Soil biodiversity

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Fig. 3. Graphical overview of the share of species living in soil. Doughnuts reflect the per- centage of species in soil versus all other ecosystems combined (e.g., marine, freshwater, built en- vironment, host organisms such as humans). The larger doughnut on top shows the total share of species, and smaller doughnuts show individual shares for the most speciose and well-known groups ordered from greatest to least specialized in soil. Illustra- tions by Michael Dandley ©.

Significance

Soil organisms mediate unique functions we rely on for food, fiber, and human and planetary health. Despite the significance of soil life, we lack a quantitative estimate of soil biodiversity, making it challenging to advocate for the importance of protecting, preserving, and restoring soil life. Here, we show that soil is likely home to 59% of life including everything from microbes to mammals, making it the singular most biodiverse habitat on Earth. Our enumeration can enable stakeholders to more quantitatively advocate for soils in the face of the biodiversity crisis.

Enumerating soil biodiversity

Research article

a,b,1 a,c a,c,1 Mark A. Anthony , S. Franz Bender , and Marcel G. A. van der Heijden

Edited by Diana Wall, Colorado State University, Fort Colliins, CO; received March 21, 2023; accepted July 2, 2023 https://doi.org/10.1073/pnas.2304663120

Soil is an immense habitat for diverse organisms across the tree of life, but just how many organisms live in soil is surprisingly unknown. Previous efforts to enumerate soil biodiversity consider only certain types of organisms (e.g., animals) or report values for diverse groups without partitioning species that live in soil versus other habitats. Here, we reviewed the biodiversity literature to show that soil is likely home to 59 ± 15% of the species on Earth. We therefore estimate an approximately two times greater soil biodiversity than previous estimates, and we include representatives from the simplest (microbial) to most complex (mammals) organisms. Enchytraeidae have the greatest percentage of species in soil (98.6%), followed by fungi (90%), Plantae (85.5%), and Isoptera (84.2%). Our results demonstrate that soil is the most biodiverse singular habitat. By using this estimate of soil biodiversity, we can more accurately and quan- titatively advocate for soil organismal conservation and restoration as a central goal of the Anthropocene.

“How many species on Earth live in soil?” Like many simple questions, it is one of the most challenging to answer. A timely account of soil biodiversity is critical as Earth faces another wave of mass extinctions (1), and evidence points to humans as the major cause (2–4). Curtailing and reversing this trend requires biodiversity monitoring and conserva- tion programs like the Endangered Species Act in the United States and the Global IUCN Redlist, the most extensive source of information on global extinction risks. Yet, these efforts largely exclude species inhabiting soil, despite the critical importance of soil organ- isms to nearly every Earth function (5–7). Only with a complete estimation of the quantity of life that lives in soil can we understand the magnitude of value for conserving and restoring soil biodiversity.

Soil organisms are indispensable drivers of ecosystem composition and function. They govern global biogeochemical fluxes and directly influence rates of climate change and human health (8–10). Without an accurate estimate of soil biodiversity, we not only overlook a fundamental component of global biodiversity but additionally lack the quantitative information essential for policy advocation (11). There has been one pre- vious estimate of global soil biodiversity based on soil animals. Decaëns et al. (12) reported that at least 25% of described animal species live in or on soil based on a rapid survey of known animal species from a limited set of encyclopedic sources (12). This value has been widely taken up in soil biodiversity research, but it has also been incor- rectly cited to comprehensively represent all species versus just animals. To systematically estimate soil biodiversity, we must synthesize species numbers across the tree of life (see organisms living in soil in Fig. 1) and overcome myriad theoretical and technical obstacles.

Fig. 1. Diversity of the major life forms found in soil. (A) bristletail (© F. Ashwood), (B) springtail (© H. Conrad), (C) nitrogen-fixing bacteria-containing nodules on clover root (© M. van der Heijden), (D) predatory mite (© H. Conrad), (E) isopod (© F. Ashwood), (F) scots pine root colonized by ectomycorrhizal fungi (yellow) (© M. Anthony), (G) earthworm (© G. Brändle), (H) nematode (© A. Murray), (I) corn root colonized by arbuscular mycorrhizal fungi (blue) (© F. Bender), (J) springtail (© F. Ashwood), (K) a common soil bacterium Bacillus (Creative Commons Attribution-Share license, photo by M. Das Murtey and P. Ramasamy), (L) horned mite (© H. Conrad), (M), pseudoscorpion (© F. Ashwood), (N) phage infecting a soil bacterium (© T. de Carvalho), (O) centipede (© F. Ashwood).

Author contributions: M.A.A., S.F.B., and M.G.A.v.d.H. designed research; M.A.A. performed research; M.A.A. analyzed data; and M.A.A., S.F.B., and M.G.A.v.d.H. wrote the paper.

The authors declare no competing interest.

This article is a PNAS Direct Submission.

Copyright © 2023 the Author(s). Published by PNAS. This article is distributed under Creative Commons Attribution-NonCommercial-NoDerivatives License 4.0 (CC BY-NC-ND).

1
To whom correspondence may be addressed. Email:

manthony5955@gmail.com or marcel.vanderheijden@ agroscope.admin.ch.

This article contains supporting information online at

https://www.pnas.org/lookup/suppl/doi:10.1073/pnas. 2304663120/-/DCSupplemental.

Published August 7, 2023.

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