Arne T. Nielsen

Devonian rise in atmospheric oxygen correlated to the radiations of terrestrial plants and large predatory fish

The evolution of Earth’s biota is intimately linked to the oxygenation of the oceans and atmosphere. We use the isotopic composition and concentration of molybdenum (Mo) in sedimentary rocks to explore this relationship. Our results indicate two episodes of global ocean oxygenation. The first coincides with the emergence of the Ediacaran fauna, including large, motile bilaterian animals, ca. 550–560 million year ago (Ma), reinforcing previous geochemical indications that Earth surface oxygenation facilitated this radiation. The second, perhaps larger, oxygenation took place around 400 Ma, well after the initial rise of animals and, therefore, suggesting that early metazoans evolved in a relatively low oxygen environment. This later oxygenation correlates with the diversification of vascular plants, which likely contributed to increased oxygenation through the enhanced burial of organic carbon in sediments. It also correlates with a pronounced radiation of large predatory fish, animals with high oxygen demand. We thereby couple the redox history of the atmosphere and oceans to major events in animal evolution.

Onset of main Phanerozoic marine radiation sparked by emerging Mid Ordovician icehouse

The Great Ordovician Biodiversification Event (GOBE) was the most rapid and sustained increase in marine Phanerozoic biodiversity. What generated this biotic response across Palaeozoic seascapes is a matter of debate; several intrinsic and extrinsic drivers have been suggested. One is Ordovician climate, which in recent years has undergone a paradigm shift from a text-book example of an extended greenhouse to an interval with transient cooling intervals – at least during the Late Ordovician. Here, we show the first unambiguous evidence for a sudden Mid Ordovician icehouse, comparable in magnitude to the Quaternary glaciations. We further demonstrate the initiation of this icehouse to coincide with the onset of the GOBE. This finding is based on both abiotic and biotic proxies obtained from the most comprehensive geochemical and palaeobiological dataset yet collected through this interval. We argue that the icehouse conditions increased latitudinal and bathymetrical temperature and oxygen gradients initiating an Early Palaeozoic Great Ocean Conveyor Belt. This fuelled the GOBE, as upwelling zones created new ecospace for the primary producers. A subsequent rise in δ13C ratios known as the Middle Darriwilian Isotopic Carbon Excursion (MDICE) may reflect a global response to increased bioproductivity encouraged by the onset of the GOBE.

A review of Ordovician agnostid genera (Trilobita)

The definition and scope of Ordovician genera of agnostid trilobites are discussed, and the generic affiliation of Ordovician species and well-illustrated material referred to in open nomenclature is assessed. Some 198 Ordovician species plus 25 species of uncertain latest Cambrian/earliest Ordovician age are enumerated, representing 24 genera and subgenera, of which Geragnostus (Novoagnostus) n.subg. and Lotagnostus (Semagnostus) n.subg. are new. Geragnostella is ranked as a subgenus of Geragnostus . The species of Arthrorhachis are tentatively divided among a tardus species group and an elspethi species group. Species of Dividuagnostus are divided among a sensu stricto group and an early species group. Pseudorhaptagnostus Lermontova, 1951 is restored as senior synonym of Neoagnostus Kobayashi, 1955. It is concluded that Ordovician agnostids were largely adapted to temperate and colder water habitats.

Nonbiomineralized carapaces in Cambrian seafloor landscapes (Sirius Passet, Greenland): Opening a new window into early Phanerozoic benthic ecology

Widespread microbial mats and the absence of significant bioturbation resulted in a poorly developed mixed layer and extensive cohesive substrates during the Ediacaran-Cambrian transition. Large nonbiomineralized arthropod carapaces overprinted with trails, interconnected burrow systems, narrow-caliber structures with dendritic terminations, and annulated burrows are abundant in the Early Cambrian Sirius Passet Lagerstatte (Greenland). Taphonomic controls were partially responsible for the pronounced association of these structures and carapaces, but ecologic conditions are envisioned as playing a significant role. Although some trace fossils (e.g., annulated structures) may be related to scavenging and/or deposit feeding, morphologic evidence suggests that this is not always the case. Interconnected burrow systems provide evidence for re-use of structures, suggesting grazing on or farming of bacteria. Sulfur bacteria were probably abundant in Cambrian sediments, requiring a source of sulfur or sulfide and limited quantities of oxygen. Anoxic sediment provided a source of hydrogen sulfide, whereas burrow outlets connected to the oxygenated sediment-water interface. Carcasses and molds of nonbiomineralized organisms may have acted as attractors, supporting a community of small invertebrates. Carapaces created a sharp boundary, further decreasing oxygen diffusion into the sediment. This steep oxygen gradient at the organic surface of the carapace promoted the growth of sulfur bacteria that could have served as the primary food for small macrofauna and meiofauna.

