Ecosystem engineers are organisms that modify their physical habitats in a way that alters resource availability and the structure of the communities they live in. The evolution of ecosystem engineers over the course of Earth history has thus been suggested to have been a driver of macroevolutionary and macroecological changes that are observed in the fossil record. However, the rise to dominance of ecosystem engineers has not been thoroughly reconstructed. Here, we investigate the history of bioturbation and reef-building (two of the most important marine ecosystem engineering behaviours today) over the Phanerozoic. Using fossil occurrences from the Paleobiology Database, we reconstruct how common communities influenced by ecosystem engineers were in the oceans, how dominant ecosystem engineers were within their own communities, and the taxonomic and ecological composition of bioturbators and reef-builders. We find that bioturbation has become an increasingly common ecosystem engineering behaviour over the Phanerozoic, while reef-building ecosystem engineers have not become more dominant since their Devonian apex. We also identify unique bioturbation and reef-building regimes that are characterized by different ecosystem engineering taxonomic groups, ecological modes, and dominance, suggesting that the nature of ecosystem engineering has at times rapidly shifted over the course of the Phanerozoic. These reconstructions will serve as important data for understanding how ecosystem engineers have driven changes in biodiversity and ecosystem structure over the course of Earth history.

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Bioturbation, the mixing of sediment through the actions of organisms, is a crucial ecosystem engineering process that controls biogeochemical cycles and helps structure marine ecosystems. Machine learning is helping to develop global maps of the intensity and depth of bioturbation.

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Mass extinctions have fundamentally altered the structure of the biosphere throughout Earth's history. The ecological severity of mass extinctions is well studied in marine ecosystems by categorizing marine taxa into functional groups based on ‘ecospace’ approaches, but the ecological response of terrestrial ecosystems to mass extinctions is less well understood due to the lack of a comparable methodology. Here, we present a new terrestrial ecospace framework that categorizes fauna into functional groups as defined by tiering, motility and feeding traits. We applied the new terrestrial and traditional marine ecospace analyses to data from the Paleobiology Database across the end-Triassic mass extinction—a time of catastrophic global warming—to compare changes between the marine and terrestrial biospheres. We found that terrestrial functional groups experienced higher extinction severity, that taxonomic and functional richness are more tightly coupled in the terrestrial, and that the terrestrial realm continued to experience high ecological dissimilarity in the wake of the extinction. Although signals of extinction severity and ecological turnover are sensitive to the quality of the terrestrial fossil record, our findings suggest greater ecological pressure from the end-Triassic mass extinction on terrestrial ecosystems than marine ecosystems, contributing to more prolonged terrestrial ecological flux.

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The radiation of bioturbation during the Ediacaran–Cambrian transition has long been hypothesized to have oxygenated sediments, triggering an expansion of the habitable benthic zone and promoting increased infaunal tiering in early Paleozoic benthic communities. However, the effects of bioturbation on sediment oxygen are underexplored with respect to the importance of biomixing and bioirrigation, two bioturbation processes which can have opposite effects on sediment redox chemistry. We categorized trace fossils from the Ediacaran and Terreneuvian as biomixing or bioirrigation fossils and integrated sedimentological proxies for bioturbation intensity with biogeochemical modeling to simulate oxygen penetration depths through the Ediacaran–Cambrian transition. Ultimately, we find that despite dramatic increases in ichnodiversity in the Terreneuvian, biomixing remains the dominant bioturbation behavior, and in contrast to traditional assumptions, Ediacaran–Cambrian bioturbation was unlikely to have resulted in extensive oxygenation of shallow marine sediments globally.

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The Permian-Triassic mass extinction severely depleted biodiversity, primarily observed in the body fossil of well-skeletonized animals. Understanding how whole ecosystems were affected and rebuilt following the crisis requires evidence from both skeletonized and soft-bodied animals; the best comprehensive information on soft-bodied animals comes from ichnofossils. We analyzed abundant trace fossils from 26 sections across the Permian-Triassic boundary in China and report key metrics of ichnodiversity, ichnodisparity, ecospace utilization, and ecosystem engineering. We find that infaunal ecologic structure was well established in the early Smithian. Decoupling of diversity between deposit feeders and suspension feeders in carbonate ramp-platform settings implies that an effect of trophic group amensalism could have delayed the recovery of nonmotile, suspension-feeding epifauna in the Early Triassic. This differential reaction of infaunal ecosystems to variable environmental controls thus played a substantial but heretofore little appreciated evolutionary and ecologic role in the overall recovery in the hot Early Triassic ocean.

