Amino acid residues in conodont elements

Thermally unaltered conodont elements, brachiopods. and vertebrates were analyzed with reverse phase high profile liquid chromatography to locate and quantify amino acid remnants of the original organic matrix in the fossils. No consistent similarities in amino acid content were found in conodont taxa. and criteria based on organic residues appear to have no taxonomic significance in the fossils tested from these localities. However, hydroxyproline. an amino acid that is found in the collagen molecules of animals. as well as in the glycoproteins in the cell walls and reproductive tissues of certain plants, is represented in most taxa. The organic matter retained in the impermeable crowns of conodont elements might have been derived originally from a form of collagen. Biochemical analyses. correlated with histochemical tests, demonstrate that organic matter is an integral part of the hyaline tissue of the element crown and not the result of surface contamination. Tests of a range of vertebrate and invertebrate fossil hard tissues produced similar results. The analyses indicate that hyaline tissue in the conodont element crown is not a form of vertebrate enamel. which contains no collagen. Albid tissue. with little or no organic content. is not a form of vertebrate bone or dentine, both based on collagen and low in mineral. Although these results do not help to determine the phylogenetic affinities of conodont animals, they indicate teat conodont elements do not contain hard tissues characteristic of vertebrate animals.

Hyaline tissue of thermally unaltered conodont elements and the enamel of vertebrates

Comparison of the ultrastructure of the hyaline tissue of conodont elements and the enamel of vertebrates provides little support for a close phylogenetic relationship between conodonts and vertebrates. Transmission and scanning electron microscopy shows that the mineralised component of the hyaline tissue of Panderodus and of Cordylodus elements consists of large, flat, oblong crystals, arranged in layers that run parallel to the long axis of the conodont. Enamel in the dentition of a living vertebrate, the lungfish Neoceratodus forsteri, has crystals of calcium hydroxyapatite, arranged in layers, and extending in groups from the dentine-enamel junction; the crystals are slender, elongate spicules perpendicular to the surface of the tooth plate, Similar crystal arrangements to those of lungfish are found in other vertebrates, but none resembles the organisation of the hyaline tissue of conodont elements, The crystals of hydroxyapatite in conodont hyaline tissue are exceptionally large, perpendicular or parallel to the surface of the element, with no trace of prisms, unlike the protoprismatic radial crystallite enamel of fish teeth and scales, or the highly organised prismatic enamel of mammals.

Oxygen isotope values from biogenic apatie (conodont elements and fish teeth) from the Lower Triassic Mianwali Formation (Salt Range, Pakistan)

Recovery from the end-Permian mass extinction is frequently described as delayed, with complex ecological communities typically not found in the fossil record until the Middle Triassic epoch. However, the taxonomic diversity of a number of marine groups, ranging from ammonoids to benthic foraminifera, peaked rapidly in the Early Triassic. These variations in biodiversity occur amidst pronounced excursions in the carbon isotope record, which are compatible with episodes of massive CO2 outgassing from the Siberian Large Igneous Province. Here we present a high-resolution Early Triassic temperature record based on the oxygen isotope composition of pristine apatite from fossil conodonts. Our reconstruction shows that the beginning of the Smithian substage of the Early Triassic was marked by a cooler climate, followed by an interval of warmth lasting until the Spathian substage boundary. Cooler conditions resumed in the Spathian. We find the greatest increases in taxonomic diversity during the cooler phases of the early Smithian and early Spathian. In contrast, a period of extreme warmth in the middle and late Smithian was associated with floral ecological change and high faunal taxonomic turnover in the ocean. We suggest that climate upheaval and carbon-cycle perturbations due to volcanic outgassing were important drivers of Early Triassic biotic recovery.

