Soil Biology & Biochemistry 37 (2005) 1898–1909

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Savanna-derived organic matter remaining in arable soils of the South

African Highveld long-term mixed cropping: Evidence
from 13C and 15N natural abundance
I. Lobea,1, R. Bolb,*, B. Ludwigc, C.C. Du Preezd, W. Amelunge
a
Institute of Soil Science and Soil Geography, University of Bayreuth, 95440 Bayreuth, Germany
b
Department of Soil; Environmental and Ecological Sciences, Institute of Grassland and Environmental Research,
North Wyke Research Station, Okehampton, Devon EX20 2SB, UK
c
Department of Environmental Chemistry, University of Kassel, 37213 Witzenhausen, Germany
d
Department of Soil, Crop and Climate Sciences, University of the Free State, P.O. Box 339, Bloemfontein 9300, South Africa
e
Institute of Soil Science and Soil Ecology, University of Bonn, 53115 Bonn, Germany
Received 9 April 2003; received in revised form 10 February 2005; accepted 10 February 2005

Abstract
Sustainable agriculture requires the formation of new humus from the crops. We utilized 13C and 15N signatures of soil organic matter to
assess how rapidly wheat/maize cropping contributed to the humus formation in coarse-textured savanna soils of the South African Highveld.
Composite samples were taken from the top 20 cm of soils (Plinthustalfs) cropped for lengths of time varying from 0 to 98 years, after
conversion from native grassland savanna (C4). We performed natural 13C and 15N abundance measurements on bulk and particle-size
fractions. The bulk soil d13C values steadily decreased from K14.6 in (C4 dominated) grassland to K16.5‰ after 90 years of arable
cropping. This d13C shift was attributable to increasing replacement of savanna-derived C by wheat crop (C3) C which dominated over maize
(C4) inputs. After calculating the annual C input from the crop yields and the output from literature data, by using a stepwise C replacement
model, we were able to correct the soil d13C data for the irregular maize inputs for a period of about one century. Within 90 years of cropping
41–89% of the remaining soil organic matter was crop-derived in the three studied agroecosystems. The surface soil C stocks after 90 years of
the wheat/maize crop rotation could accurately be described with the Rothamsted Carbon Model, but modelled C inputs to the soil were very
low. The coarse sand fraction reflected temporal fluctuations in 13C of the last C3 or C4 cropping and the silt fraction evidenced selective
erosion loss of old savanna-derived C. Bulk soil 15N did not change with increasing cropping length. Decreasing d15N values caused by
fertilizer N inputs with prolonged arable cropping were only detected for the coarse sand fraction. This indicated that the present N
fertilization was not retained in stable soil C pool. Clearly, conventional cropping practices on the South African highlands neither contribute
to the preservation of old savanna C and N, nor the effective humus reformation by the crops.
q 2005 Elsevier Ltd. All rights reserved.

Keywords: 13C and 15N; Duration of arable cropping; Particle-size fractionation; Alternating C3/C4 vegetation; SOM turnover; C model; Roth-C Model

