Two-phase increase in the maximum size of life over 3.5 billion years reflects biological innovation and environmental opportunity
- aDepartment of Geological and Environmental Sciences, Stanford University, 450 Serra Mall, Building 320, Stanford, CA 94305;
- bDepartment of Biology, University of New Mexico, Albuquerque, NM 87131;
- cDepartment of Geosciences, Virginia Polytechnic Institute and State University, Blacksburg, VA 24061;
- dMuseum für Naturkunde der Humboldt–Universität zu Berlin, D-10115, Berlin, Germany;
- eDepartment of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560;
- fMonterey Bay Aquarium Research Institute, Moss Landing, CA 95039;
- gDepartment of Biology, Box 90338, Duke University, Durham, NC 27708;
- hDepartment of Geosciences, University of West Georgia, Carrollton, GA 30118;
- iDepartment of Geological Sciences, University of Colorado, Boulder, CO 80309; and
- jDepartment of Mathematics and Statistics, Swarthmore College, 500 College Avenue, Swarthmore, PA 19081
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Edited by James W. Valentine, University of California, Berkeley, CA, and approved November 14, 2008 (received for review July 1, 2008)

Abstract
The maximum size of organisms has increased enormously since the initial appearance of life >3.5 billion years ago (Gya), but the pattern and timing of this size increase is poorly known. Consequently, controls underlying the size spectrum of the global biota have been difficult to evaluate. Our period-level compilation of the largest known fossil organisms demonstrates that maximum size increased by 16 orders of magnitude since life first appeared in the fossil record. The great majority of the increase is accounted for by 2 discrete steps of approximately equal magnitude: the first in the middle of the Paleoproterozoic Era (≈1.9 Gya) and the second during the late Neoproterozoic and early Paleozoic eras (0.6–0.45 Gya). Each size step required a major innovation in organismal complexity—first the eukaryotic cell and later eukaryotic multicellularity. These size steps coincide with, or slightly postdate, increases in the concentration of atmospheric oxygen, suggesting latent evolutionary potential was realized soon after environmental limitations were removed.
Footnotes
- 1To whom correspondence should be addressed. E-mail: jlpayne{at}stanford.edu
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Author contributions: J.L.P., A.G.B., J.H.B., S.F., M.K., R.A.K., S.K.L., C.R.M., D.W.M., P.M.N.-G., F.A.S., J.A.S., and S.C.W. designed research, performed research, analyzed data, and wrote the paper.
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The authors declare no conflict of interest.
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This article is a PNAS Direct Submission.
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This article contains supporting information online at www.pnas.org/cgi/content/full/0806314106/DCSupplemental.
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Freely available online through the PNAS open access option.
- © 2008 by The National Academy of Sciences of the USA