<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD Journal Publishing with OASIS Tables v3.0 20080202//EN" "journalpub-oasis3.dtd">
<article xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:oasis="http://docs.oasis-open.org/ns/oasis-exchange/table" xml:lang="en" dtd-version="3.0"><?xmltex \makeatother\@nolinetrue\makeatletter?>
  <front>
    <journal-meta><journal-id journal-id-type="publisher">BG</journal-id><journal-title-group>
    <journal-title>Biogeosciences</journal-title>
    <abbrev-journal-title abbrev-type="publisher">BG</abbrev-journal-title><abbrev-journal-title abbrev-type="nlm-ta">Biogeosciences</abbrev-journal-title>
  </journal-title-group><issn pub-type="epub">1726-4189</issn><publisher>
    <publisher-name>Copernicus Publications</publisher-name>
    <publisher-loc>Göttingen, Germany</publisher-loc>
  </publisher></journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.5194/bg-18-2063-2021</article-id><title-group><article-title>Similar importance of edaphic and climatic factors for controlling soil organic carbon stocks of the world</article-title><alt-title>Global SOC stocks as impacted by edaphic and climatic factors</alt-title>
      </title-group><?xmltex \runningtitle{Global SOC stocks as impacted by edaphic and climatic factors}?><?xmltex \runningauthor{Z. Luo et al.}?>
      <contrib-group>
        <contrib contrib-type="author" corresp="yes" rid="aff1">
          <name><surname>Luo</surname><given-names>Zhongkui</given-names></name>
          <email>luozk@zju.edu.cn</email>
        <ext-link>https://orcid.org/0000-0002-6744-6491</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff2">
          <name><surname>Viscarra-Rossel</surname><given-names>Raphael A.</given-names></name>
          
        <ext-link>https://orcid.org/0000-0003-1540-4748</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Qian</surname><given-names>Tian</given-names></name>
          
        <ext-link>https://orcid.org/0000-0003-2754-1475</ext-link></contrib>
        <aff id="aff1"><label>1</label><institution>College of Environmental and Resource Sciences, Zhejiang University,
Hangzhou, Zhejiang 310058, China</institution>
        </aff>
        <aff id="aff2"><label>2</label><institution>Soil and Landscape Science, School of Molecular and Life Sciences,
Curtin University, Perth, WA 6845, Australia</institution>
        </aff>
      </contrib-group>
      <author-notes><corresp id="corr1">Zhongkui Luo (luozk@zju.edu.cn)</corresp></author-notes><pub-date><day>22</day><month>March</month><year>2021</year></pub-date>
      
      <volume>18</volume>
      <issue>6</issue>
      <fpage>2063</fpage><lpage>2073</lpage>
      <history>
        <date date-type="received"><day>1</day><month>August</month><year>2020</year></date>
           <date date-type="rev-request"><day>11</day><month>August</month><year>2020</year></date>
           <date date-type="rev-recd"><day>2</day><month>February</month><year>2021</year></date>
           <date date-type="accepted"><day>15</day><month>February</month><year>2021</year></date>
      </history>
      <permissions>
        <copyright-statement>Copyright: © 2021 Zhongkui Luo et al.</copyright-statement>
        <copyright-year>2021</copyright-year>
      <license license-type="open-access"><license-p>This work is licensed under the Creative Commons Attribution 4.0 International License. To view a copy of this licence, visit <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">https://creativecommons.org/licenses/by/4.0/</ext-link></license-p></license></permissions><self-uri xlink:href="https://bg.copernicus.org/articles/18/2063/2021/bg-18-2063-2021.html">This article is available from https://bg.copernicus.org/articles/18/2063/2021/bg-18-2063-2021.html</self-uri><self-uri xlink:href="https://bg.copernicus.org/articles/18/2063/2021/bg-18-2063-2021.pdf">The full text article is available as a PDF file from https://bg.copernicus.org/articles/18/2063/2021/bg-18-2063-2021.pdf</self-uri>
      <abstract><title>Abstract</title>
    <p id="d1e105">Soil organic carbon (SOC) accounts for two-thirds of terrestrial
carbon. Yet, the role of soil physicochemical properties in regulating SOC
stocks is unclear, inhibiting reliable SOC predictions under land use and
climatic changes. Using legacy observations from 141 584 soil profiles
worldwide, we disentangle the effects of biotic, climatic and edaphic
factors (a total of 31 variables) on the global spatial distribution of SOC
stocks in four sequential soil layers down to 2 m. The results indicate that
the 31 variables can explain 60 %–70 % of the global variance of SOC in the
four layers, to which climatic variables and edaphic properties each
contribute <inline-formula><mml:math id="M1" display="inline"><mml:mrow><mml:mo>∼</mml:mo><mml:mn mathvariant="normal">35</mml:mn></mml:mrow></mml:math></inline-formula> % except in the top 20 cm soil. In the top
0–20 cm soil, climate contributes much more than soil properties (43 % vs.
31 %), while climate and soil properties show the similar importance in
the 20–50, 50–100 and 100–200 cm soil layers. However, the most important
individual controls are consistently soil-related and include soil texture,
hydraulic properties (e.g. field capacity) and pH. Overall, soil properties
and climate are the two dominant controls. Apparent carbon inputs
represented by net primary production, biome type and agricultural
cultivation are secondary, and their relative contributions were
<inline-formula><mml:math id="M2" display="inline"><mml:mrow><mml:mo>∼</mml:mo><mml:mn mathvariant="normal">10</mml:mn></mml:mrow></mml:math></inline-formula> % in all soil depths. This dominant effect of
individual soil properties challenges the current climate-driven framework of SOC dynamics and needs to be considered to reliably project SOC changes for
effective carbon management and climate change mitigation.</p>
  </abstract>
    </article-meta>
  </front>
<body>
      