Triassic stratigraphic evolution of the Arabian-Greater India embayment of the southern Tethys margin

An exceptional, tectonically remarkably unaffected, nearly 200 m-thick continuous section of hemipelagic and turbiditic sediments, covering most of the Triassic is described from the Batain Complex of north-eastern Oman. According to conodont and radiolarian data the sequence spans the late Scythian to the early Norian, a time period of nearly 30 M. Coupled with a high resolution stratigraphy, the lithostratigraphy, sedimentology, as well as sequence and isotope stratigraphy of the section are documented. For the Triassic of the Batain Plain we propose the new name Sal Formation, which replaces the formerly used Matbat Formation, and subdivide it into three new members. The Sal Formation was deposited on the proximal continental margin of northeastern Arabia and records various depositional environments. The lower member is interpreted as the distal part of a homoclinal ramp which evolves to a distally steepened ramp during time of deposition of the middle member. The upper member displays a toe of slope position which is indicated by an increase of proximal turbidites. These sediments form part of a segment of the Neo-Tethyan embayment between Arabia and India. The stratigraphic analysis indicates highly varying sedimentation rates from a minimum of 2 m/M gamma around the Anisian/Ladinian boundary up to 15 m/M gamma during the Lower and Upper Triassic. Sequence-stratigraphically, the Sal section is subdivided into six third order cycles which are biochronologically well integrated into the global Triassic cycle chart. The mixed siliciclastic-calcareous upper member of the Sal Formation typically shows highstand related carbonate shedding. It is, therefore, an important test case for sequence-stratigraphic controlled carbonate export to mixed basin fills. The well developed sequence stratigraphic cycles are mirrored in the isotope patterns. Additionally, the carbon and oxygen isotope data from the Sal Formation record the same chemostratigraphic marker at the Spathian/Anisian boundary known from other Tethyan sections.

Geology and correlation of the Mersin mélanges, southern Turkey

Our paper aims to give a thorough description of the infra-ophiolitic melanges associated with the Mersin ophiolite. We propose new regional correlations of the Mersin melanges with other melange-like units or similar series, located both in southern Turkey and adjacent regions. The palaeotectonic implications of the correlations are also discussed. The main results may be summarized as follows: the infra-ophiolitic melange is subdivided into two units, the Upper Cretaceous Sorgun ophiolitic melange and the Ladinian-Carnian Hacialani melange. The Mersin melanges, together with the Antalya and Mamonia domains, are represented by a series of exotic units now found south of the main Taurus range, and are characteristic of the South-Taurides Exotic Units. These melanges clearly show the mixed origin of the different blocks and broken formations. Some components have a Palaeotethyan origin and are characterized by Pennsylvanian and Lower to Middle Permian pelagic and slope deposits. These Palaeotethyan remnants, found exclusively in the Hacialani melange, were reworked as major olistostromes in the Neotethys basin during the Eo-Cimmerian orogenic event. Neotethyan elements are represented by Middle Triassic seamounts and by broken formations containing typical Neotethyan conodont faunas such as Metapolygnathus mersinensis Kozur & Moix and M. primitius s. s., both present in the latest Carnian interval, as well as the occurrence of the middle Norian Epigondolella praeslovakensis Kozur, Masset & Moix. Other elements are clearly derived from the former north Anatolian passive margin and are represented by Huglu-type series including the Upper Triassic syn-rift volcanic event. These sequences attributed to the Huglu-Pindos back-arc ocean were displaced southward during the Late Cretaceous obduction event. The Tauric elements are represented by Eo-Cimmerian flysch-like and molasse sequences intercalated in Neotethyan series. Additionally, some shallow-water blocks might be derived from the Bolkardag para-autochthonous and the Taurus-Beydaglari marginal sequences.

The alkaline intraplate volcanism of the Antalya nappes (Turkey): a Late Triassic remnant of the Neotethys

Late Triassic submarine alkali basalts and hawaiites were collected from two superimposed tectonic slices belonging to the Kara Dere - Sayrun unit of the Middle Antalya nappes, southwestern Turkey. New determinations on conodont faunas allow to date this sequence to the Lower Carnian (Julian). The volcanic rocks show rather homogeneous compositions, with high TiO2 and relatively low MgO and Ni contents which suggest olivine fractionation. Their primitive mantle-normalised multi-elements plots show Nb and Ta enrichments relative to La, Pb negative anomalies and heavy rare earth element and Y depletions typical of intraplate ocean island basalts. These characteristics are consistent with the major and trace element compositions of their primary clinopyroxene phenocrysts, which do not show any feature ascribable to crustal contamination. The studied lavas display a restricted range of epsilon Nd (+4.6 to +5.2) which falls within the range of ocean island basalts. Their initial (Nd-143/Nd-144)i ratios are too low to be explained by a simple mixing line between depleted MORB mantle (DMM) and HIMU components. Their Pb and Nd isotopic compositions plot along a mixing line between HIMU component and an enriched mantle, the composition of which could be the result of the addition of about 5 to 8% of an EM2 component (recycled marine sediments) to DMM. The lack of evidence for any continental crustal component. in their genesis could be consistent with their emplacement in an intra-oceanic setting.

