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  <front>
    <journal-meta>
<journal-id journal-id-type="publisher">ESD</journal-id>
<journal-title-group>
<journal-title>Earth System Dynamics</journal-title>
<abbrev-journal-title abbrev-type="publisher">ESD</abbrev-journal-title>
<abbrev-journal-title abbrev-type="nlm-ta">Earth Syst. Dynam.</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">2190-4987</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/esd-7-745-2016</article-id><title-group><article-title>Impacts of land-use history on the recovery of ecosystems after agricultural abandonment</article-title>
      </title-group><?xmltex \runningtitle{Impacts of land-use history on the recovery of ecosystems after agricultural abandonment}?><?xmltex \runningauthor{A.~Krause et al.}?>
      <contrib-group>
        <contrib contrib-type="author" corresp="yes" rid="aff1">
          <name><surname>Krause</surname><given-names>Andreas</given-names></name>
          <email>andreas.krause@kit.edu</email>
        <ext-link>https://orcid.org/0000-0003-3345-2989</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1 aff2">
          <name><surname>Pugh</surname><given-names>Thomas A. M.</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Bayer</surname><given-names>Anita D.</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff3">
          <name><surname>Lindeskog</surname><given-names>Mats</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Arneth</surname><given-names>Almut</given-names></name>
          
        </contrib>
        <aff id="aff1"><label>1</label><institution>Karlsruhe Institute of Technology, Institute of Meteorology and Climate Research – Atmospheric Environmental Research (IMK-IFU), Kreuzeckbahnstr. 19, 82467 Garmisch-Partenkirchen, Germany</institution>
        </aff>
        <aff id="aff2"><label>2</label><institution>School of Geography, Earth &amp; Environmental Science and Birmingham Institute of Forest Research, University of Birmingham, Birmingham, B15 2TT, UK</institution>
        </aff>
        <aff id="aff3"><label>3</label><institution>Department of Physical Geography and Ecosystem Science, Lund University, 22362 Lund, Sweden</institution>
        </aff>
      </contrib-group>
      <author-notes><corresp id="corr1">Andreas Krause (andreas.krause@kit.edu)</corresp></author-notes><pub-date><day>15</day><month>September</month><year>2016</year></pub-date>
      
      <volume>7</volume>
      <issue>3</issue>
      <fpage>745</fpage><lpage>766</lpage>
      <history>
        <date date-type="received"><day>16</day><month>March</month><year>2016</year></date>
           <date date-type="rev-request"><day>14</day><month>April</month><year>2016</year></date>
           <date date-type="rev-recd"><day>26</day><month>August</month><year>2016</year></date>
           <date date-type="accepted"><day>29</day><month>August</month><year>2016</year></date>
      </history>
      <permissions>
<license license-type="open-access">
<license-p>This work is licensed under a Creative Commons Attribution 3.0 Unported License. To view a copy of this license, visit <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/3.0/">http://creativecommons.org/licenses/by/3.0/</ext-link></license-p>
</license>
</permissions><self-uri xlink:href="https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016.html">This article is available from https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016.html</self-uri>
<self-uri xlink:href="https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016.pdf">The full text article is available as a PDF file from https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016.pdf</self-uri>


      <abstract>
    <p>Land-use changes have been shown to have large effects on climate and
biogeochemical cycles, but so far most studies have focused on the effects of
conversion of natural vegetation to croplands and pastures. By contrast, relatively little is known about the long-term influence of past agriculture on vegetation regrowth and carbon sequestration following land abandonment. We used the LPJ-GUESS dynamic vegetation
model to study the legacy effects of different land-use histories (in terms
of type and duration) across a range of ecosystems. To this end, we performed
six idealized simulations for Europe and Africa in which we made a transition
from natural vegetation to either pasture or cropland, followed by a
transition back to natural vegetation after 20, 60 or 100 years. The
simulations identified substantial differences in recovery trajectories of
four key variables (vegetation composition, vegetation carbon, soil carbon,
net biome productivity) after agricultural cessation. Vegetation carbon and
composition typically recovered faster than soil carbon in subtropical,
temperate and boreal regions, and vice versa in the tropics. While the
effects of different land-use histories on recovery periods of soil carbon
stocks often differed by centuries across our simulations, differences in
recovery times across simulations were typically small for net biome
productivity (a few decades) and modest for vegetation carbon and composition
(several decades). Spatially, we found the greatest sensitivity of recovery
times to prior land use in boreal forests and subtropical grasslands, where
post-agricultural productivity was strongly affected by prior land
management. Our results suggest that land-use history is a relevant factor
affecting ecosystems long after agricultural cessation, and it should be
considered not only when assessing historical or future changes in
simulations of the terrestrial carbon cycle but also when establishing
long-term monitoring networks and interpreting data derived therefrom,
including analysis of a broad range of ecosystem properties or local climate
effects related to land cover changes.</p>
  </abstract>
    </article-meta>
  </front>
<body>
      

