Biofilm Harvesters in Coastal Settings of the Early Palaeozoic

The ichnogenera Syringomorpha and Daedalus are here interpreted as products of infaunal biofilm harvesters. This study investigated: (1) Syringomorpha nilssoni and Syringomorpha isp. from the Cambrian Series 2‐Miaolingian Campanario Formation, northwest Argentina; and (2) Daedalus halli from the Floian Grès et Schistes de la Cluse de l’Orb Formation, Montagne Noire, France. Syringomorpha nilssoni occurs in sandy to mixed intertidal to lower shoreface deposits, whereas Syringomorpha isp. in the lower intertidal zone. Daedalus halli occurs in a lagoon and intertidal to lower shoreface sands of a barrier island. Syringomorpha and Daedalus comprise a vertical J‐shaped causative burrow and deep spreite. These ichnotaxa form monospecific assemblages (bioturbation index BI = 3–5) in quartzose medium‐ to fine‐grained sandstone, recording colonization in high‐energy tide‐and wave‐dominated settings. Lower abundances (BI = 1–2) are observed in silty sandstone. The abundance of both ichnogenera in mature sandstone is inconsistent with a classic deposit‐feeding strategy because ‘clean’ sediments are commonly impoverished of organic detritus, this being particularly true in Cambro‐Ordovician littoral settings lacking terrestrial plant detritus. Based on morphology, host sediment properties and comparison with modern structures, such those produced on intertidal and shallow subtidal setting by Arenicola marina and Paraonis fulgens, it is suggested that the diet of Syringomorpha and Daedalus producers may have consisted of biofilms colonising sand grains, associated eukaryotic microbes, and possibly meiofauna. Whereas Syringomorpha is a product of the Cambrian explosion, Daedalus is associated with the Ordovician Radiation. In contrast to most ichnotaxa, which display long temporal ranges, these two ichnogenera are restricted to the Cambrian and Ordovician‐Silurian, respectively. The underlying reasons for the relatively restricted stratigraphic ranges of these ichnotaxa are unclear, but space competition, and increased predation pressure may have played a role. The feeding strategy of the Daedalus and Syringomorpha producers was less efficient than suspension feeding and passive predation, trophic types epitomized by the dominant macroinfauna that persisted in water‐agitated nearshore sands during the rest of the Phanerozoic

Sedimentology and Ichnology of Paleozoic Estuarine and Shoreface Reservoirs, Morrow Sandstone, Lower Pennsylvanian of Southwest Kansas, USA

Integration of facies and trace-fossil evidence tests and refines depositional models constructed solely on the basis of physical sedimentology. In recent years, the petroleum industry has increasingly used trace-fossil analysis of cores as an aid in reservoir characterization. In particular, ichnologic data have been instrumental in the recognition of estuarine deposits and their distinction from open-marine facies (e.g., MacEachern and Pemberton, 1994). Previous ichnologic analyses of cores, however, have concentrated on post-Paleozoic reservoirs (e.g., Bockelie, 1991; Pemberton, 1992; Taylor and Gawthorpe, 1993; Howell et al., 1996; Martin and Pollard, 1996; MacEachern and Pemberton, 1997). The present study represents one of the first attempts to apply trace-fossil analysis to cores from Paleozoic reservoirs. The Lower Pennsylvanian Morrow Sandstone contains oil and gas reservoirs in a wide variety of shallow and marginal-marine depositional environments. Delta-front, shoreface, and estuarine valley-fill reservoir sandstones are encased in offshore and estuarine mudstones (Sonnenberg, 1985; Krystinik and Blakeney, 1990; Sonnenberg et al., 1990; Wheeler et al., 1990). An integrated stratigraphic, sedimentologic, and ichnologic study provides a more accurate characterization of reservoir facies and geometry. This study allows distinction between marine-shoreface and estuarine valley-fill sandstones from four cores of the lower Morrow in southwestern Kansas. Core analysis subsequently was integrated with well-log information. Previous studies have emphasized the presence of estuarine valley-fills in the upper Morrow (Wheeler et al., 1990). Our integrated approach extends the estuarine valley interpretation into the lower Morrow. Within the midcontinent, trace fossils are useful in distinguishing different facies in estuarine incised valleys and marine shorefaces. Detailed study of biogenic structures provides high-resolution information to solve problems in facies, stratigraphic, and reservoir modeling. In some cases, they represent the only evidence available to develop a reasonable picture of depositional conditions and to estimate reservoir heterogeneity. The present study provides a detailed analysis of the sedimentary facies, documents the associated trace fossils, and illustrates how trace fossils are used to refine environmental interpretations of the lower Morrow sandstone reservoirs

