Chitinozoan biostratigraphy of the Upper Ordovician Fågelsång GSSP, Scania, southern Sweden

In the Fågelsång section, the new Global Stratotype Section and Point (GSSP) for the base of the Upper Ordovician Series, 22 samples from the E14 (a, b and c) and E15 outcrops have been studied for chitinozoans. They yield rich and rather diverse species assemblages of this microfossil group. The approximately 16-m-thick sampled part of the section can be subdivided, from bottom to top, into two biozones and a subbiozone: the Laufeldochitina stentor zone, the Eisenackitina rhenana subzone and the Lagenochitina dalbyensis zone. The boundary between the lowest biozone and the subzone is situated 1.7 m below the marker “Fågelsång Phosphorite” bed, thus only slightly lower (0.3 m) than the base of the Nemagraptus gracilis graptolite biozone, which defines the base of the Upper Ordovician Series. The base of the L. dalbyensis zone is located just above the Fågelsång Phosphorite, remarkably low compared to the graptolite biostratigraphy. It is evident that the chitinozoan biozonation provides an additional tool to recognise the base of the Upper Ordovician in this section. Furthermore, a distinct faunal change is shown across the phosphorite bed, possibly indicating a hiatus.     

Revision of the Zone 13 graptolite biostratigraphy in the Marathon, Texas, standard succession and its bearing on Upper Ordovician graptolite biogeography

One of the long-standing problems in North American graptolite biostratigraphy is the distinct differences in assemblages of post-Climacograptus bicornis age between the classical graptolite sequences in the New York - Quebec and Marathon, west Texas, regions. These have been attributed either to faunal provincialism or to the presence of a major hiatus between the Woods Hollow and Maravillas formations in Texas. New collections from the key Marathon Picnic Grounds section contain diagnostic Late Ordovician graptolites that confirm the existence of a major stratigraphic gap below the Maravillas Formation. The lower Maravillas Formation (Zone 13) has a Late Ordovician, low-diversity Pacific Province graptolite fauna that includes the biostratigraphically diagnostic species Climacograptus nevadensis, C. tubuliferus, Orthograptus fastigatus and Dicellograptus ornatus. Zone 13 graptolite assemblages from the Marathon region correlate with the C. tubuliferus to D. ornatus zones in the Trail Creek, Idaho, succession, the Ea4-Bo2 interval in Victoria, the O. fastigatus Zone in the Canadian Arctic Islands, the O. quadrimucronatus to D. ornatus zones of the Canadian Cordillera, and the D. complanatus to D. anceps zones in Scotland. The hiatus between the Woods Hollow and Maravillas formations spans an interval corresponding to at least the Eal Ea3 interval in Australia, the C. americanus to upper A. manitoulinensis zones in the New York - Quebec succession, and the D. clingani and P. linearis zones in Scotland. These results agree with the magnitude of the hiatus previously indicated by conodont biostratigraphy. Late Ordovician graptolite distribution patterns in North America can be explained by an extension of Cooper, Fortey & Lindholm's (1991; Lethaia 24) Lower Ordovician graptolite biofacies model into the Upper Ordovician, which incorporates both lateral water-mass specificity and depth stratification. Using this model, we recognize in Laurentia two separate biofacies among tropical-zone Late Ordovician Pacific Province graptolite faunas, a cosmopolitan Oceanic biofacies, and a cratonic Laurentian biofacies. The lower Maravillas Formation graptolite fauna is clearly part of the Oceanic biofacies, whereas the coeval Appalachian faunas represent the Laurentian biofacies. □Graptolites, Ordovician, biostratigraphy, Texas, biofacies, biogeography.     

The Upper Tremadocian (Ordovician) graptolite Bryograptus: taxonomy, biostratigraphy and biogeography

Abstract:  The taxonomy, biostratigraphical and palaeogeographical distribution of the Lower Ordovician graptolite genus Bryograptus is evaluated. Bryograptus is recognized as a distinct triradiate anisograptid with a multiramous, pendent rhabdosome. The species of the genus Bryograptus can be interpreted as shallow water faunal elements with a strongly limited biogeographical distribution to the Atlantic Faunal Realm. Bryograptus is restricted to a narrow interval in the Upper Tremadocian, the Bryograptus Biozone of Scandinavia and South America (Argentina), making it a taxon with a high potential for precise biostratigraphical correlations. The proximal end development can be used to differentiate the genus Bryograptus from other pendent multiramous graptoloid genera with a homoplastic rhabdosome development. Characteristics of the proximal end development and structure easily differentiate these genera in relief specimens, but not in flattened material.     