Upper Cambrian (Furongian) biostratigraphy in Scandinavia revisited: definition of superzones

The traditional Furongian trilobite biozones of Scandinavia, recently proposed abandoned due to inconsistent boundary definitions [Terfelt, F., Eriksson, M.E., Ahlberg, P. & Babcock, L.E., 2008: Furongian Series (Cambrian) biostratigraphy of Scandinavia – a revision. Norwegian Journal of Geology88, 73–87], are resurrected and elevated to superzonal rank. These superzones are usually readily recognized in the field, even by non-specialists, and for general correlation and mapping the more wide-ranging biozones are considerably more practicable than the very detailed zonation (formerly subzonation) introduced by Terfelt et al. (2008). Formal definition of the superzones is outlined including designation of stratotype sections. The superzones are each defined by the FAD of a characterizing species group and upwards delimited by the base of the succeeding superzone. The long used Olenus, Parabolina and Leptoplastus (super)zones as well as the recently introduced Acerocarina Superzone ( = Acerocare Zone of older literature) are maintained and formalized. The Protopeltura praecursor, Peltura minor and Peltura scarabaeoides zones are abandoned and replaced by two new units, named the Protopeltura and Peltura superzones, respectively. Accordingly, all Furongian superzones have a uniform naming style referring to a characteristic genus. The six Furongian superzones currently comprise 27 trilobite zones. The Paradoxides forchhammeri Superzone (Cambrian Series 3) is extended upwards to the base of the Olenus Superzone, thereby formally including the Agnostus pisiformis Zone.

Ecostratigraphical interpretation of lower Middle Ordovician East Baltic sections based on brachiopods

A detailed ecostratigraphical framework is established for the lower Middle Ordovician Kundan regional stage of the East Baltic area corresponding to the Asaphus expansus , A. raniceps and A. eichwaldi trilobite zones (lower Darriwilian). The study is based on approximately 6200 brachiopods collected bed by bed from limestone sections in northern Estonia (Harku Trench and Saka) and western Russia (Putilovo Quarry, Lava River canyon and Lynna River valley) with, in addition, the first detailed systematic assessment of the Kundan brachiopods of the East Baltic. These sections represent an oblique depth transect some 400 kilometres long, deepening eastwards. Five biofacies associations have been recognized using detrended correspondence and cluster analyses: a shallow-water Lycophoria association, a transitional Gonambonites association and two deeper-water associations, the soft-substrate Orthis callactis and the hard-substrate Orthambonites associations. A separate, fifth soft-substrate association is present in the marl beds at the main locality of Putilovo Quarry. The associations reflect a combination of palaeo-water depth and substrate. The biofacies facilitate an ecostratigraphical correlation along the transect, and third and fourth order sea-level curves are reconstructed, reflecting mainly eustasy. The sea-level was relatively low, early in the Kundan, but then rose significantly into the A. raniceps Biozone. This corroborates the recent discovery of possible small early Darriwilian ice caps on Gondwana.

A highly diverse trilobite fauna with Avalonian affinities from the Middle Cambrian Acidusus atavus Zone (Drumian Stage) of Bornholm, Denmark

A newly collected trilobite fauna from the lowermost part of the Alum Shale Formation at Øleå, Bornholm, Denmark, demonstrates the presence of a thin but richly fossiliferous Middle Cambrian Acidusus atavus Zone (Paradoxides paradoxissimus Superzone). The lower part of the zone (the Tomagnostus fissus–Ptychagnostus atavus Zone of older Scandinavian literature) is represented by a discontinuous limestone bed, up to 20 cm thick, whereas the upper part (Hypagnostus parvifrons Zone of older literature) is represented by shale, less than 80 cm thick, containing lenses of bituminous limestone. A total of 39 agnostid and 21 polymerid species are recorded from the A. atavus Zone (including 12 taxa treated under open nomenclature). One additional polymerid species collected from an ice-rafted boulder derived from the Bornholm area is also treated. The fauna is described and illustrated, with designation and reillustration of relevant lectotypes. The abundance of polymerid trilobites in comparison with nearby Scania, southern Sweden, where 25 agnostid and nine polymerid species have been reported from equivalent strata, is indicative of a less dysoxic environment in the Bornholm area, which was probably uplifted. The trilobite fauna resembles the coeval assemblages described from England, Wales, eastern Newfoundland and eastern Siberia, sharing several species not reported previously from Baltica, including Anopolenus sp., Agraulos longicephalus, Bailiella ornata, Clarella impar, Solenopleura? applanata, Acadagnostus aff. bulkurensis and Phalagnostus ovalis. A few faunal elements originally described from Bohemia and Australia are also present, viz. Hydrocephalus aff. carens, Phalagnostus nudus, Skryjagnostus pompeckji, Hypagnostus aff. clipeus, Euagnostus aff. interstrictus and Euagnostus? aff. glandifer.

The nature of Ordovician limestone-marl alternations in the Oslo-Asker District (Norway): witnesses of primary glacio-eustasy or diagenetic rhythms?