A unique Permian-Triassic boundary section from the Neotethyan Hawasina Basin, Central Oman Mountains

In the Wadi Wasit area (Central Oman Mountains), Dienerian breccias are widespread. These breccias consist mostly of Guadalupian reefal blocks, often dolomitised, and some rare small-sized blocks of lowermost Triassic bivalve-bearing limestones. A unique block, with a size of about 200 m(3), including Permian and earliest Triassic faunas has been studied in detail. The so-called Wadi Wasit block consists of three major lithological units. A basal unstratified grey limestone is rich in various reef-building organisms (rugose corals, calcareous sponges, stromatoporoids) and has been dated as Middle Permian. It is disconformably overlain by well- and thin-bedded light grey to yellowish coloured limestones rich in molluscs. Two major lithologies (Coquina Limestone respectively Bioclastic Limestone unit) characterise the shelly limestones, their contact seems gradual. These two units are well-dated; they are of Griesbachian age and contain three conodont zones, the Parvus Zone, the Staeschei Zone and the Sosioensis Zone, and two ammonoid zones, the Ophiceras tibeticum Zone and an 'unnamed interval'. The third unit consists of a grey marly limestone containing Neospathodus kummeli (basal Dienerian). It is the first record of well-dated basal Triassic sediments in the Arabian Peninsula. The Coquina Limestone is dominated by the bivalve Promyalina with some Claraia and Eumorphotis. This bivalve association is interpreted as a pioneering opportunistic assemblage. Towards the top of the Bioclastic Limestone unit, the faunal diversity increases and contains probably more than 20 taxa of bivalves, microgastropods, crinoids, brachiopods, ammonoids, echinoid spines, ostracods and conodonts. The generic diversity of this biofacies exceeds by far any other Griesbachian assemblage known. Our data give new evidence for the geodynamical history for the distal carbonate shelf bordering the Hawasina Basin. A break in the sedimentation characterises the Late Permian. The basal Triassic shows a steady transgression and the breccias may record a distinct gravitational collapse of platform margins linked with sea-level low stand at the end of Induan time (late Dienerian-basal Smithian). delta(13)C(carb) isotopic analyses were performed and yield typical Permian values of around 4parts per thousand for the Reefal Limestone, with a strong negative shift across the Permian-Triassic boundary. During the Griesbachian values shift positively from 0.5 to 3.1parts per thousand parallel to an increase in faunal diversity and probably primary productivity. The detailed faunal analysis and the discovery of an unexpected diversity give,us a new understanding of the recovery of the Early Triassic marine ecosystem.

Paragenetic history of the Ordovician Trenton Group carbonates, Southwestern Ontario