Changing patterns of sediment accumulation in a small lake in Scania,southern Sweden

Bioassays with water of eutrophic Lake Tjeukemeer were carried out in the laboratory, and, in the lake itself, by placing plexiglass tubes (? 9 cm) firmly in the mud. In the laboratory nitrogen had a strong growth-promoting effect on the concentrations of chlorophyll a, N-cell and COD-cell. In the bioassays in situ nitrogen increased the concentration of chlorophyll a, but not those of cellular N and cellular COD. It is argued that chlorophyll a is not a good indicator for algal growth in bioassays and that the extrapolation of laboratory experiments — where nitrogen limited algal growth — to the field situation — where light limited algal growth — often leads to erroneous results, especially in shallow eutrophic lakes. The main reason is probably that in bioassays in the laboratory the nutrient supply — both from external loading and from sediments — is often different and that processes like denitrification occur in the lake, but not in the bioassays.     

A refined graptolite biostratigraphy for the late Ordovician–early Silurian of central Wales

Morphometric analysis of graptolites from the persculptus and acuminatus biozones of central Wales identifies four successive morphospecies of normalograptids. These graptolites can be used for biostratigraphical subdivision of these strata as follows: (i) an early persculptus Biozone interval containing broad forms with geniculate thecae that have the morphology of Persculptograptus persculptus with an early insertion point for the full median septum (theca 1¹); (ii) a supra‐adjacent level of early persculptus Biozone age, with narrower, parallel‐sided forms, that have been referred to as Normalograptus? aff. parvulus and have a slightly later insertion point for the full median septum (theca 1²); (iii) a third interval, encompassing the later part of the persculptus Biozone to the early acuminatus Biozone, with Normalograptus? cf. parvulus, which has the full median septum delayed to the level of theca 3–7; and (iv) a younger interval, in the mid‐acuminatus Biozone with Persculptograptus cf. persculptus specimens that do not display the median septum on its reverse side. These taxa can be used for refined biostratigraphy and correlation in the late Ordovician and early Silurian of central Wales. The progressive delay in the insertion of the median septum in these taxa may have wider application for the correlation of the interval immediately after the Hirnantian glacial maximum.     

The Rhaetian flora of Rögla,northern Scania,Sweden

Abstract: Rögla is the northernmost locality yielding Mesozoic plant fossils in Scania, southern Sweden, and is one of the northernmost Rhaetian assemblages in Europe. The assemblage consists of over 500 specimens collected 50–60 years ago, of which 139 yielded identifiable plant remains referable to 15 plant species; another 19 specimens are tentatively assigned to four species because of their fragmentary preservation. The flora includes sphenophytes, ferns, cycads, bennettitaleans, seed ferns of uncertain alliance, conifers and some leaf remains that are tentatively assigned to ginkgophytes based on their epidermal anatomy. The species‐level composition of the assemblage is consistent with a Rhaetian age and is similar to well‐known floras from nearby Höganäs and Bjuv, except for the absence of cycads belonging to Nilssonia, which are very common in most other Scanian floras. The fossil assemblage is interpreted to derive from multi‐storey vegetation occupying moist habitats on a coastal plain. Strong affinities are evident with the coeval floras of Jameson Land, Greenland, reinforcing the concept of a distinctive North Atlantic floristic sub‐province at the close of the Triassic.     

Cambrian high-resolution biostratigraphy and carbon isotope chemostratigraphy in Scania, Sweden: first record of the SPICE and DICE excursions in Scandinavia