      <?xmltex \hack{\newpage}?>
<sec id="Ch1.S1" sec-type="intro">
  <label>1</label><title>Introduction</title>
      <p id="d1e139">Soil organic carbon (SOC) represents the largest pool of terrestrial carbon (Le Quéré et al., 2016; Batjes, 2016) and plays a key role in
combating climate change and ensuring soil productivity. To better manage
land for maintaining SOC levels or enhancing carbon sequestration, it is
vital to elucidate controlling factors of SOC stabilization and stock. As an
important soil property, it is reasonable to expect that SOC might be
integrally influenced by five predominant factors controlling soil
development and formation – namely, climate, organisms, topography, parent
materials and time (Jenny, 1941). However, climate is usually prioritized
and considered to be critical (Carvalhais et al., 2014)
because of its direct effect on soil carbon inputs via photosynthetic carbon
assimilation and output via microbial decomposition. But climate-driven
predictions of SOC dynamics (e.g. using Earth system models) remain largely
uncertain, particularly across large extents (Todd-Brown et al.,
2013; Bradford et al., 2016; Luo et al., 2017).</p>
      <p id="d1e142">A primary source of the uncertainty is our poor understanding of how edaphic
properties regulate SOC stabilization and stock in soil (Davidson and Janssens, 2006; Dungait et al., 2012). For
example, SOC can be physically protected from decomposition via occlusion
within soil aggregates and adsorption onto minerals (Six et al.,
2000), which create physical barriers preventing microorganisms from decomposing carbon sources (Doetterl et al., 2015; Schimel and Schaeffer, 2012)
regardless of climate conditions, but how this protection influences global
SOC stocks is unclear. Additionally, the soil physicochemical environment
controls the supply of water, nutrients, oxygen and other resources,<?pagebreak page2064?> which
are required for microbial communities to utilize SOC as well as for plant
carbon assimilation to replenish soil carbon pool. Considering the large
spatial variability of soil properties globally, we need to understand the
edaphic controls of SOC better. By explicitly considering the effect of soil
physicochemical properties, we hope to promote a review of climate-driven
frameworks of SOC dynamics.</p>
      <p id="d1e145">In addition to our incomplete understanding of the general importance of
soil properties in regulating SOC stocks, whether and how their effects vary
with soil depth are also unclear. Most studies focus on topsoil layers
(e.g. 0–30 cm), even though, globally, deeper soil layers (below 30 cm)
store more carbon than topsoils (Jobbágy and Jackson, 2000; Batjes,
2016). Like the topsoil SOC pool, the subsoil SOC pool may actively respond
to climate and land use changes. Studies of whole-soil profiles have
observed increased loss of subsoil SOC under warming (Pries et al.,
2017; Melillo et al., 2017; Zhou et al., 2018) as well as under additional
supply of fresh carbon (Fontaine et al., 2007). Land uses such as
cropping and grazing can also induce substantial subsoil SOC loss (Sanderman et al., 2017), which is concerning because of the potential
adverse effects of climate and land use changes. It is therefore imperative
that we better understand the controlling factors of SOC in deep soil layers
as this will help to develop unbiased strategies to effectively manage
whole-soil profile carbon.</p>
      <p id="d1e148">Here, we aim to disentangle the relative importance of climatic, biotic and
edaphic controls on SOC stocks globally in different soil layers. To do so,
we assessed data from 141 584 whole-soil profiles across the globe including
measurements of SOC and other soil physicochemical properties, collated by
the World Soil Information Service (WoSIS) (Batjes et al.,
2017). For each profile, 19 climate-related covariates reflecting
seasonality, intra- and inter-annual variability of climate were obtained
from the WorldClim database (Fick and Hijmans, 2017); the MODIS NPP (net
primary productivity) product (Zhao and Running, 2010) was used to infer
apparent carbon input into soil; and the MODIS land cover product (Friedl et al., 2010) was used to obtain land use information. Using these datasets, we disentangled the relative importance of biotic, climatic and
edaphic covariates (a total of 31 variables, Table 1) in controlling the
spatial variance in SOC stocks worldwide in four sequential soil layers
(i.e. 0–20, 20–50, 50–100 and 100–200 cm) and identified the
correlations between SOC stock and the most important variables.</p>

<?xmltex \floatpos{t}?><table-wrap id="Ch1.T1" specific-use="star"><?xmltex \currentcnt{1}?><label>Table 1</label><caption><p id="d1e155">Covariates used in the modelling of soil carbon stocks across the
globe.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="5">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="left"/>
     <oasis:colspec colnum="3" colname="col3" align="left"/>
     <oasis:colspec colnum="4" colname="col4" align="left"/>
     <oasis:colspec colnum="5" colname="col5" align="left"/>
     <oasis:thead>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1">Covariates</oasis:entry>
         <oasis:entry colname="col2">Code</oasis:entry>
         <oasis:entry colname="col3">Description</oasis:entry>
         <oasis:entry colname="col4">Unit</oasis:entry>
         <oasis:entry colname="col5">Data sources</oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>
         <oasis:entry colname="col1">Soil</oasis:entry>
         <oasis:entry colname="col2">TCEQ</oasis:entry>
         <oasis:entry colname="col3">Calcium carbonate content</oasis:entry>
         <oasis:entry colname="col4">g kg<inline-formula><mml:math id="M3" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col5">Batjes et al. (2017)</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">properties</oasis:entry>
         <oasis:entry colname="col2">ECEC</oasis:entry>
         <oasis:entry colname="col3">Effective cation exchange capacity</oasis:entry>
         <oasis:entry colname="col4">cmol kg<inline-formula><mml:math id="M4" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2">ELCO</oasis:entry>
         <oasis:entry colname="col3">Electrical conductivity</oasis:entry>
         <oasis:entry colname="col4">dS m<inline-formula><mml:math id="M5" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2">Clay</oasis:entry>
         <oasis:entry colname="col3">Clay content</oasis:entry>
         <oasis:entry colname="col4">%</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2">Sand</oasis:entry>
         <oasis:entry colname="col3">Sand content</oasis:entry>
         <oasis:entry colname="col4">%</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2">Silt</oasis:entry>
         <oasis:entry colname="col3">Silt content</oasis:entry>
         <oasis:entry colname="col4">%</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2">pH</oasis:entry>
         <oasis:entry colname="col3">pH measured in H<inline-formula><mml:math id="M6" display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula>O</oasis:entry>
         <oasis:entry colname="col4">–</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2">LL15</oasis:entry>
         <oasis:entry colname="col3">Lower limit obtained at a matric potential of 1500 kPa</oasis:entry>
         <oasis:entry colname="col4">%</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2">DUL</oasis:entry>
         <oasis:entry colname="col3">Drained upper limit obtained at a matric potential of 33 kPa</oasis:entry>