Geochemical examination of the rock matrix and cements from core material extracted from four oil wells within southwestern Ontario suggest various stages of diagenetic alteration and preservation of the Trenton Group carbonates. The geochemical compositions of Middle Ordovician (LMC) brachiopods reflect the physicochemical water conditions of the ambient depositional environment. The sediments appear to have been altered in the presence of mixed waters during burial in a relatively open diagenetic microenvironment. Conodont CAl determination suggests that the maturation levels of the Trenton Group carbonates are low and proceeded at temperatures of about 30 - 50°C within the shallow burial environment. The Trenton Group carbonates are characterized by two distinct stages of dolomitization which proceeded at elevated temperatures. Preexisting fracture patterns, and block faulting controlled the initial dolomitization of the precursor carbonate matrix. Dolomitization progressed In the presence of warm fluids (60 75°C) with physicochemical conditions characteristic of a progressively depleted basinal water. The matrix is mostly Idiotopic-S and Idiotopic-E dolomite, with Xenotopic-A dolomite dominating the matrix where fractures occur. The second stage of dolomitization involved hydrothermal basinal fluid(s) with temperatures of about 60 - 70°C. These are the postulated source for the saddle dolomite and blocky calcite cements occurring in pore space and fractures. Rock porosity was partly occluded by Idiotopic-E type dolomite. Late stage saddle dolomite, calcite, anhydrite, pyrite, marcasite and minor sphalerite and celestite cements effectively fill any remaining porosity within specific horizons. Based on cathode luminescence, precipitation of the different diagenetic phases probably proceeded in open diagenetic systems from chemically homogeneous fluids. Ultraviolet fluorescence of 11 the matrix and cements demonstrated that hydrocarbons were present during the earliest formation of saddle dolomite. Oxygen isotope values of -7.6 to -8.5 %0 (PDB), and carbon isotope values of - 0.5 and -3.0 %0 (PDB) from the latest stage dog-tooth calcite cement suggest that meteoric water was introduced into the system during their formation. This is estimated to have occurred at temperatures of about 25 - 40°C. Specific facies associations within the Trenton Group carbonates exhibit good hydrocarbon generating potential based on organic carbon preservation (1-3.5%). Thermal maturation and Lopatin burial-history evaluations suggest that hydrocarbons were generated within the Trenton Group carbonates some time after 300 Ma . Progressively depleted vanadium trends measured from hydrocarbon samples within southwestern Ontario suggests its potential use as a hydrocarbon migration indicator on local (within an oilfield) and on regional scales.

Distribution, genesis and age of the Schlagwasser breccia (Warstein anticline, Rhenish massif)

The name "Schlagwasser breccia" is a synopsis of several debris flows in the Warstein area, which can be derived from the Warstein carbonate platform and the Scharfenberg reef. Though only locally developed, the breccia is important for the understanding of paleogeography and sedimentology in the Eastern Sauerland. Considering this breccia some gravitational-resedimentary slide movements between a high, consisting of reef carbonates, and a basin with flinz beds can be pointed out. From the uppermost Middle Devonian to the lowermost Lower Carboniferous several slides yielded the sedimentary components building up the 30 to 50 m thick polymict breccia. Some breccias were redeposited repeatedly as can be verified by different conodont maxima in single samples. Supplying area was the western part of the Warstein high, from which the slide masses glided off to the East and Southeast, more seldom to the West and Westsouthwest. All conodont zones from the upper Middle Devonian up to the lowermost Carboniferous could be identified in the Schlagwasser breccia. Therefore, an uninterrupted continuous sedimentation must have been prevalent in the supplying area; today this area nearly is denuded of flinz beds and cephalopod limestones. The slide masses spread transgressively to the East up to a substratum consisting of different units as massive limestone, flinz beds and cephalopod limestone; they are overlapped by Hangenberg beds, alum schists and siliceous rocks of the Lower Carboniferous. Parts of the substratum were transported during the progress of the slide masses. Proximal and distal parts of the flow masses can be distinguished by the diameter of the pebbles. Graded bedding and banking structures are marked only rarely. Way of transport was up to 3 km. Differently aged slide masses do not always overlap, but are placed side by side, too. Usually the slide masses do not spread out upon a greater area during sedimentation, but form closely limited debris flows. Synsedimentary fracturing and tilting of the reef platform, epirogenetic movements and seaquakes caused the slides. The entire formation period of the breccia includes about 20 millions of years. The longevity of the events points to solid paleomorphological situations around the eastern margin of the carbonate platform.

Selected Devonian/Carboniferous boundary profiles in the Sauerland area (Rhenish massif)