A core drilling (Andrarum‐3), from the classical locality at Andrarum, Scania, southernmost Sweden, penetrated a 28.90‐m‐thick Cambrian succession. The core comprises dark grey to black, finely laminated mudstones and shales with early concretionary carbonate lenses (stinkstones or orsten) and a few primary carbonate beds. The middle Cambrian (provisional Series 3) part of the core comprises 17.35 m, whereas the Furongian Series (upper Cambrian) part covers the remaining 11.55 m. Nineteen trilobite and two phosphatocopine genera are present in the middle Cambrian, whereas the less diverse Furongian interval yielded four trilobite and three phosphatocopine genera. Other, less frequent, faunal elements include conodonts (s. l.), brachiopods, sponge spicules, bradoriids, and coprolites. Trilobites and phosphatocopines were used to subdivide the core into seven biozones ranging from the Ptychagnostus atavus Zone to the Parabolina spinulosa Zone (P. spinulosa Subzone). Carbon isotopic analyses (δ¹³C_org) through the core show two important excursions, the negative DrumIan Carbon isotope Excursion (DICE) in the Pt. atavus Zone, and the Steptoean Positive Carbon Isotope Excursion (SPICE) beginning near the first appearance of Glyptagnostus reticulatus and extending upward into the Olenus and Agnostus (Homagnostus) obesus Zone. The DICE displays a peak value, in the samples at hand, of –30.45‰δ¹³C_org in the lower part of the P. atavus Zone. The δ¹³C_org values increase through the overlying L. laevigata and A. pisiformis zones and display peak values of c. –28.00‰δ¹³C_org in the lowermost Furongian Olenus wahlenbergi and O. attenuatus subzones. Thereafter the values decrease significantly through the O. scanicus Subzone. Both isotopic excursions have been documented from several palaeocontinents, but never before from Baltica. Moreover, for the first time these excursions are recorded from organic matter in an alum shale setting. The recorded shift of +1.50–2.00‰δ¹³C_org is approximately half the magnitude of the SPICE documented from other regions. This discrepancy may be related to temporal variations in the type, origin, or diagenesis of the organic fraction analysed.     

Stratigraphical distribution of the Ordovician graptolite Azygograptus Nicholson & Lapworth in the Central Andean Basin (northwestern Argentina and southern Bolivia)

We analyzed new occurrences of Azygograptus lapworthi from the Cordillera Oriental, Argentina. The bearer sandstones levels, corresponding to the Acoite Formation, are overlying the deposits, in which the Didymograptellus bifidus Biozone (Lower Ordovician, upper Floian, Fl3) was previously recognized, and are overlain by younger pelitic levels yielding Xiphograptus lofuensis (Middle Ordovician, early Dapingian, Dp2). Previous records from the Central Andean Basin are also reviewed in detail and accurately correlated, allowing us to conclude that the Azygograptus lapworthi Biozone corresponds to the Middle Ordovician (lower Dapingian, Dp1). This biostratigraphic framework documents that the transition between the Lower and Middle Ordovician deposits occurs in the uppermost levels of the Acoite Formation in the Argentine Cordillera Oriental. It is additionally integrated with up to date conodont records to establish a high-resolution regional correlation, with equivalent deposits from the Puna of northwestern Argentina and Cordillera Oriental of Bolivia, and to discuss new insights for global correlation.     

The identity of the Ordovician (Darriwilian) graptolite Fucoides dentatus Brongniart, 1828

Abstract: The biostratigraphically important Middle Ordovician (Darriwilian) biserial graptolite Fucoides dentatus Brongniart, 1828 is redescribed and illustrated from its type material and from additional specimens collected at the type locality – Lévis, Quebec, Canada. It is referred to Levisograptus gen. nov., which includes also the austrodentatus group of early axonophoran graptolites. The species has previously been confused with a younger, mid‐Darriwilian species, now referred to as Eoglyptograptus gerhardi sp. nov., and recognized as the type species of the genus Eoglyptograptus Mitchell. Both species can be differentiated easily by their respective proximal development types and show nonoverlapping biostratigraphical ranges. Levisograptus dentatus (Brongniart) is an important biostratigraphical index species in the early Darriwilian. Eoglyptograptus species are found in the higher Darriwilian and are biogeographically restricted to the Atlantic Faunal Realm.     

Proximal structure and development in the Ordovician graptolite Parisograptus Chen and Zhang, 1996

The proximal development of Parisograptus Chen and Zhang is described from three–dimensionally preserved specimens. The unique development features an origin of proximal thecae like a string of pearls vertically upon each other on the reverse side of the rhabdosome with the dorsal sides of the initial stipes placed side by side. The development differs strongly from that found in the superficially similar Arienigraptus in which the first thecal pairs grow downwards side by side, even though the rhabdosome shapes are quite similar. It represents a first step towards the development of a completely biserial rhabdosome and eventually leads to the biserial, monopleural glossograptids.     

The phosphorus economy of a hypertrophic seepage lake in Scania,south Sweden groundwater influence

The phosphorus dynamics and economy of Lake Bysjön, a hypertrophic seepage lake in Scania, southern Sweden, were investigated during 1973–1977. The mean dissolved inorganic phosphorus concentration (1973–1977) was 580 µg · l^–1. There were no correlations between dissolved inorganic P, total organic P, dissolved organic P, particulate P and phytoplankton biomass. Groundwater inflow and lake water outflow through the ground are the most important factors for maintaining a constant water volume. Groundwater seepage is also important for water quality. Groundwater inflow, together with planktonic activity, keeps the P concentration high in the lake water.     

