RESUMO
The mandibular apparatus of batoids (skates, electric rays, guitarfishes, stingrays, and sawfishes) is composed of a few skeletal elements to which the muscular bundles, responsible for all movements involved in the feeding mechanism, are inserted. The description of the different mandibular morphologies can help to understand the different feeding guilds in this group. In this study, we examined the cranio-mandibular myology of adult Rostroraja velezi, Narcine entemedor, and Zapteryx exasperata, three species of rays that coexist in the Southern Gulf of California, Mexico. This study described the muscles on the ventral and the dorsal surfaces for each species, identified the origins and insertions of these muscles, as well as the general characteristics of muscle morphology. There were 17 and 18 muscle bundles attached to the feeding apparatus, including five on the dorsal surface. Only the levator rostri, which elevates the rostrum during feeding, showed considerable differences in shape and size among species. The muscles of the adductor complex showed the greatest differences in size among the three species. N. entemedor presented the exclusive muscle X in the lower mandibular area and the extreme reduction of the coracohyoideus in the pharyngeal area derived from the absence of the basihyal cartilage. The information generated in our study supports the morphological specialization of electric rays (N. entemedor) for an almost exclusive suction feeding strategy.
Assuntos
Rajidae , Animais , México , Rajidae/anatomia & histologia , Mandíbula/anatomia & histologia , Torpedo/anatomia & histologia , Músculo Esquelético/anatomia & histologiaRESUMO
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.
Assuntos
Estruturas Animais/anatomia & histologia , Filogenia , Torpedo/anatomia & histologia , Torpedo/classificação , Animais , Cartilagem/anatomia & histologiaRESUMO
The majority of batoids are listed as Threatened (20.4%) or Data Deficient (41%) by the IUCN Red List. A key challenge to assessing Data-Deficient species is obtaining estimates of key life-history characteristics. Here, a Bayesian approach was used to estimate derived life-history characteristics from a growth model applied to the Data-Deficient Brazilian electric ray Narcine brasiliensis. The age of 170 specimens (107 females, 63 males) was estimated from vertebral centra, and total length, disc width, total weight and birth size were used in a joint estimation of sex-specific length-weight models and two-dimensional von Bertalanffy growth models. Estimates of age at length zero, age at maturity, longevity and mortality at age were derived simultaneously. The Bayesian joint modelling approach was robust to small sample sizes by adding a likelihood to constrain L0 and sharing parameters, such as Brody growth coefficient between length measurements. The median growth parameter estimates were a shared L0 = 38.8 mm, female L∞ = 515 mm, ð = 0.125 and male L∞ = 387 mm, ð = 0.194. Age at maturity was estimated to be 7.40-7.49 years for females and 4.45-4.47 years for males, whereas longevity was 22.5-22.6 years for females and 14.2 years for males depending on length measurement. Age-1 natural mortality was estimated to be 0.199-0.207 for females and 0.211-0.213 for males. The derived life-history characteristics indicate N. brasiliensis is earlier maturing, but slower growing relative to other Torpediniformes. These characteristics along with the species' endemism to southern Brazil and high by-catch rates indicate that one of the IUCN Red List threatened categories may be more appropriate for the currently Data-Deficient status. The Bayesian approach used for N. brasiliensis can prove useful for utilizing limited age-growth data in other Data-Deficient batoid species to inform necessary life characteristics for conservation and management.
Assuntos
Características de História de Vida , Modelos Biológicos , Torpedo/fisiologia , Animais , Teorema de Bayes , Tamanho Corporal , Peso Corporal , Brasil , Feminino , Longevidade , Masculino , Coluna Vertebral/crescimento & desenvolvimento , Torpedo/anatomia & histologia , Torpedo/crescimento & desenvolvimentoRESUMO
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.
