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Synopsis The invasion of the land was a complex, protracted process, punctuated by mass extinctions, that involved multiple routes from marine environments. We integrate paleobiology, ichnology, sedimentology, and geomorphology to reconstruct Paleozoic terrestrialization. Cambrian landscapes were dominated by laterally mobile rivers with unstable banks in the absence of significant vegetation. Temporary incursions by arthropods and worm-like organisms into coastal environments apparently did not result in establishment of continental communities. Contemporaneous lacustrine faunas may have been inhibited by limited nutrient delivery and high sediment loads. The Ordovician appearance of early land plants triggered a shift in the primary locus of the global clay mineral factory, increasing the amount of mudrock on the continents. The Silurian-Devonian rise of vascular land plants, including the first forests and extensive root systems, was instrumental in further retaining fine sediment on alluvial plains. These innovations led to increased architectural complexity of braided and meandering rivers. Landscape changes were synchronous with establishment of freshwater and terrestrial arthropod faunas in overbank areas, abandoned fluvial channels, lake margins, ephemeral lakes, and inland deserts. Silurian-Devonian lakes experienced improved nutrient availability, due to increased phosphate weathering and terrestrial humic matter. All these changes favoured frequent invasions to permament establishment of jawless and jawed fishes in freshwater habitats and the subsequent tetrapod colonization of the land. The Carboniferous saw rapid diversification of tetrapods, mostly linked to aquatic reproduction, and land plants, including gymnosperms. Deeper root systems promoted further riverbank stabilization, contributing to the rise of anabranching rivers and braided systems with vegetated islands. New lineages of aquatic insects developed and expanded novel feeding modes, including herbivory. Late Paleozoic soils commonly contain pervasive root and millipede traces. Lacustrine animal communities diversified, accompanied by increased food-web complexity and improved food delivery which may have favored permanent colonization of offshore and deep-water lake environments. These trends continued in the Permian, but progressive aridification favored formation of hypersaline lakes, which were stressful for colonization. The Capitanian and end-Permian extinctions affected lacustrine and fluvial biotas, particularly the invertebrate infauna, although burrowing may have allowed some tetrapods to survive associated global warming and increased aridification.
Synopsis The invasion of the land was a complex, protracted process, punctuated by mass extinctions, that involved multiple routes from marine environments. We integrate paleobiology, ichnology, sedimentology, and geomorphology to reconstruct Paleozoic terrestrialization. Cambrian landscapes were dominated by laterally mobile rivers with unstable banks in the absence of significant vegetation. Temporary incursions by arthropods and worm-like organisms into coastal environments apparently did not result in establishment of continental communities. Contemporaneous lacustrine faunas may have been inhibited by limited nutrient delivery and high sediment loads. The Ordovician appearance of early land plants triggered a shift in the primary locus of the global clay mineral factory, increasing the amount of mudrock on the continents. The Silurian-Devonian rise of vascular land plants, including the first forests and extensive root systems, was instrumental in further retaining fine sediment on alluvial plains. These innovations led to increased architectural complexity of braided and meandering rivers. Landscape changes were synchronous with establishment of freshwater and terrestrial arthropod faunas in overbank areas, abandoned fluvial channels, lake margins, ephemeral lakes, and inland deserts. Silurian-Devonian lakes experienced improved nutrient availability, due to increased phosphate weathering and terrestrial humic matter. All these changes favoured frequent invasions to permament establishment of jawless and jawed fishes in freshwater habitats and the subsequent tetrapod colonization of the land. The Carboniferous saw rapid diversification of tetrapods, mostly linked to aquatic reproduction, and land plants, including gymnosperms. Deeper root systems promoted further riverbank stabilization, contributing to the rise of anabranching rivers and braided systems with vegetated islands. New lineages of aquatic insects developed and expanded novel feeding modes, including herbivory. Late Paleozoic soils commonly contain pervasive root and millipede traces. Lacustrine animal communities diversified, accompanied by increased food-web complexity and improved food delivery which may have favored permanent colonization of offshore and deep-water lake environments. These trends continued in the Permian, but progressive aridification favored formation of hypersaline lakes, which were stressful for colonization. The Capitanian and end-Permian extinctions affected lacustrine and fluvial biotas, particularly the invertebrate infauna, although burrowing may have allowed some tetrapods to survive associated global warming and increased aridification.
Biogenic sedimentary structures offer a unique perspective for understanding the role of the biosphere in the interaction with other Earth subsystems and the building up of our planet. The record of their ancient equivalents provides a wealth of information for reconstructing the role of bioturbators as ecosystem engineers using multiple ichnological proxies and methods. In this study, we present an overview of how bioturbation has worked across a broad range of spatial and temporal scales from the perspective of ecosystem engineering. Comprehensive and systematic datasets allow analyses at regional and global spatial extents, and especially over long temporal scales where sampling intensity and rock record biases can be considered. Our results underscore the significance of the Cambrian Explosion in the establishment of modern‐style shallow marine ecosystems and of the Ordovician Radiation for their deep marine counterparts, as well as the continuous ecosystem impact of bioturbation during Palaeozoic terrestrialization. Comparable datasets for the rest of the Phanerozoic have not yet been compiled. However, preliminary information indicates that colonization of ultra‐deep tiers, the rise to prominence of regenerators, increased burrowing efficiency, and increased compartmentalization of the endobenthic ecospace, were products of the Mesozoic Marine Revolution. The trace‐fossil record offers hard data to evaluate bioturbation as a driving force in ecosystem re‐structuring and as a key factor in geobiological cycles. Models assessing these fundamental issues should be rooted empirically at different scales, from both autoecological and synecological to macroecological, making the best possible use of the rich and rapidly developing ichnological toolbox.
The Triassic Katberg Formation has played a central role in interpreting the end-Permian ecosystem crisis, as part of a hypothesis of aridification, vegetation loss, and sediment release in continental settings. We use drone images of an inaccessible cliff near Bethulie to investigate the Swartberg member, a braided-fluvial body 45 m thick, describing remote outcrop facies to identify geomorphic units and using spatial analysis to estimate their proportions in 2-D sections. Here the Swartberg member comprises three channel belts within shallow valleys, the lowermost of which is ∼500 m wide and incised into lacustrine deposits. The component channel bodies consist mainly of trough cross-bedded sand sheets (48%) and channel-scour fills (28%). Recognizable bars (15%) comprise unit bars with high-angle slipfaces and mounded bar cores (components of mid-channel compound bars), bars built around vegetation, and bank-attached bars in discrete, probably low-sinuosity conduits. Abandoned channels constitute 8% and 16% of flow-parallel and -transverse sections, respectively. When corrected for compaction, the average thalweg depth of the larger channels is 3.9 m, with an average bankfull width of 84 m, scaling broadly with the relief of the bars and comparable in scale to the Platte and South Saskatchewan rivers of North America. The fluvial style implies perennial but seasonably variable flow in a vegetated landscape with a humid paleoclimate. The northward paleoflow accords with regional paleoflow patterns and deposition on a megafan sourced in the Cape Fold Belt, where the Swartberg member represents the avulsion of a major transverse-flowing river. U-Pb dating of in situ and reworked pedogenic carbonate nodules from below the base of the Swartberg member yielded Anisian to Ladinian ages (Middle Triassic), younger than the previously assumed Early Triassic age and implying that considerable gaps in time exist in the succession. An assessment of the interval spanning the lower to mid Katberg Formation is needed to reevaluate the inferred unidirectional trend in fluvial style, aridification, and fossil distributions in this condensed, disjunct succession.
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