         <oasis:entry colname="col4">%</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Climatic</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M7" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>1</oasis:entry>
         <oasis:entry colname="col3">Annual mean temperature</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M8" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C</oasis:entry>
         <oasis:entry colname="col5">WorldClim (Fick</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">variables</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M9" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>2</oasis:entry>
         <oasis:entry colname="col3">Mean diurnal range</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M10" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C</oasis:entry>
         <oasis:entry colname="col5">and Hijmans, 2017)</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M11" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>3</oasis:entry>
         <oasis:entry colname="col3">Isothermality (<inline-formula><mml:math id="M12" display="inline"><mml:mrow><mml:mi>T</mml:mi><mml:mn mathvariant="normal">2</mml:mn><mml:mo>/</mml:mo><mml:mi>T</mml:mi><mml:mn mathvariant="normal">7</mml:mn><mml:mo>×</mml:mo><mml:mn mathvariant="normal">100</mml:mn></mml:mrow></mml:math></inline-formula>)</oasis:entry>
         <oasis:entry colname="col4">%</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M13" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>4</oasis:entry>
         <oasis:entry colname="col3">Temperature seasonality (standard deviation <inline-formula><mml:math id="M14" display="inline"><mml:mrow><mml:mo>×</mml:mo><mml:mn mathvariant="normal">100</mml:mn></mml:mrow></mml:math></inline-formula>)</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M15" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M16" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>5</oasis:entry>
         <oasis:entry colname="col3">Max temperature of warmest month</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M17" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M18" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>6</oasis:entry>
         <oasis:entry colname="col3">Min temperature of coldest month</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M19" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M20" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>7</oasis:entry>
         <oasis:entry colname="col3">Temperature annual range (<inline-formula><mml:math id="M21" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>5–<inline-formula><mml:math id="M22" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>6)</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M23" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M24" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>8</oasis:entry>
         <oasis:entry colname="col3">Mean temperature of wettest quarter</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M25" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M26" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>9</oasis:entry>
         <oasis:entry colname="col3">Mean temperature of direst quarter</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M27" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M28" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>10</oasis:entry>
         <oasis:entry colname="col3">Mean temperature of warmest quarter</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M29" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M30" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>11</oasis:entry>
         <oasis:entry colname="col3">Mean temperature of coldest quarter</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M31" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M32" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>1</oasis:entry>
         <oasis:entry colname="col3">Annual precipitation</oasis:entry>
         <oasis:entry colname="col4">mm</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M33" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>2</oasis:entry>
         <oasis:entry colname="col3">Precipitation of wettest month</oasis:entry>
         <oasis:entry colname="col4">mm</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M34" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>3</oasis:entry>
         <oasis:entry colname="col3">Precipitation of driest month</oasis:entry>
         <oasis:entry colname="col4">mm</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M35" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>4</oasis:entry>
         <oasis:entry colname="col3">Precipitation seasonality (coefficient of variation)</oasis:entry>
         <oasis:entry colname="col4">%</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M36" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>5</oasis:entry>
         <oasis:entry colname="col3">Precipitation of wettest quarter</oasis:entry>
         <oasis:entry colname="col4">mm</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M37" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>6</oasis:entry>
         <oasis:entry colname="col3">Precipitation of driest quarter</oasis:entry>
         <oasis:entry colname="col4">mm</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M38" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>7</oasis:entry>
         <oasis:entry colname="col3">Precipitation of warmest quarter</oasis:entry>
         <oasis:entry colname="col4">mm</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M39" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>8</oasis:entry>
         <oasis:entry colname="col3">Precipitation of coldest quarter</oasis:entry>
         <oasis:entry colname="col4">mm</oasis:entry>
         <oasis:entry colname="col5"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Other</oasis:entry>
         <oasis:entry colname="col2">Biome</oasis:entry>
         <oasis:entry colname="col3">Biome type</oasis:entry>
         <oasis:entry colname="col4">–</oasis:entry>
         <oasis:entry colname="col5">WWF (Olson et</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry rowsep="1" colname="col2"/>
         <oasis:entry rowsep="1" colname="col3"/>
         <oasis:entry rowsep="1" colname="col4"/>
         <oasis:entry rowsep="1" colname="col5">al., 2001)</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2">NPP</oasis:entry>
         <oasis:entry colname="col3">Net primary productivity</oasis:entry>
         <oasis:entry colname="col4">kg C m<inline-formula><mml:math id="M40" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula> yr<inline-formula><mml:math id="M41" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col5">MODIS (Zhao and</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry rowsep="1" colname="col2"/>
         <oasis:entry rowsep="1" colname="col3"/>
         <oasis:entry rowsep="1" colname="col4"/>
         <oasis:entry rowsep="1" colname="col5">Running, 2010)</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2">Cultivation</oasis:entry>
         <oasis:entry colname="col3">Whether the land is cultivated (yes or no)</oasis:entry>
         <oasis:entry colname="col4">–</oasis:entry>
         <oasis:entry colname="col5">MODIS (Friedl et</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2"/>
         <oasis:entry colname="col3"/>
         <oasis:entry colname="col4"/>
         <oasis:entry colname="col5">al., 2010)</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>