Assuntos
Animais , Torpedo/anatomia & histologia , Filogenia , Osso e Ossos/anatomia & histologia , Brasil , Torpedo/classificação , Tamanho Corporal , Filogeografia , Fósseis/anatomia & histologiaRESUMO
This study provides information on the reproductive biology of Narcine brasiliensis based on 105 individuals (72 females and 33 males) sampled in São Paulo State, Brazil. The total length at maturity for females was 318·9 mm and for males was 279·8 mm; pregnant females were observed only during summer and autumn. The peak of the gonado-somatic index for females and condition factor for males in the spring suggest a preparation for pregnancy and a mating period during this season. The capture of immature individuals indicates a need for management of the species in this region.
Assuntos
Reprodução , Torpedo/fisiologia , Animais , Brasil , Conservação dos Recursos Naturais , Biologia do Desenvolvimento , Feminino , Masculino , Gravidez , Estações do Ano , Comportamento Sexual Animal , Maturidade Sexual , Torpedo/anatomia & histologiaRESUMO
The dental anatomy of elasmobranch fishes (sharks, rays and relatives) creates a functional system that is more dynamic than that of mammalian dentition. Continuous dental replacement (where new teeth are moved rostrally to replace older ones) and indirect fibrous attachment of the dentition to the jaw allow teeth to reorient relative to the jaw over both long- and short-term scales, respectively. In this study, we examine the processing behavior and dental anatomy of the lesser electric ray Narcine brasiliensis (Olfers, 1831) to illustrate that the freedom of movement of elasmobranch dentition allows a functional flexibility that can be important for complex prey processing behaviors. From static manipulations of dissected jaws and observations of feeding events in live animals, we show that the teeth rotate during jaw protrusion, resulting in a secondary grasping mechanism that likely serves to hold prey while the buccal cavity is flushed free of sediment. The function of teeth is not always readily apparent from morphology; in addition to short-term reorientation, the long-term dental reorientation during replacement allows a given tooth to serve multiple functions during tooth ontogeny. Unlike teeth inside the mouth, the cusps of external teeth (on the portion of the tooth pad that extends past the occlusal plane) lay flat, such that the labial faces act as a functional battering surface, protecting the jaws during prey excavation.
Assuntos
Comportamento Alimentar/fisiologia , Dente/anatomia & histologia , Dente/fisiologia , Torpedo/anatomia & histologia , Torpedo/fisiologia , Animais , Ingestão de Alimentos/fisiologia , Arcada Osseodentária/anatomia & histologia , Arcada Osseodentária/fisiologia , Boca/anatomia & histologia , Boca/fisiologia , FilogeniaRESUMO
The design of minimum-weight structures that retain their integrity under dynamic loading regimes has long challenged engineers. One solution to this problem found in both human and biological design is the optimization of weight and strength by hollowing a structure and replacing its inner core with supportive struts. In animals, this design is observed in sand dollar test, avian beak, and the cancellous bone of tetrapod limbs. Additionally, within the elasmobranch fishes, mineralized trabeculae (struts) have been reported in the jaws of durophagous myliobatid stingrays (Elasmobranchii: Batoidea), but were believed to be absent in basal members of the batoid clade. This study, however, presents an additional case of batoid trabeculation in the lesser electric ray, Narcine brasiliensis (Torpediniformes). The trabeculae in these species likely play different functional roles. Stingrays use their reinforced jaws to crush bivalves, yet N. brasiliensis feeds by ballistically protruding its jaws into the sediment to capture polychaetes. In N. brasiliensis, trabeculae are localized to areas likely to experience the highest load: the quadratomandibular jaw joints, hyomandibular-cranial joint, and the thinnest sections of the jaws immediately lateral to the symphyses. However, the supports perform different functions dependent on location. In regions where the jaws are loaded transversely (as in durophagous rays), "load leading" trabeculae distribute compressive forces from the cortex through the lumen of the jaws. In the parasymphyseal regions of the jaws, "truss" trabeculae form cross-braces perpendicular to the long axes of the jaws. At peak protrusion, the jaw arch is medially compressed and the jaw loaded axially such that these trabeculae are positioned to resist buckling associated with excavation forces. "Truss" trabeculae function to maintain the second moment of area in the thinnest regions of the jaws, illustrating a novel function for batoid trabeculation. Thus, this method of structural support appears to have arisen twice independently in batoids and performs strikingly different ecological functions associated with the distribution of extreme loading environments.