Ordovician conodont biostratigraphy of the Willara Formation in the Canning Basin,Western Australia

The conodont fauna from the Willara Formation, a carbonate-dominated stratigraphic unit widely distributed in the subsurface Canning Basin of Western Australia, is represented by 41 species, including a new species, Erraticodon neopatu Zhen n. sp. The Jumudontus gananda and Histiodella altifrons biozones are recognized in the lower and upper parts, respectively, of the Willara Formation. Deposited primarily in shallow nearshore settings, the Willara Formation is characterized by the occurrence of predominantly long-range coniform species of Triangulodus, Scalpellodus, Drepanoistodus, Drepanodus, and Kirkupodus. Several widely distributed age-diagnostic species, including Histiodella altifrons, Histiodella holodentata, Histiodella serrata, and Jumudontus gananda, serve as keys for biostratigraphic analysis and correlation. Our study also shows that the basal and top boundaries of the Willara Formation are diachronous across the basin, extending from the middle Floian (Oepikodus communis Biozone) to middle Darriwilian (Histiodella holodentata-Eoplacognathus pseudoplanus Biozone). This contribution provides crucial new biostratigraphic data for precise correlation of the Willara Formation with its time equivalents regionally and internationally.     

Lunnomidinium scaniense Lindström, gen. et sp. nov., a new suessiacean dinoflagellate cyst from the Rhaetian of Scania, southern Sweden

The dinoflagellate cyst Lunnomidinium scaniense gen. et sp. nov. is present in the lower part of a thin sequence of Rhaetian sedimentary rocks exposed in the Lunnom Coal and Clay Pit in NW Scania, southern Sweden. It occurs in diverse, Rhaetian palynomorph assemblages, dominated by spores and pollen, but with rare specimens of the dinoflagellate cysts Rhaetogonyaulax rhaetica (Sarjeant) Loeblich and Loeblich 1968, Shublikodinium sp. and Beaumontella? caminuspina (Wall) Below 1987. Lunnomidinium scaniense is characterized by an epicystal {tAtItP} archeopyle, a large number of paraplates arranged in seven or eight latitudinal series, and intratabular ornamentation in some but not all of the paraplate series. Thus, L. scaniense is assignable to the family Suessiaceae. Lunnomidinium scaniense can be subdivided into two different morphological varieties, based on the autophragm ornamentation and cyst size.     

Orthoconic cephalopods and associated fauna from the late Ordovician Soom Shale Lagersta¨tte, South Africa

ABSTRACT. Orthoconic cephalopods from the Soom Shale Member (Ashgill) are exceptionally preserved and are colonized by lingulate brachiopods and cornulitids. Other fossils commonly associated with orthocones include myodocopid ostracodes and chitinozoans. Size distribution analysis of the brachiopods on one orthocone indicates that it was colonized in vivo. Four orthocone radulae are preserved extending the record of these structures 50 My back to the late Ordovician. Orthocone radula configuration is more similar to that of ammonoids and coleoids than to that of nautiloids.     

Bioerosional innovation for living in carbonate hardgrounds in the Early Ordovician of Sweden

Some of the world's oldest macroborings occur in hardgrounds in lower Ordovician (Arenig) limestones exposed on the island of Öland, southern Sweden. The trace fossils, which are described here as Gastrochaenolites oelandicus isp. nov., appear to be dwelling structures excavated in the indurated substrate by invertebrates of unknown taxonomic affinity. They are the oldest examples of this ichnogenus. The appearance of a macroboring life habit at this early time represents a revolutionary new adaptive strategy for inhabiting carbonate hardgrounds. However, this innovative strategy apparently was not successful for the long term, because this particular macroboring taxon seems to have disappeared shortly after its early Ordovician appearance.     

The earliest myodocopes: ostracodes from the late Ordovician Soom Shale Lagerstätte of South Africa

The late Ordovician Soom Shale Lagerstätte of South Africa has yielded Myodoprimigenia fistuca n. gen. and n. sp., the earliest and only known Ordovician occurrence of myodocopes, one of the major groups of ostracodes. M. fistuca is a likely sister group of the Upper Silurian 'cypridinid' myodocopes and allied forms. It had a thin, lightly mineralized and flexible shell with microstructures resulting from in vivo calcification processes. It probably fed on cephalopod carrion, thus extending evidence for a carnivorous scavenging lifestyle in ostracodes back by 200  Ma. The species was probably nektobenthic and thus consistent with the notion that the origin of the late Silurian pelagic myodocopes - and therefore of pelagic ostracodes - is to be charted in a benthic to pelagic ecological shift in the group.     