</sec>
<sec id="Ch1.S2">
  <label>2</label><title>Materials and methods</title>
<sec id="Ch1.S2.SS1">
  <label>2.1</label><title>Observed soil profile data and harmonization</title>
      <p id="d1e1070">The World Soil Information Service (WoSIS) collates and manages the largest
database of explicit soil profile observations across the globe (Batjes et al., 2017) which forms the foundation of a
series of digital soil mapping products such as the global SoilGrids (Hengl et al., 2017). The WoSIS dataset is still growing. When we
visited the dataset last on 25 March 2019, there were a total of 141 584 profiles which were used in this study. These profile observations were
quality-assessed and standardized, using consistent procedures (Batjes et al., 2017). In each soil profile, multiple
layers were sampled for determining SOC content and/or other soil
properties. A total of 48 soil properties were recorded with multiple
variates of the same property (e.g. pH measured in H<inline-formula><mml:math id="M42" display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula>O, CaCl<inline-formula><mml:math id="M43" display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula>,
KCl). In the data assessment, we considered nine principal soil
physicochemical properties other than SOC itself in the data analysis (Table 1). Taking the advantage of all measurements, however, other soil properties
were used for missing data imputation (see the Sect. 2.2). The layer
depths are inconsistent between soil profiles. We harmonized all soil
properties including SOC to four standard depths (i.e. 0–20, 20–50,
50–100 and 100–200 cm) using mass-preserving splines (Bishop et
al., 1999; Malone et al., 2009). This harmonization enables the calculation
of SOC stock in the defined standard layers, making it possible to directly
compare among soil profiles.</p>
</sec>
<sec id="Ch1.S2.SS2">
  <label>2.2</label><title>SOC stock calculation and filling missing values</title>
      <p id="d1e1099">We calculated SOC stock (<inline-formula><mml:math id="M44" display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="normal">SOC</mml:mi><mml:mi mathvariant="normal">s</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>, kg C m<inline-formula><mml:math id="M45" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>) in each standard depth as
            <disp-formula id="Ch1.E1" content-type="numbered"><label>1</label><mml:math id="M46" display="block"><mml:mrow><mml:msub><mml:mi mathvariant="normal">SOC</mml:mi><mml:mi mathvariant="normal">s</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mi mathvariant="normal">OC</mml:mi><mml:mn mathvariant="normal">100</mml:mn></mml:mfrac></mml:mstyle><mml:mo>⋅</mml:mo><mml:mi>D</mml:mi><mml:mo>⋅</mml:mo><mml:mi mathvariant="normal">BD</mml:mi><mml:mo>⋅</mml:mo><mml:mfenced open="(" close=")"><mml:mrow><mml:mn mathvariant="normal">1</mml:mn><mml:mo>-</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mi>G</mml:mi><mml:mn mathvariant="normal">100</mml:mn></mml:mfrac></mml:mstyle></mml:mrow></mml:mfenced><mml:mo>,</mml:mo></mml:mrow></mml:math></disp-formula>
          where OC is the weight percentage SOC content in the fine earth fraction <inline-formula><mml:math id="M47" display="inline"><mml:mrow><mml:mi mathvariant="italic">&lt;</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:math></inline-formula> mm, <inline-formula><mml:math id="M48" display="inline"><mml:mi>D</mml:mi></mml:math></inline-formula> the soil depth (i.e. 0.2, 0.3, 0.5 or 1 m in this study),
BD the bulk density of the fine earth fraction <inline-formula><mml:math id="M49" display="inline"><mml:mrow><mml:mi mathvariant="italic">&lt;</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:math></inline-formula> mm (kg m<inline-formula><mml:math id="M50" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula>)
and <inline-formula><mml:math id="M51" display="inline"><mml:mi>G</mml:mi></mml:math></inline-formula> the volume percentage gravel content (<inline-formula><mml:math id="M52" display="inline"><mml:mrow><mml:mi mathvariant="italic">&gt;</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:math></inline-formula> mm) of soil.
Amongst the 141 584 soil profiles, unfortunately, only 9672 profiles have
all the measurements of OC, <inline-formula><mml:math id="M53" display="inline"><mml:mi>D</mml:mi></mml:math></inline-formula>, BD and <inline-formula><mml:math id="M54" display="inline"><mml:mi>G</mml:mi></mml:math></inline-formula> to enable direct calculation of SOC stock.
We call these profiles “stock profiles”.</p>
      <?pagebreak page2065?><p id="d1e1239">Another 82 734 profiles have measured OC (i.e. the weight percentage SOC
content), but BD and/or <inline-formula><mml:math id="M55" display="inline"><mml:mi>G</mml:mi></mml:math></inline-formula> are missing. We call these profiles “content
profiles”. To utilize and take advantage of all OC measurements, we used
generalized boosted regression modelling (GBM) to perform imputations (i.e.
fill missing data). As such, SOC<inline-formula><mml:math id="M56" display="inline"><mml:msub><mml:mi/><mml:mi mathvariant="normal">s</mml:mi></mml:msub></mml:math></inline-formula> can be estimated. To do so, for BD and <inline-formula><mml:math id="M57" display="inline"><mml:mi>G</mml:mi></mml:math></inline-formula> in
each standard soil depth, GBM was developed based on all measurements of
that property (e.g. BD) in the 141 584 profiles with 32 other soil
properties. Total carbon which includes organic and inorganic carbon and
another nine soil properties (Table 1) which were used as predictors of SOC
stocks were excluded as covariates (i.e. predictors). The final GBM model
was validated using 10-fold cross validation repeated 10 times and applied
to predict missing values of BD and G. A total of 92 406 SOC profiles
including stock profiles and content profiles with relevant
measurements of nine soil properties (Table 1) was obtained and used
to assess the effects of various variables on SOC<inline-formula><mml:math id="M58" display="inline"><mml:msub><mml:mi/><mml:mi mathvariant="normal">s</mml:mi></mml:msub></mml:math></inline-formula> (Sect. 2.4). These soil profiles cover 13 major biome groups although the profile numbers
vary from 472 in flooded grasslands and savannas to 24 382 in temperate
broadleaf and mixed forests (Fig. 1). The profiles also cover various
climate conditions across the globe with the mean annual temperature ranging
from <inline-formula><mml:math id="M59" display="inline"><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">19.6</mml:mn></mml:mrow></mml:math></inline-formula> to 30.7 <inline-formula><mml:math id="M60" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C and mean precipitation ranging from 0 to
667.4 cm yr<inline-formula><mml:math id="M61" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:math></inline-formula> (Fig. 1). The prediction error of the GBM was propagated into the calculation of SOC<inline-formula><mml:math id="M62" display="inline"><mml:msub><mml:mi/><mml:mi mathvariant="normal">s</mml:mi></mml:msub></mml:math></inline-formula> to account for uncertainty resulting from data imputation (see Sect. 2.4).</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F1"><?xmltex \currentcnt{1}?><?xmltex \def\figurename{Figure}?><label>Figure 1</label><caption><p id="d1e1317">Distribution of soil profiles with soil carbon data in relation to
mean annual air temperature and precipitation. Different colours show the
biome type to which the soil profile belongs to. Numbers in parentheses show
the number of soil profiles in the relevant biome. Some soil profiles (1382)
were not included as climate and/or biome type could not be identified for
them.</p></caption>
          <?xmltex \igopts{width=241.848425pt}?><graphic xlink:href="https://bg.copernicus.org/articles/18/2063/2021/bg-18-2063-2021-f01.png"/>