<sec id="Ch1.S1" sec-type="intro">
  <title>Introduction</title>
      <p>Historically, many natural forests or grasslands on Earth have been cleared
or cultivated for grazing, timber, food production, mining or settlements.
However, land-use change (LUC) in these areas has rarely been continuous,
and land cover and management have often changed for a variety of reasons
(Burgi and Turner, 2002). Based on the HYDE dataset, Campbell et al. (2008)
estimated that 269 Mha of cropland and 479 Mha of pasture
have been abandoned between 1700 and 2000. Recently, agricultural cessation
rates have risen globally, especially in the temperate region. For example,
during the last decades, large areas in Europe previously used for pasture
or crop cultivation have been abandoned (e.g., Schierhorn et al., 2013; Smith
et al., 2005). Following agricultural abandonment, and in the absence of
further anthropogenic influence, natural vegetation recolonizes in a typical
succession from herbaceous vegetation to shrubland and forests, if
environmental conditions are suitable for tree growth. These secondary
forests act as an important carbon (C) sink during the years of regrowth,
thereby reducing the growth rate of global atmospheric CO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula>
concentration (Pan et al., 2011).</p>
      <p>The immediate effects of land-use (LU) practices on C fluxes and nutrient
cycles have been studied in some detail over recent decades. Generally,
agriculture significantly reduces C and, in the absence of supplementary
sources, nitrogen (N) pools due to initial deforestation, reduced soil
litter input, and accelerated soil decomposition and erosion (Davidson
and Ackerman, 1993; Fujisaki et al., 2015; Guo and Gifford, 2002;
McLauchlan, 2006; Murty et al., 2002). Pasture soils can be an exception as
they have been found to accumulate C, depending on location and management
(McSherry and Ritchie, 2013; Milchunas and Lauenroth, 1993). The
long-term importance of past LU on ecosystems, however, was recognized only
recently, and much less effort has been put so far into the investigation of
legacy effects of LU history on ecosystem processes, how long these effects
persist, or whether they may even be irreversible (Chazdon, 2014; Compton and
Boone, 2000; Cramer et al., 2008; Hobbs et al., 2009; McLauchlan, 2006).
This is important not only for understanding present-day ecological systems but
also because, due to demographical, social, technological, economic and
environmental changes, LUC and land abandonment will continue to occur in
the future (Hurtt et al., 2011).</p>
      <p>Most observational studies that looked at the recovery of ecosystems after
agricultural cessation focused on the first years of succession. Analyses of
the long-term effects of historical LU are often limited by the availability
of adequate LU information and the absence of undisturbed ecosystems, and
usually rely on chronosequences (Chazdon, 2003; Knops and Tilman, 2000).
Only a few long-term observational study plots like the one maintained at
the Rothamsted Experimental Station (e.g., Poulton et
al., 2003) exist. Differences between (near) pristine and post-agricultural
forests or grasslands have been reported to persist for decades or centuries
after agricultural abandonment for various variables, including soil pH
(Falkengren-Grerup et al., 2006); microbial communities (Fichtner et
al., 2014); soil C, N and phosphorus (Compton and Boone, 2000); and other
nutrients (Wall and Hytonen, 2005). Furthermore, aboveground (ag) biomass
(Wandelli and Fearnside, 2015), percentage vegetation cover (Lesschen
et al., 2008), biodiversity (Vellend, 2004), species composition (Aide
et al., 2000) and structure (Bellemare et al., 2002) remained affected
for years to decades, or even longer. These effects have consequences, not
only for the C sink capacity of the ecosystem but also for water and energy
exchange between the land and the atmosphere (Foley et al.,
2003), which also has important, albeit still highly uncertain, implications
for regional climate change (e.g., Arora and Montenegro, 2011; Brovkin et
al., 2013; de Noblet-Ducoudre et al., 2012). Some studies have detected an
influence of ancient agriculture on forest composition and diversity even
thousands of years later (Dambrine et al., 2007; Dupouey et al., 2002;
Willis et al., 2004). However, the persistence of legacy effects varies
considerably with former LU, geographical location, sampling methods and
examined variables, making recovery trajectories often hard to predict
(Cramer et al., 2008; Foster et al., 2003; Guariguata and Ostertag, 2001;
Norden et al., 2015; Post and Kwon, 2000; Suding et al., 2004).</p>
      <p>In this study, we performed idealized simulations with the LPJ-GUESS dynamic
global vegetation model (DGVM) to explore the importance of agricultural LU
history in terms of type and duration for the regeneration of ecosystems and
C stocks and fluxes under a range of environmental conditions. We converted
natural vegetation to either pasture or cropland, followed by a
re-transition to natural vegetation after time periods of 20, 60 and
100 years. While there are numerous variables suitable to measure recovery
(Chazdon, 2003; Martin et al., 2016), we analyzed recovery times for
vegetation composition (represented here by the dominant plant functional
type), vegetation C, soil C, and net biome productivity to evaluate the
longevity of the effects of LU history on the C cycle component of
ecosystems and to ascertain whether the system eventually recovers to its
pre-disturbance state.</p>
</sec>
<sec id="Ch1.S2">
  <title>Methods</title>
<sec id="Ch1.S2.SS1">
  <title>LPJ-GUESS</title>
      <p>LPJ-GUESS is a process-based DGVM that is driven by climate, atmospheric
CO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula> concentration and N input (Smith et al., 2014). Plants are
attributed to one of 11 plant functional types (PFTs, nine groups of tree
species and two grasses) which are distinguished, for instance, in terms of
their climate preferences for establishment and survival, photosynthetic
pathways, growth rates, and growth strategies (see Table A1 for PFT acronyms
and names). Vegetation dynamics and composition at a given location result
from competition between plants for light and soil resources in a number of
independent replicate patches (50 in this study), averaged per
0.5<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> <inline-formula><mml:math display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 0.5<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> grid cell. Wildfire is included in the model
and, additionally, stochastic disturbances kill all the biomass in a patch,
representing, for example, storm or insect damages, with a typical return period of
100 years (Smith et al., 2014). Recent model updates include the
representation of LUC (Lindeskog et al., 2013) and the implementation of
the N cycle in natural vegetation and grasses (Smith et al., 2014). The
representation of the N cycle is crucial for this study because previous
agricultural N dynamics, such as extraction through harvest and input
through fertilization, can greatly affect ecosystems even after many decades
(Richter et al., 2000).</p>
      <p><?xmltex \hack{\newpage}?>Conversion of natural to managed land in LPJ-GUESS is characterized by the
initial killing of all living vegetation in the affected area. The
corresponding woody biomass is partly oxidized immediately (67–76 %) and
partly transferred to the product (21 %) or litter (3–12 %) pool.
Ten percent of the leaves are oxidized, while the rest of the leaves and the fine
roots enter the litter pool. Only the litter thus remains in the ecosystem
subsequent to land conversion. Pastures are represented by preventing tree
establishment and wildfires and by splitting the aboveground biomass of the
grasses equally between atmosphere (harvest) and litter at the end of each
year. Crops were represented by grass PFTs modified to mimic aspects of
cropland important for the C and N cycles. Settings for croplands and
pastures were as follows:
<list list-type="order"><list-item>
      <p>For transitions from natural vegetation to cropland, we transferred only
3 % of the cleared woody biomass to the litter instead of 12 % for
natural vegetation–pasture transitions. This accounts for the practice
that farmers would try to remove as many coarse roots as possible before
planting of crops.</p></list-item><list-item>
      <p>Harvest efficiency (in this study: fraction of aboveground biomass that is
oxidized) was 0.5 yr<inline-formula><mml:math 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> for pasture, representing the net effect of
grazing processes (Lindeskog et al., 2013). For crop simulations we
changed the harvest efficiency to 0.8 yr<inline-formula><mml:math 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>, representing simplified
crop harvest, as in Lindeskog et al. (2013).</p></list-item><list-item>
      <p>While we removed 100 % of harvested N biomass for croplands, we changed
this value to 65 % for pastures. That accounts for significant urine N
regain from animals fed on pastures (Dean et al., 1975; Lauenroth and Milchunas, 1992).</p></list-item><list-item>
      <p>Root turnover rate was 0.7 yr<inline-formula><mml:math 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> for pasture and was adapted to
1.0 yr<inline-formula><mml:math 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> for croplands to represent the annual plant types used in most croplands.</p></list-item><list-item>
      <p>In croplands we estimated tillage effects by increasing heterotrophic
respiration by a factor of 1.94 (Pugh et al., 2015).</p></list-item><list-item>
      <p>We simulated N fertilization in croplands by applying 75 kg ha<inline-formula><mml:math 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> yr<inline-formula><mml:math 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>
equally throughout the year to sustain crop productivity with time. This value
represents a compromise between higher values presently found in parts of Europe
and lower values in most of Africa (e.g., Potter et al., 2010).</p></list-item></list>
After patch-destroying disturbances or managed land converting back to
natural vegetation, there is a typical succession from grasses to
light-demanding pioneer trees, eventually followed in many ecosystems by the
establishment of shade-tolerant PFTs. It has been shown that LPJ-GUESS is
able to realistically simulate observed succession pathways and species
variations (Hickler et al., 2004; Smith et al., 2014).</p><?xmltex \hack{\newpage}?>
</sec>
<sec id="Ch1.S2.SS2">
  <title>Simulation setup</title>
      <p>During spin-up (500 years) and the simulation period (900 years), we forced
LPJ-GUESS with temperature-detrended, repeated 1981–2000 climate from the
University of East Anglia Climate Research Unit 3.21 dataset
(CRU, 2013), 1990s mean N deposition (Lamarque et al.,
2013) and a fixed atmospheric CO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula> mixing ratio of 356 ppmv. We ran the
model for Europe and Africa (33<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> E to 55<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> W), covering a
wide range of environmental conditions. These regions include all major
biomes (Smith et al., 2014). We chose Africa and Europe for the
simulation domain because the original LU version of the model was evaluated
against observations in Africa (Lindeskog et al., 2013) and to limit the
computational expense of the simulations. We did not intend to realistically
represent typical crop and pasture management across the domain (i.e., the
spatial variability in fertilizer use, multiple cropping systems, or
irrigation). For all simulations we used potential natural vegetation cover
to spin up the model, followed by a transition to either pasture or
croplands directly after spin-up and a transition back to natural vegetation
after time periods of 20, 60 and 100 years. This resulted in three pasture (P20,
P60, P100) and three cropland (C20, C60, C100) simulations.
Additionally, we performed a reference simulation in which natural
vegetation was retained throughout the whole simulation period.</p>
</sec>
<sec id="Ch1.S2.SS3">
  <title>Analyzed grid cells and biome classification</title>
      <p>To facilitate the interpretation, we classified each grid cell to one biome.
We used the same classification rules as Smith et al. (2014), aggregated
to eight biomes as in Bayer et al. (2015). Afterwards, we excluded grid cells
from the analyses which were classified as desert or tundra, had a mean net
primary productivity (NPP) below 0.1 kg C m<inline-formula><mml:math 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 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>, or were
located above 62.5<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> N, making the assumption that the relevance of
these low-production areas for agriculture is negligible.</p>
</sec>
<sec id="Ch1.S2.SS4">
  <title>Analyzed variables and definition of recovery</title>
      <p>We studied the influence of LU history on ecosystems by analyzing four key
variables: dominant PFT, vegetation C, soil C (excluding litter) and net
biome productivity (NBP). NBP is the net atmosphere–land carbon flux after
C losses associated with respiratory fluxes, fire, harvest, land clearing and
decomposition of LUC product pools are subtracted from gross primary
productivity. We investigated the legacy effects of LU history by
calculating a recovery time for each variable, simulation and grid cell
after the conversion back to natural vegetation. For vegetation C, soil C
and NBP, recovery time was defined as the year in which the 20-year running
mean of the variable exceeded the threshold of one standard deviation (<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula>)
below the mean of the reference simulation (full simulation
period) for the first time after agricultural abandonment. <inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula> was
calculated on the 20-year running mean of the reference simulation. To avoid
“false-positive” identifications of recovery in cases for which the variable
of interest was initially within 1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula> but then exhibited dynamics
taking it outside this range (e.g., soil C in Fig. A1), we applied an
additional criterion of whether the minimum after the transition to natural
vegetation occurred in the first 200 years and if it was below the mean
minus 1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula> threshold. If that was the case, the condition was
expanded so that the variable could only be defined as recovered after the
year in which the minimum occurred (“minimum rule”). A 200-year window was
chosen because the minimum occurred within the first 200 years for all biome
averages of all variables and simulations. If the minimum was located after
200 years, we assumed the minimum to be a result of natural variability and
recovery was achieved as soon as the variable in question exceeded the
threshold of 1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula> below the reference mean. Figure A1 shows an
example of how soil C recovery was calculated for one site.</p>
      <p>For the dominant PFT recovery, we first identified which PFT dominates each
grid cell in the reference simulation based on the annual maximum leaf area
index (LAI) amongst PFTs. We then checked for dominant PFT recovery in the same
way as we did for vegetation C, soil C and NBP (i.e., whether its LAI exceeded the
threshold of 1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula> below the reference simulation mean; condition 1)
but additionally checked whether its LAI was also larger than the LAI of any
other PFT in the same simulation and year (i.e., the dominant PFT is the same
as in the reference simulation, condition 2). Thus, dominant PFT recovery
was only possible if both conditions were fulfilled. For example, if the
temperate broadleaved evergreen (TeBS) tree was the dominant PFT in the
reference simulation (with an average maximum LAI of, for example, 3.0 and standard
deviation of <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula>0.2), dominant PFT recovery in a specific LU simulation
(e.g., P20) would occur once the LAI of TeBS in this simulation (a) hits the
threshold of 2.8 (3.0 <inline-formula><mml:math display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula> 0.2, condition 1) and (b) is larger than the LAI of
any other PFT in P20 in the specific year – i.e., TeBS is the dominant PFT in
the grid cell (condition 2). For all variables, the recovery time was capped
at 800 years after reconversion to natural vegetation, the point when
simulations ended. Recovery times of 800 years thus represent a lower limit.
However, the actual recovery time in these cases could theoretically lie
between 801 years and infinity.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F1" specific-use="star"><caption><p>Vegetation C (kg C m<inline-formula><mml:math 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>) for the reference simulation,
averaged over the whole simulation period of 900 years (upper left panel), soil C
(kg C m<inline-formula><mml:math 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>) (lower left panel), dominant PFT (upper right panel), and
corresponding biomes (lower right panel). Grid cells with a NPP below
0.1 kg C m<inline-formula><mml:math 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 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>, deserts and tundra, and latitudes above
62.5<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> N are masked in grey. PFT abbreviations are given in Table A1.</p></caption>
          <?xmltex \igopts{width=341.433071pt}?><graphic xlink:href="https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016-f01.png"/>