Ordovician limestone-marl alternations in the Oslo-Asker District have been interpreted as signaling glacio-eustatic lowstands, which would support a prolonged “Early Palaeozoic Icehouse”. However, these rhythmites could alternatively reflect differential diagenesis, without sedimentary trigger. Here, we test both hypotheses through one Darriwilian and three Katian sections. Our methodology consists of a bed-by-bed analysis of palynological (chitinozoan) and geochemical (XRF) data, to evaluate whether the limestone/marl couplets reflect an original cyclic signal. The results reveal similar palynomorph assemblages in limestones and marls. Exceptions, which could be interpreted as reflecting palaeoclimatological fluctuations, exist at the species level: Ancyrochitina bornholmensis seems to be more abundant in the marl samples from the lower Frognerkilen Formation on Nakkholmen Island. However, these rare cases where chitinozoans differ between limestone/marl facies are deemed insufficient for the identification of original cyclicity. The geochemical data show a near-perfect correlation between insoluble elements in the limestone and the marls, which indicates a similar composition of the potential precursor sediment, also in the Frognerkilen Formation. This is consistent with the palynological data. Although an original cyclic pattern could still be recorded by other, uninvestigated parameters, our palaeontological and geochemical data combined do not support the presence of such a signal.

The late Cambrian (Furongian) Acerocarina Superzone (new name) on Kinnekulle, Västergötland, Sweden

The traditional Furongian biozones of Scandinavia, based on olenid trilobites, are proposed elevated to superzonal rank. For the time being they are treated informally, except that the designation Acerocarina Superzone is proposed as replacement name for the uppermost superzone, formerly referred to as the Acerocare Zone. It comprises four zones. The lowest of these zones, hitherto referred to as the Peltura transiens Zone, is renamed as the Acerocarina granulata Zone. The distribution of the Acerocarina Superzone in Baltoscandia is mapped. A newly discovered 1.4 m thick succession on Kinnekulle, Västergötland, yielded Parabolina heres heres, P. transiens, Peltura costata, Peltura scarabaeoides westergaardi and Sphaerophthalmus alatus. In addition, A. granulata, Pelturina punctifera and Leptoplastides? sp. were collected from loose boulders deriving from the same interval. This is the first unambiguous record of P. costata and P. punctifera in Sweden. The fossil assemblage, assigned to the P. costata Zone, contains species hitherto considered characteristic of five different biozones, viz. the Ctenopyge tumida Zone, the Parabolina lobata Zone, the P. transiens Zone, the P. costata Zone and the Westergaardia scanica Zone. P. transiens, P. scarabaeoides westergaardi and S. alatus are assumed to have been reworked, whereas P. punctifera appears one zone earlier than in Norway. Parabolina (Parabolina) heres lata, not found during this study, may turn up in the P. costata Zone or even earlier. The Acerocarina Superzone is resting on the P. lobata Zone, and the intervening zone, characterized by Peltura paradoxa (uppermost “Peltura scarabaeoides” superzone), is missing in the succession on Kinnekulle.

The Middle Cambrian Paradoxides paradoxissimus Superzone on Öland, Sweden

Trilobites from the Middle Cambrian Paradoxides paradoxissimus Superzone on Öland are reviewed, including species found in glacial erratic boulders in Germany and Denmark. The fauna recorded contains 20 species including 10 agnostids. The succession, up to 70 m thick, comprises the Äleklinta Member of the Borgholm Formation (previously known as the Paradoxissimus sandstone or siltstone) and a thin overlying conglomerate, here informally referred to as the Mörbylilla conglomerate, forming the base of the Alum Shale Formation. The thin Granulata Conglomerate, including a limestone layer previously correlated with the Exsulans Limestone, forms the base of the Äleklinta Member. It has yielded several species characteristic of the Triplagnostus gibbus Zone, e.g. Ctenocephalus exsulans, Bailiella tenuicincta, Solenopleura parva, Parasolenopleura aculeata and T. gibbus. The Äleklinta Member contains T. gibbus, Ellipsocephalus lejostracus and P. aculeata, all indicative of the T. gibbus Zone. The overlying Mörbylilla conglomerate contains reworked (?) specimens of E. lejostracus and T. gibbus associated with Acidusus atavus, Tomagnostus fissus, Ptychagnostus affinis, Onymagnostus hybridus, Tomagnostella cf. truncata, Hypagnostus parvifrons and H. mammillatus. Most of these agnostids are found in small stinkstone pockets within the conglomerate. Tomagnostus bothrus? is recorded in Scandinavia for the first time. The fauna shows that the conglomerate represents the A. atavus Zone. The stinkstone pockets indicate that the conglomerate was deposited under dysoxic “Alum Shale” conditions. In the beach section at Mörbylilla, the 0.16 m thick Mörbylilla conglomerate is overlain by an anthraconite bed, up to 0.6 m thick, representing the Exporrecta Conglomerate Bed.