Trace fossils, water table and depositional evolution in eolian systems (Cretaceous Mulichinco Formation, subsurface of western Argentina)

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Continental invertebrate and plant trace fossils in space and time: State of the art and prospects

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Late Cambrian – Tremadocian faunas and events from Angosto del Moreno Section, Eastern Cordillera, Argentina

The Santa Victoria Group (SVG, late Upper Cambrian – Caradocian) comprises pre–Ashgillian Ordovician deposits of the Argentinean Eastern Cordillera. The most significant section of the SVG in the western flank of the Eastern Cordillera is located in the Angosto del Moreno area (Figure 1a, b). At this locality, the SVG unconformably overlies the Mesón Group (Cambrian s.l.), and unconformably underlies Cretaceous rocks (Yacoraite Formation). Upper Cambrian to lower Lower Ordovician units are separated from upper Lower to Middle Ordovician units (Parcha and Sepulturas formations) by the Tumbaya unconformity (Figure 2). The Angosto del Moreno Section of the SVG is exceptional in terms of the quality of exposures, continuity of deposits, richness of fossils, and accessibility. Diverse aspects concerning the Ordovician geology of this study area have been discussed by Moya et al. (1994, 1998), Moya and Albanesi (2000), Moya and Monteros (2000), Malanca and Brandán (2000), and Gómez Martínez et al. (2002). Previous data and recent paleontological collections enable a preliminary biostratigraphic scheme (Figure 2) for the Upper Cambrian to Tremadocian units of the SVG. A synthesis of the sequence stratigraphy and depositional environments of these units is given by Buatois et al. (this volume).Fil: Moya, Maria Cristina. Universidad Nacional de Salta. Consejo de Investigacion; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Salta; ArgentinaFil: Malanca, Susana. Universidad Nacional de Salta. Consejo de Investigacion; ArgentinaFil: Monteros, Julio A.. Universidad Nacional de Salta. Consejo de Investigacion; ArgentinaFil: Albanesi, Guillermo Luis. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas, Físicas y Naturales. Museo de Paleontología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba; ArgentinaFil: Ortega, Gladys del Carmen. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas, Físicas y Naturales. Museo de Paleontología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba; ArgentinaFil: Buatois, Luis Alberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Tucumán. Instituto Superior de Correlación Geológica. Universidad Nacional de Tucumán. Facultad de Ciencias Naturales e Instituto Miguel Lillo. Departamento de Geología. Cátedra Geología Estructural. Instituto Superior de Correlación Geológica; Argentin

The Angosto del Moreno area, Eastern Cordillera, Jujuy province

Ordovician and Silurian rocks exposed in the western belt of the Eastern Cordillera of Jujuy Province will be discussed during the journey through the provincial road 16, which connects the Purmamarca Village with the Angosto del Moreno locality. Following stops refer to diverse geological aspects of the region.Fil: Moya, Maria Cristina. Universidad Nacional de Salta. Consejo de Investigacion; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Salta; ArgentinaFil: Malanca, Susana. Universidad Nacional de Salta. Consejo de Investigacion; ArgentinaFil: Monteros, Julio A.. Universidad Nacional de Salta. Consejo de Investigacion; ArgentinaFil: Albanesi, Guillermo Luis. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas, Físicas y Naturales. Museo de Paleontología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba; ArgentinaFil: Ortega, Gladys del Carmen. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas, Físicas y Naturales. Museo de Paleontología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba; ArgentinaFil: Buatois, Luis Alberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Tucumán. Instituto Superior de Correlación Geológica. Universidad Nacional de Tucumán. Facultad de Ciencias Naturales e Instituto Miguel Lillo. Departamento de Geología. Cátedra Geología Estructural. Instituto Superior de Correlación Geológica; Argentin

The ichnologic signature of deep-sea colonization during the Ordovician radiation