        </fig>

</sec>
</sec>
<sec id="Ch1.S3">
  <title>Results</title>
<sec id="Ch1.S3.SS1">
  <title>Reference simulation</title>
      <p>Maps of simulated vegetation and soil C, as well as dominant PFT and biomes
derived from PFT composition for the reference simulation, are shown in Fig. 1.
The salient features of biome and C storage distribution at the regional
scale are captured (Haxeltine and Prentice, 1996; Scharlemann et al.,
2014). Vegetation C reaches its highest values in tropical forests of
central Africa and decreases towards the deserts of southern and northern
Africa. Patterns are more homogeneous in Europe, where most areas store
5–10 kg C m<inline-formula><mml:math 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>. Similar to vegetation C, soil C in the (sub)tropics also
decreases with drier conditions; however, the differences are small, with
typical values of 5–10 kg C m<inline-formula><mml:math 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>. Soils in the temperate and southern
boreal ecosystems of Europe generally store more C (usually <inline-formula><mml:math display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 10 kg C m<inline-formula><mml:math 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>),
especially in colder environments. While Europe is mostly
dominated by woody PFTs (e.g., TeBS is the acronym for temperate broadleaved
summergreen tree), in Africa there is a shift from C<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">3</mml:mn></mml:msub></mml:math></inline-formula> and C<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">4</mml:mn></mml:msub></mml:math></inline-formula> grasses in the
dry regions to trees in the humid tropics. This gradient also appears in the
corresponding biome map: in Africa and the Arabian Peninsula, LPJ-GUESS
reproduces the transition from grasslands to savannas and tropical forests (TrFo)
as the Equator is approached. Europe is mostly classified as
temperate forests (TeFo), with some boreal forests (BoFo) in the north and
some shrublands/savannas in the south.</p>
</sec>
<sec id="Ch1.S3.SS2">
  <title>Dominant PFT recovery</title>
      <p>The LAI of the dominant PFT recovers on average within around one century
for all LU histories (Fig. 2). Maps of the recovery time (Fig. 3) show
distinct geographical patterns which occur in all simulations. Most
subtropical grasslands and savannas, and parts of the temperate and boreal
forests, recover within several decades, some grasslands even within 5
years. In contrast, recovery times are clearly longer (<inline-formula><mml:math display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 100 years)
in other parts of the temperate forests and in the tropical forests.
Long recovery is associated with woody successional vegetation dynamics, as
slow-recovering areas are usually dominated by temperate broadleaved
summergreen and tropical broadleaved evergreen forests (compare PFT
distribution in Fig. 1). These are shade-tolerant PFTs that establish only
slowly after disturbances. For 84 % of all analyzed grid cells, condition 1
(LAI recovery) was the delaying condition for dominant PFT recovery
(numbers exemplified for the P60 simulation), compared to only 3 % for
condition 2 (dominance recovery). For the remaining grid cells, both
conditions were fulfilled in the same year.</p>
      <p>Overall, differences across simulations of different LU histories are
moderate, with generally only small differences in temperate forests,
savannas and shrublands (Fig. 3; see also biome averages in Table 1 and the
histogram in Fig. A2). Areas of major differences are central Africa, where
P20 recovers faster than other simulations because post-agricultural net
mineralization rates are higher in this region for P20 than for the other
simulations (Fig. 4), thereby relatively increasing post-agricultural
N availability compared to the other simulations (Fig. 5), and the African
Mediterranean coast, where croplands recover much faster because the reduced
C : N ratio in the soil (not shown) enhances N mineralization and thus plant
N availability compared to pastures. Furthermore, in parts of the boreal zone
recovery takes several hundred years for C100 instead of a few decades for
the other simulations because lower available N levels relatively reduce the
growth of IBS (the dominant PFT in this region) compared to other woody
PFTs. Figure 6 shows the maximum differences between recovery times across
all simulations per biome (black dots), as well as across a subset of
simulations (colored squares and triangles). The differences were first
calculated for each grid cell and only then averaged over biomes, thereby
providing an estimate of the relative importance of former LU duration
versus former LU type on recovery times. While substantial differences occur
across the pasture simulations (P20, P60, P100) in tropical forests,
savannas and grasslands, and across cropland simulations (C20, C60, C100) in
boreal forests (emphasizing the importance of LU duration in these regions),
major differences between P100 and C100 occur in boreal forests and
grasslands (emphasizing the importance of LU type if agricultural duration
was long). On the other hand, in our simulations, dominant PFT recovery in
temperate forests is hardly influenced by the type of former LU or,
conversely, pasture duration has negligible effects on boreal forest
recovery. Interestingly, temperate forests recover faster for P100 and C100
then for P20 and C20. This pattern is generally restricted to areas where
the TeBS PFT dominates. We interpret this behavior as reduced soil N
favoring TeBS in the competition with other tree PFTs, thereby reaching its
background LAI levels earlier.</p>
</sec>
<sec id="Ch1.S3.SS3">
  <title>Vegetation C recovery</title>
      <p>Compared to dominant PFT, recovery occurs slightly later for vegetation C
(Fig. 2, Table 1). Spatial patterns look more homogeneous than for the dominant
PFT (Fig. 3). While most grasslands recover within a few decades for all
simulations, in particular so for post-cropland recovery, recovery occurs
only after several decades or centuries in forest ecosystems. Lower standard
deviations for the mean differences in vegetation C recovery times compared
to the standard deviations for the mean differences in dominant PFT recovery
times for most biomes (Fig. 6a and b) reflect the spatially more uniform
response of vegetation C recovery. Exceptions are tropical forests and
grasslands, where the standard deviation is higher for vegetation C recovery
compared to dominant PFT recovery.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F2" specific-use="star"><caption><p>Time series (20-year running mean) of dominant PFT, vegetation C,
soil C and NBP for the different experiments, starting from the time of
reconversion to natural vegetation and area-averaged over all grid cells.
Dominant PFT, vegetation C and soil C are shown in relative values compared
to reference simulation mean, while NBP is shown as absolute values
(kg C m<inline-formula><mml:math 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 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>)
because values cannot be presented relative to a zero background. The cyan-shaded
area corresponds to reference simulation mean <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula>. Note the
different scales on the <inline-formula><mml:math display="inline"><mml:mi>y</mml:mi></mml:math></inline-formula> axes.</p></caption>
          <?xmltex \igopts{width=341.433071pt}?><graphic xlink:href="https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016-f02.png"/>

        </fig>

      <?xmltex \floatpos{t}?><fig id="Ch1.F3" specific-use="star"><caption><p>Maps of recovery times in years for the dominant PFT, vegetation C,
soil C, and NBP for the P20, P100, C20, and C100 simulations.</p></caption>
          <?xmltex \igopts{width=341.433071pt}?><graphic xlink:href="https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016-f03.png"/>

        </fig>

      <?xmltex \floatpos{t}?><fig id="Ch1.F4"><caption><p>Average net N mineralization rates (kg N ha<inline-formula><mml:math 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> yr<inline-formula><mml:math 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>) in
the soil for the reference simulation (full simulation period) and during
the first 100 years of regrowth for the P20, P100, C20, and
C100 simulations.</p></caption>
          <?xmltex \igopts{width=236.157874pt}?><graphic xlink:href="https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016-f04.png"/>

        </fig>

      <p><?xmltex \hack{\newpage}?>Significant differences in recovery times occur between simulations of
different LU types that have the same duration, and between simulations of
the same LU type but with different duration. For example, in the grasslands
and savannas of southern, eastern and northern Africa, former croplands
recover much faster than former pastures (see also Table 1 and Fig. A2)
because post-agricultural N availability is enhanced in these regions
(Fig. 5). In former croplands in these environments, the combined effect of
fertilizing and harvest is a net N flux to the ecosystem (not shown) and
mineralization rates are enhanced after cropland abandonment (Fig. 4). This
net N flux can partially be explained by high levels of water stress in
these savannas and grasslands, resulting in greater C and N allocation to
roots relative to leaves and thereby decreased harvest removal in this
region (Fig. A3). Conversely, recovery in northern European forests is
delayed for C60 and, to an even greater extent, C100 because in this region
N removal by annual harvest exceeds N addition through fertilization during
the agricultural period (not shown) and post-agricultural N mineralization
rates in this region are substantially reduced compared to the other
simulations many decades or even a few centuries after abandonment (Fig. 4).
Differences in vegetation recovery times resulting from agricultural
duration are mostly found in temperate and boreal forests for the cropland
simulations (here longer durations result in longer recovery times due to
reduced N availability, Fig. 5) and in tropical forests and shrublands for
the pasture simulations, emphasizing the importance of agricultural duration
in these regions (see also Fig. 6b).</p>
</sec>
<sec id="Ch1.S3.SS4">
  <title>Soil C recovery</title>
      <p>Relative depletion of soil C content under crop and pasture LU is not as
large (loss of 0–11 % compared to the reference simulation) as for
vegetation C (Fig. 2). However, regeneration proceeds over longer timescales due to slower C accumulation in soils than in vegetation. C depletion
is generally more pronounced for former crops than for pastures due to the
greater harvest efficiency, which leads to more biomass removed each year,
and the effect of tillage enhancing soil respiration (Sect. 2.1). Upon
re-conversion, soil C accumulation is delayed for the pasture simulations
compared to the cropland simulations, especially for P20, where the residual
roots and other litter left after the original deforestation event continue
to decay and soil C decreases for some decades. The general delay for
pastures is associated with larger heterotrophic respiration rates (not
shown) compared to rates calculated in recovering croplands.</p>
      <p>Soil C recovery rates are highly latitude-dependent (Fig. 3), being much
slower in temperate (<inline-formula><mml:math display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 250 years) and boreal forests
(<inline-formula><mml:math display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 400 years) than in the tropics (<inline-formula><mml:math display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 100 years,
sometimes even within 5 years). Initial soil C depletions are larger in
higher latitudes, while these regions also suffer from low productivity,
thereby reducing C input to the soil upon regrowth. Additionally, in the
intensive LU simulations (P100, C60, C100), vegetation productivity in the
boreal region is further reduced compared to the reference simulation in the
first 200 years of regrowth (not shown) due to N limitation (Smith et
al., 2014), reducing litter input to the soil even further.</p>
      <p>Soil C recovery times differ substantially between simulations in many
areas. LU type is particularly important in grasslands and non-tropical
forests. While croplands tend to recover faster than pastures in grasslands
of southern and northern Africa, the opposite occurs in most temperate and
boreal forests but also the northern Sahel, where soil C after
re-conversion from croplands does not recover at all. Post-agricultural
N availability is enhanced in parts of the Sahel for the cropland simulations
due to increased N mineralization rates (Figs. 4 and 5), and trees
benefit more than grasses, leading to a shift in the equilibrium vegetation
state towards woody species (not shown), which results in an overall lower
soil C pool size. It should be noted that even though some regions do not
recover within 800 years, a large fraction of the original C loss is already
replenished after a few centuries, thereby limiting implications for the
C cycle. Counter to a priori expectations, for tropical and
temperate forests and for shrublands, the difference between P20 and C20 is
usually higher than between P60 and C60 or P100 and C100 (Fig. 6c). Pasture
duration is relevant for speed of soil C recovery in most ecosystems and,
apart from in the tropics, a longer duration usually delays recovery, mainly
due to substantial initial depletions after long pasture durations (Fig. 2).
For croplands, longer durations tend to delay recovery in temperate and
boreal forests but accelerate soil C recovery in the (sub)tropics. This is
somewhat unexpected for the tropical forest biome, where longer cropland
durations usually do not increase N availability upon abandonment in our
simulations (Fig. 5). However, while tropical soils lose large amounts of C
during the first decades of cropland use, slow C accumulation takes place
thereafter, resulting in higher soil C values at the end of the agricultural
period for C100 than for C20 in large parts of eastern Africa. This occurs
because tillage-driven C losses in more labile soil pools, which dominate
the system's response during the first decades, are eventually supplanted as
the dominant process by accumulation in more stable pools. This is different
to temperate and boreal forest, where soil C decreases throughout the entire
cropland period. Overall, the greatest sensitivity of soil C recovery times
to different LU histories is found in boreal forests and grasslands, where
maximum differences across simulations are often several centuries (Fig. 6c).
The maximum differences across all simulations
(P20/P60/P100/C20/C60/C100) in boreal forests are mainly due to differences
across simulations of same LU type but different duration (e.g., P20/P60/P100),
whereas the sensitivity of grasslands mainly reflects
differences across simulations of different LU type but same duration
(e.g., P100/C100), emphasizing the importance of duration and type of agriculture
in a range of biomes.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F5"><caption><p>Average N limitation on vegetation RuBisCO capacity (and thus on
gross primary production) for the reference simulation (full simulation period) and during the
first 100 years of regrowth for the P20, P100, C20, and
C100 simulations. N limitation is a number scaling from 0 (completely N-limited)
to 1 (no N limitation) (Smith et al., 2014).</p></caption>
          <?xmltex \igopts{width=236.157874pt}?><graphic xlink:href="https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016-f05.png"/>

