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New chondrichthyan microremains from several Frasnian–Famennian sections in the Holy Cross Mountains and Dębnik area (Southern Poland) are investigated and compared to previous data. The reaction of different groups of chondrichthyans to environmental changes during the Kellwasser Event is analysed. Following the extinction of phoebodont sharks of Phoebodus bifurcatus group before the end of the Frasnian, only two chondrichthyan species, viz. Protacrodus vetustus Jaekel, 1921 and Stethacanthus resistens sp. nov. (possibly closely related to “Cladodus” wildungensis Jaekel, 1921), occur in the upper part of Frasnian Palmatolepis linguiformis conodont Zone and persist into the Famennian. Global cooling is considered a possible cause of the extinction of Frasnian subtropical phoebodonts on Laurussian margins.
A diverse vertebrate fauna, dominated by elasmobranch taxa, was collected from the upper Oligocene (Chattian) Chandler Bridge Formation in Summerville, Dorchester County, South Carolina. Nearly 3,500 teeth and dermal denticles are assigned to 29 species of sharks and rays, and our sample includes the oldest known occurrence of the whale shark, Rhincodon, as well as a new skate, Raja mccollumi sp. nov. The Chandler Bridge elasmobranch assemblage is comparable in species diversity to Chattian assemblages of Virginia and North Carolina, USA, and Germany. Notable absences from Germany include Rhincodon, Hemipristis, and Sphyrna zygaena, likely reflecting the influence of colder water on the North Sea Basin during the Chattian. Squaloids, pristiophoroids, and hexanchoids are known from Chattian deposits of the Albemarle Embayment (North Carolina), Salisbury Embayment (Virginia), and North Sea Basin, but these taxa are absent from the Chandler Bridge assemblage, perhaps because of shallow, warm water (20 to 25°C) conditions within the more southerly Charleston Embayment.
Whereas cosmopolitan distribution patterns are established for many Late Devonian vertebrates (e.g., placoderms, onychodontiforms), few palaeobiogeographic studies have considered chondrichthyans. Recent discoveries of shark material demonstrate that some chondrichthyans were cosmopolitan by the Middle Devonian. Abundant Givetian microremains have been recovered from the Cairo quarry in eastern New York State, USA. These include teeth of two shark species with Gondwanan affinities, the omalodontid Portalodus mannoliniae sp. nov. and the antarctilamnid Wellerodus priscus. Abundant teeth of P. mannoliniae sp. nov. are characterized by a smooth diplodont crown, polarized cusps, and a labially oriented base. The teeth demonstrate monognathic heterodonty. The juvenile morph is distinguished from the adult by smaller size, slender cusps, and variation in the shape of the base. W. priscus is represented by rare juvenile teeth. Two groups of scales that show affinity to material from northern (Spain) and East Gondwana (Antarctica) are tentatively attributed to the two described species. Antarctilamnid distribution suggests a north Gondwanan origin and a colonization of the margin of the landmass before dispersing to Laurentia by the Middle Devonian. This material further indicates that vertebrate global dispersal was initiated by the Middle Devonian, and emphasizes earlier palaeogeographic interpretations that the Middle Devonian “Hamilton fauna” of North American Laurussia originated in the Early Devonian in South American Gondwana.
The teeth of a well known late Palaeozoic cladodont chondrichthyan, “Cladodus” occidentalis from Russia, USA, and England are restudied and a new generic name, Glikmanius gen. nov., is proposed for this species. Yet another tooth−based species, formerly described as ?Symmorium myachkovensis, occurring on the Russian Platform and in Nebraska, is considered to belong to the newly erected genus. Although there is no direct evidence that Glikmanius possessed fin spines, the broad similarity between its teeth and those of Ctenacanthus compressus suggests it had a ctenacanthiform affinity. The possible relationships between Glikmanius, Cladodus sensu stricto, “Ctenacanthus” costellatus, and Heslerodus, are suggested. However, the proposition put forward by an earlier author that the teeth of Heslerodus might represent the lower jaw dentition of G. myachkovensis, is rejected. The overall resemblance of Glikmanius teeth and those of Cladoselache and Squatinactis is recognised as convergent.