The fossil record of deep-marine environments is notoriously poor in comparison with that of their shallow-marine counterparts. Notably, deep-marine deposits are typically host to diverse and abundant trace-fossil assemblages, providing evidence of the ancient deep-sea benthos. To analyze the early colonization of the deep sea, we constructed a global dataset of trace-fossil occurrences from a survey of EdiacaranâDevonian stratigraphic units. This analysis highlights the importance of the Ordovician radiation as a pivotal time in the colonization of the deep sea. Ediacaran deep-marine trace fossils consist of very simple trails and burrows. Global and alpha ichnodiversity, as well as ichnodisparity, were extremely low. Nonspecialized grazing trails reveal the exploitation of microbial mats. These strategies persisted in the Cambrian, although with an increase in ichnodiversity (both global and alpha) and ichnodisparity. An increase in the complexity of morphologic patterns, as illustrated by the undermat mining ichnogenus Oldhamia, is apparent during the Cambrian. The face of the deep sea started to change during the end of the Cambrian and beginning of the Ordovician with the protracted expansion of farming and trapping strategies. The main architectural designs of deep-marine trace fossils (e.g. regular networks, delicate spiral burrows, guided meandering graphoglyptids) were established in the deep sea by the Early Ordovician, recording the first appearance of the Nereites Ichnofacies. Lower to Middle Ordovician deep-marine ichnofaunas are moderately diverse, and fodinichnia commonly dominates rather than graphoglyptids. A significant ichnodiversity and ichnodisparity increase occurred in the Late Ordovicianâearly Silurian, with ichnofaunas recording higher proportions of graphoglyptids and evidencing the establishment of a deep-marine ecosystem of modern aspect. The distinction between the Nereites and Paleodictyon ichnosubfacies, with the former characterized by the dominance of feeding traces in muddy turbidites and the later by the dominance of graphoglyptids in sandy turbidites, can also be tracked back to the Ordovician radiation. This trend of increased colonization of the deep sea continued through all the Silurian and the Devonian. However, colonization of carbonate turbidites may have lagged behind that of siliciclastic turbidites. The progressive increase in abundance and diversity of graphoglyptids resulted in an increased role of gallery biodiffusers. This faunal turnover in the deep sea was coincident with an increase in oxygenation in slope and base-of-slope settings, which is thought to have been a driver of Ordovician biodiversifications. The formation of permanent open burrows in the deep sea may have increased bioirrigation in the uppermost zone of the deep-sea sediment, therefore increasing ventilation and potentially generating a feedback loop between bioturbation and oxygenation, with the endobenthos engineering its environment

Sedimentology and ichnology of the middle interval of the Agua del Jagüel Formation, Carboniferous of Mendoza Precordillera: paleoenvironmental implications of the postglacial transgression

Se describen las características paleoambientales e icnológicas del tramo medio de la Formación Agua del Jagüel (Carbonífero, Precordillera de Mendoza). Fueron reconocidas cinco facies agrupadas en tres asociaciones de facies sedimentarias, las que caracterizan la evolución de un sistema fluvio-estuarino. La asociación de facies 1, formada por conglomerados y areniscas conglomerádicas de origen fluvial, se dispone sobre una superficie erosiva que los separa de niveles postglaciales correspondientes a la parte basal de la Formación Agua de Jagüel. La asociación de facies 2 incluye en su base a depósitos finos (fangolitas) que indican la inundación del sistema, seguida por sedimentación arenosacorrespondiente a ambientes de boca de estuario indicando una progresiva transgresión dentro del estuario. La asociación de facies 3 corresponde a dos intervalos de areniscas36 muy gruesas fluviales, separados por areniscas muy finas y fangolitas depositadas durante un breve evento transgresivo. Paralelamente a los estudios sedimentológicos se analizaron las estructuras biogénicas provenientes de fangolitas y areniscas muy finas a finas de la facies 2. Las trazas fósiles reconocidas incluyen a Cochlichnus anguineus, Gordia marina, Helminthoidichnites tenuis, Helminthopsis tenuis, Mermia carickensis, Treptichnus bifurcus,T. pollardi, y trazas en rosario. Esta asociación icnológica muestra moderada diversidad y baja icnodisparidad que sugieren condiciones de estrés ambiental, probablemente vinculadas a importantes fluctuaciones de salinidad dentro del estuario. La presencia de la icnofacies de Mermia en estos depósitos es una nueva evidencia de que los sistemas depositacionales marino marginales postglaciales estuvieron fuertemente afectados por significativas descargas de agua dulce al producirse la deglaciación del Paleozoico tardío.The paleoenvironmental and ichnological characteristics of the middle section of the Agua del Jagüel Formation (Carboniferous) are described in this paper. Five sedimentary facies, grouped into three facies associations illustrating the evolution of a fluvio-estuarine system, were identified. Facies association 1, composed of fluvial conglomerates and coarse-grained sandstones, rests on an erosive surface carved into postglacial diamictites, sandstones, and shales belonging to the lower part of the Agua del Jagüel Formation. Facies association 2 comprises at the base mudstones and shales that indicate a marine the flooding of the system, followed by sandy sedimentation corresponding to estuary mouth settings suggesting a progressive transgression within the estuary. Finally, facies association 3 comprises two fluvial intervals of coarse-grained sandstones and conglomerates, separated by fine-grained sandstones and shales deposited during a short transgressive event. Together with the sedimentological studies, biogenic structures were analyzed from shales, mudstones, and very fine- to fine-grained sandstones of facies 2. The trace fossils identified include Cochlichnus anguineus, Gordia marina, Helminthoidichnites tenuis, Helminthopsis tenuis, Mermia carickensis, ?Treptichnus bifurcus,?T. pollardi and rosary-like traces. This ichnological association shows moderate diversity and low ichnodisparity suggesting the presence of environmental stress, probably linked to important salinity fluctuations within the estuary. The presence of the Mermia ichnofacies in these deposits represents new evidence that postglacial marginal-marine environments were affected by high discharge of freshwater during the late Paleozoic deglaciation.Fil: Alonso Muruaga, Pablo Joaquin. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Geociencias Básicas, Aplicadas y Ambientales de Buenos Aires. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Geociencias Básicas, Aplicadas y Ambientales de Buenos Aires; ArgentinaFil: Limarino, Carlos Oscar. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Geociencias Básicas, Aplicadas y Ambientales de Buenos Aires. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Geociencias Básicas, Aplicadas y Ambientales de Buenos Aires; ArgentinaFil: Buatois, Luis A.. University of Saskatchewan; CanadáFil: Pirrone, Cecilia Anabel. Universidad Nacional de Cuyo. Facultad de Ciencias Exactas y Naturales; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentin

A novel tool to untangle the ecology and fossil preservation knot in exceptionally preserved biotas

Understanding the functioning of extinct ecosystems is a complicated knot of ecological, evolutionary, and preservational strands that must be untangled. For instance, anatomical and behavioral differences can profoundly alter fossilization pathways. This is particularly true in exceptionally preserved soft-bodied biotas that record the earliest phases of animal evolution during the Cambrian Explosion and the Ordovician Radiation. Herein, a novel method of data partitioning based on probabilistic modelling is developed to examine these processes for the Walcott Quarry, Burgess Shale, Canada (510Ma), and the Fezouata Shale, Morocco (c. 475Ma). The modelling shows that the mechanism for soft-tissue preservation in the Walcott Quarry is ecologically selective, favoring the endobenthos. This is not found in the Fezouata Shale. Taken in concert with bioturbation data, a new model of comparative preservation is developed based on sedimentary flow dynamics. This suggests that during the Cambrian Explosion and Ordovician Radiation the most exceptional fossils sites must still be calibrated against each other to understand the unfolding evolutionary events and the ecological structuring of ancient animal communities

Trilobite expansion into estuarine environments during the Ordovician radiation

Trilobites have traditionally been considered fully marine. Through the integration of ichnological, palaeobiological and sedimentological datasets within a sequence-stratigraphic and strati­graphic palaeobiology framework, we challenge this assumption. This analysis is based on the study of incised fluvio-estuarine valley deposits from the Furongian Tilcara Member (TM) and the latest Furongian Pico de Halcón Member (PHM) of the Santa Rosita Formation, the early late Tremadocian Cardonal Formation (CF), and the DapingianâDarriwilian Alto del Cóndor Formation (ACF), from Cordillera Oriental of northwest Argentina. These valleys were incised into wave-dominated shallow-marine strata and filled with transgressive deposits that accumulated in tide-dominated estuaries. Whereas the TM lacks any body or trace fossil evidence of the presence of trilobites in estuarine settings, the other three units reveal that trilobites were able to inhabit these settings. The PHM and CF are host to trilobite trace fossils in outer estuarine facies, both containing various ichnospecies of Cruziana (e.g., C. omanica and C. semiplicata in the TM) and Rusophycus (e.g., R. latus in both units). In addition, the PHM also contains body fossils of the olenid trilobite Neoparabolina frequens argentina in the same deposits in which the trace fossils are preserved, as well as from middle estuarine facies. The ACF displays trilobite trace fossils of the C. rugosa group in inner, middle, and outer estuarine deposits, illustrating further landward incursions. This unit also contains body fossils of the asaphid trilobite Ogyginus sp. Accordingly, our data indicate two attempts of landward exploration via brackish water: phase 1 in which the outer to middle portion of estuaries were colonized by olenids (Furongianâearly late Tremadocian) and phase 2 involving exploration of the inner, middle, and outer estuarine zones by asaphids (DapingianâDarriwilian). Our study indicates that these trilobites were tolerant to salinity stress and able to make use of the ecological advantages offered by marginal-marine environments migrating up-estuary, following salt wedges either reflecting amphidromy or as euryhaline marine wanderers. It is suggested that tolerance to salinity stress arose independently among different trilobite groups as a result of the broad array of behaviors and adaptations of trilobites during the Ordovician radiation. We speculate that the assumption that all trilobites were stenohaline may have resulted in the misinterpretation of some tide-dominated estuarine deposits as fully marine