        </fig>

      <?xmltex \floatpos{t}?><fig id="Ch1.F6" specific-use="star"><caption><p>Maximum difference in recovery time (longest recovery time minus
shortest recovery time of all selected simulations) for the dominant PFT,
vegetation C, soil C, and NBP. Black dots show maximum differences across
all six simulations (P20, P60, P100, C20, C60, C100), green squares
differences across 20-year pasture and cropland simulations (P20, C20), blue
squares differences across 60-year pasture and cropland simulations (P60, C60),
red squares differences across 100-year pasture and cropland simulations (P100,
C100), orange triangles differences across pasture simulations (P20, P60,
P100), and purple triangles differences across cropland simulations (C20, C60,
C100). Background colors indicate associated biomes, arrows one standard
deviation, and the dashed line 0 years' difference. Thus, the black dots show the
sensitivity of recovery times to LU history across all simulations for each
biome. The red, blue and green squares indicate the relative contribution of
LU type for a specific LU duration to this sensitivity, and the orange and
purple squares indicate the relative contributions of pasture and of cropland
duration. For example, if recovery times for one variable in one grid cell
were to be 50, 60, 65, 90, 100, 110 years (for P20, P60, P100, C20, C60,
C100), the maximum difference in recovery time across all simulations
(black) would be 60 years, across the 20-year simulations (green) 40 years,
across the 60-year simulations (blue) 40 years, across the 100-year
simulations (red) 45 years, across the pasture simulations (orange) 15 years
and across the cropland simulations (purple) 20 years.</p></caption>
          <?xmltex \igopts{width=398.338583pt}?><graphic xlink:href="https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016-f06.png"/>

        </fig>

</sec>
<sec id="Ch1.S3.SS5">
  <title>NBP recovery</title>
      <p>NBP switches from being a C source to the atmosphere during the period of
land management to a C sink after reconversion to natural vegetation (Fig. 2).
The sink capacity of the recovering ecosystem is greatest during the
first decades and then gradually returns to the NBP levels of the reference
simulation. P20 and, to a lesser extent, C20 act as a smaller sink than the
other simulations at least during the first 100 years of regrowth. Recovery
generally occurs slower in temperate and boreal regions than in the tropics
for all simulations (Fig. 3). Apart from boreal forests, standard deviations
of mean differences in recovery times are very small in all biomes compared
to the other variables (Fig. 6d). Recovery times are often somewhat lower
than those which would be expected from vegetation and soil C recovery
times. This is because the greater standard deviation of NBP in our
reference simulation (Fig. 2) reduces the threshold value in our recovery
definition, thereby making it easier to reach recovery levels for NBP. We
discuss the implications of this further in Sect. 4.2.</p>
      <p>Differences in NBP recovery times between simulations are relatively small
(typically a few years to decades; see Table 1). The largest differences
in recovery times are found in the boreal forests between the cropland
simulations, and, as for soil C, the differences are often greater between
P20 and C20 than between P100 and C100 (Fig. 6d).</p>

<?xmltex \floatpos{t}?><table-wrap id="Ch1.T1" specific-use="star"><caption><p>Average recovery times and standard deviations per biome and for each
simulation. Recovery times are depicted in Fig. 3.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="7">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="right"/>
     <oasis:colspec colnum="3" colname="col3" align="right"/>
     <oasis:colspec colnum="4" colname="col4" align="right"/>
     <oasis:colspec colnum="5" colname="col5" align="right"/>
     <oasis:colspec colnum="6" colname="col6" align="right"/>
     <oasis:colspec colnum="7" colname="col7" align="right"/>
     <oasis:thead>
       <oasis:row>  
         <oasis:entry colname="col1">Biome</oasis:entry>  
         <oasis:entry rowsep="1" namest="col2" nameend="col7" align="center">Simulation </oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">P20</oasis:entry>  
         <oasis:entry colname="col3">P60</oasis:entry>  
         <oasis:entry colname="col4">P100</oasis:entry>  
         <oasis:entry colname="col5">C20</oasis:entry>  
         <oasis:entry colname="col6">C60</oasis:entry>  
         <oasis:entry colname="col7">C100</oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry namest="col2" nameend="col7" align="center">Dominant PFT recovery time, averaged per biome </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Tropical forest</oasis:entry>  
         <oasis:entry colname="col2">90 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 55</oasis:entry>  
         <oasis:entry colname="col3">112 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 48</oasis:entry>  
         <oasis:entry colname="col4">121 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 50</oasis:entry>  
         <oasis:entry colname="col5">113 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 54</oasis:entry>  
         <oasis:entry colname="col6">125 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 52</oasis:entry>  
         <oasis:entry colname="col7">126 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 51</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Temperate forest</oasis:entry>  
         <oasis:entry colname="col2">102 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 74</oasis:entry>  
         <oasis:entry colname="col3">96 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 63</oasis:entry>  
         <oasis:entry colname="col4">93 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 57</oasis:entry>  
         <oasis:entry colname="col5">99 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 71</oasis:entry>  
         <oasis:entry colname="col6">89 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 61</oasis:entry>  
         <oasis:entry colname="col7">92 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 69</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Boreal forest</oasis:entry>  
         <oasis:entry colname="col2">47 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 89</oasis:entry>  
         <oasis:entry colname="col3">52 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 97</oasis:entry>  
         <oasis:entry colname="col4">53 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 90</oasis:entry>  
         <oasis:entry colname="col5">47 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 95</oasis:entry>  
         <oasis:entry colname="col6">60 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 111</oasis:entry>  
         <oasis:entry colname="col7">145 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 178</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Savanna</oasis:entry>  
         <oasis:entry colname="col2">47 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 71</oasis:entry>  
         <oasis:entry colname="col3">57 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 74</oasis:entry>  
         <oasis:entry colname="col4">62 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 77</oasis:entry>  
         <oasis:entry colname="col5">50 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 65</oasis:entry>  
         <oasis:entry colname="col6">57 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 73</oasis:entry>  
         <oasis:entry colname="col7">59 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 76</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Shrub</oasis:entry>  
         <oasis:entry colname="col2">95 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 93</oasis:entry>  
         <oasis:entry colname="col3">104 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 101</oasis:entry>  
         <oasis:entry colname="col4">108 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 100</oasis:entry>  
         <oasis:entry colname="col5">103 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 100</oasis:entry>  
         <oasis:entry colname="col6">109 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 112</oasis:entry>  
         <oasis:entry colname="col7">109 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 112</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Grassland</oasis:entry>  
         <oasis:entry colname="col2">76 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 108</oasis:entry>  
         <oasis:entry colname="col3">102 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 109</oasis:entry>  
         <oasis:entry colname="col4">115 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 109</oasis:entry>  
         <oasis:entry colname="col5">45 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 77</oasis:entry>  
         <oasis:entry colname="col6">55 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 97</oasis:entry>  
         <oasis:entry colname="col7">58 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 100</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1">Total</oasis:entry>  
         <oasis:entry colname="col2">80 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 85</oasis:entry>  
         <oasis:entry colname="col3">93 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 84</oasis:entry>  
         <oasis:entry colname="col4">99 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 84</oasis:entry>  
         <oasis:entry colname="col5">77 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 78</oasis:entry>  
         <oasis:entry colname="col6">83 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 85</oasis:entry>  
         <oasis:entry colname="col7">90 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 95</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry namest="col2" nameend="col7" align="center">Vegetation C recovery time, averaged per biome </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Tropical forest</oasis:entry>  
         <oasis:entry colname="col2">106 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 50</oasis:entry>  
         <oasis:entry colname="col3">137 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 61</oasis:entry>  
         <oasis:entry colname="col4">150 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 65</oasis:entry>  
         <oasis:entry colname="col5">121 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 65</oasis:entry>  
         <oasis:entry colname="col6">138 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 73</oasis:entry>  
         <oasis:entry colname="col7">139 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 74</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Temperate forest</oasis:entry>  
         <oasis:entry colname="col2">84 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 24</oasis:entry>  
         <oasis:entry colname="col3">93 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 31</oasis:entry>  
         <oasis:entry colname="col4">108 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 46</oasis:entry>  
         <oasis:entry colname="col5">91 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 29</oasis:entry>  
         <oasis:entry colname="col6">124 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 59</oasis:entry>  
         <oasis:entry colname="col7">149 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 79</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Boreal forest</oasis:entry>  
         <oasis:entry colname="col2">102 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 47</oasis:entry>  
         <oasis:entry colname="col3">113 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 57</oasis:entry>  
         <oasis:entry colname="col4">127 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 71</oasis:entry>  
         <oasis:entry colname="col5">111 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 55</oasis:entry>  
         <oasis:entry colname="col6">144 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 79</oasis:entry>  
         <oasis:entry colname="col7">187 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 107</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Savanna</oasis:entry>  
         <oasis:entry colname="col2">49 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 37</oasis:entry>  
         <oasis:entry colname="col3">61 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 44</oasis:entry>  
         <oasis:entry colname="col4">66 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 46</oasis:entry>  
         <oasis:entry colname="col5">35 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 40</oasis:entry>  
         <oasis:entry colname="col6">42 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 43</oasis:entry>  
         <oasis:entry colname="col7">43 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 44</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Shrub</oasis:entry>  
         <oasis:entry colname="col2">73 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 40</oasis:entry>  
         <oasis:entry colname="col3">86 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 48</oasis:entry>  
         <oasis:entry colname="col4">96 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 51</oasis:entry>  
         <oasis:entry colname="col5">60 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 38</oasis:entry>  
         <oasis:entry colname="col6">69 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 48</oasis:entry>  
         <oasis:entry colname="col7">73 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 54</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Grassland</oasis:entry>  
         <oasis:entry colname="col2">96 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 136</oasis:entry>  
         <oasis:entry colname="col3">119 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 140</oasis:entry>  
         <oasis:entry colname="col4">126 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 138</oasis:entry>  
         <oasis:entry colname="col5">40 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 98</oasis:entry>  
         <oasis:entry colname="col6">43 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 102</oasis:entry>  
         <oasis:entry colname="col7">45 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 105</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1">Total</oasis:entry>  
         <oasis:entry colname="col2">88 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 80</oasis:entry>  
         <oasis:entry colname="col3">106 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 87</oasis:entry>  
         <oasis:entry colname="col4">117 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 90</oasis:entry>  
         <oasis:entry colname="col5">75 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 74</oasis:entry>  
         <oasis:entry colname="col6">92 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 87</oasis:entry>  
         <oasis:entry colname="col7">101 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 98</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry namest="col2" nameend="col7" align="center">Soil C recovery time, averaged per biome </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Tropical forest</oasis:entry>  
         <oasis:entry colname="col2">74 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 60</oasis:entry>  
         <oasis:entry colname="col3">69 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 43</oasis:entry>  
         <oasis:entry colname="col4">66 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 45</oasis:entry>  
         <oasis:entry colname="col5">80 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 46</oasis:entry>  
         <oasis:entry colname="col6">64 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 46</oasis:entry>  
         <oasis:entry colname="col7">49 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 43</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Temperate forest</oasis:entry>  
         <oasis:entry colname="col2">207 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 98</oasis:entry>  
         <oasis:entry colname="col3">229 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 105</oasis:entry>  
         <oasis:entry colname="col4">241 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 117</oasis:entry>  
         <oasis:entry colname="col5">237 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 108</oasis:entry>  
         <oasis:entry colname="col6">261 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 133</oasis:entry>  
         <oasis:entry colname="col7">260 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 144</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Boreal forest</oasis:entry>  
         <oasis:entry colname="col2">327 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 107</oasis:entry>  
         <oasis:entry colname="col3">381 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 122</oasis:entry>  
         <oasis:entry colname="col4">421 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 140</oasis:entry>  
         <oasis:entry colname="col5">362 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 112</oasis:entry>  
         <oasis:entry colname="col6">425 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 132</oasis:entry>  
         <oasis:entry colname="col7">454 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 161</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Savanna</oasis:entry>  
         <oasis:entry colname="col2">84 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 132</oasis:entry>  
         <oasis:entry colname="col3">132 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 191</oasis:entry>  
         <oasis:entry colname="col4">162 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 233</oasis:entry>  
         <oasis:entry colname="col5">85 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 112</oasis:entry>  
         <oasis:entry colname="col6">83 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 125</oasis:entry>  
         <oasis:entry colname="col7">74 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 126</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Shrub</oasis:entry>  
         <oasis:entry colname="col2">107 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 140</oasis:entry>  
         <oasis:entry colname="col3">129 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 161</oasis:entry>  
         <oasis:entry colname="col4">135 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 168</oasis:entry>  
         <oasis:entry colname="col5">137 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 173</oasis:entry>  
         <oasis:entry colname="col6">139 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 183</oasis:entry>  
         <oasis:entry colname="col7">125 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 183</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Grassland</oasis:entry>  
         <oasis:entry colname="col2">286 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 234</oasis:entry>  
         <oasis:entry colname="col3">366 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 262</oasis:entry>  
         <oasis:entry colname="col4">422 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 283</oasis:entry>  
         <oasis:entry colname="col5">239 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 227</oasis:entry>  
         <oasis:entry colname="col6">219 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 229</oasis:entry>  
         <oasis:entry colname="col7">198 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 228</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1">Total</oasis:entry>  
         <oasis:entry colname="col2">182 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 176</oasis:entry>  
         <oasis:entry colname="col3">220 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 209</oasis:entry>  
         <oasis:entry colname="col4">245 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 236</oasis:entry>  
         <oasis:entry colname="col5">182 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 171</oasis:entry>  
         <oasis:entry colname="col6">183 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 186</oasis:entry>  
         <oasis:entry colname="col7">174 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 194</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry namest="col2" nameend="col7" align="center">NBP recovery time, averaged per biome </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Tropical forest</oasis:entry>  
         <oasis:entry colname="col2">57 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 37</oasis:entry>  
         <oasis:entry colname="col3">65 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 26</oasis:entry>  
         <oasis:entry colname="col4">71 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 27</oasis:entry>  
         <oasis:entry colname="col5">56 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 28</oasis:entry>  
         <oasis:entry colname="col6">64 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 24</oasis:entry>  
         <oasis:entry colname="col7">65 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 24</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Temperate forest</oasis:entry>  
         <oasis:entry colname="col2">97 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 29</oasis:entry>  
         <oasis:entry colname="col3">108 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 29</oasis:entry>  
         <oasis:entry colname="col4">113 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 31</oasis:entry>  
         <oasis:entry colname="col5">102 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 30</oasis:entry>  
         <oasis:entry colname="col6">112 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 31</oasis:entry>  
         <oasis:entry colname="col7">119 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 36</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Boreal forest</oasis:entry>  
         <oasis:entry colname="col2">136 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 55</oasis:entry>  
         <oasis:entry colname="col3">146 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 56</oasis:entry>  
         <oasis:entry colname="col4">152 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 58</oasis:entry>  
         <oasis:entry colname="col5">139 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 54</oasis:entry>  
         <oasis:entry colname="col6">151 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 59</oasis:entry>  
         <oasis:entry colname="col7">169 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 71</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Savanna</oasis:entry>  
         <oasis:entry colname="col2">31 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 40</oasis:entry>  
         <oasis:entry colname="col3">34 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 30</oasis:entry>  
         <oasis:entry colname="col4">36 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 26</oasis:entry>  
         <oasis:entry colname="col5">29 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 18</oasis:entry>  
         <oasis:entry colname="col6">32 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 17</oasis:entry>  
         <oasis:entry colname="col7">33 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 17</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Shrub</oasis:entry>  
         <oasis:entry colname="col2">51 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 37</oasis:entry>  
         <oasis:entry colname="col3">58 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 31</oasis:entry>  
         <oasis:entry colname="col4">59 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 29</oasis:entry>  
         <oasis:entry colname="col5">52 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 27</oasis:entry>  
         <oasis:entry colname="col6">58 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 26</oasis:entry>  
         <oasis:entry colname="col7">59 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 25</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Grassland</oasis:entry>  
         <oasis:entry colname="col2">25 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 37</oasis:entry>  
         <oasis:entry colname="col3">31 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 31</oasis:entry>  
         <oasis:entry colname="col4">35 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 30</oasis:entry>  
         <oasis:entry colname="col5">27 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 15</oasis:entry>  
         <oasis:entry colname="col6">34 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 20</oasis:entry>  
         <oasis:entry colname="col7">36 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 22</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Total</oasis:entry>  
         <oasis:entry colname="col2">59 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 51</oasis:entry>  
         <oasis:entry colname="col3">66 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 49</oasis:entry>  
         <oasis:entry colname="col4">71 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 49</oasis:entry>  
         <oasis:entry colname="col5">60 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 45</oasis:entry>  
         <oasis:entry colname="col6">68 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 47</oasis:entry>  
         <oasis:entry colname="col7">72 <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 52</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>

