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Impacts of spatial and environmental differentiation on early Palaeozoic marine biodiversity


The unprecedented diversifications in the fossil record of the early Palaeozoic (541–419 million years ago) increased both within-sample (α) and global (γ) diversity, generating considerable ecological complexity. Faunal difference (β diversity), including spatial heterogeneity, is thought to have played a major role in early Palaeozoic marine diversification, although α diversity is the major determinant of γ diversity through the Phanerozoic. Drivers for this Phanerozoic shift from β to α diversity are not yet resolved. Here, we evaluate the impacts of environmental and faunal heterogeneity on diversity patterns using a global spatial grid. We present early Palaeozoic genus-level α, β and γ diversity curves for molluscs, brachiopods, trilobites and echinoderms and compare them with measures of spatial lithological heterogeneity, which is our proxy for environmental heterogeneity. We find that α and β diversity are associated with increased lithological heterogeneity, and that β diversity declines over time while α increases. We suggest that the enhanced dispersal of marine taxa from the Middle Ordovician onwards facilitated increases in α diversity by encouraging the occupation of narrow niches and increasing the prevalence of transient species, simultaneously reducing spatial β diversity. This may have contributed to a shift from β to α diversity as the major determinant of γ diversity increase over this critical evolutionary interval.

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Fig. 1: Diversity curves from 540–419 million years ago.
Fig. 2: Heterogeneity of lithological units within grids, and nestedness from 540-419 Ma.
Fig. 3: Relationship between within-grid α diversity and within-grid number of formations.

Data availability

All data used in this work can be downloaded from the PBDB ( The direct links for retrieving the data used to generate these results are available in the accompanying code (see the code availability).

Code availability

The complete code and relevant results are recorded in R code and can be downloaded via Zeonodo (


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This study was part of the Academy of Finland-funded project ‘Ecological engineering as a biodiversity driver in deep time’. We thank S. Scholze for data entry into the PBDB over the course of this study, M. Wale for help with running sensitivity analyses and R. Hofmann for helpful discussions on β diversity in the Palaeozoic. This is a contribution to IGCP 653 (The onset of the Great Ordovician Biodiversification Event).

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Authors and Affiliations



A.P. and B.K. devised the research, agreed on analytical techniques and wrote the paper. A.P. calculated β diversity and checked correlations. B.K. downloaded and formatted data from the PBDB, calculated α and γ diversity, and drew the figures.

Corresponding author

Correspondence to Amelia Penny.

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Extended data

Extended Data Fig. 1 Raw beta diversity through time.

Raw (non RAC) beta diversity through time, showing high values interpreted to be caused by incomplete sampling.

Extended Data Fig. 2 Sensitivity analysis of the effect of grid size on beta diversity.

RAC beta diversity curves made using a global grid of hexagons with side (a) 55 km and (b) 222 km. Kolmogorov-Smirnov tests show no significant difference between these curves and the curve made with a 111 km grid. (a) D = 0.15, p = 0.99 for grids of side 55km; (b) D = 0.08, p = 1 for grids of side 222 km.

Extended Data Fig. 3 Sensitivity analysis of the effect of standardization coverage on beta diversity.

RAC beta diversity curves made using a standardization coverage of (a) 0.2 and (b) 0.5. Kolmogorov-Smirnov tests show no significant difference between these curves and the curve with a standardization coverage of 0.4 (a) D = 0.23, p = 0.90 for a standardization coverage of 0.2; (b) D = 0.19, p = 0.94 for a standardization coverage of 0.5.

Extended Data Fig. 4 Beta diversity curves made using the Simpson and Bray-Curtis dissimilarities.

RAC beta diversity curves showing (a) the Bray-Curtis and (b) the Simpson dissimilarity through time. Kolmogorov-Smirnov tests show no significant difference between these curves and the RAC Sørensen dissimilarity curve. (a) D = 0.23, p = 0.90 for the Bray-Curtis curve; (b) D = 0.38, p = 0.30 for the Simpson dissimilarity curve.

Extended Data Fig. 5 Autocorrelation function plots for all variables for which correlations were tested.

Autocorrelation function plots for variables tested for correlation in this study. No variable shows significant autocorrelation, which would be indicated by columns passing the dashed blue horizontal lines at lags greater than 0.

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Penny, A., Kröger, B. Impacts of spatial and environmental differentiation on early Palaeozoic marine biodiversity. Nat Ecol Evol 3, 1655–1660 (2019).

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