        </fig>

</sec>
<sec id="Ch1.S2.SS3">
  <label>2.3</label><title>Biotic and climatic covariates</title>
      <p id="d1e1334">For each SOC profile, NPP was extracted from the MODIS NPP product (Zhao
and Running, 2010). The NPP product includes the annual NPP from 2001 to
2015 at the resolution of 1 km<inline-formula><mml:math id="M63" display="inline"><mml:msup><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:math></inline-formula>, which was estimated by analysing
satellite data from MODIS using the global MODIS NPP algorithm (Zhao et
al., 2005; Zhao and Running, 2010). NPP is the net carbon gained by plants
(i.e. the difference between gross primary productivity and autotrophic
respiration). If assuming a steady state of the vegetation (i.e. no
long-term directional change of carbon biomass in plants), NPP will end up
in soil via rhizodeposition and litter fall and will be equal to total carbon
input into soil. Here we calculated the average NPP based on the data from
2001 to 2015 and called this average NPP the apparent carbon input to soil,
acknowledging that not all ecosystems are at strict steady state,
particularly those ecosystems (e.g. croplands) actively interacting with human activities. The WWF (World Wildlife Fund) map of the terrestrial ecoregions
of the world (Olson et al., 2001;<?pagebreak page2066?> <uri>https://www.worldwildlife.org/publications/terrestrial-ecoregions-of-the-world</uri>, last access: 18 March 2021)
was used to extract the biome type for each location. The MODIS land cover
map (Friedl et al., 2010) at the same resolution of NPP databases was
used to identify whether or not the land is cultivated (i.e. land cover type of
croplands and cropland/natural vegetation mosaic) at the location of each
soil profile.</p>
      <p id="d1e1349">In addition to NPP, land cover and biome type, 19 climatic variables (Table 1) for each SOC profile were obtained from the WorldClim version 2 (Fick and Hijmans, 2017). The WorldClim version 2 calculates biologically
meaningful variables using monthly temperature and precipitation during the
period 1970–2000. The data at the same spatial resolution of the NPP data
(i.e. <inline-formula><mml:math id="M64" display="inline"><mml:mrow><mml:mo>∼</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:math></inline-formula> km<inline-formula><mml:math id="M65" display="inline"><mml:msup><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:math></inline-formula>) were used in this study. Eleven of the 19
climatic variables are temperature-related (Table 1), and eight are
precipitation-related (Table 1). These variables reflect the seasonality,
intra- and inter-annual variability of climate, which would have both a direct
(via decomposition thus carbon outputs from soil) and an indirect (via carbon
assimilation thus carbon inputs to soil) effect on SOC stock.</p>
</sec>
<sec id="Ch1.S2.SS4">
  <label>2.4</label><title>Data analysis</title>
      <p id="d1e1379">A machine learning-based statistical model – boosted regression trees (BRTs)
– was performed to explain the variability of SOC<inline-formula><mml:math id="M66" display="inline"><mml:msub><mml:mi/><mml:mi mathvariant="normal">s</mml:mi></mml:msub></mml:math></inline-formula> across the globe and
identify important controlling factors. A big advantage of the BRT model is
its ability to model high-dimensional dataset, taking into account
non-linearities and interplay (Elith et al., 2008). Using the BRT model,
we modelled SOC<inline-formula><mml:math id="M67" display="inline"><mml:msub><mml:mi/><mml:mi mathvariant="normal">s</mml:mi></mml:msub></mml:math></inline-formula> in each standard depth as a function of edaphic variables
in that depth, climatic and biotic variables (Table 1):
            <disp-formula id="Ch1.E2" content-type="numbered"><label>2</label><mml:math id="M68" display="block"><mml:mrow><mml:msub><mml:mi mathvariant="normal">SOC</mml:mi><mml:mi mathvariant="normal">s</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mi>f</mml:mi><mml:mfenced close=")" open="("><mml:mrow><mml:mi mathvariant="normal">edaphic</mml:mi><mml:mo>,</mml:mo><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">climatic</mml:mi><mml:mo>,</mml:mo><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">biome</mml:mi><mml:mo>,</mml:mo><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">NPP</mml:mi><mml:mo>,</mml:mo><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">cultivation</mml:mi></mml:mrow></mml:mfenced><mml:mo>.</mml:mo></mml:mrow></mml:math></disp-formula>
          We used a 10-fold cross validation to constrain the BRT model in R 3.6.1 (R
Core Team, 2019) using algorithms implemented in the R package <italic>dismo</italic>. The amount
of variance in SOC<inline-formula><mml:math id="M69" display="inline"><mml:msub><mml:mi/><mml:mi mathvariant="normal">s</mml:mi></mml:msub></mml:math></inline-formula> explained by the model was assessed by the
coefficient of determination (<inline-formula><mml:math id="M70" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>). To assess the potential uncertainty
induced by the uneven distribution of soil profiles across the globe as well
as the imputation of missing BD and <inline-formula><mml:math id="M71" display="inline"><mml:mi>G</mml:mi></mml:math></inline-formula> for estimating SOC<inline-formula><mml:math id="M72" display="inline"><mml:msub><mml:mi/><mml:mi mathvariant="normal">s</mml:mi></mml:msub></mml:math></inline-formula>, we conducted
200 bootstrapping simulations (i.e. resample all soil profiles with
replacement). For each bootstrap sample, SOC<inline-formula><mml:math id="M73" display="inline"><mml:msub><mml:mi/><mml:mi mathvariant="normal">s</mml:mi></mml:msub></mml:math></inline-formula>, if BD and <inline-formula><mml:math id="M74" display="inline"><mml:mi>G</mml:mi></mml:math></inline-formula> are missing,
was recalculated using BD and <inline-formula><mml:math id="M75" display="inline"><mml:mi>G</mml:mi></mml:math></inline-formula> imputed by GBM plus an error randomly
sampled from the distribution of imputation error. Using the new SOC<inline-formula><mml:math id="M76" display="inline"><mml:msub><mml:mi/><mml:mi mathvariant="normal">s</mml:mi></mml:msub></mml:math></inline-formula>
estimations, then, a new BRT model was fitted.</p>
      <p id="d1e1515">Considering the potential collinearity in the 19 climatic variables as well
as in the nine soil properties, the BRT model was conducted using their
principal components. That is, a principal component analysis (PCA) was
performed to eliminate potential correlations in the soil and climatic
variables, respectively. The important principal components (PCs) with
variances of greater than 1 were retained in the BRT model based on Kaiser's
criterion (Kaiser, 1960). The PCA was performed using the function prcomp in
the package stats in R 3.6.1 (R Core Team, 2019). In addition, in order to
demonstrate the importance of soil properties, we fitted another set of BRT
models without soil properties. The model performance with and without soil
properties were compared in terms of explaining the variance of SOC stocks
across the globe.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F2" specific-use="star"><?xmltex \currentcnt{2}?><?xmltex \def\figurename{Figure}?><label>Figure 2</label><caption><p id="d1e1520">Loadings of 19 climatic variables <bold>(a)</bold> and nine soil properties <bold>(b)</bold> to
the two most important principal components. <inline-formula><mml:math id="M77" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>1, annual mean temperature;
<inline-formula><mml:math id="M78" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>2, mean diurnal range; <inline-formula><mml:math id="M79" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>3, isothermality; <inline-formula><mml:math id="M80" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>4, temperature seasonality; <inline-formula><mml:math id="M81" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>5,
max temperature of warmest month; <inline-formula><mml:math id="M82" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>6, min temperature of coldest month; <inline-formula><mml:math id="M83" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>7,
temperature annual range; <inline-formula><mml:math id="M84" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>8, mean temperature of wettest quarter; <inline-formula><mml:math id="M85" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>9, mean
temperature of driest quarter; <inline-formula><mml:math id="M86" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>10, mean temperature of warmest quarter;
<inline-formula><mml:math id="M87" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>11, mean temperature of coldest quarter; <inline-formula><mml:math id="M88" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>1, annual precipitation; <inline-formula><mml:math id="M89" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>2,
precipitation of wettest month; <inline-formula><mml:math id="M90" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>3, precipitation of driest month; <inline-formula><mml:math id="M91" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>4,
precipitation seasonality; <inline-formula><mml:math id="M92" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>5, precipitation of wettest quarter; <inline-formula><mml:math id="M93" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>6,
precipitation of driest quarter; <inline-formula><mml:math id="M94" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>7, precipitation of warmest quarter; <inline-formula><mml:math id="M95" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>8, precipitation of coldest quarter. DUL, drained upper limit of soil; LL15,
lower limit of soil; ELCO, electrical conductivity; ECEC, effective cation
exchange capacity; TCEQ, calcium carbonate content; sand, silt and clay, the
fraction of sand, silt and clay content of soil; pH, soil pH. See Table 1
for more details about the variables.</p></caption>
          <?xmltex \igopts{width=341.433071pt}?><graphic xlink:href="https://bg.copernicus.org/articles/18/2063/2021/bg-18-2063-2021-f02.png"/>