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The first Devonian holocephalian tooth from Poland

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A recently found “bradyodont” holocephalian tooth from bituminous shales of the Kowala Quarry, south−western Holy Cross Mountains, Poland, dated as the middle Famennian Palmatolepis trachytera conodont Zone, is described. In spite of its resemblance to the forms often attributed to Helodus, the tooth is referred to as Psephodus cf. magnus (Agassiz, 1838), and supposed to represent the anterior part of the dentition, based on a partly articulated specimen of Psephodus from the Carboniferous of Scotland. The analysis of early helodonts and psephodonts, and other Famennian chondrichthyan crushing teeth, shows numerous similarities in tooth−base structure, such as the reduction of lingual basal extension, loss of articulation devices, development of numerous nutritive foramina, and the tendency to fusion between the teeth in a tooth−family. Based on these shared characters, close phylogenetic relationships between the Protacrodontoidea, Hybodontoidea, and the Holocephali are postulated.
A new record of the chondrichthyan hybodontoid genus Lissodus is presented from two localities within the Mississippian (Tournaisian) rocks of Ireland. Five morphotypes of the genus are described within each of which occurs morphological variance. Specimens recovered and described herein are from crinoidal limestones whose palaeoenvironments are interpreted as ranging from a moderately shallow high−energy carbonate shelf, to relatively deep off−shore. The richest fauna recovered from the high−energy carbonate shelf, contains all five morphotypes raising the possibility that they may have been derived from a single species of shark. A discussion on the relationship between the five morphotypes and other Carboniferous Lissodus teeth is offered and it is argued that although the morphotypes differ slightly from other Carboniferous Lissodus teeth, they may belong to a closely related species not formally named until additional evidence is obtained. A mouth reconstruction using the teeth recovered from the Lower Carboniferous of Ireland is proposed.
The revision of shark teeth from the Pennsylvanian black shales of central USA, ascribed to “Phoebodus heslerorum” Williams, 1985 and Symmorium reniforme Cope, 1893, shows that “Ph. heslerorum” is a junior synonym of “Cladodus” divergens Trautschold, 1879. This species belongs neither to Phoebodus nor to Cladodus, so a new genus Heslerodus is proposed. Very common, robust cladodont teeth with a deep labio−basal depression and two buttons, often referred to as S. reniforme, do not belong to the latter species, but to “Cladodus” occidentalis Leidy, 1859. The generic affinity of “C.” occidentalis is yet undetermined, but it is possible that it represents ctenacanthoids.
Cenozoic lamniform sharks are mostly represented by isolated teeth and vertebrae, whereas articulated skeletal remains are usually very scarce. Here, we describe a partial skeleton of an extinct lamniform shark consisting of 42 slightly disarticulated teeth, 49 vertebrae, and additional unidentifiable cranial and postcranial remains. The specimen originates from the Miocene mica-clay of Groß Pampau (North Germany), which is of late Langenfeldian age (= Serravallian-Tortonian boundary; middle-late Miocene). A total of 13 measurements of each tooth, as well as morphological features, were used to reconstruct the dentition of this specimen and to provide detailed taxonomic information. Additionally, the total body size and age at death were established using methodologies based on vertebral and tooth measurements and vertebral centra growth ring counts, respectively. The specimen undoubtedly represents the most complete individual of "Carcharodon (= Isurus) escheri", previously known only from a few isolated teeth. The dental pattern (e.g., marked dignathic and monognathic heterodonty patterns; only slightly labio-lingually compressed upper teeth; upper teeth slender with distally inclined or curved main cusps; massive, hook-like upper intermediate tooth; main cusps with crenulated cutting edges; lateral cusplets in teeth of all ontogenetic stages) clearly separates this shark from all hitherto known Cenozoic and Recent lamnids and a new genus, Carcharomodus, consequently is introduced. Carcharomodus escheri comb. nov. is a characteristic element of late early Miocene to the Pliocene Western and Central European fish faunas. All previously identified Pacific occurrences represent a different taxon. We estimate that the specimen had a total body length of about 4 m and that it was older than 10 years and thus might have reached maturity before death, as indicated by all available evidence.
