Phylogenetic significance of clasper morphology of electric rays (Chondrichthyes: Batoidea: Torpediniformes).

Torpediniformes (electric rays) is a monophyletic group strongly supported by morphological and molecular phylogenetic studies. The claspers of electric rays, however, are poorly documented in comparation to the clasper of other batoids, especially skates, and the knowledge of their anatomical variation is restricted to the description of a few species. The present article analyzes the external and skeletal clasper anatomy of electric rays and reports newly discovered characters that can be useful for taxonomic diagnoses and higher-level systematic studies. The family Torpedinidae exclusively presents the integumental flap, a poorly calcified clasper skeleton, and a dorsal marginal cartilage with a medial flange on its distal portion. Derived or diagnostic characters were not found in the clasper of the reportedly nonmonophyletic families Narcinidae and Narkidae; however, the claspers of species and genera of narcinids and narkids present different anatomical patterns that can be useful for taxonomic and phylogenetic studies.

Descripción y comparación de la línea lateral de tres especies de rayas eléctricas del género Narcine (Torpediniformes: Narcinidae) / Description and comparison of the lateral line of three species of electric rays of the genus Narcine (Torpediniformes: Narcinidae)

Resumen Se presenta la descripción de la composición y arreglo estructural del sistema de la línea lateral de tres especies de rayas eléctricas (Narcine bancroftii, N. entemedor y N. vermiculatus). Las tres especies muestran los canales supraorbital (SO), infraorbital (IO), hiomandibular (HYO) y de la línea lateral posterior (LLP), menos ramificados que en otros batoideos. Narcine entemedor difiere de sus congéneres por la gran ramificación de los canales SO e IO, los cuales convergen en la región inter-orbital; en N. vermiculatus estos canales presentan una condición similar, pero con una menor ramificación. En contraste N. bancroftii presenta los canales divergentes (dirigidos hacia afuera de la región inter-orbital). Los canales HYO están localizados en la periferia de los órganos eléctricos en estas especies; en N. bancroftii las ramificaciones se extienden hacia la región posterior, mientras que en las demás especies están reducidas (N. entemedor) o casi ausentes (N. vermiculatus). La LLP es de forma curva y está dirigida hacia el urostilo, asimismo muestra un gran número de ramificaciones, excepto en N. vermiculatus. La información sobre el arreglo y el número de poros sobre el sistema de canales de la LL en Narcine, puede ser de utilidad para la distinción taxonómica de sus especies, tal como ha sido corroborado para otros Torpediniformes.

Abstract This study describes the composition and structural arrangement of the lateral line system of three electric ray species (Narcine bancroftii, N. entemedor and N. vermiculatus). All three species showed the supraorbital (SO), infraorbital (IO), hyomandibular (HYO) and posterior lateral line (PLL) canals, less branched compared to other batoids. Narcine entemedor differs from its congeners by the extensive branching of the SO and IO canals, which converge in the inter-orbital region; in N. vermiculatus these canals present a similar condition, but are less branched. In contrast, N. bancroftii has divergent canals (directed outwards from the inter-orbital region). HYO canals are located on the periphery of the electric organs in these species; in N. bancroftii the branches extend towards the posterior region, whereas in the other species are reduced (N. entemedor) or almost absent (N. vermiculatus). The PLL is curved and projected towards the urostyle, it also shows a large number of ramifications, except in N. vermiculatus. Information about the LL canal system arrangement and pore number in Narcine can be useful for the taxonomic distinction of its species, as it has been corroborated in other Torpediniformes. Rev. Biol. Trop. 66(2): 586-592. Epub 2018 June 01.

Reassessment and Relationships of Scutatuspinosus itapagipensis (Teleostei, Clupeomorpha, Ellimmichthyiformes) from the Neocomian of Recôncavo Basin, Northeastern Brazil

