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In the Podolian Dniester Basin (southwestern Ukraine) the Lower Devonian marine deposits are represented by about 530 m thick continuous sequence of interlaminated carbonate and schale outcrops at several localities. Conodonts occur in most of the carbonate layers of the whole Lochkovian but are not abundant and their ramiform elements are mostly broken or lacking. Therefore, only the pectiniform, Pa elements of twenty five stratigraphically important conodont species occurring in the region are discussed and two new species, Caudicriodus schoenlaubi and Pandorinellina? parva are proposed. The hypothetical phyletic relationships within the main representatives of the icriodontid and spathognathodontid genera, Caudicriodus, Zieglerodina, and Pandorinellina? are traced. Comparison of the previously published and newly obtained data revealed discrepancies in the hitherto used interpretation of some of the conodont taxa and their stratigraphic ranges. Contrary to the earlier reports, Caudicriodus postwoschmidti does not occur in the lower Lochkovian but only in the middle part of the Chortkiv Formation, high above the Monograptus uniformis Zone. Based on new material and verification of the previous determinations, a modified scheme of the Lochkovian conodont zonation in Podolia is proposed. Conodont zones: Caudicriodus hesperius, C. transiens, C. postwoschmidti, C. serus, and ?Caudicriodus steinachensis are distinguished. The zones are correlated with conodont zonations in other regions—Barrandian, Cantabrian Mountains, Pyrenees, and Nevada. Biostratigraphy of the Siluro−Devonian transition and Lochkovian is integrated with the carbon isotope stratigraphy.
A cosmopolitan spathognathodontid taxon “Ancyrodelloides carlsi” is not regarded in this paper as one of the earliest known representatives of Ancyrodelloides according to former concepts, but is considered as conspicuous member of the genus Lanea. The taxon shows distinct morphological innovation of the upper surface of the Pa element within the Lanea lineage which makes it easily recognizable worldwide and suitable for global correlation. Presented stratigraphic correlation using conodonts and other faunal groups qualifies a short−lived Lanea carlsi as probably the best marker of the middle Lochkovian base. This paper presents reconstruction of the apparatus of L. carlsi which is the first complete reconstruction in the genus Lanea. The comparison of the Lanea apparatus and assumed apparatus of early Ancyrodelloides from the Požáry Quarries (Barrandian, Czech Republic) resulted to proposal of an alternative concept of evolution of these two genera. The relatively short−lived genus Ancyrodelloides is considered to split off from the Lanea clade in the late middle Lochkovian by the entry of Ancncyrodelloides transitans.
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Conodonts, a large group of tiny extinct marine animals ranging in age from the Late Cambrian to Late Triassic (ca. 500 to 200 Mya), are usually considered as jawless vertebrates. Their only commonly occurring fossilized remains are minute, phosphatic, teeth−like elements of their feeding apparatuses. In most of the early conodonts the elements were conical and strongly elongated. Many of them are characterized by possession of a deep, longitudinal groove, usually associated with sharp edges or ridges. A comparative study of the grooved elements and venomous teeth and spines of living and extinct vertebrates strongly suggests that the groove in conodonts was also used for delivery of venom. Structural convergence of the conodont apparatus Panderodus with the grasping apparatus of chaetognaths, a group of extant, venomous invertebrate predators of similarly ancient origin, provides additional support for this conclusion.
Multidisciplinary study of the Early–Middle Frasnian boundary in the north−western East European Platform (Main Devonian Field) allows evaluation of changes in facies, brachiopod, ostracod, and conodont associations, as well as carbon isotope composition. Brachiopod and ostracod faunas, characterized by predominance of rhynchonellids, spiriferids, podocopids, and platycopids in the Early Frasnian, and by predominance of spiriferids, palaeocopids, and kloedenellocopids in the Middle Frasinan, demonstrate significant changes in dominance and diversity likely caused by regressiontransgression couplets in the extremely shallow−water environment. Changes in diversity of the conodont associations, represented mainly by shallow−water polygnathids and spathognathids, are controlled by sea−level fluctuations as well; however significant evolutionary turnover is absent. Deepening pulses lead to an increasing in diversity of both the benthic and nectic groups, whilst regressions cause diversity fall due to progressive habitat reduction. Most prominent decreasing in the fauna diversity is observed in the late Early Frasnian (Dubnik time) coinciding with the regressive phase of the Late Givetian to Early Frasnian eustatic cycle. Positive−negative δ¹³C excursion, detected in brachiopod calcite from the early interval of the Middle Frasnian, can be correlated with global isotopic perturbations near the Frasnian substage boundary: the positive 2.6‰ δ¹³C excursion is probably linked with enhanced primary production in high−nutrient regimes in the epeiric sea.