New specimens, including the first record of lower dental plates, of the extinct myliobatid Myliobatis wurnoensis were recovered from the Maastrichtian (Late Cretaceous) of the Iullemmeden Basin, Mali, and are the oldest record of the taxon. We evaluated the phylogenetic position of this taxon with reference to other myliobatids (extinct and extant) using osteology and dentition. Our results indicate that Myliobatinae and Myliobatis are each paraphyletic, and that Aetobatus and Rhinoptera are monophyletic. We also found that taxa known only from the Cretaceous, Brachyrhizodus and Igdabatis, are highly nested within Myliobatidae. The phylogenetic position of these taxa unambiguously extends the origin of Myliobatidae and most of its representative taxa into the Mesozoic.
The shallow water assemblage of chondrichthyan microremains, teeth, tooth plates and scales, from the middle Tournaisian (Mississippian) of the vicinity of Muhua village, Guizhou province, southern China, is thus far the richest and most diverse association of this age collected from a single locality and horizon, and represents a chondrichthyan community very restricted in time and space. It was recovered from a small bioclastic limestone lens, MH−1, occurring among basinal marls near the base of the Muhua Formation, and dated as to the Siphonodella crenulata conodont Zone. The majority of the fauna presented here consists of teeth with euselachian−type bases and crushing crowns belonging to bottom−dwelling durophagous chondrichthyans, most probably feeding on shelly invertebrates such as the abundant brachiopods. We assigned most of these teeth to Euselachii (six species, among them Cassisodus margaritae gen. et sp. nov.), Petalodontiformes (two species), Holocephali (five species), and Euchondrocephali incertae sedis (Cristatodens sigmoidalis gen. et sp. nov.). We also identified primitive polycuspid, clutching teeth representing Phoebodontiformes (Thrinacodus bicuspidatus sp. nov.), Symmoriiformes, and Ctenacanthiformes. The scales are typical growing, compound forms of the protacrodont, ctenacanth, and hybodont types. Two problematic denticulated plates were found, one of which resembles mandibular or palatal plates of Sibyrhynchus (Iniopterygii). Several of the identified chondrichthyan taxa have hitherto been known only from Laurussia, especially from the British Isles and central USA. In particular we found the first record of Chondrenchelyssp. and Diclitodus denshumani outside their type locality. Th. bicuspidatus sp. nov., also known from Nevada, Iran, and NW Australia, appears to be a cosmopolitan, middle Tournaisian index fossil.
Three genera of xenacanths, based on isolated teeth, occur in the lepospondyl (amphibian)−dominated fauna from the upper Black Prince Limestone (late Bashkirian). Orthacanthus donnelljohnsi sp. nov. teeth, with carinae lacking serrations on the compressed principal cusps, and only one intermediate cusp, represent both adult and juvenile teeth. Heterodonty occurs in both adult and juvenile dentitions. The absence of serrations is unique among Pennsylvanian species of Orthacanthus. Teeth with often highly asymmetrical bases with an aborally−flexed lingual marginal flange (= anterolingual shelf) and a single intermediate cusp are assigned to Triodus elpia sp. nov. A central foramen occurs in the base, unlike most other species; the moderately compressed principal cusps bear generally straight cristae. They represent the first reported occurrence of Triodus in the Paleozoic of North America. Five teeth, with cristae extending from the cusps onto their bases, belong to Bransonella. Two are questionably assigned to Bransonella nebraskensis, one to B. ?lingulata with its labio−lingually elongated apical button and smaller than normal intermediate cusp, and one each to Bransonella sp. “A” and “B”. Bransonella sp. “A” has a base wider (labio−lingual) than long, the reverse of the other Bransonella teeth. Bransonella sp. “B” is distinctly different, as it lacks an intermediate cusp (as in some B. lingulata teeth), and the basal tubercle is beneath one of the cusps (with no evidence of deformity).