Selected sections, containing Devonian/Carboniferous boundary beds, are described from the northern and northeastern margin of the Rhenish massif, especially from the Seiler region near Iserlohn and the Warstein area. These sections are from prospecting trenches, quarries and road cuts. The dominantly carbonate sequences were investigated in regard to the development of conodonts. The Devonian/Carboniferous boundary could be placed precisely in both areas by means of the phylogenetic transition from Siphonodella praesulcata to S. sulcata. Compared investigations lead to the following conclusions: - The basal part of the Hangenberg limestone is heterochronous. - The Devonian/Carboniferous boundary lies distinctly below the Hangenberg limestone, i. e. at the same stratigraphical level as the Stockum limestone. - The Imitoceras limestone lens of Stockum and the Stockum limestone represent a special facies within the Hangenberg schists. 80th belong either to the praesulcata- and sulcata-zone or are restricted only to the sulcata-zone. - Protognathodus kuehni appears together with Siphonodella sulcata. Where S. sulcata is lacking, P. kuehni may be considered as a valid index conodont indicating the beginning of the Carboniferous. - The upper part of the Wocklum beds, following above the Wocklum limestone, usually consists up to the lower Carbonilerous boundary in a more or less consistent facies, that of the Hangenberg schists. Only in the section 01 the northeastern wall of the eastern Provincial Quarry at Drewer and in the road profile Rüthen - Nuttlar, the Devonian/Carboniferous boundary is to be placed in a continuous carbonate sequence. - The eastern Provincial Quarry at Drewer is therefore proposed as a new candidate section for the Devonian/Carboniferous boundary stratotype. - In many places the carbonates at the Devonian / Carboniferous boundary and the Hangenberg limestone are characterized by an impoverished conodont fauna. - Using platform conodonts, biofacies models are developed, permitting to conclude on the position of the respective setting 01 sedimentation area, either close to a rise or a basin.

Neptunian dikes and their fillings in carbonates of the Warstein area (northeastern Rhenish massif)

Neptunian dikes and cavities as weil as their fillings are described from Middle to Upper Devonian carbonates of the Warstein area. The genesis of the pre-Upper Carboniferous dikes is due to pre-orogenic synsedimentary tensional movements. Lifting, subsidence and tilting caused joints and cracks, which are enlarged to dikes and cavities on submarine conditions. The post-Upper Carboniferous dikes are based on the orogenesis during Upper Carboniferous time, causing numerous tectonical divisional planes in the sediments. Along these planes a far-reaching karstification took place since mesozoic time. According to their size the cavities are subdivided into macro-, mega- and microdikes. With the exception of one macrodike all the others are limited to the massive limestone. Megadikes especially occur in Upper Devonian cephalopod limestone and in the Erdbach limestone, microdikes can be found in all carbonatic rocks. The dikes follow pre-orogenic, tectonical and sedimentary divisional planes and are orientated to ac-, bc- as well as bedding planes and diagonal directions. The fillings happened down from above either in a solitary event or repeatedly in long-lived dikes during a span of several ten millions of years. More seldom the fillings took place laterally or upside from beneath. The dikes contain - without regard to autochthonous conodont faunas - older and/or younger mixed faunas, too. Occasionally they were used as life district by a trilobite fauna adapted to the dikes. The dikes represent sedimentary pitfalls and conserve sediments eroded in other places. Therefore, by aid of the fillings, it can be demonstrated, that stratigraphic gaps are not absolutely due to primary interruptions of sedimentation, but were caused by reworking. Some dikes contain the distal offsets of slides and suspension streams. Relations between condensation and development of dikes could not be derived in the Warstein area. However, an increase of the frequency of dikes towards east to the eastern margin of the Warstein carbonate platform could be pointed out. This margin is a slope, persisting more than 10 millions of years, between a block and a basin. Evidently cracks and dikes, which were caused by settlements, slides and earth quakes, occured there frequently. The Warstein dikes and cavities, caused by karstification, are filled with terrestrial Lower Cretaceous, marine Upper Cretaceous and terrestrial Pleistocene to Holocene sediments. Tertiary sediments could not be detected.

Palynostratigraphy of Ordovician strata, Canning Basin, Western Australia. Part One: acritarchs and prasinophytes