The influence of denitrifying seawater on graptolite extinction and diversification during the Hirnantian (latest Ordovician) mass extinction event

A continuous trench exposure within the uppermost type Vinini Formation at Vinini Creek, Roberts Mountains, Nevada, provides an unparalleled opportunity to examine the fate of graptolites, prominent Paleozoic zooplankton, during most of the Hirnantian mass extinction event. On the basis of a detailed biostratigraphic and sedimentological dataset, the relatively complete extinction record is examined in the context of ecological constraints, and it is found to reflect an ecological collapse driven by glacio-eustatic sea-level fall and associated changes in oceanic circulation. Diverse graptolite populations of the Dicranograptidae-Diplograptidae-Orthograptidae (DDO) fauna, which flourished in denitrifying waters within the oceanic oxygen-minimum zone (OMZ) during sea-level highstand, largely vanished with the loss of these conditions during glacio-eustatic sea-level fall. However, populations of one clade, the normalograptids, which inhabited the oxygenated waters of the photic zone, not only survived but diversified. These survivors gave rise to rapid recolonization and diversification with re-establishment of the oxygen-minimum and denitrifying conditions during post-Hirnantian sea-level rise. This ecological model also applies globally to other well-documented coeval stratigraphic intervals, representing both oceanic and platform sea settings.     

Late Ordovician palaeoceanographic changes as reflected in the Hirnantian–early Llandovery succession of Jämtland, Sweden

A study of the Upper Ordovician–Lower Silurian strata in Jämtland, central Sweden, shows that large-scale changes in shelf deposition took place close to the systems boundary. These changes include unconformity development and the replacement of a siliciclastic shelf with a carbonate-dominated shelf, suggesting the interaction of allocyclic controls such as changing eustatic sea-level and climate. The 6-m-thick Ede Formation is a key lithosome for interpretation of this transition. Its sediments were deposited in the Caledonian foreland basin, situated east of the closing Iapetus Ocean on the western margin of the Baltic craton. A major part of the late Caradoc to late Ashgill (into the Hirnantian) was characterised by continuous and uniform deposition over wide areas (Kogsta Formation), whereas erosional surfaces and complex lateral facies relationships characterise the Ordovician–Silurian boundary strata (Ede Formation and lateral equivalents). The Ede Formation represents the end of terrigenous deposition, which in the middle Aeronian was followed by regional expansion of carbonate deposition (Berge Formation). A syn-sedimentary erosional surface, with at least 1 m of relief locally, forms the lower boundary of the Ede Formation. This surface is overlain by two types of conglomerate. Lower parts of the Ede Formation consist of medium to thick-bedded quartzites. A second erosional surface with only minor (few centimetres) relief occurs on top of these quartzites. The upper parts of the Ede Formation consist of a thin, basal favositid biostrome overlain by thin bedded, calcareous sandstones, limestones and intensely bioturbated shales. Analysis of stratigraphic boundaries and the facies succession suggests that the lower Ede Formation represents a major downward shift in coastal onlap and by-pass sedimentation that created the lower erosional surface. The erosional surface in the middle of the Ede Formation is inferred to have formed during the subsequent maximum lowstand or as a ravinement surface, and is interpreted as an unconformity. The succession is subdivided into four facies associations, each corresponding to a specific systems tract: (a) a Shale–Siltstone Association (uppermost Kogsta Formation), deposited during a highstand situation in mid-outer shelf areas; (b) a Quartzite Association (the lower Ede Formation), deposited during forced regression in a shoreface environment; (c) a Mixed Carbonate–Siliciclastic Association (the upper Ede Formation), deposited during transgression in a wave-dominated, proximal shelf environment when clastic supply was reduced; and (d) a Micritic Limestone Association (lowermost Berge Formation), deposited during a second highstand situation in a low-energy, offshore environment.

Conodont data, together with a previously reported Hirnantia fauna, constrain the position of the Ordovician–Silurian boundary to the lower 1.65 m of the Ede Formation, or less likely, to the uppermost metre of the underlying Kogsta Formation, i.e., within a 2.65-m-thick uncertainty interval. The base of the Berge Formation is about 4 m above the top of the uncertainty interval, and is dated as being mid-Aeronian in age, suggesting condensation and/or a hiatus close to, or at, the Ordovician–Silurian boundary. These data tie the unconformity and the regional facies change from a siliciclastic to a carbonate-dominated shelf to Late Ordovician–Early Silurian eustatic and climatic changes.