</sec>
</sec>
<sec id="Ch1.S4">
  <title>Discussion</title>
<sec id="Ch1.S4.SS1">
  <title>Comparison to observations and previous studies</title>
      <p>The effects of forest conversion to croplands or pastures are relatively
well studied. Tilled croplands typically show large depletions of soil C
compared to natural forest vegetation, but the picture for pasture is more
diverse (Davidson and Ackerman, 1993; Don et al., 2011; Guo and Gifford,
2002). Table 2 summarizes recent reviews about observed soil C changes in
agriculture compared to our results. LPJ-GUESS tends to simulate lower
C loss in croplands than commonly reported in observations. We attribute this
to a combination of the observation's focus on the top soil (while in
LPJ-GUESS soil C is implicitly averaged over the whole soil column) and our
relatively high fertilizer rates increasing productivity and thereby C input
to the soil. Pugh et al. (2015) studied the C dynamics of soils in
managed lands in LPJ-GUESS and found C accumulation even after 100 years of
grazed pasture at some locations, especially for low atmospheric CO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula>
concentrations. However, they used the C-only version of the model, thereby
neglecting C–N interactions and increased N limitation on grass growth with
time due to N removal by harvest. Croplands were explicitly represented by a
number of managed, but unfertilized, crop functional types in Pugh et al. (2015).
They found soil C reductions in Europe and Africa of
<inline-formula><mml:math display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 50 % after 100 years of cultivation, whereas in our study
C losses were much smaller (<inline-formula><mml:math display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 12 %), possibly partly due to
different tillage effects in the two soil models applied.</p>
      <p>In contrast to studies of LU effects compared to previously natural
ecosystems, the regeneration of ecosystems after agricultural abandonment
has been studied less, and a direct comparison to our simulations is
challenging, either because limited information about former LU or reference
conditions was provided in these studies or because there are important
differences from our setup in terms of management and LU duration or other
site-specific characteristics. Additionally, most of the available studies
were conducted in Amazonia or North America (Don et al., 2011) and there
is large variability in physical and biotic characteristics as well as in
land management (Kauffman et al., 2009). Many studies focus on the
recovery of biodiversity or species richness (Cramer et al., 2008;
Queiroz et al., 2014), but these variables cannot be adequately captured by
our large-scale PFT approach. It is often assumed that the ecosystem will
gradually return to its previous state and that intensive LU delays recovery
but the timescales are widely unknown and differ across variables and
regions, e.g., tropical species composition recovers much slower than forest
structure and soil nutrients (Chazdon, 2003). Different recovery
processes are strongly interlinked, e.g., vegetation accumulation and
turnover are key factors in the replenishment of soil quality and nutrients
which in turn determine plant productivity, and post-agricultural soil C and
N dynamics have been shown to correlate during the regeneration of
ecosystems (Knops and Tilman, 2000; Li et al., 2012).</p>
      <p>Table 2 includes several studies about ecosystem vegetation and soil
recovery after agricultural abandonment. Overall, the studies that looked at
vegetation recovery upon abandonment indicate that biomass accumulation
slows down after some decades and that accumulation rates correlate
negatively with agricultural duration. Our simulations show that the rate of
vegetation C sequestration indeed declines over time and that longer LU
durations delay recovery in each of the analyzed biomes. Observations also
indicate that use of land for pasture delays recovery in the tropics upon
pasture abandonment compared to cropping, but in our simulations this seems
to be the case only after long agricultural durations. For studies about
soil C dynamics after agricultural abandonment, interpretation is often
hindered by combining different soil layers or aggregating different LU
types (Li et al., 2012) and by large variations observed across studies
(Post and Kwon, 2000). Nevertheless, most of the observed
patterns are reproduced in our simulations, suggesting that LPJ-GUESS
captures the salient processes: after abandonment, croplands accumulate C
faster than pastures, and recovery often takes more than a century. The
impact of LU duration has rarely been studied; however, our results suggest that
even though longer agricultural durations mostly result in greater initial
soil C depletions, recovery can occur at similar or even faster speed in the
subtropics and tropics. In temperate and boreal forests long LU durations
tend to delay recovery.</p>