ABSTRACT Clupeomorphs are found in many assemblages of the northeastern Brazil, whose ages range from the Neocomian to Paleogene. Ten species were described. Among them †Scutatuspinosus itapagipensis, a torpedo-like fish found in shales of the Marfim Formation from Bahia, remains poorly known. At first it was positioned within †Scutatuspinosinae and indicated as closely related to †Diplomystus. Diagnoses for the subfamily and genus were based on certain characters masking relationships. Recently it was placed in †Ellimmichthyiformes, but its systematic position remains controversial. We furnish additional data and restorations together with a parsimony analysis with TNT program, using 60 unordered and unweighted characters from selected species of 13 genera. Our analysis produced 12 trees with 158 steps, a consistency index (CI) of 0.44 and retention index (RI) of 0.52. Low support indices still indicate insufficient data for many taxa and uncertain status for clades. According to the majority rule consensus, †Scutatuspinosus itapagipensis is placed within a "†paraclupeine" group with †Ezkutuberezi carmenae, †Ellimma branneri, †Ellimmichthys longicostatus, and †Paraclupea chetunguensis. Its minimum age and position among †paraclupeids indicate an early history of the group older than it was thought to be, supporting a biogeographical hypothesis based on generalized track from China to northeastern Brazil during the Lower Cretaceous.

Narcine baliensis, a new species of electric ray from southeast Asia (Chondrichthyes: Torpediniformes).

A new species of numbfish, Narcine baliensis, sp. nov., is described from the tropical eastern Indian Ocean from Indonesia. It is superficially similar to N. brevilabiata and N. atzi in aspects of its color pattern, but is distinguished from both congeners in details of its color pattern, in tooth band morphology, and in proportions of its dorsal fins, among other features. Narcine baliensis, sp. nov., is unique in having a dorsal color pattern composed of large, circular, ovoid or elongate dark brown spots or blotches on dorsal disc along with more numerous small (about eye-sized or slightly greater) brownish, subcircular spots, with large blotches and small spots surrounded by a very slender creamy-white pattern, as well as in having broadly circular upper and lower tooth bands of about the same width and shape. The genus Narcine is now composed of 20 valid species, but uncertainty remains concerning the identification and morphological variation of some of its species in the tropical Indo-West Pacific region.

Tetronarce cowleyi, sp. nov., a new species of electric ray from southern Africa (Chondrichthyes: Torpediniformes: Torpedinidae).

A new species of torpedo ray, Tetronarce cowleyi, sp. nov., is described from specimens collected from the southeastern Atlantic Ocean. The new species is placed in the genus Tetronarce based on a uniform dorsal coloration and absence of papillae around the spiracles. The new species is distinguished from its closest congeners, the North Atlantic Tetronarce nobiliana Bonnaparte, 1835, and southwestern Atlantic Tetronarce puelcha Lahille, 1926, by a combination of morphological characteristics including a shorter spiracular length, a proportionally greater head length as measured between snout margin and fifth gill openings, a proportionally greater preoral snout length, a uniform shiny black or dark gray dorsal surface, lacking any prominent markings, and a creamy white ventral color with dark edges in juveniles but fading with growth. Teteronarce cowleyi, sp. nov., is further distinguished from T. nobiliana by its more circular anterior disc shape (vs. relatively straight in T. nobiliana), fewer tooth rows (32/28 vs. 38-53/38-52 in T. nobiliana), greater mouth width (1.5-1.7 times as great as interorbital width vs. 0.5-0.6 times interorbital width in T. nobiliana), smaller distance between second dorsal and caudal fins (3.5-4.9% vs. 6.6-6.8% in T. nobiliana), and a clasper length extending nearly to lower caudal fin origin (claspers in T. nobiliana that extend only two-thirds distance between second dorsal and caudal fins). Teteronarce cowleyi, sp. nov., is known from Walvis Bay, Namibia to Algoa Bay, Eastern Cape, South Africa, at depths of 110 to 457 m.

The impacts of comparative anatomy of electric rays (Batoidea: Torpediniformes) on their systematic hypotheses.