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The Ediacaran–Cambrian transition marks one of the most important geobiological revolutions in Earth History, including multiple waves of evolutionary radiation and successive episodes of apparent mass extinction. Among the proposed drivers of these events (in particular the extinction of the latest Neoproterozoic ‘Ediacara biota’) is the emergence of complex metazoans and their associated behaviors. Many metazoans are thought to have crucial geobiological impacts on both resource availability and the character of the physical environment – ‘ecosystem engineering’ – biological processes best preserved in the geological record as trace fossils. Here, we review this model using the trace fossil record of the Ediacaran to Cambrian Nama Group of southern Namibia, combining previous published accounts with the results of our own field investigations. We produce a revised ichnostratigraphy for the Nama Group that catalogues new forms, eliminates others, and brings the trace fossil record of the Nama into much closer alignment with what is known from other Ediacaran sections worldwide. We provide evidence for a link between sequence stratigraphy, oxygen, and the emergence of more complex bilaterian behaviors. Lastly, we show that observed patterns of extinction and survival over pulses of Ediacaran extinction are hard to ally with any one specific source of ecological stress associated with bioturbation, and thus a biologically-driven extinction of the Ediacara biota, if it occurred, was more likely to have been driven by some combination of these factors, rather than any single one.

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Ediacaran fossils from the Nama Group (Kuibis Subgroup) of southern Namibia have a long history of scientific scrutiny, however many of the fossil localities still require investigation from a sedimentary facies and sequence stratigraphic standpoint. Detailed sedimentary analyses utilizing chemostratigraphy and facies-based approaches resulted in five proposed facies and two paleoenvironmental settings that allow for interpretation of the fossiliferous sections as a nearshore to a protected shallow marine paleoenvironment. The classic Ediacaran taxon Ernietta was restricted to the Kliphoek Member of the Nama Group, which limits the stratigraphic range of these organisms to the younger depositional sequence of the Kuibis Subgroup. The paleoenvironment has been interpreted as a mixed carbonate-siliciclastic, protected shallow marine environment at Farm Hansburg, suggesting that these organisms thrived in environments with medium to high flow velocities and periodic clastic sediment supply. Studies that utilize detailed paleoenvironmental reconstructions may aid in constraining phylogenetic affinities of the Ediacara biota by placing reasonable bounds on the local habitat.

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The end-Permian mass extinction was the most severe mass extinction event of the Phanerozoic and was followed by a several million-year delay in benthic ecosystem recovery. While much work has been done to understand biotic recovery in both the body and trace fossil records of the Early Triassic, almost no focus has previously been given to analyzing patterns in ecosystem engineering complexity as a result of the extinction drivers. Bioturbation is a key ecosystem engineering behavior in marine environments, as it results in changes to resource flows and the physical environment. Thus, the trace fossil record can be used to examine the effect of the end-Permian mass extinction on bioturbating ecosystem engineers. We present a dataset compiled from previously published literature to analyze burrowing ecosystem engineering behaviors through the Permian-Triassic boundary. We report two key observations: first, that there is no loss in bioturbation ecosystem engineering behaviors after the mass extinction, and second, that these persisting behaviors include deep tier, high-impact, complex ecosystem engineering. These findings suggest that while environmental conditions may have limited deeper burrowing, complex ecosystem engineering behaviors were able to persist in the Early Triassic. Furthermore, the persistence of deep tier bioirrigated three-dimensional network burrows implies that benthic biogeochemical cycling could have been maintained at pre-extinction states in some local environments, stimulating ecosystem productivity and promoting biotic recovery in the Early Triassic.

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The disappearance of the soft-bodied Ediacara biota at the Ediacaran–Cambrian boundary potentially represents the earliest mass extinction of complex life, although the precise driver(s) of this extinction remain unresolved. The ‘biotic replacement’ model proposes that an evolutionary radiation of metazoan ecosystem engineers in the latest Ediacaran profoundly altered marine palaeoenvironments, resulting in the extinction of Ediacara biota and setting the stage for the subsequent Cambrian Explosion. However, metazoan ecosystem engineering across the Ediacaran–Cambrian transition has yet to be quantified. Here, we test this key tenet of the biotic replacement model by characterizing the intensity of metazoan bioturbation and ecosystem engineering in trace fossil assemblages throughout the latest Ediacaran Nama Group in southern Namibia. The results illustrate a dramatic increase in both bioturbation and ecosystem engineering intensity in the latest Ediacaran, prior to the Cambrian boundary. Moreover, our analyses demonstrate that the highest-impact ecosystem engineering behaviours were present well before the onset of the Cambrian. These data provide the first support for a fundamental prediction of the biotic replacement model, and evidence for a direct link between the early evolution of ecosystem engineering and the extinction of the Ediacara biota.

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