Ecosystem engineers are keystone taxa whose behaviours alter the habitability of their environments for themselves and other organisms by directly influencing the availability of resources in their ecosystems. From a deep time perspective, ecosystem engineers are hypothesized to have played a major role in the co-evolution of life and the Earth systems, as many major ecosystem engineering activities directly modulate the cycling of key nutrients. Moreover, ecosystem engineers are thought to have impacted diversity by increasing environmental heterogeneity, and so their evolution may drive some of the biodiversity dynamics observed in the fossil record. Here, we investigate the impact of two groups of marine ecosystem engineers – bioturbators and reef-builders – on biodiversity through the Phanerozoic. Using fossil occurrence data from the Paleobiology Database, we calculate the effect size of bioturbating and reef-building ecosystem engineers on various biodiversity metrics for each stage through the Phanerozoic. Most broadly, we find that ecosystem engineers had a positive impact on biodiversity within the environments where they live during the Phanerozoic. We also find clear taxonomic differences between environments with and without ecosystem engineers, suggesting ecosystem engineers create a unique set of environmental characteristics to which taxa of specific ecological characteristics become adapted. These results emphasize the important role of ecosystem engineers in influencing key aspects of the Earth systems on a variety of scales that manifest in changes in biodiversity.
Phanerozoic marine biodiversity dynamics have been shaped by continuous environmental change and biotic interactions. Although ecosystem engineers – animals whose behaviours modify resource availability – have established impacts on modern community ecology and diversity, their impacts on ancient ecosystems over geologic time have remained quantitatively underexplored. Here, we investigate the impacts of reef-building and bioturbating marine ecosystem engineers on biodiversity at each stage of the Phanerozoic, including their ability to maintain biodiversity over mass extinctions, and identify extrinsic conditions that modulate their impacts. We analysed fossil communities with and without ecosystem engineers to calculate effect sizes, thereby quantifying effects of the presence of ecosystem engineers on Shannon’s Diversity (H) at each stage in the Phanerozoic. We show that ecosystem engineers are significantly associated with increased biodiversity during the majority of the Phanerozoic. We also show that, over the last ~250 million years, the effectiveness of modern-type reef-builders is more sensitive to climate-driven stress than for modern-style bioturbators. Finally, our results reveal that climate can modulate ecosystem engineer impacts, with both bioturbators and reef-builders exhibiting strongest effect sizes within an optimal moderate temperature range. These results underscore ecosystem engineers’ crucial role in contributing to fluctuating biodiversity dynamics over the Phanerozoic.
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.
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.
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.
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.
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.