This paper is the initial part of a comprehensive bipartite monograph of palynomorphs (viz., acritarchs, prasinophyte phycomata, and chitinozoans) that are represented profusely in marine lower Palaeozoic strata of the Canning Basin, Western Australia. The prime aim is to establish a palynologically based zonal scheme for the Ordovician sequence as represented in five cored boreholes drilled through the Lower to Middle Ordovician strata of the central-northeastern Canning Basin. These strata embrace the Oepikodus communis through Phragmodus-Plectodina conodont zonal interval and comprise (in ascending order) the Willara, Goldwyer, and Nita formations, of inferred early Arenig to Llanvirn age. All three formations contain moderately diverse and variably preserved palynomorphs. The palynomorph taxa, detailed systematically in the current Part One of this monograph, comprise 66 species of acritarchs and six of prasinophytes. Of these, two species of prasinophytes and 11 of acritarchs are newly established: Cymatiosphaera meandrica and Pterospermella franciniae; Aremoricanium hyalinum, A. solaris, Baltisphaeridium tenuicomatum, Gorgonisphaeridium crebrum, Lophosphaeridium aequalium, L. aspersum, Micrhystridium infrequens, Pylantios hadrus, Sertulidium amplexum, Striatotheca indistincta, and Tribulidium globosum. Pylantios (typified by P. hadrus), Sertulidium (typified by S. amplexum), and Tribulidium (typified by T globosum); are defined as new acritarch genera. Three new combinations are instituted: Baltisphaeridium pugiatum (PLAYFORD & MARTIN 1984), Polygonium canningianum (COMRAZ & PENIGUEL 1972), and Sacculidium furtivum (PLAYFORD & MARTIN 1984); and Ammonidium macilentum PLAYFORD & MARTIN 1984 and Sacculidium furtivum (PLAYFORD & MARTIN 1984) are emended. An appreciable number of palynomorph species are not formally named owing to lack of sufficient or adequately preserved specimens; others are compared but not positively identified with previously instituted species. The ensuing Part Two of this study will complete the systematic-descriptive documentation, i.e., chitinozoans, and evaluate the Canning Basin palynoflora in terms of its chronological and stratigraphic-correlative significance.

Palynostratigraphy of Ordovician strata, Canning Basin, Western Australia. Part Two: chitinozoans and biostratigraphy

This second and concluding part of a comprehensive palynological study of the Lower to Middle Ordovician succession of the central-northeastern Canning Basin completes the systematic documentation of the palynomorphs, i.e., chitinozoans, and formulates a palynostratigraphic zonation scheme embracing all three constituent formations of this investigation, viz., the Willara, Goldwyer, and Nita formations. A total of 21 species of chitinozoans (five genera), detailed systematically herein, are identified. Although chitinozoan recovery per sample proved variable, the following species occur fairly persistently in the productive samples: Belonechitina micracantha, Conochitina subcylindrica, C. poumoti, C. langei, Calpichitina windjana, and Rhabdochitina magna. Five, stratigraphically successive acritarch/prasinophyte assemblage zones, ranging in age from early Arenig through late Llanvirn, are proposed as follows (ascending order): Athabascaella rossii Assemblage Zone (corresponding to the lower Willara Formation; and dated as early-mid Arenig); Comasphaeridium setaricum Assemblage Zone (upper Willara and lowermost Goldwyer; late Arenig-earliest Llanvirn); Sacculidium aduncum Assemblage Zone (lower Goldwyer; early Llanvirn); Aremorica-nium solaris Assemblage Zone (middle and lower upper Goldwyer; mid Llanvirn); and Dactylofusa striatogranulata Assemblage Zone (upper Goldwyer and lower Nita; late Llanvirn). Four chitinozoan assemblage zones, stratigraphically coinciding (within the limits of sampling) with the acritarch/prasinophyte zones, comprise (in ascending order): Lagenochitina combazi Assemblage Zone (equivalent to the A. rossii and L. heterorhabda Assemblage Zones); Conochitina langei Assemblage Zone; Conocbitina subcylindrica Assemblage Zone; and Belonecbitina micracantha Assemblage Zone. Chronostratigraphic assignments are based principally on associated conodont and graptolite faunas. Whereas the acritarch/prasinophyte zones bear scant similarities to those established globally elsewhere, the chitinozoan zones show significant affiliations with those known from Laurentia.

Late Devonian (Famennian) jaws: the icriodid multielement apparatus and its function

In this study the conodont multielement apparatus of Late Devonian (Famennian) Icriodus altematus is described which has been reconstructed from clustered group findings and separated elements. This apparatus is markedly different from classical ozarkodinid apparatuses and needs further consideration of its functional morphology. Since bedding plane assemblages of Icriodus altematus are yet unknown, a spatial reconstruction of this apparatus and a feeding mechanism are proposed which are based on the oropharyngal apparatus of recent lampreys. Though the extant representatives of petromyzontoids are not close phylogenetic relatives of extinct conodonts, there exist intriguing analogies concerning the morphology of the tooth types and the presumed spatial distribution within the oral cavity of both taxa.