<?xmltex \floatpos{p}?><table-wrap id="Ch1.T2" specific-use="star"><caption><p>Observations and LPJ-GUESS results of soil C changes during
agriculture (cropland and/or pasture) and vegetation and soil C recovery
after abandonment.</p></caption><oasis:table frame="topbot"><?xmltex \begin{scaleboxenv}{.95}[.95]?><oasis:tgroup cols="6">
     <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:colspec colnum="6" colname="col6" align="left"/>
     <oasis:thead>
       <oasis:row>  
         <oasis:entry colname="col1">Observation</oasis:entry>  
         <oasis:entry colname="col2">Biome</oasis:entry>  
         <oasis:entry colname="col3">Observation value</oasis:entry>  
         <oasis:entry colname="col4">Closest</oasis:entry>  
         <oasis:entry colname="col5">Average model value</oasis:entry>  
         <oasis:entry colname="col6">Reference</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">type</oasis:entry>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4">simulations</oasis:entry>  
         <oasis:entry colname="col5">for the specific biome</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4">in terms of</oasis:entry>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4">LU history</oasis:entry>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row rowsep="1">  
         <oasis:entry namest="col1" nameend="col6" align="center">Soil C changes during agriculture </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Soil C change</oasis:entry>  
         <oasis:entry colname="col2">global</oasis:entry>  
         <oasis:entry colname="col3">42 % loss for forest–</oasis:entry>  
         <oasis:entry colname="col4">P20, P60,</oasis:entry>  
         <oasis:entry colname="col5">7–17 % loss in forest</oasis:entry>  
         <oasis:entry colname="col6">Guo and Gifford</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">averaged over</oasis:entry>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">cropland conversions,</oasis:entry>  
         <oasis:entry colname="col4">P100, C20,</oasis:entry>  
         <oasis:entry colname="col5">biomes for croplands,</oasis:entry>  
         <oasis:entry colname="col6">(2002)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">different depths</oasis:entry>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">8 % gain for forest–</oasis:entry>  
         <oasis:entry colname="col4">C60, C100</oasis:entry>  
         <oasis:entry colname="col5">2 % gain to 7 % loss</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">pasture conversions</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">for pastures</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Soil C change at</oasis:entry>  
         <oasis:entry colname="col2">tropical</oasis:entry>  
         <oasis:entry colname="col3">25 % loss for cropland,</oasis:entry>  
         <oasis:entry colname="col4">C20, C60,</oasis:entry>  
         <oasis:entry colname="col5">11–12 % loss for</oasis:entry>  
         <oasis:entry colname="col6">Don et al. (2011)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">36 cm</oasis:entry>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3">12 % loss for pasture/</oasis:entry>  
         <oasis:entry colname="col4">P20, P60</oasis:entry>  
         <oasis:entry colname="col5">croplands, 2 % gain to</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">grassland</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">4 % loss for pastures</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Soil C change at</oasis:entry>  
         <oasis:entry colname="col2">temperate</oasis:entry>  
         <oasis:entry colname="col3">new equilibrium after</oasis:entry>  
         <oasis:entry colname="col4">C100</oasis:entry>  
         <oasis:entry colname="col5">C loss throughout the</oasis:entry>  
         <oasis:entry colname="col6">Poeplau et al. (2011)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">29 cm</oasis:entry>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3">23 years</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">entire cropland</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">duration</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry namest="col1" nameend="col6" align="center">Vegetation recovery after agricultural abandonment </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">ag<inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>*</mml:mo></mml:msup></mml:math></inline-formula> vegetation</oasis:entry>  
         <oasis:entry colname="col2">tropical</oasis:entry>  
         <oasis:entry colname="col3">189 years</oasis:entry>  
         <oasis:entry colname="col4">C20</oasis:entry>  
         <oasis:entry colname="col5">121 years</oasis:entry>  
         <oasis:entry colname="col6">Saldarriaga et al.</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1">recovery time</oasis:entry>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6">(1988)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">ag vegetation</oasis:entry>  
         <oasis:entry colname="col2">tropical</oasis:entry>  
         <oasis:entry colname="col3">slowdown with time,</oasis:entry>  
         <oasis:entry colname="col4">P20, P60,</oasis:entry>  
         <oasis:entry colname="col5">(slight) slowdown,</oasis:entry>  
         <oasis:entry colname="col6">Silver et al. (2000)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">recovery rate</oasis:entry>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3">recovery slower for</oasis:entry>  
         <oasis:entry colname="col4">P100, C20,</oasis:entry>  
         <oasis:entry colname="col5">pasture recovery</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">pasture than for</oasis:entry>  
         <oasis:entry colname="col4">C60, C100</oasis:entry>  
         <oasis:entry colname="col5">slower only for long</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">cropland</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">durations (P100/C100)</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Total and</oasis:entry>  
         <oasis:entry colname="col2">temperate</oasis:entry>  
         <oasis:entry colname="col3">linear with time</oasis:entry>  
         <oasis:entry colname="col4">P60, P100</oasis:entry>  
         <oasis:entry colname="col5">(slight) slowdown</oasis:entry>  
         <oasis:entry colname="col6">Hooker and Compton</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">vegetation C</oasis:entry>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6">(2003)</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1">recovery rate</oasis:entry>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Vegetation</oasis:entry>  
         <oasis:entry colname="col2">temperate</oasis:entry>  
         <oasis:entry colname="col3">linear with time</oasis:entry>  
         <oasis:entry colname="col4">C20, C60</oasis:entry>  
         <oasis:entry colname="col5">(slight) slowdown</oasis:entry>  
         <oasis:entry colname="col6">Poulton et al. (2003)</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1">recovery rate</oasis:entry>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">ag vegetation</oasis:entry>  
         <oasis:entry colname="col2">tropical</oasis:entry>  
         <oasis:entry colname="col3">recovery speed</oasis:entry>  
         <oasis:entry colname="col4">P20, P60,</oasis:entry>  
         <oasis:entry colname="col5">recovery speed</oasis:entry>  
         <oasis:entry colname="col6">Uhl et al. (1988)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">recovery rate</oasis:entry>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3">inversely related</oasis:entry>  
         <oasis:entry colname="col4">P100</oasis:entry>  
         <oasis:entry colname="col5">inversely related</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">to LU duration</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">to LU duration</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">ag vegetation</oasis:entry>  
         <oasis:entry colname="col2">tropical</oasis:entry>  
         <oasis:entry colname="col3">73 years, recovery</oasis:entry>  
         <oasis:entry colname="col4">C20, C60,</oasis:entry>  
         <oasis:entry colname="col5">121–139 years,</oasis:entry>  
         <oasis:entry colname="col6">Hughes et al. (1999)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">recovery rate</oasis:entry>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3">speed inversely related</oasis:entry>  
         <oasis:entry colname="col4">C100</oasis:entry>  
         <oasis:entry colname="col5">recovery speed</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">and time</oasis:entry>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">to LU duration</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">inversely related</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">to LU duration</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Maximum tree</oasis:entry>  
         <oasis:entry colname="col2">tropical</oasis:entry>  
         <oasis:entry colname="col3">recovery speed</oasis:entry>  
         <oasis:entry colname="col4">C20, C60,</oasis:entry>  
         <oasis:entry colname="col5">recovery speed</oasis:entry>  
         <oasis:entry colname="col6">Randriamalala et al.</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">height recovery</oasis:entry>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3">inversely related</oasis:entry>  
         <oasis:entry colname="col4">C100</oasis:entry>  
         <oasis:entry colname="col5">inversely related</oasis:entry>  
         <oasis:entry colname="col6">(2012)</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1">rate</oasis:entry>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">to LU duration</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">to LU duration</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Vegetation</oasis:entry>  
         <oasis:entry colname="col2">tropical</oasis:entry>  
         <oasis:entry colname="col3">slower for pasture</oasis:entry>  
         <oasis:entry colname="col4">P20, P60,</oasis:entry>  
         <oasis:entry colname="col5">slower only for long</oasis:entry>  
         <oasis:entry colname="col6">Moran et al. (2000)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">height recovery</oasis:entry>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3">than for cropland</oasis:entry>  
         <oasis:entry colname="col4">P100, C20,</oasis:entry>  
         <oasis:entry colname="col5">durations (C100/P100)</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1">rate</oasis:entry>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4">C60, C100</oasis:entry>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">ag vegetation</oasis:entry>  
         <oasis:entry colname="col2">tropical</oasis:entry>  
         <oasis:entry colname="col3">slower for pasture</oasis:entry>  
         <oasis:entry colname="col4">P20, C20</oasis:entry>  
         <oasis:entry colname="col5">faster for P20 than for</oasis:entry>  
         <oasis:entry colname="col6">Wandelli and</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1">recovery rate</oasis:entry>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3">than for cropland</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">C20</oasis:entry>  
         <oasis:entry colname="col6">Fearnside (2015)</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry namest="col1" nameend="col6" align="center">Soil C recovery after agricultural abandonment </oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Soil C recovery</oasis:entry>  
         <oasis:entry colname="col2">global</oasis:entry>  
         <oasis:entry colname="col3">large variation across</oasis:entry>  
         <oasis:entry colname="col4">P20, P60,</oasis:entry>  
         <oasis:entry colname="col5">tendency to lose C in</oasis:entry>  
         <oasis:entry colname="col6">Paul et al. (2002)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">at up to 30 cm</oasis:entry>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">studies, tendency to</oasis:entry>  
         <oasis:entry colname="col4">P100, C20,</oasis:entry>  
         <oasis:entry colname="col5">the first years for</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">lose C in the first years</oasis:entry>  
         <oasis:entry colname="col4">C60, C100</oasis:entry>  
         <oasis:entry colname="col5">pastures, immediate</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">for pastures, immediate</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">accumulation for</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">accumulation for</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">croplands</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">croplands</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup><?xmltex \end{scaleboxenv}?></oasis:table><?xmltex \begin{scaleboxenv}{.95}[.95]?><table-wrap-foot><p><inline-formula><mml:math display="inline"><mml:msup><mml:mi/><mml:mo>*</mml:mo></mml:msup></mml:math></inline-formula> ag <inline-formula><mml:math display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> aboveground.</p></table-wrap-foot><?xmltex \end{scaleboxenv}?></table-wrap>