        </fig>

      <p id="d1e1672">The BRT model allows the estimation of the relative influence of each
individual variable in predicting SOC<inline-formula><mml:math id="M96" display="inline"><mml:msub><mml:mi/><mml:mi mathvariant="normal">s</mml:mi></mml:msub></mml:math></inline-formula>, i.e. the percentage contribution
of variables in the model. The relative influence is calculated based on the
times a variable selected for splitting when growing a tree, weighted by
squared model improvement due to that splitting, and then averaged over all
fitted trees which were determined by the algorithm when adding more trees
cannot reduce prediction residuals (Elith et al., 2008; Friedman and
Meulman, 2003). As such, the larger the relative influence of a variable,
the stronger the effect on SOC<inline-formula><mml:math id="M97" display="inline"><mml:msub><mml:mi/><mml:mi mathvariant="normal">s</mml:mi></mml:msub></mml:math></inline-formula>. In addition, we also calculated the
95 % confidence intervals as the 2.5 % and 97.5 quantiles of the
relative influence estimated by 200 bootstrapping simulations, which
represent the uncertainty in the importance of variables. To facilitate
interpretation, the relative influence of each variable is scaled so that
the sum of the influence of all variables is equal to 100. The overall
relative influences of<?pagebreak page2067?> edaphic (i.e. the sum relative importance of all
soil-related variables) and climatic (i.e. the sum relative importance of
all climate-related variables) variables as well as biome type, NPP and
cultivation were also calculated and compared. As we have 200 estimations
(i.e. 200 bootstraps) of the relative influence, we calculated a weighted
average relative influence for each variable with weights based on the
<inline-formula><mml:math id="M98" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> of each BRT model.</p>
</sec>
</sec>
<sec id="Ch1.S3">
  <label>3</label><title>Results</title>
      <p id="d1e1713">The 19 climatic variables could be represented by four principal components
(PCs, i.e. Climate1–4, which were selected by Kaiser's criterion) which
could explain 88 % of their variance (Fig. 2; only the first two PCs were
shown); and 72 % of the variance in nine soil properties could be
explained by three PCs (i.e. Soil1–3, Fig. 2). For Climate1–4, the most
important contributing variables were <inline-formula><mml:math id="M99" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>11 (mean temperature of coldest
quarter), <inline-formula><mml:math id="M100" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>6 (precipitation of driest quarter), <inline-formula><mml:math id="M101" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>5 (max temperature of
warmest month) and <inline-formula><mml:math id="M102" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>7 (precipitation of warmest quarter), respectively. For
Soil1–3, the most important contributing variables were sand content, pH and
silt content, respectively (Fig. 2). Using Climate1–4, NPP, biome type and
cultivation as predictors, the BRT model could explain 53 %, 46 %,
42 % and 49 % of the variance of SOC stocks in the 0–20, 20–50, 50–100
and 100–200 cm soil layers across the globe, respectively (Fig. 3). If
Soil1–3 were included, an additional 18 %, 18 %, 20 % and 13 % of the
variance could be explained in the four layers, respectively (Fig. 3). This
result demonstrated that soil properties must be considered in order to
explain the spatial variability of SOC stocks across the globe. However, it
is noteworthy that the fitted model overestimated low SOC stocks and
underestimated high SOC stocks. This bias of model performance at both
ends of observed SOC stocks is common across all four depths (Fig. 3).</p>
      <p id="d1e1744">The results of the BRT model including soil properties (i.e. Soil1–3)
indicated that Soil1 (i.e. the first PC of soil properties) was
consistently the most important individual control of SOC stocks in the
three deeper soil layers (i.e. 20–50, 50–100 and 100–200 cm; Fig. 4). On
average, Soil1 alone contributed 21 % (with 95 % confidence intervals
ranging from 17 %–24 %), 23 % (20 %–28 %) and 22 % (18 %–26 %) to the
explained variance of SOC stocks in the three deeper soil layers,
respectively (Fig. 4). In the top 20 cm soil layer, Climate2 was the most
important, contributing 19 % (15 %–23 %) to the explained variance of SOC
stocks, and Soil1 was the second most important and contributed 18 %
(16 %–20 %). In the three deeper layers, the second most important
contributors were NPP, biome type and Climate3, respectively (Fig. 4).</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F3"><?xmltex \currentcnt{3}?><?xmltex \def\figurename{Figure}?><label>Figure 3</label><caption><p id="d1e1749">An example of the performance of boosted regression trees in
explaining soil organic carbon stocks in four standard soil depths across
the globe. <bold>(a)</bold> 0–20 cm, <bold>(b)</bold> 20–50 cm, <bold>(c)</bold> 50–100 cm and <bold>(d)</bold> 100–200 cm.
The data were natural-logarithm-transformed. The dashed line shows the <inline-formula><mml:math id="M103" display="inline"><mml:mrow><mml:mn mathvariant="normal">1</mml:mn><mml:mo>:</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:math></inline-formula>
line. Chocolate and green circles show the results without and with
predictors of soil properties, respectively.</p></caption>
        <?xmltex \igopts{width=241.848425pt}?><graphic xlink:href="https://bg.copernicus.org/articles/18/2063/2021/bg-18-2063-2021-f03.png"/>

      </fig>

      <?xmltex \floatpos{t}?><fig id="Ch1.F4"><?xmltex \currentcnt{4}?><?xmltex \def\figurename{Figure}?><label>Figure 4</label><caption><p id="d1e1785">The relative influence of individual biotic, climatic and edaphic
variables influencing global soil organic carbon stocks. Upper limit <bold>(a)</bold>, mean <bold>(b)</bold> and lower limit <bold>(c)</bold> show the
97.5 %, average and 2.5 % quantiles of 200 bootstrapping simulations,
respectively. Biome, biome type; Climate1–4, the four most important
principal components of 19 climatic variables; cultivation, whether or not
the land is cultivated (yes or no); NPP, net primary production; Soil1–3,
the three most important principal components of nine soil properties.</p></caption>
        <?xmltex \igopts{width=199.169291pt}?><graphic xlink:href="https://bg.copernicus.org/articles/18/2063/2021/bg-18-2063-2021-f04.png"/>

      </fig>

      <p id="d1e1803">Summing the relative importance of individual variables, the overall effect
of soil properties was relatively consistent among the four layers,
accounting for 30 %–40 % of the overall influence of all assessed variables
respectively, but they were more important in the deepest two layers than in the
first top layer (Fig. 5). The overall relative influence of climate was
significantly higher than that of soil in the top 20 cm soil layer (43 %
vs. 31 %; Fig. 5). In the three deeper soil layers, the overall influences
of climatic<?pagebreak page2068?> variables and soil properties were comparable and did not show a
significant difference. Overall, climatic variables accounted for 43 %
(38 %–47 %), 36 % (32 %–40 %), 33 % (28 %–37 %) and 35 % (31 %–39 %)
in the four layers, respectively; and soil properties accounted for 30 %
(27 %–33 %), 35 % (30 %–39 %), 39 % (35 %–43 %) and 37 % (33 %–41 %),
respectively (Fig. 5). The relevant influence of the remaining three variables (i.e. NPP, biome type and cultivation) was secondary and marginal
(<inline-formula><mml:math id="M104" display="inline"><mml:mrow><mml:mo>∼</mml:mo><mml:mn mathvariant="normal">10</mml:mn></mml:mrow></mml:math></inline-formula> % in terms of relevant influence) compared to climate
and soil variables and together accounted for the remaining <inline-formula><mml:math id="M105" display="inline"><mml:mrow><mml:mo>∼</mml:mo><mml:mn mathvariant="normal">30</mml:mn></mml:mrow></mml:math></inline-formula> % of the explained variance. With increasing soil depth, in general,
the relevant influence of climate was decreased, while the influence of soil
was increased. However, the overall influences of climate and soil remained
relatively stable at the level of 70 %. These results demonstrate the
comparable and primary effects of climate and soil properties on SOC stocks.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F5"><?xmltex \currentcnt{5}?><?xmltex \def\figurename{Figure}?><label>Figure 5</label><caption><p id="d1e1828">The overall relative influence of edaphic, climatic and biotic
variables on soil organic carbon stocks in four soil depths across the
globe. The overall relative influence for climate and soil is calculated as
the sum of the relative influence of their individual variables (which is
shown in Fig. 4). Error bars show the 95 % confidence interval estimated
based 200 bootstrapping simulations.</p></caption>
        <?xmltex \igopts{width=199.169291pt}?><graphic xlink:href="https://bg.copernicus.org/articles/18/2063/2021/bg-18-2063-2021-f05.png"/>

