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11 June 2024 The Fezouata Shale Formation biota is typical for the high latitudes of the Early Ordovician—a quantitative approach
Jared C. Richards, Karma Nanglu, Javier Ortega-Hernández
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Abstract

The Fezouata Shale Formation in present-day Morocco is a site of exceptional fossil preservation from the Lower Ordovician that provides a unique view of animal life before one of the most important radiation events in Earth's history, the great Ordovician biodiversification event (GOBE). Previous work on the fossil diversity of the Fezouata Shale has suggested that there are faunistic differences between the two major intervals with exceptional preservation and that the overall shelly biota of the Fezouata Shale is comparable to other Lower Ordovician sites that reflect open-marine conditions. In this study, we make the first comprehensive quantitative comparison between the Fezouata Shale Formation and other high-latitude Early Ordovician sites based on their shelly fossil biotas with publicly available fossil occurrence information from the Paleobiology Database. We find that the fossil subassemblages of the stratigraphically older lower Fezouata Shale are more heterogeneous than those of the younger upper Fezouata Shale. The fossil biota preserved in the lower Fezouata Shale is most similar to those found in other high-latitude deposits from the Lower Ordovician. We also find that there are differences in faunal composition between Tremadocian- and Floian-aged deposits. Our work provides the first quantitative support for faunistic differences between the lower and upper Fezouata Shale Formation and indicate that the lower Fezouata Shale conventional fossil biota is typical for the Tremadocian, further contextualizing the ecology of the polar regions before the GOBE as informed by this major site of exceptional fossil preservation.

The Fezouata Shale Formation has dramatically impacted our understanding of Early Ordovician marine ecosystems before the great Ordovician biodiversification event (GOBE), thanks to the abundance and quality of exceptionally preserved animals within it. Systematic work has noted that the shelly fossil subassemblages of the Fezouata Shale biota are typical of open-marine deposits from the Lower Ordovician, but no studies have tested the quantitative validity of this statement. We extracted 491 occurrences of recalcitrant fossil genera from the Paleobiology Database to reconstruct 31 subassemblages to explore the paleoecology of the Fezouata Shale and other contemporary, high-latitude (66°S–90°S) deposits from the Lower Ordovician (485.4–470 Ma) and test the interpretation that the Fezouata Shale biota is typical for an Ordovician open-marine environment. Sørensen's dissimilarity metrics and Wilcoxon tests indicate that the subassemblages of the Tremadocian-aged lower Fezouata Shale are approximately 20% more heterogenous than the Floian-aged upper Fezouata Shale. Dissimilarity metrics and visualization suggest that while the lower Fezouata and upper Fezouata share faunal components, the two sections have distinct faunas. We find that the faunal composition of the lower Fezouata Shale is comparable with other Tremadocian-aged subassemblages from high latitudes, suggesting that it is typical for an Early Ordovician open-marine environment. We also find differences in faunal composition between Tremadocian- and Floian-aged deposits. Our results corroborate previous field-based and qualitative systematic studies that concluded that the shelly assemblages of the Fezouata Shale are comparable with those of other Lower Ordovician deposits from high latitudes. This establishes the first quantitative baseline for examining the composition and variability within the assemblages of the Fezouata Shale and will be key to future studies attempting to discern the degree to which it can inform our understanding of marine ecosystems just before the start of the GOBE.

Jared C. Richards, Karma Nanglu, and Javier Ortega-Hernández "The Fezouata Shale Formation biota is typical for the high latitudes of the Early Ordovician—a quantitative approach," Paleobiology 50(2), 226-238, (11 June 2024). https://doi.org/10.1017/pab.2024.7
Received: 26 July 2023; Accepted: 21 February 2024; Published: 11 June 2024
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