert & Clerkin, 2015, by the presence of numerous large, partially composite papillae on the tongue and roof of the mouth (vs. papillae lacking), plain light coloration (vs. medium to dark brown ground color, light fin edges and a distinctly dark dusky-colored snout), only slightly enlarged dermal denticles on the anterior upper caudal-fin margin (vs. dermal denticles distinctly enlarged), a higher diversity in dermal denticle morphology in general, and smaller maximum size and size at maturity. The distinction of both species is also supported by molecular results. The new species differs from all other congeners in the western Indian Ocean in the stout body shape of large specimens, coloration, larger size, as well as several morphometrics, including larger claspers, longer eyes and dorsal fins, and shorter pelvic-anal and pelvic-caudal spaces. The genus is reviewed, a key to its species given.

Thorns and dermal denticles of skates Atlantoraja cyclophora and A. castelnaui: Microscopic features and functional implications.

Some batoid species are covered with dermal denticles (or placoid scales) that occasionally develop into thorns. In sexually mature males, sharp teeth and alar thorns found on the apex of the lateral disc are used to hold the female during copulation. This study set out to analyze microscopic features of modified dermal denticles and thorns and to investigate sexual dimorphism in Atlantoraja cyclophora and A. castelnaui species. Skin samples collected from areas covered with thorns were fixed in 10% formaldehyde, processed and analyzed using scanning electron microscopy. Alar thorn morphology varied within species, while caudal thorn, rostral and caudal dermal denticle morphology varied within and between species. These structures play an important role in the protection and reproduction of the species studied and constitute important taxonomic information, given they are often the only elements preserved in archaeological sites and fossil records.

Assessing the fidelity of marine vertebrate microfossil δ18O signatures and their potential for palaeo-ecological and -climatic reconstructions

Conodont biogenic apatite has become a preferred analytical target for oxygen isotope studies investigating ocean temperature and palaeoclimate changes in the Palaeozoic. Despite the growing application in geochemically-based palaeoenvironmental reconstructions, the paucity or absence of conodont fossils in certain facies necessitates greater flexibility in selection of robust oxygen-bearing compounds for analysis. Vertebrate microfossils (teeth, dermal denticles, spines) offer a potential substitute for conodonts from the middle Palaeozoic. Vertebrate bioapatite is particularly advantageous given a fossil record extending to the present with representatives across freshwater to fully marine environments, thus widening the scope of oxygen isotope studies on bioapatite. However, significant tissue heterogeneity within vertebrates and differential susceptibility of these tissues to diagenetic alteration have been raised as potential problems affecting the reliability of the oxygen isotope ratios as palaeoclimatic proxies. Well-preserved vertebrate microfossils and co-occurring conodont fossils from the Upper Devonian and Lower Carboniferous of the Lennard Shelf, Canning Basin, Western Australia, were analysed using bulk (gas isotope ratio mass spectrometry, GIRMS) and in-situ (secondary ion mass spectrometry, SIMS) methodologies, with the latter technique allowing investigation of specific tissues within vertebrate elements. The δ18Oconodont results may be interpreted in terms of palaeolatitudinally and environmentally sensible palaeo-salinity and -temperature and provide a baseline standard for comparison against vertebrate microfossil δ18O values. Despite an absence of obvious diagenetic modification, GIRMS of vertebrate denticles yielded δ18O values depleted in 18O by 2–4‰ relative to co-occurring conodonts. SIMS analysis of dentine tissues exhibited significant heterogeneity, while hypermineralised tissues in both scales and teeth produced δ18O values comparable with those of associated conodonts. The susceptibility of permeable phosphatic fossil tissues to microbial activity, fluid interaction and introduction of mineral precipitates post-formation is demonstrated in the dentine of vertebrate microfossils, which showed significant heterogeneity and consistent depletion in 18O relative to conodonts. The hypermineralised tissues present in both teeth and scales appear resistant to many diagenetic processes and indicate potential for palaeoclimatic reconstructions and palaeoecological investigations.

A new subfamily, Styracurinae, and new genus, Styracura, for Himanturaschmardae (Werner, 1904) and Himantura pacifica (Beebe & Tee-Van, 1941) (Chondrichthyes: Myliobatiformes).

A new subfamily (Styracurinae, subfam. nov.) and genus (Styracura, gen. nov.) are erected for Trygon schmardae Werner, 1904 (type species) and Dasyatis pacificus Beebe & Tee-Van, 1941, species placed in Himantura Müller & Henle, 1837 since 1953 but which differ significantly from it and all dasyatid genera in many morphological features. Recent molecular phylogenetic hypotheses based on the entire protein-coding mitochondrial genome have recovered Styracura as the sister group of the Neotropical freshwater stingrays (Potamotrygonidae), agreeing with previous morphological and molecular phylogenies. The Styracurinae, subfam. nov., is therefore placed in the Potamotrygonidae to reflect the current phylogenetic view. Morphological characters of phylogenetic relevance of the dermal denticles, ventral lateral-line canals, hyoid arch, cranial musculature, and pectoral and pelvic girdles are discussed and compared to other stingrays, in particular to dasyatids and potamotrygonins (now ranked as a subfamily). Inferred derived characters of Styracura and potamotrygonins require homoplasy (they are absent from Paratrygon and Heliotrygon), and ventral lateral-line canal patterns and morphology of the scapular process and pelvic girdle generally reveal an affinity to different dasyatid genera, but one most likely based on plesiomorphies or independent evolution; Styracura is considered more closely related to potamotrygonins.