Cardabiodon ricki and Cardabiodon venator were large lamniform sharks with a patchy but global distribution in the Cenomanian and Turonian. Their teeth are generally rare and skeletal elements are less common. The centra of Cardabiodon ricki can be distinguished from those of other lamniforms by their unique combination of characteristics: medium length, round articulating outline with a very thick corpus calcareum, a corpus calcareum with a laterally flat rim, robust radial lamellae, thick radial lamellae that occur in low density, concentric lamellae absent, small circular or subovate pores concentrated next to each corpus calcareum, and papillose circular ridges on the surface of the corpus calcareum. The large diameter and robustness of the centra of two examined specimens suggest that Cardabiodon was large, had a rigid vertebral column, and was a fast swimmer. The sectioned corpora calcarea show both individuals deposited 13 bands (assumed to represent annual increments) after the birth ring. The identification of the birth ring is supported in the holotype of Cardabiodon ricki as the back-calculated tooth size at age 0 is nearly equal to the size of the smallest known isolated tooth of this species. The birth ring size (5-6.6 mm radial distance [RD]) overlaps with that of Archaeolamna kopingensis (5.4 mm RD) and the range of variation of Cretoxyrhina mantelli (6-11.6 mm RD) from the Smoky Hill Chalk, Niobrara Formation. The revised, reconstructed lower jaw dentition of the holotype of Cardabiodon ricki contains four anterior and 12 lateroposterior files. Total body length is estimated at 5.5 m based on 746 mm lower jaw bite circumference reconstructed from associated teeth of the holotype.
Detailed studies on Carboniferous species of the xenacanth Orthacanthus have shown that the xenacanth dorsal fin spine can be used for skeletochronological analyses and provides valuable information about development, growth and environmental life conditions of those extinct sharks. We report here for the first time the histology and skeletochronology of Permian specimens, dorsal spines of Orthacanthus platypternus from the Craddock Bone Bed (lower Clear Fork Formation; Early Permian, Leonardian age) of northern Baylor County (north-central Texas, USA). Twelve dorsal spines of O. platypternus preserve a highly vascularized wall mainly composed of centrifugally growing dentine in a succession of dentine layers, probably deposited with an annual periodicity. As expected, spines of individuals with 1–2 dentine layers, presumably juveniles, present the smallest sizes. However, spines of individuals showing at least 3–4 dentine layers and interpreted to be subadults/young adults, are distributed in two spine-size clusters corresponding to females (probably the largest spines) and males, in agreement with the hypothesis of sexual size dimorphism proposed in a previous biometric analysis. Our comparative study of O. platypternus and the Stephanian species O. meridionalis further suggests that spine denticulation can be useful for distinguishing between species of Orthacanthus and sexually dimorphic forms (juvenile to adults) in each species. Total body length estimations of O. platypternus from the Craddock Bone Bed point to relatively large juveniles and small subadults/young adults (less than 2 m in total length), living as opportunistic predators in the pond-channel coastal plain environments represented by the bone bed deposits. The comparative analyses of the ontogenetic stages of the recorded specimens of O. platypternus and their distribution along different facies and localities indicate that this species was euryhaline, diadromous with a catadromous life-cycle which was strongly regulated by the semi-arid, seasonally dry tropical climate affecting western Pangaea during the Early Permian.
During the last two decades, an abundant selachian assemblage has been collected from the late Ypresian (NP12) fossiliferous sands of Prémontré (Aisne, northern France) but has received little attention. Sharks of the family Triakidae (Carcharhiniformes) are particularly well represented and all are described and figured herein. Among them, two new species of the genus Galeorhinus are described: G. duchaussoisi sp. nov. and G. louisi sp. nov.; these are compared to the common Paleogene G. ypresiensis which is refigured. Another triakid taxon, the genus Gomphogaleus gen. nov., is described. Most of the triakids have been recorded elsewhere in the North Atlantic region, suggesting a wider distribution than expected for these small sharks during the Paleogene. The present paper updates the list of selachians from Prémontré, bringing the number of taxa from 19 to 33 (including 22 sharks and 11 batoids) and improving our knowledge of the ancient North Atlantic Ypresian selachian fauna. Despite this vastly improved record, it is clear that fossil data are still very incomplete and insufficient for calibrating phylogenetic hypotheses of living forms. Review of the Prémontré fauna shows that the Triakidae were much more diverse and broadly distributed than at present, suggesting that the limited distribution and low diversity of living forms is probably a recent phenomenon.
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