Assuntos
Evolução Biológica , Mandíbula/anatomia & histologia , Mandíbula/fisiologia , Torpedo/anatomia & histologia , Animais , Fenômenos Biomecânicos , Calcificação Fisiológica , Masculino , Torpedo/fisiologiaRESUMO
Protrusion of the jaws during feeding is common in Batoidea (rays, skates, sawfishes, and guitarfishes), members of which possess a highly modified jaw suspension. The lesser electric ray, Narcine brasiliensis, preys primarily on polychaete annelids using a peculiar and highly derived mechanism for jaw protraction. The ray captures its prey by protruding its jaws beneath the substrate and generating subambient buccal pressure to suck worms into its mouth. Initiation of this protrusion is similar to that proposed for other batoids, in that the swing of the distal ends of the hyomandibulae is transmitted to Meckel's cartilage. A "scissor-jack" model of jaw protrusion is proposed for Narcine, in which the coupling of the upper and lower jaws, and extremely flexible symphyses, allow medial compression of the entire jaw complex. This results in a shortening of the distance between the right and left sides of the jaw arch and ventral extension of the jaws. Motion of the skeletal elements involved in this extreme jaw protrusion is convergent with that described for the wobbegong shark, Orectolobus maculatus. Narcine also exhibits asymmetrical protrusion of the jaws from the midline during processing, accomplished by unequal depression of the hyomandibulae. Lower jaw versatility is a functional motif in the batoid feeding mechanism. The pronounced jaw kinesis of N. brasiliensis is partly a function of common batoid characteristics: euhyostylic jaw suspension (decoupling the jaws from the hyoid arch) and complex and subdivided cranial musculature, affording fine motor control. However, this mechanism would not be possible without the loss of the basihyal in narcinid electric rays. The highly protrusible jaw of N. brasiliensis is a versatile and maneuverable feeding apparatus well-suited for the animal's benthic feeding lifestyle.
Assuntos
Ingestão de Alimentos/fisiologia , Arcada Osseodentária/anatomia & histologia , Arcada Osseodentária/fisiologia , Boca/anatomia & histologia , Boca/fisiologia , Torpedo/anatomia & histologia , Torpedo/fisiologia , Animais , Feminino , Masculino , FilogeniaRESUMO
1. The distribution of phospholipids between the two leaflets of the lipid bilayer in acetylcholine receptor (AChR)-rich membranes from T. marmorata has been examined with two complementary techniques: chemical derivatization with the membrane-impermeable reagent trinitrobenzenesulphonate (TNBS) and B.cereus phospholipase C hydrolysis. 2. AChR-membranes were reacted with TNBS at 0-4 and 37 degrees C and the accessibility of their aminophospholipids was compared to that of rod outer segment and erythrocyte membranes. The results indicate that more of the total ethanolamine glycerophospholipid (EGP) than of the total phosphatidylserine (PS) is located in the outer monolayer. 3. Nearly half the phospholipid content of AChR membranes is hydrolyzed by phospholipase C with a half-time of ca. 1.6 min at 25 degrees C. Consistent with the TNBS results, more of the total EGP than of the total PS is degraded. Beyond 3 min the reaction slows down, relatively smaller additional amounts of lipids are hydrolyzed, and all phospholipid classes are attacked to a similar extent, indicating that after half the lipid is removed all phospholipids become accessible to the enzyme. 4. The results indicate that the outer leaflet of the bilayer is richer in ethanolamine and choline glycerophospholipids, whereas phosphatidylinositol, most of the sphingomyelin, and ca 65% of the PS are located on the inner leaflet.