A rich conodont fauna from two Neotethyan sections, Pizzo Mondello (western Sicily, Italy) and Pignola 2 sections (southern Apennines, Italy) includes conodonts described herein as a new species. Metapolygnathus praecommunisti sp. nov. is transitional between Paragondolella noah and Metapolygnathus communisti. The genus Metapolygnathus (including M. praecommunisti) is now characterised by posterior prolongation of the keel termination, associated with a centrally located pit and with a weak ornamentation confined to the anterior part of the platform margins. The establishment of M. praecommunisti addresses the problems related to the origin and the peculiar, probably facies−controlled, distribution of its descendant species M. communisti. Since M. praecommunisti occurs in the entire Tethys and in North America, we propose the species as a good guide fossil for global correlations, characterised by a short temporal range limited to the uppermost Tuvalian (upper Carnian). The stratigraphic occurrence of the genus Metapolygnathus is restricted to the Tuvalian–Lacian (upper Carnian–lower Norian), excluding its presence in the Julian substage (lower Carnian).
Throughout their history, species had to face environmental variations spatially and temporally. How both levels of variation interact will be of key importance in conditioning their response to major perturbations. We addressed this question by focusing on a period in Earth’s history marked by dramatic environmental and faunal changes, the Late Devonian Frasnian/Famennian boundary. From a paleogeographic point of view, this period is characterized by a cosmopolitanism of the faunas across a large ocean, the Prototethys. We considered the biotic reaction at a seldom considered scale, namely within a single subgenus of conodont, Palmatolepis (Manticolepis). Patterns of spatial and temporal differentiation were quantified using morphometrics of its platform element. The recognized cosmopolitanism of the faunas was confirmed at this scale of variation since temporal records gathered in distant areas around the Prototethys, including the seldom documented regions located nowadays in South−East Asia, displayed similar morphological trends in response to the major F/F crisis. Beyond this overall cosmopolitanism, subtle geographic structure was evidenced but was not stable through time. Geographic differentiation was maximal shortly before the F/F crisis, suggesting that despite high sea−level, tectonics leaded to complex submarine landscapes promoting differentiation. In contrast any geographic structure was swamped out after the crisis, possibly due to a global recolonization from few favorable patches.
The oxygen isotopic composition of conodont apatite derived from the Late Triassic (Carnian to lower Norian), Pignola 2 and Sasso di Castalda sections in the Lagonegro Basin (Southern Apennines, Italy) was studied in order to constrain the habitat of Late Triassic conodont animals. Oxygen isotope ratios of conodonts range from 18.5 to 20.8‰ V−SMOW, which translate to palaeotemperatures ranging from 22 to 31ºC, assuming a δ18O of Triassic subtropical sea water of −0.12‰ V−SMOW. These warm temperatures, which are well comparable to those of modern subtropical−tropical oceans, along with the body features of the conodont animal suggest that conodont δ18O values reflect surface water temperatures, that the studied conodont taxa lived in near−surface waters, and that δ18O values of Late Triassic conodonts can be used for palaeoclimatic reconstructions.
Two global isotopic events, the early Sheinwoodian (early Wenlock) and that at the Silurian–Devonian transition, have been comprehensively studied in representative carbonate successions at Kytayhorod and Dnistrove, respectively, in Podolia, Ukraine, to compare geochemistry and biotic changes related correspondingly to the Ireviken and Klonk events. These two large−scale isotope excursions reveal different regional ecosystem tendencies. The well−defined increasing trend across the Llandovery–Wenlock boundary in siliciclastic input, redox states and, supposedly, bioproductivity, was without strict correlative relations to the major ¹³C enrichment event. The environmental and biotic evolution was forced by eustatic sea−level fluctuations and two−step climate change toward a glaciation episode, but strongly modified by regional epeirogeny movements due to location near the mobile Teisseyre−Törnquist Fault Zone. Thus, the global early Sheinwoodian biogeochemical perturbation was of minor depositional significance in this epeiric sea, as in many other Laurussian domains. Conversely, the Podolian sedimentary record of the Klonk Event exhibits temporal links to the abrupt δ¹³C anomaly, overprinted by a tectonically driven deepening pulse in the crucial S–D boundary interval. This carbon cycling turnover was reflected in the regional carbonate crisis and cooling episodes, paired with a tendency towards eutrophication and recurrent oxygen deficiency, but also with major storms and possible upwelling. Faunal responses in both Podolian sections follow some characters of the Silurian pattern worldwide, as manifested by conodont changeover prior to the major early Sheinwoodian isotopic/climatic anomaly. This contrasts with the relative brachiopod and chitinozoan resistances in the course of the Ireviken Event. Also, during the Klonk Event, a moderate faunal turnover, both in benthic and pelagic groups, occurred only near the very beginning of the prolonged ¹³C−enriched timespan across the system boundary, possibly due to progressive dysoxia and temperature drop. The characters point to a peculiarity of the Klonk Event by comparison with the Silurian global events, and some similarity already to the succeeding Devonian transgressive/anoxic episodes.