      </fig>

</sec>
<sec id="Ch1.S4">
  <label>4</label><title>Discussion</title>
<sec id="Ch1.S4.SS1">
  <label>4.1</label><title>The importance of soil properties</title>
      <?pagebreak page2069?><p id="d1e1853">A series of soil properties may directly or indirectly affect SOC dynamic
processes via influencing carbon inputs to soil, microbial activity and
accessibility of carbon substrates to microbes and thereby SOC stocks. Sand
content (which is the most important contributor to the first PC of soil),
for example, has significant effects on the formation and transformation of
soil aggregates which regulate the stability of SOC as well as soil porosity and thereby oxygen availability for microbial decomposition of SOC (Dungait et
al., 2012; Six et al., 2002). In addition, soil properties such as LL15 and
DUL have dominant control over soil water dynamics, which further influence
water availability for plant growth. Theoretically, LL15 is close to the permanent plant wilting point; it thus may strongly
regulate plant growth therefore carbon inputs into soil and final SOC
stocks. Together with DUL (i.e. drained upper limit – soil water content
obtained at the matric potential of 33 kPa), LL15 determines the available
water capacity of soil (AWC, i.e. the difference between DUL and LL15), and
thus LL15 would affect SOC stock via its determination on soil AWC, while
AWC couples with a series of soil hydrological processes such as runoff and
drainage, which have direct effects on the vertical/horizontal translocation
of SOC (Luo et al., 2020; Kaiser and Kalbitz, 2012). Soil properties are more
important for controlling SOC stocks in deeper layers than in upper layers.
This phenomenon may be due to the fact that soil structure may have substantial effects on water and oxygen diffusion in deeper layers. Potentially more frequent
waterlogging and low oxygen in subsoil result in additional environmental
constraints inhibiting microbial decomposition of SOC (Huang et
al., 2020).</p>
      <p id="d1e1856">Our results demonstrate the primary control of soil properties on SOC stocks
in the whole-soil profile across the globe. Indeed, the results suggested
that soil-related principal components were consistently the most important
individual influential variables in three deeper soil layers except in the
assessed 0–20 cm soil layer. Soil physical and chemical properties directly
determine the activity of the decomposer community which mediates the
decomposition of soil carbon (Derrien et al., 2014; Foesel et al.,
2014; Bernard et al., 2012). More importantly, soil carbon can be physically
protected from decomposition via occlusion with soil aggregates and binding
with minerals (Lehmann and Kleber, 2015; Dungait et al., 2012; Schmidt et
al., 2011), while the protection capacity is largely determined by soil
physicochemical properties (Six et al., 2000). These physical protection processes may lead to soil-dependent stabilization/destabilization of different soil carbon substrates (Waldrop and Firestone, 2004; Keiluweit et al., 2015; Six
et al., 2002). However, it should be noted that complex interplays of
various soil properties are involved in SOC stabilization and
destabilization processes. It is also difficult to obtain a cause–effect
conclusion on the relationship between a particular soil physicochemical
property and SOC stocks.</p>
</sec>
<sec id="Ch1.S4.SS2">
  <label>4.2</label><title>The importance of climate</title>
      <p id="d1e1867">Few studies have paid particular attention to the dynamics of SOC in
subsoils across large scales. One might expect greater importance of climate
in surface soils as topsoil is at the frontline of the interaction with the
atmosphere. But our results do not show a clearly decreasing importance of
climate with soil depth. Rather, the overall influence of climatic variables
on SOC stocks is statistically similar in all soil layers. In a forest soil,
a recent study found that SOC in the whole-soil profile down to 1 m is
sensitive to warming (Pries et al., 2017). This sensitivity may be
general across the globe. However, it is noteworthy that neither mean annual
temperature nor mean annual precipitation was the most important individual
climatic variable. Rather, climatic variables reflecting seasonal
variability were more important. This result may suggest that, except for
average change trend, it is important to understand the change patterns of
temperature and precipitation under climate change. For example, a number of
studies have demonstrated that extreme climate events (e.g. drought and
heatwaves) have significant effects on the carbon cycle, including soil carbon,
due to their dramatic influence on the transport and availability of water
and energy as well as ecosystem functional processes (Reichstein
et al., 2013). Field observations, particularly via manipulative experiments
of whole-soil profile, are certainly needed to detect how deep soil carbon
responds to climate change as the result may have significant implications
on the fate of deep soil carbon under future climatic conditions.</p>
</sec>
<sec id="Ch1.S4.SS3">
  <label>4.3</label><title>Secondary role of carbon inputs in determining spatial variability of SOC stocks</title>
      <p id="d1e1878">The effect of apparent carbon input, NPP, on SOC stock is generally small in
all assessed soil layers (Fig. 5). This result is in line with findings from
a continental-scale study across sub-Saharan Africa where climate and
geochemistry are more important predictors of SOC content than aboveground
carbon inputs (von Fromm et al.,
2020). The importance of NPP may largely depend on how much NPP ends up in
the soil and how it is translocated to different depths (Wang
et al., 2021). Total NPP may not be a useful indicator of actual carbon
inputs into different soil depths, particularly in deeper layers.
Cultivation, for example, may substantially change the fate of plant
biomass – a large fraction of plant biomass may be harvested as yield or
consumed by livestock and thus does not contribute to soil carbon. This
could explain the phenomena that cultivation (cultivated vs. non-cultivated
in this study) and NPP show the similar importance in general. In addition,
the final importance of carbon inputs may also depend on their quality
(e.g. carbon-to-nitrogen ratio), while NPP alone does not bring such
information. The quality of carbon inputs represented by their nutrient
content and chemical structure plays a vital role in SOC formation and
transformation (Hessen et al., 2004; Jastrow et al., 2007). In our
study, biome type (which shows similar importance to NPP) would partially
reflect the importance of carbon input quality as different biome types have
distinct carbon biomass quality (e.g. wood vs. leaf litter, which<?pagebreak page2070?> are the
main component of NPP). However, here we must to point out that the minor
role of carbon inputs in determining the global spatial distribution of SOC
stocks does not mean that they are not important for local carbon
management. Under the same climatic and edaphic conditions, indeed, carbon
inputs should be the predominant factor controlling if the soil is a carbon
sink or source (Luo et al., 2017).</p>
</sec>
<sec id="Ch1.S4.SS4">
  <label>4.4</label><title>Limitations and future research</title>
      <p id="d1e1889">Although we have used a diverse and representative dataset across the globe
for the analysis, there are still some limitations in the datasets and
assessment. First, our study did not bring detailed land use history and
intensity (such as the time length of cropping and the intensity of grazing)
into the analysis, which may significantly affect SOC stabilization
processes and thus SOC stocks in managed landscapes (Sanderman et al.,
2017). As anthropogenic land use may change from year to year, it is
challenging to accurately explain SOC stock changes in those systems that
experience intensive human disturbances across large extents. Second, all
soil properties including SOC were treated as constant. In reality, however,
some soil properties, particularly chemical variables such as pH, may
actively respond to external disturbance including human activities.
Treating these variables as constant may result in under- or
overestimations of the variable importance if a variable shows marked
temporal variability. Third, in managed systems, the apparent carbon input
represented by NPP may not accurately reflect the real carbon input into
soil (Luo et al., 2018; Pausch and Kuzyakov, 2018) as
discussed above, leading to biased estimation of the importance of C inputs.
In cropping areas, for example, yield harvesting and crop residue removal
certainly reduce the fraction of NPP ending up in the soil. Fourth, we would
like to point out that, albeit edaphic factors appear to be the dominant
individual controls on SOC stock, climate might have an impact on those
edaphic factors and hence SOC stocks in the long term (Jenny, 1941). Indeed,
Luo et al. (2017) have provided evidence that climate not only directly but also indirectly (via its effect on edaphic factors) exerts significant effect on
SOC dynamics. All these limitations should be overcome to provide more
robust predictions on the role of different factors in SOC stabilization and
stock, which will be particularly important for understanding long-term SOC
dynamics in managed systems. Fourth, we would like to note that this study
focused on the controls over the global spatial pattern of SOC stocks and
did not explicitly assess the potential variability of controls at small
scales. Under different land use types, for example, factors controlling SOC
stock would change. A recent study focused on SOC component fractions has
found that continental drivers of SOC stocks were modulated by regional
environmental factors (Viscarra Rossel et al.,
2019). In order to better understand regional-scale factors controlling SOC
dynamics, we should further explore the controls over different spatial
scales. Considering that we only included limited soil properties in our
assessment and different soil properties may play different roles at
different scales, scale-dependent understanding of controls over SOC stocks
is important to make site-specific management practices for sustainable soil
use and carbon management. Finally, soil samples are still limited in some
areas (e.g. tundra and flooded grasslands and savannas) (Batjes,
2016). We do not know much about whether some of the relationships we find
between SOC stocks and predictor variables are universal or maybe
fundamentally different in less studied soils. The uneven distribution of
soil samples may also help to explain the model bias in explaining low and
high SOC stocks (Fig. 3). Our results indicate that there are large
uncertainties in the relative importance of climate and soil depending on
the data used to fit the model (Fig. 5).</p>
</sec>
</sec>
<sec id="Ch1.S5" sec-type="conclusions">
  <label>5</label><title>Conclusions</title>
      <p id="d1e1901">Quantitatively, we have demonstrated the primary role of soil properties
together with climate in regulating SOC stock in the whole-soil profile
across the globe. This result has important implications for understanding
mechanisms of SOC stabilization and destabilization. Previous modelling and
experimental efforts have mostly focused on climatic and biotic aspects, and
many of the studies are over smaller scales. We argue that soil
physicochemical characteristics define the boundary conditions for the
climatic and biotic factors. That is, climatic and biotic factors (e.g.
carbon inputs) can regulate the rate of SOC of shifting from one capacity to
another, but a soil's physicochemical properties (e.g. soil structure) may
inherently determine the SOC stock capacity of soil. It is thus critical to
understand how soil processes mediated by different soil properties in
different soil layers respond to those climatic and biotic factors and land
management practices and feed this information into the prediction of SOC
stock capacity in the whole-soil profile. However, individual soil variables
work together involving complex interactions and non-linear relationships
with each other as well as with climate to regulate SOC stock (Figs. 2 and
4). We need more and better quality data (e.g. following the same soil
sampling and measuring procedure and using a novel approach for monitoring of
soil properties) and innovative methods (Viscarra Rossel et al., 2017)
for representing soil heterogeneity to facilitate robust prediction of SOC
dynamics over large extents. Results of this study further demonstrate that
globally the influence of individual climatic variables on SOC stock is
weaker than the influence of individual soil properties regardless of soil
depth. Current Earth system models are mostly driven by climate, with few
cases having approximated the regulation of soil properties on carbon
stabilization and destabilization (Tang and Riley,
2014; Riley et al., 2014). Undoubtedly, climate has direct effect on plant
growth and thus potential carbon inputs to the soil, but our results
demonstrate that soil properties are also primary controls of global SOC
stocks.<?pagebreak page2071?> Our research highlights the urgent need to consider soil properties
and their interactions with climate to provide more reliable predictions of
SOC stock and changes under climatic and land use changes.</p>
</sec>