<?xmltex \floatpos{t}?><table-wrap id="Ch1.T3" specific-use="star"><caption><p>Observations and LPJ-GUESS results of soil C changes during
agriculture (cropland and/or pasture) and vegetation and soil C recovery
after abandonment.</p></caption><oasis:table frame="topbot"><?xmltex \begin{scaleboxenv}{.95}[.95]?><oasis:tgroup cols="6">
     <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:colspec colnum="6" colname="col6" align="left"/>
     <oasis:thead>
       <oasis:row>  
         <oasis:entry colname="col1">Observation</oasis:entry>  
         <oasis:entry colname="col2">Biome</oasis:entry>  
         <oasis:entry colname="col3">Observation value</oasis:entry>  
         <oasis:entry colname="col4">Closest</oasis:entry>  
         <oasis:entry colname="col5">Average model value</oasis:entry>  
         <oasis:entry colname="col6">Reference</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">type</oasis:entry>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4">simulations</oasis:entry>  
         <oasis:entry colname="col5">for the specific biome</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4">in terms of</oasis:entry>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4">LU history</oasis:entry>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>  
         <oasis:entry colname="col1">Soil C recovery</oasis:entry>  
         <oasis:entry colname="col2">global</oasis:entry>  
         <oasis:entry colname="col3">more accumulation for</oasis:entry>  
         <oasis:entry colname="col4">P20, P60,</oasis:entry>  
         <oasis:entry colname="col5">more accumulation for</oasis:entry>  
         <oasis:entry colname="col6">Laganiere et al.</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">at 34 cm</oasis:entry>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">croplands than for</oasis:entry>  
         <oasis:entry colname="col4">P100, C20,</oasis:entry>  
         <oasis:entry colname="col5">croplands than for</oasis:entry>  
         <oasis:entry colname="col6">(2010)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">pastures, no</oasis:entry>  
         <oasis:entry colname="col4">C60, C100</oasis:entry>  
         <oasis:entry colname="col5">pastures, slower</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">accumulation in boreal</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">accumulation in boreal</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">zone</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">zone</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Soil C recovery</oasis:entry>  
         <oasis:entry colname="col2">temperate</oasis:entry>  
         <oasis:entry colname="col3">linear accumulation, no</oasis:entry>  
         <oasis:entry colname="col4">C20</oasis:entry>  
         <oasis:entry colname="col5">linear accumulation, no</oasis:entry>  
         <oasis:entry colname="col6">Poeplau et al. (2011)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">at 28/40 cm</oasis:entry>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3">equilibrium after</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">equilibrium after</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3">120 years</oasis:entry>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">120 years</oasis:entry>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Soil C recovery</oasis:entry>  
         <oasis:entry colname="col2">grassland</oasis:entry>  
         <oasis:entry colname="col3">158 years</oasis:entry>  
         <oasis:entry colname="col4">C100</oasis:entry>  
         <oasis:entry colname="col5">198 years</oasis:entry>  
         <oasis:entry colname="col6">Potter et al. (1999)</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1">time at 0–60 cm</oasis:entry>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Soil C recovery</oasis:entry>  
         <oasis:entry colname="col2">savanna/</oasis:entry>  
         <oasis:entry colname="col3">230 years</oasis:entry>  
         <oasis:entry colname="col4">C20</oasis:entry>  
         <oasis:entry colname="col5">85 (savanna)/237</oasis:entry>  
         <oasis:entry colname="col6">Knops and Tilman</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">time at 0–60 cm</oasis:entry>  
         <oasis:entry colname="col2">temperate</oasis:entry>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5">(temperate forest) years</oasis:entry>  
         <oasis:entry colname="col6">(2000)</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4"/>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Soil C recovery</oasis:entry>  
         <oasis:entry colname="col2">temperate</oasis:entry>  
         <oasis:entry colname="col3"><inline-formula><mml:math display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 100 years</oasis:entry>  
         <oasis:entry colname="col4">C20, C60,</oasis:entry>  
         <oasis:entry colname="col5">237–261 years</oasis:entry>  
         <oasis:entry colname="col6">Foote and Grogan</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">  
         <oasis:entry colname="col1">time 0–10 cm</oasis:entry>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4">C100</oasis:entry>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6">(2010)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">Soil C recovery</oasis:entry>  
         <oasis:entry colname="col2">tropical</oasis:entry>  
         <oasis:entry colname="col3">50–60 years</oasis:entry>  
         <oasis:entry colname="col4">P20, P60,</oasis:entry>  
         <oasis:entry colname="col5">49–80 years</oasis:entry>  
         <oasis:entry colname="col6">Silver et al. (2000)</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">time 0–25 cm</oasis:entry>  
         <oasis:entry colname="col2">forest</oasis:entry>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4">P100, C20,</oasis:entry>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1"/>  
         <oasis:entry colname="col2"/>  
         <oasis:entry colname="col3"/>  
         <oasis:entry colname="col4">C60, C100</oasis:entry>  
         <oasis:entry colname="col5"/>  
         <oasis:entry colname="col6"/>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup><?xmltex \end{scaleboxenv}?></oasis:table></table-wrap>

      <p>The LPJ-GUESS model has been successfully tested against a range of
observations and observation-based products, including vegetation
distribution and dynamics and soil C response to changes in vegetation
cover (Hickler et al., 2004; Miller et al., 2008; Pugh et al., 2015;
Smith et al., 2014). In our simulations, we used only two different
agricultural land cover types (intensive grazing and fertilized, tilled
crops). Our analysis would therefore not identify effects of, for instance,
clearing technique (e.g., burning compared to mechanical removal) or
different land management practices (e.g., repeated burning or irrigation)
within one land cover type. For example, recovery of species richness and
maximum tree height of secondary forests occurs faster under no tillage
compared to heavy tillage (Randriamalala et al., 2012).</p>
      <p>Our study is intended as an idealized experiment to highlight the importance
of LU history on ecosystem state and fluxes across biomes. Still, some
processes with the potential to affect post-agricultural ecosystem recovery, at least regionally, are not currently included in LPJ-GUESS. One aspect
is the phosphorus cycle, which is not implemented in LPJ-GUESS, even though
it can be significantly altered by LUC (MacDonald et al., 2012;
McLauchlan, 2006). Moreover, while C and N cycles interact in LPJ-GUESS (Smith
et al., 2014), the uniform annual fertilizer rate we applied in this study
might be realistic in some regions, such as parts of Europe, but exceeds
present-day fertilizer use in Africa (Potter et al., 2010). Seed
availability, remnant trees and resprouting from surviving roots are
important factors during initial stages of tree colonization following
agricultural cessation (Bellemare et al., 2002; Cramer et al., 2008).
While LPJ-GUESS does not account for these effects explicitly, seedling
establishment is limited by a suitable growth environment, such that effects
like re-sprouting or remnant trees as seed sources are mimicked. The model
has been shown to, for example, reproduce vegetation recolonization in northern
Europe during the Holocene well (Miller et al., 2008), as well as canopy
structural changes as a function of forest age (Smith et al., 2014). What
is more, by using a prescribed climate in our simulations, hydrological
biosphere–atmosphere interactions and feedbacks are not captured (Eltahir
and Bras, 1996; Giambelluca, 2002), which could alter regional climate in
response to land cover change, potentially affecting recovery rates,
especially in tropical regions. Biophysical effects are not restricted to
modifications of the water cycle but also include changes in surface albedo
and roughness length as a function of ecosystem structure and composition,
thereby affecting air mixing and heat transfer. While forests generally
absorb more sunlight than grasslands (e.g., Culf et al.,
1995), differences amongst tree species and age classes exist as well.
Substantial impacts related to realistic land-use have been found on
local-to-regional scales (Alkama and Cescatti, 2016; Peng et al., 2014).
Whether or not the locally observed changes translate to a significant
global radiative forcing is still debated as the direction of change differs
across regions in some climate models, which may cancel when integrated
globally (Pielke et al., 2011). Additionally, while we focus on C sequestration rates in our
analysis, there might be biogeochemical
implications beyond C. For instance, the emissions of biogenic volatile
organic compounds (BVOCs) to the atmosphere vary greatly amongst plant
species (Kesselmeier and Staudt, 1999). BVOCs affect atmospheric
composition and climate via ozone production, lengthening the lifetime of
atmospheric methane, and contributing to secondary organic aerosol formation
(Penuelas and Staudt, 2010; Wu et al., 2012). BVOC emission factors might
also be drastically influenced by wildfires (Ciccioli et al., 2014),
which in turn are driven by species composition and vegetation density.
Thus, different successional trajectories of ecosystem structure and
composition recovery have the potential to directly modify air quality and
climatic conditions under which regrowth occurs, potentially creating
positive or negative climate system feedbacks.</p>
</sec>
<sec id="Ch1.S4.SS2">
  <title>Implications of recovery definition</title>
      <p>The term recovery is subjective and, in the absence of a universal
definition amongst ecologists, several definitions currently exist. The
definition used in this study examines recovery from a C sequestration
perspective which does not capture situations, for example, where the system
approaches a new equilibrium (as soil C did in some regions in the cropland
simulations). In order to obtain a better understanding of the uncertainties
related to our definition we therefore explored four alternative plausible
recovery definitions.</p>
      <p>When applying a mean minus 2<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula> threshold (instead of a mean minus
1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula> threshold), recovery times are generally shorter, e.g., on average
75 instead of 106 years for vegetation C in P60, but the overall geographic
patterns are very consistent across both definitions (not shown). For all
variables and simulations, notable differences between both definitions
occur in regions with longest recovery times, especially for subtropical
soil C in the pasture simulations.</p>
      <p>Recovery based on percentage change (Fig. A4) results in more heterogeneous
patterns across variables when compared to our standard recovery definition.
Applying a threshold of 95 % of the mean, instead of a mean minus
1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula> threshold, produces slightly longer dominant PFT recovery times in
parts of the temperate and tropical forests, and shorter recovery times in
grasslands, especially for the pasture simulations. Vegetation C shows
similar patterns to the dominant PFT; however, the differences to our
standard definition are more pronounced. Soil C recovery times generally
decrease dramatically, especially outside the tropics. NBP recovery times
generally increase, particularly in forest ecosystems.</p>
      <p>By expanding our standard recovery definition by an upper threshold
(reference mean plus 1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula>), and with the “minimum rule” also
applied to the maximum (see Sect. 2.4), one can test whether some ecosystems
recover from higher rather than lower values than in the reference
simulation. Mostly grasslands are affected by this alternative definition
(Fig. A5). Dominant PFT recovery under this definition takes slightly longer
throughout the African grasslands for the pasture simulations, and
considerably longer in parts of northern and southern Africa for the
cropland simulations. Patterns are similar for vegetation C, but the increase
in vegetation C recovery times is often larger than the increase in dominant
PFT recovery times, especially for croplands. Soil C recovery is notably
longer in subtropical and eastern African grasslands. The recovery times of
NBP are hardly affected. However, we do not use an upper threshold in the
primary definition used in this study because in this case the ecosystem is
already operating at a level of service above that which the undisturbed
ecosystem would have provided and our aim here was to investigate recovery
from a depletion perspective.</p>
      <p>Finally, when using the mean <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula> definition and additionally
checking whether the variable is still in the mean <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula> range at
the end of the simulation period (not shown), many grid cells did not
recover even within the set maximum cut-off of 800 years. Elements of random
fluctuations due to natural variability arising from stochastic processes
and disturbances and responding C, N, and water dynamics made a clear
identification of recovery period difficult in that case. In particular for
soil C, no recovery is found for parts of eastern and subtropical Africa.
The system converges towards a new equilibrium state in these regions which
lies above reference values. NBP stays within background levels everywhere.</p>
      <p>Altogether, the alternative recovery definitions agree on the general
findings when applying our standard definition, especially in terms of
relative recovery rates. For all definitions, vegetation C and dominant PFT
recover faster in grasslands than in forest-dominated ecosystems, and soil C
recovery takes much longer in higher latitudes. However, some areas,
especially in the subtropics, “recover” from values higher than in the
reference simulation, and these cases are not captured by our standard
definition. Additionally, in the tropics, soil C accumulation sometimes does
not stop once background values are reached and soil C leaves the reference
range. When recovery is defined based on standard deviation, NBP recovery is
often quicker than recovery of the C pools. This inconsistency emphasizes
the importance of both recovery definition and selected variables when
studying the recovery of ecosystems (Jones and Schmitz, 2009).
This is particularly relevant for flux tower measurements, where an
underlying long-term trend caused by the recovery from previous, often
unquantified or unknown LU change, might be overlooked due to a large
interannual variability in net ecosystem exchange.</p>
</sec>
</sec>
<sec id="Ch1.S5" sec-type="conclusions">
  <title>Conclusions</title>
      <p>Most studies which have explored the effects of distant human activities on
present-day ecosystems were restricted by sampling difficulties, small
spatial scales, short time periods since abandonment, and little information
about background conditions or the specific LU history of the site. Here, we
use a model-based approach to study the legacy effects of agricultural LU
history (type and duration) on ecosystem regeneration and C sink capacity
after the cessation of agriculture in a range of biomes across Europe and
Africa. The model reproduces qualitatively the response found at study
locations, including distinct differences in recovery between different
variables of the terrestrial carbon cycle. Long-lasting legacy effects of
former agricultural intensity emerge as important for present-day ecosystem
functions. These findings have implications for various scientific
applications:
<list list-type="order"><list-item>
      <p>Long-term monitoring sites (e.g., FLUXNET) and Earth observation systems need
to collect and maintain detailed information about past and present land
cover and land management to adequately interpret their data.</p></list-item><list-item>
      <p>Assessments of trends in data from sites that seek to identify impacts of
climate change and/or increasing atmospheric CO<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:math></inline-formula> concentration need to
make sure that legacy effects of past LU are not confounding the observed trends.</p></list-item><list-item>
      <p>Simulation experiments need to move beyond deforestation but also
represent, in a more detailed manner, re-growth dynamics following agricultural
abandonment at the sub-grid level. At the moment a few DGVMs have started to
do so (Shevliakova et al., 2009; Stocker et al., 2014; Wilkenskjeld et
al., 2014) based on model products of tropical shifting cultivation
(Hurtt et al., 2011), but accounting for gross land cover changes is also
important in other regions like Europe (Fuchs et al., 2015). Failure to
consider LU history may lead to errors in the simulation of vegetation
properties, potentially resulting in biases in carbon sequestration or
energy balance calculations, with subsequent implications for simulations of
regional and global climate. Our study suggests that, for vegetation and soil C
studies, accounting for LUC over the last 100–150 years is sufficient in
the tropics, while more than 200 years
might be necessary in the temperate and boreal zone; studies restricted to vegetation should not have to
account for LUC more than 150 years ago in any major climatic zone.
<?xmltex \hack{\newpage}?></p></list-item><list-item>
      <p>Assessing the efficiency of climate mitigation through large-scale
reforestation or afforestation projects will require knowledge about the
type and duration of previous LU. Our simulations suggest that the potential
to rapidly sequester C in biomass and soil is greatest in tropical forests
following short periods of cropland, while boreal forests accumulate C
slowest, especially when previously used for pasture. Special attention
should be paid to monitoring changes in belowground C, as in most places
the accumulation of soil C is much more sensitive to LU history than
C accumulation in re-growing trees.</p></list-item><list-item>
      <p>In terms of soil C, our results suggest that some subtropical regions might
not recover at all on timescales relevant for humans. However, given the low
absolute amounts of C “missing” in these soils, implications for the global
C cycle are expected to be small.</p></list-item></list></p>
</sec>
<sec id="Ch1.S6">
  <title>Data availability</title>
      <p>Researchers interested in the LPJ-GUESS source code can contact the model
developers (<uri>http://iis4.nateko.lu.se/lpj-guess/contact.html</uri>). The CRU TS 3.21
climate data can be downloaded from <uri>http://browse.ceda.ac.uk/browse/badc/cru/data/cru_ts/cru_ts_3.21</uri>. The
LPJ-GUESS simulation data are stored at the IMK-IFU computing facilities and
can be obtained on request (andreas.krause@kit.edu).</p><?xmltex \hack{\clearpage}?>
</sec>