An earliest Famennian (Late Devonian) shell bed was discovered in the Hanover Shale Member of the Java Formation 1.4 meters above the Frasnian–Famennian (F–F) boundary in western New York. The invertebrate shelly fauna of the shell bed (Lower Palmatolepis triangularis conodont Zone), provides information on taxonomy of an outer shelf benthic association during the survival interval of recovery in the Appalachian foreland basin soon after the terminal Upper Kellwasser event marking the F–F mass extinction. Shelly invertebrates are extremely rare in the upper Hanover immediately above and below the shell bed. Abundance of brachiopod valves and remains of other groups in the shell bed reach 80–100 valves/100 cm2. Elongate valves of the linguloid brachiopod Barroisella cf. B. campbelli have preferred alignments roughly parallel to direction of down−slope flow in the deep−water foreland basin depositional setting. The brachiopod fauna is dominated by the representatives of Retichonetes, Barroisella, Cyrtospirifer, Tylothyris, and Praewaagenoconcha. Rare elements include species of Thiemella, Schizophoria, Ripidiorhynchus?, Chapinella?, an indeterminate rhynchonellid, Ambocoelia, and extremely rare Orbiculoidea. Forms including Cyrtospirifer hornellensis, Tylothyris mesacostalis, Praewaagenoconcha speciosa, and few others are late Frasnian carryovers. The range inception of Thiemella leonensis is just above the F–F boundary (Upper Kellwasser horizon) in the upper Hanover Shale shell bed in the western Appalachian foreland basin.
The dominant factor in faunal succession of conodonts in the Frasnian of Poland is the apparent immigration of species originating allopatrically in other regions. Each immigration event usually changes the population variability of a local species (character displacement). Only a few lineages show their phyletic evolution within the studied area. Attempts to distinguish conodont species on the basis of platform element shape failed in some of the latest Frasnian palmatolepidids. Even at the apparatus−based generic level, certain ramiform elements of the apparatus appear much more diagnostic than the platforms. Correlative value of the late Frasnian palmatolepidids of unknown apparatus structure is thus questionable. The evolution of platform elements in Ancyrodella offers a more solid basis for age determination although their great population variability makes resolution rather low and requires the population approach. The panderodontids Belodella(?) tenuiserrata sp., B. minutidentata sp. nov., B. robustidentata sp. nov., prioniodontid Icriodus kielcensis sp. nov., enigmatic monospecific Playfordiidae fam. nov., prioniodinids Dyminodina planidentata gen. et sp. nov., D. anterodenticulata sp. nov., D. kovalensis sp. nov., Pluckidina kielcensis gen. et sp. nov., P. slupiensis sp. nov., P. robustipegmata sp. nov., and P. lagoviensis sp. nov., derived polygnathid Avignathus bifurcatus sp. nov., probably secondarily simplified polygnathid Nicollidina gen. nov., and palmatolepidids Kielcelepis gen. nov., Lagovilepis gen. nov. and Klapperilepis gen. nov. are proposed.
Scolecodonts and conodonts are described from five samples of three cores from the Svalis Dome, central Barents Sea. The conodont species Mesogondolella rosenkrantzi and Neospathodus svalbardensis confirm latest Permian (Dzulfian) and earliest Triassic (Dienerian) ages for the investigated intervals. Correlation with adjacent conodont distribution indicates a shorter depositional gap in the late Permian than is evident on Svalbard. The conodont elements show little thermal alteration with CAI (colour alteration index) values between 1.0 and 1.5 indicating a maximum burial temperature of 50—90°C. The scolecodonts described herein have important evolutionary and distribution implications for polychaetes: the order Eunicida shows a richer diversification in the Permian than previously known; the family Paulinitidae survived at least until the Late Permian; the family Hartmaniellidae diversified already in the late Palaeozoic; some forms are considered as representatives of unknown species and genera; the order Phyllodocida originated already in the latest Palaeozoic.
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An earlier hypothesis concerning the origin of chaetognaths from protoconodonts found additional support in new discoveries and in recent structural, chemical and molecular investigations. The new findings show that the head armature of protoconodonts was composed not only of grasping spines but also of much smaller spicules corresponding in size and shape to the chaetognath teeth. Grasping spines of protoconodonts were originally built mainly of an organic substance. Their original composition was changed by secondary phosphatisation. The thickest layer of the protoconodont spines was originally constructed of organic fibrils, similar to those in the corresponding layer of chaetognaths. Recent molecular investigations show that the chaetognath lineage separated in the early stage of metazoan radiation, which fits the presented hypothesis. Described are some previously unknown structural details of chaetognath grasping spines, including composition of the outer layer and the origin of their distinctive tips.
The Lower Ordovician conodont Scolopodus striatus Pander, 1856 (= Scolopodus rex Lindström, 1955) has an apparatus composed of five element morphotypes: acontiodiform, subrounded, compressed paltodiform, paltodiform, and scandodiform. The identification of the morphotypes is based on the general asymmetry of elements and shape of their bases. The elements are variable within each morphological group and form a continuous transition series generally reflected in differences in the depth of the basal cavity, height and degree of lateral compression of the base. The new collection from the localities near St. Petersburg, the type area of the first investigations on conodonts by Christian Pander in 1856, was examined and species of Scolopodus named by him are revised. All Pander’s species with the exception of the type species Scolopodus sublaevis are identified as a single species of Scolopodus that was named 99 years later as Scolopodus rex. The S. sublaevis sensu formae was not recognised in collections studied and its validity is questionable. S. striatus is the most easily recognisable among Pander’s species of Scolopodus and is here proposed to be a senior synonym of S. rex.
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