The fossil record provides one of the strongest tests of the hypothesis that diversity within local communities is constrained over geological timescales. Constraints to diversity are particularly controversial in modern terrestrial ecosystems, yet long-term patterns are poorly understood. Here we document patterns of local richness in Phanerozoic terrestrial tetrapods using a global data set comprising 145,332 taxon occurrences from 27,531 collections. We show that the local richness of non-flying terrestrial tetrapods has risen asymptotically since their initial colonization of land, increasing at most threefold over the last 300 million years. Statistical comparisons support phase-shift models, with most increases in local richness occurring: (1) during the colonization of land by vertebrates, concluding by the late Carboniferous; and (2) across the Cretaceous/Paleogene boundary. Individual groups, such as mammals, lepidosaurs and dinosaurs also experienced early increases followed by periods of stasis often lasting tens of millions of years. Mammal local richness abruptly tripled across the Cretaceous/Paleogene boundary, but did not increase over the next 66 million years. These patterns are consistent with the hypothesis that diversity is constrained at the local-community scale.

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Data availability

The data used in this study were downloaded from the PaleoDB (http://www.paleobiodb.org) and have been archived, together with all custom analysis scripts, on Dryad (https://doi.org/10.5061/dryad.3v0p84v).

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We thank all contributors to the PaleoDB. This is PaleoDB official publication #334. This research was funded by the European Union’s Horizon 2020 research and innovation programme under grant agreement no. 637483 (ERC Starting Grant TERRA, R.J.B.). P.D.M. was supported by a Leverhulme Trust Early Career Fellowship (no. ECF-2014-662) and a Royal Society University Research Fellowship (no. UF160216).

Author information


  1. School of Geography, Earth and Environmental Sciences, University of Birmingham, Birmingham, UK

    • Roger A. Close
    • , Juan Benito
    • , Terri J. Cleary
    • , Emma M. Dunne
    •  & Richard J. Butler
  2. Department of Earth Sciences, University of Oxford, Oxford, UK

    • Roger B. J. Benson
  3. Department of Biological Sciences, Macquarie University, Sydney, NSW, Australia

    • John Alroy
  4. Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA

    • Anna K. Behrensmeyer
    •  & Matthew T. Carrano
  5. Department of Biology and Biochemistry, Faculty of Science, University of Bath, Bath, UK

    • Juan Benito
  6. Department of Earth Sciences, Natural History Museum, London, UK

    • Terri J. Cleary
  7. Department of Earth Science and Engineering, Imperial College London, London, UK

    • Philip D. Mannion
  8. Department of Atmospheric Oceanic and Earth Sciences, George Mason University, Fairfax, VA, USA

    • Mark D. Uhen


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R.A.C., R.B.J.B. and R.J.B. conceived the study. J.A., A.K.B., J.B., R.B.J.B., R.J.B., M.T.C., T.J.C., E.M.D., P.D.M. and M.D.U. contributed to the data set. R.A.C. designed and conducted the analyses and wrote the manuscript. R.B.J.B., J.A. and M.T.C. provided methodological advice. R.J.B. and R.B.J.B. drafted portions of the manuscript. All authors provided critical comments on the manuscript.

Competing interests

The authors declare no competing interests.

Corresponding author

Correspondence to Roger A. Close.

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