      
      </body>
    <back><notes notes-type="codeavailability"><title>Code availability</title>

      <p id="d1e1908">Code used to generate the
results in this study can be reasonably requested from the corresponding author.</p>
  </notes><notes notes-type="dataavailability"><title>Data availability</title>

      <p id="d1e1914">Soil data including soil organic carbon and the considered soil properties from WoSIS are available at <uri>http://www.isric.org/explore/wosis/accessing-wosis-derived-datasets</uri>, last access: 18 March 2021.
Climate data including 19 bioclimatic variables from WorldClim are available
at <uri>https://www.worldclim.org/data/worldclim21.html</uri>, last access: 18 March 2021.</p>
  </notes><notes notes-type="authorcontribution"><title>Author contributions</title>

      <p id="d1e1926">ZL conceived the study and assessed the data; ZL interpreted the results
and wrote the manuscript with the contribution of RAVR; TQ contributed
to data assessment and creating figures.</p>
  </notes><notes notes-type="competinginterests"><title>Competing interests</title>

      <p id="d1e1932">The authors declare that they have no conflict of interest.</p>
  </notes><ack><title>Acknowledgements</title><p id="d1e1938">We thank the people who originally collected the data and made this
invaluable data publicly available.</p></ack><notes notes-type="financialsupport"><title>Financial support</title>

      <p id="d1e1943">This research has been supported by the National Natural Science Foundation of
China (grant no. 41930754), Fundamental Research Funds for the Central Universities (grant no. 2020FZZX001-06), and Research Innovation Foundation for Young Scholars (grant no. K20200203).</p>
  </notes><notes notes-type="reviewstatement"><title>Review statement</title>

      <p id="d1e1949">This paper was edited by Michael Bahn and reviewed by two anonymous referees.</p>
  </notes><ref-list>
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<abstract-html><p>Soil organic carbon (SOC) accounts for two-thirds of terrestrial
carbon. Yet, the role of soil physicochemical properties in regulating SOC
stocks is unclear, inhibiting reliable SOC predictions under land use and
climatic changes. Using legacy observations from 141&thinsp;584 soil profiles
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31&thinsp;%), while climate and soil properties show the similar importance in
the 20–50, 50–100 and 100–200&thinsp;cm soil layers. However, the most important
individual controls are consistently soil-related and include soil texture,
hydraulic properties (e.g. field capacity) and pH. Overall, soil properties
and climate are the two dominant controls. Apparent carbon inputs
represented by net primary production, biome type and agricultural
cultivation are secondary, and their relative contributions were
 ∼ 10&thinsp;% in all soil depths. This dominant effect of
individual soil properties challenges the current climate-driven framework of SOC dynamics and needs to be considered to reliably project SOC changes for
effective carbon management and climate change mitigation.</p></abstract-html>
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