The Comparative anatomy of the 11 recognized genera within Torpediniformes is described, systematically categorized, and illustrated in a comprehensive photo-atlas. Data are compiled into a character matrix and cladistically analyzed using parsimony to test hypotheses about the previously recognized subfamilies, while reconstructing the possible evolutionary history of Torpediniformes. Results are consistent with the previous rank-based classifications, regardless of the parsimony criteria used to generate the phylogenetic hypothesis, with one notable exception: a monophyletic Narcininae was never recovered. Torpedinoidea (=Hypnos + Torpedo) is supported by the presence of long, slender, flexible jaw cartilages, absence of a large rostral fontanelle, presence of suprascapular antimeres that are each shorter than the scapular process of the scapulocoracoid, antorbital cartilages that articulate on the anterior aspect of the nasal capsules and absence of a frontoparietal fontanelle. Subfamilial names Hypninae and Torpedininae are redundant with the genus names Hypnos and Torpedo and are not adopted here. Narcinoidea (=nontorpedinoid torpediniforms) is supported by unambiguous character transformations to the presence of a divided lower lip, labial cartilages, laterolingually compressed palatoquadrates, bifurcated antorbital cartilages, a rostral fontanelle, ventrally projecting nasal capsules, a dorsal rim of the synarcual mouth posterior to occipital condyle, posteriorly positioned lateral stays, and obtuse anterior margins of lateral stays. Narkidae is supported by unambiguous character transformations to the presence of an uncovered eye that protrudes above dorsal surface, a shared rim between the spiracle and the eye, an anterior nasal turret that projects ventrally, a nasal curtain that covers the upper lip and dentition when the mouth is closed, tab-like prepelvic processes, a mesopterygium that is shorter than propterygium but longer than metapterygium, a slender median rostral cartilage, and a basibranchial cartilage with an anterior margin that is depressed medially and a posterior margin that tapers.

Sphingomyelin composition and physical asymmetries in native acetylcholine receptor-rich membranes.

Selective enzymatic hydrolysis, lipid compositional analyses, and fluorescence studies have been carried out on acetylcholine receptor (AChR)-rich membranes from Torpedinidae to investigate the topology of sphingomyelin (SM) in the native membrane and its relationship with the AChR protein. Controlled sphingomyelinase hydrolysis of native membranes showed that SM is predominantly (approximately 60%) localized in the outer half of the lipid bilayer. Differences were also observed in the distribution of SM fatty acid molecular species in the two bilayer leaflets. A fluorescent SM derivative ( N-[10-(1-pyrenyl)decanoyl]sphingomyelin; Py-SM) was used to study protein-lipid interactions in the AChR-rich membrane and in affinity-purified Torpedo AChR reconstituted in liposomes made from Torpedo electrocyte lipid extracts. The efficiency of Förster resonance energy transfer (FRET) from the protein to the pyrenyl-labeled lipid as a function of acceptor surface density was used to estimate distances and topography of the SM derivative relative to the protein. The dynamics of the lipid acyl chains were explored by measuring the thermal dependence of Py-SM excimer formation, sensitive to the fluidity of the membrane. Differences were observed in the concentration dependence of excimer/monomer pyrenyl fluorescence when measured by direct excitation of the probe as against under FRET conditions, indicating differences in the intermolecular collisional frequency of the fluorophores between bulk and protein-vicinal lipid environments, respectively. Py-SM exhibited a moderate selectivity for the protein-vicinal lipid domain, with a calculated relative affinity K(r) approximately 0.55. Upon sphingomyelinase digestion of the membrane, FRET efficiency increased by about 50%, indicating that the resulting pyrenyl-ceramide species have higher affinity for the protein than the parental SM derivative.

Brain variation and phylogenetic trends in elasmobranch fishes.

A cladistic analysis utilizing the out-group criterion to establish the phylogenetic polarity of neural features among living chondrichthyans leads to the conclusion that the earliest chondrichthyans possessed relatively small brains. Relative brain size appears to have increased independently in some myliobatiforms and in the common ancestor of heterodontid and galeomorph sharks. Furthermore, the brains of the earliest chondrichthyans must have exhibited a wide rhomboid fossa of the medulla, an unfoliated cerebellar corpus divided into anterior and posterior lobes, a bilobed and laminated optic tectum with the majority of the cell bodies located periventricularly, a moderately developed midbrain octavolateralis complex, and a telencephalon consisting of sessile olfactory bulbs and paired cerebral hemispheres with extensive lateral ventricles and thin walls. The corpus of the cerebellum has increased dramatically in size and has become complexly foliated independently in some batoids and galeomorph sharks. Similarly, the majority of tectal neurons have migrated away from a periventricular zone in all living elasmobranchs resulting in two distinctly derived patterns, compared to that in holocephalimorphs, but the polarity of these patterns among elasmobranchs is uncertain. The midbrain octavolateralis complex is uniquely derived in all living batoids, but two patterns appear to occur in other chondrichthyans, and, again, their polarity is uncertain. Different portions of the telencephalic roof appear to have hypertrophied in holocephalimorphs, in some batoids, and in some galeomorphs. An examination of pertinent details of chondrichthyan general biology reveals that the biological significance of most chondrichthyan neural trends is unknown.