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The biomass distribution on Earth

Bar-On, Yinon M. and Phillips, Rob and Milo, Ron (2018) The biomass distribution on Earth. Proceedings of the National Academy of Sciences of the United States of America, 115 (25). pp. 6506-6511. ISSN 0027-8424. PMCID PMC6016768. doi:10.1073/pnas.1711842115.

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A census of the biomass on Earth is key for understanding the structure and dynamics of the biosphere. However, a global, quantitative view of how the biomass of different taxa compare with one another is still lacking. Here, we assemble the overall biomass composition of the biosphere, establishing a census of the ≈550 gigatons of carbon (Gt C) of biomass distributed among all of the kingdoms of life. We find that the kingdoms of life concentrate at different locations on the planet; plants (≈450 Gt C, the dominant kingdom) are primarily terrestrial, whereas animals (≈2 Gt C) are mainly marine, and bacteria (≈70 Gt C) and archaea (≈7 Gt C) are predominantly located in deep subsurface environments. We show that terrestrial biomass is about two orders of magnitude higher than marine biomass and estimate a total of ≈6 Gt C of marine biota, doubling the previous estimated quantity. Our analysis reveals that the global marine biomass pyramid contains more consumers than producers, thus increasing the scope of previous observations on inverse food pyramids. Finally, we highlight that the mass of humans is an order of magnitude higher than that of all wild mammals combined and report the historical impact of humanity on the global biomass of prominent taxa, including mammals, fish, and plants.

Item Type:Article
Related URLs:
URLURL TypeDescription Information and code CentralArticle
Bar-On, Yinon M.0000-0001-8477-609X
Phillips, Rob0000-0003-3082-2809
Milo, Ron0000-0003-1641-2299
Additional Information:© 2018 the Author(s). Published by PNAS. This open access article is distributed under Creative Commons Attribution-NonCommercial-NoDerivatives License 4.0 (CC BY-NC-ND). Edited by Paul G. Falkowski, Rutgers, The State University of New Jersey, New Brunswick, NJ, and approved April 13, 2018 (received for review July 3, 2017) published ahead of print May 21, 2018. We thank Shai Meiri for help with estimating the biomass of wild mammals, birds, and reptiles and Arren Bar-Even, Oded Beja, Jorg Bernhardt, Tristan Biard, Chris Bowler, Nuno Carvalhais, Otto Coredero, Gidon Eshel, Ofer Feinerman, Noah Fierer, Daniel Fisher, Avi Flamholtz, Assaf Gal, José Grünzweig, Marcel van der Heijden, Dina Hochhauser, Julie Huber, Qusheng Jin, Bo Barker Jørgensen, Jens Kallmeyer, Tamir Klein, Christian Koerner, Daniel Madar, Fabrice Not, Katherine O’Donnell, Gal Ofir, Victoria Orphan, Noam Prywes, John Raven, Dave Savage, Einat Segev, Maya Shamir, Izak Smit, Rotem Sorek, Ofer Steinitz, Miri Tsalyuk, Assaf Vardi, Colomban de Vargas, Joshua Weitz, Yossi Yovel, Yonatan Zegman, and two anonymous reviewers for productive feedback on this manuscript. This research was supported by the European Research Council (project NOVCARBFIX 646827), the Israel Science Foundation (Grant 740/16), the ISF-NRF Singapore Joint Research Program (Grant 7662712), the Beck Canadian Center for Alternative Energy Research, Dana and Yossie Hollander, the Ullmann Family Foundation, the Helmsley Charitable Foundation, the Larson Charitable Foundation, the Wolfson Family Charitable Trust, Charles Rothschild, and Selmo Nussenbaum. This study was also supported by the NIH through Grant 1R35 GM118043-01 (MIRA). R.M. is the Charles and Louise Gartner Professional Chair. Author contributions: Y.M.B.-O., R.P., and R.M. designed research; Y.M.B.-O. and R.M. performed research; Y.M.B.-O. and R.M. analyzed data; and Y.M.B.-O., R.P., and R.M. wrote the paper. The authors declare no conflict of interest. This article is a PNAS Direct Submission. Data deposition: All of the data used to generate our estimates, as well as the code used for analysis, are available on GitHub at This article contains supporting information online at
Funding AgencyGrant Number
European Research Council (ERC)646827 NOVCARBFIX
Israel Science Foundation740/16
ISF-NRF Singapore Joint Research Program7662712
Beck Canadian Center for Alternative Energy ResearchUNSPECIFIED
Dana and Yossie HollanderUNSPECIFIED
Ullmann Family FoundationUNSPECIFIED
Helmsley Charitable FoundationUNSPECIFIED
Larson Charitable FoundationUNSPECIFIED
Wolfson Family Charitable TrustUNSPECIFIED
Charles RothschildUNSPECIFIED
Selmo NussenbaumUNSPECIFIED
NIH1R35 GM118043-01
Charles and Louise Gartner Professional ChairUNSPECIFIED
Subject Keywords:ecology; biomass; biosphere; quantitative biology
Issue or Number:25
PubMed Central ID:PMC6016768
Record Number:CaltechAUTHORS:20180521-155526693
Persistent URL:
Official Citation:The biomass distribution on Earth Yinon M. Bar-On, Rob Phillips, Ron Milo Proceedings of the National Academy of Sciences Jun 2018, 115 (25) 6506-6511; DOI: 10.1073/pnas.1711842115
Usage Policy:No commercial reproduction, distribution, display or performance rights in this work are provided.
ID Code:86531
Deposited By: George Porter
Deposited On:21 May 2018 23:23
Last Modified:10 Mar 2022 17:07

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