      
      </body>
    <back><app-group>

<app id="App1.Ch1.S1">
  <title>Additional tables and figures</title>

<?xmltex \floatpos{h!}?><table-wrap id="App1.Ch1.T1"><?xmltex \hack{\hsize\textwidth}?><caption><p>Plant functional types used in this study.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="2">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="left"/>
     <oasis:tbody>
       <oasis:row>  
         <oasis:entry colname="col1">BNE</oasis:entry>  
         <oasis:entry colname="col2">Boreal needleleaved evergreen tree</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">BINE</oasis:entry>  
         <oasis:entry colname="col2">Boreal shade-intolerant needleleaved evergreen tree</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">BNS</oasis:entry>  
         <oasis:entry colname="col2">Boreal needleleaved summergreen tree</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">TeBS</oasis:entry>  
         <oasis:entry colname="col2">Shade-tolerant temperate broadleaved summergreen tree</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">IBS</oasis:entry>  
         <oasis:entry colname="col2">Shade-intolerant broadleaved summergreen tree</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">TeBE</oasis:entry>  
         <oasis:entry colname="col2">Temperate broadleaved evergreen tree</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">TrBE</oasis:entry>  
         <oasis:entry colname="col2">Tropical broadleaved evergreen tree</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">TrIBE</oasis:entry>  
         <oasis:entry colname="col2">Tropical shade-intolerant broadleaved evergreen tree</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">TrBR</oasis:entry>  
         <oasis:entry colname="col2">Tropical broadleaved raingreen tree</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">C3G</oasis:entry>  
         <oasis:entry colname="col2">Cool C<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">3</mml:mn></mml:msub></mml:math></inline-formula> grass</oasis:entry>
       </oasis:row>
       <oasis:row>  
         <oasis:entry colname="col1">C4G</oasis:entry>  
         <oasis:entry colname="col2">Warm C<inline-formula><mml:math display="inline"><mml:msub><mml:mi/><mml:mn mathvariant="normal">4</mml:mn></mml:msub></mml:math></inline-formula> grass</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>

      <?xmltex \floatpos{h!}?><fig id="App1.Ch1.F1"><caption><p>Soil C for the six simulations after conversion to natural
vegetation at one single example site to illustrate how recovery time was
calculated according to our definition. The cyan-shaded area corresponds to
reference simulation mean <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula>. When soil C exceeds the mean <inline-formula><mml:math display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula> 1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula>
threshold and the time of the minimum (which in this case is
located in the first 200 years and below the mean <inline-formula><mml:math display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula> 1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula> threshold
for all six simulations) is passed, recovery is achieved.</p></caption>
        <?xmltex \hack{\hsize\textwidth}?>
        <?xmltex \igopts{width=236.157874pt}?><graphic xlink:href="https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016-f07.png"/>

      </fig>

<?xmltex \hack{\clearpage}?><?xmltex \floatpos{h!}?><fig id="App1.Ch1.F2"><caption><p>Histograms of recovery times for the dominant PFT, vegetation C, soil C,
and NBP for the six experiments. Colors indicate different biomes.</p></caption>
        <?xmltex \hack{\hsize\textwidth}?>
        <?xmltex \igopts{width=369.885827pt}?><graphic xlink:href="https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016-f08.png"/>

      </fig>

<?xmltex \hack{\clearpage}?><?xmltex \floatpos{h!}?><fig id="App1.Ch1.F3"><caption><p>Annual ratio of C removed by harvest and C stored in vegetation,
averaged over the whole agricultural period and for P60. As only
aboveground biomass is harvested, lower values indicate increased C
allocation to roots compared to leaves due to limited water supply.</p></caption>
        <?xmltex \hack{\hsize\textwidth}?>
        <?xmltex \igopts{width=170.716535pt}?><graphic xlink:href="https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016-f09.png"/>

      </fig>

      <?xmltex \floatpos{h!}?><fig id="App1.Ch1.F4"><caption><p>Maps of recovery time for the dominant PFT, vegetation C, soil C and
NBP with an alternative recovery definition for the P60 and C60
simulations. The definition is the same as our standard definition but with a
mean <inline-formula><mml:math display="inline"><mml:mo>⋅</mml:mo></mml:math></inline-formula> 0.95 threshold instead of mean <inline-formula><mml:math display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula> 1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula>.</p></caption>
        <?xmltex \hack{\hsize\textwidth}?>
        <?xmltex \igopts{width=497.923228pt}?><graphic xlink:href="https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016-f10.png"/>

      </fig>

<?xmltex \hack{\clearpage}?><?xmltex \floatpos{h!}?><fig id="App1.Ch1.F5"><caption><p>Maps of recovery time for the dominant PFT, vegetation C, soil C and
NBP with an alternative recovery definition for the P60 and C60
simulations. The definition is the same as our standard definition but with a
mean <inline-formula><mml:math display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula> threshold and the minimum check also applied to the
maximum instead of a mean <inline-formula><mml:math display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula> 1<inline-formula><mml:math display="inline"><mml:mi mathvariant="italic">σ</mml:mi></mml:math></inline-formula> threshold and only checking the minimum.</p></caption>
        <?xmltex \hack{\hsize\textwidth}?>
        <?xmltex \igopts{width=497.923228pt}?><graphic xlink:href="https://esd.copernicus.org/articles/7/745/2016/esd-7-745-2016-f11.png"/>

      </fig>

<?xmltex \hack{\clearpage}?>
</app>
  </app-group><ack><title>Acknowledgements</title><p>This work was funded by the Helmholtz Association through the International
Research Group CLUCIE and by the European Commission's 7th Framework
Programme, under grant agreement number 603542 (LUC4C). A. D. Bayer acknowledges
support by the European Commission's 7th Framework Programme, under grant
agreement number 308393 (OPERAs). This work was supported, in part, by the
German Federal Ministry of Education and Research (BMBF), through the
Helmholtz Association and its research program ATMO. It also represents paper
number 20 of the Birmingham Institute of Forest Research. <?xmltex \hack{\newline}?><?xmltex \hack{\newline}?>
The article processing charges for this open-access <?xmltex \hack{\newline}?> publication
were covered by a Research <?xmltex \hack{\newline}?> Centre of the Helmholtz Association. <?xmltex \hack{\newline}?><?xmltex \hack{\newline}?>
Edited by: V. Arora <?xmltex \hack{\newline}?>
two anonymous referees</p></ack><ref-list>
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<abstract-html><p class="p">Land-use changes have been shown to have large effects on climate and
biogeochemical cycles, but so far most studies have focused on the effects of
conversion of natural vegetation to croplands and pastures. By contrast, relatively little is known about the long-term influence of past agriculture on vegetation regrowth and carbon sequestration following land abandonment. We used the LPJ-GUESS dynamic vegetation
model to study the legacy effects of different land-use histories (in terms
of type and duration) across a range of ecosystems. To this end, we performed
six idealized simulations for Europe and Africa in which we made a transition
from natural vegetation to either pasture or cropland, followed by a
transition back to natural vegetation after 20, 60 or 100 years. The
simulations identified substantial differences in recovery trajectories of
four key variables (vegetation composition, vegetation carbon, soil carbon,
net biome productivity) after agricultural cessation. Vegetation carbon and
composition typically recovered faster than soil carbon in subtropical,
temperate and boreal regions, and vice versa in the tropics. While the
effects of different land-use histories on recovery periods of soil carbon
stocks often differed by centuries across our simulations, differences in
recovery times across simulations were typically small for net biome
productivity (a few decades) and modest for vegetation carbon and composition
(several decades). Spatially, we found the greatest sensitivity of recovery
times to prior land use in boreal forests and subtropical grasslands, where
post-agricultural productivity was strongly affected by prior land
management. Our results suggest that land-use history is a relevant factor
affecting ecosystems long after agricultural cessation, and it should be
considered not only when assessing historical or future changes in
simulations of the terrestrial carbon cycle but also when establishing
long-term monitoring networks and interpreting data derived therefrom,
including analysis of a broad range of ecosystem properties or local climate
effects related to land cover changes.</p></abstract-html>
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