1. Introduction can be reduced through appropriate management strategies,

Long-term arable cropping results in a decline of soil
organic matter (SOM) and soil inherent fertility. This decline
* Corresponding author. Tel.: C44 1837 883500; fax: C44 1837 82139.
E-mail address: roland.bol@bbsrc.ac.uk (R. Bol).
1 improve our understanding of the processes involved in this
Present address: UFZ, Centre for Environmental Research Leipzig–
Halle, Department of River Ecology, Brueckst. 3a, 39114 Mogdeburg,
Germany.
0038-0717/$ - see front matter q 2005 Elsevier Ltd. All rights reserved.
doi:10.1016/j.soilbio.2005.02.030
e.g. crop rotations, inorganic fertilizers (Campbell et al., 1991;
Paustian et al., 1997). In the South African Highveld, the C and
N concentrations of sandy soils had decreased to 35 and 45%
of the original values of the grassland after 90 years of
cropping (Lobe et al., 2001). Despite regular fertilizer
applications, yields also decreased in the long-term. To
reduction in soil C and N content and the associated reduction
in crop yield, we first quantified the rates of organic carbon and
total N losses from different particle-size pools (Lobe et al.,
2001). We hypothesised that soil inherent fertility was lost,
 I. Lobe et al. / Soil Biology & Biochemistry 37 (2005) 1898–1909
because silt-associated SOM was lost by wind-erosion.
However, we could not clarify whether or not these losses
selectively affected ‘old’ grassland-derived SOM. Further-
more, no differentiation could be made between crop- and
savanna-derived C, hence also real turnover rates remained
unknown. The application of the natural isotope tracer
technique can assist to elucidate these questions.
Changes in natural isotope ratios of organic matter in
arable soils have been used to determine the turnover of the
organic matter by the change of C3 to C4 vegetation or vice
versa in temperate and tropical climates (Balesdent et al.,
1987; Vitorello et al., 1989; Trouve et al., 1994; Veldkamp,
1994; Balesdent and Mariotti, 1996; Follett et al., 1997;
Liang et al., 1998). Less is known about the velocity of the
turnover of the organic matter in subtropical climates (e.g.
Boutton et al., 1998), as also exemplified by South African
savanna ecosystems.
The vegetation of the extensively grazed grassland in the
South African Highveld, which was used as a reference of
soils with 0 years of arable cropping, consisted predomi-
nantly of C4 grasses. These discriminate less against the
heavier 13C isotope during photosynthesis than the C3 plants
(Vogel, 1980; Boutton, 1996). Therefore, the C4 grasses are
enriched in 13C relative to the C3 plants, i.e. they show
higher d13C values than, e.g. wheat or sunflowers (C3)
which were predominantly grown on the arable land. When
‘old’ C4 organic material of the grassland is substituted by
material that is derived from C3 plants, the d13C values of
the soil decrease with increasing duration of cultivation. In
soils with an ideal and sustained change of cultivation from
C4 to C3 plant mono-culture it is possible to calculate the
turnover rates of the native C4 and newly incorporated C3
carbon present in the soil.
However, the investigated arable soils in South Africa,
with various length of cropping, were not cultivated in
mono-culture with wheat. The crops rotated between wheat
(C3), sunflower (C3), and maize (C4, Lobe et al., 2001). The
proportion of the ‘new’ (derived from the crops) or the ‘old’
C (derived from the native C4 grasses) of the soil organic
matter can therefore not be calculated directly, simply from
the change of the isotope ratio since the start of arable
cropping. However, the cropping history of most sites is
well known and therefore the C input proportion of C3
wheat and sunflower to C4 maize plants into the arable soils.
If we assume that maize, wheat, or sunflower residues are
Table 1
Climatic data and soil properties of the three agro-ecosystems at Harrismith, Kroonstad and Tweespruit, (Lobe et al., 2001)
Area Clay (%) Maximum of Mean annual
cultivation temperature
(years) (8C)
Harrismith 13–19 90 13.8
Kroonstad 10–15 98 16.6
Tweespruit 10–16 90 16.0
CECpot, potential cation exchange capacity.
1899
transformed in the soil at a similar rate, then we are able to
correct the isotope data for the maize input.
The isotope technique might also be used to trace the fate
of the inorganic fertilizer N. As soil N transformations
result, on average, in a discrimination of 15N, soil 15N values
are enriched in 15N relative to mineral N of the fertilizers
(Yoneyama, 1996), i.e. the long-term application of
inorganic fertilizers should result in a decrease of soil
d15N values. Yet, it is uncertain whether this process can
really be seen in the environment, because various N
transformations influence the soil d15N values in different
directions. The chronosequences provide a set of sites with
comparable conditions, reducing the various parameters
which alter the d15N values. As our samples were taken at
the same time within the dry season, it seemed reasonable to
assume that different water conditions in the soil may not
govern soil N transformations as indicated by the change in
the natural abundance of 15N.
It was the aim of this study, to use the 13C and 15N natural
abundance tracer technique as a tool: (i) to quantify the
turnover of savanna-derived and cropping-derived C in this
mixed-cropping ecosystem; (ii) to examine the origin of silt-
associated C resistant to erosion; and (iii) to elucidate the
fate of fertilizer N within soil-textural C and N pools. To
account for mixed C input from various C3 and C4 plants,
the conventional isotope-mixing model of Balesdent and
Mariotti (1996) was modified. We also aimed to verify if the
changes in soil C stocks during 90 years of the mixed C3
and C4 crop rotations could be described by the Rothamsted
Carbon Model.
2. Materials and methods
2.1. Sites
We sampled the soil at nine sites in each of the three
agro-ecosystems close to Harrismith, Kroonstad and
Tweespruit in the Free State Province of South Africa.
The characteristics of the sites are given in Lobe et al.
(2001) (see also Table 1). All three agro-ecosystems are
in the summer rainfall region (6 months from October
to March with O50 mm rainfall), with 516–625 mm
mean annual rainfall, and 13.8–16.6 8C mean annual
temperature. The nine individual sampling sites in each
agro-ecosystem comprised arable land, with varied
Mean annual pH CECpot
precipitation (mmolc kgK1)
H2O KCl
(mm)
625 4.6–5.7 3.8–4.6 63–130
563 5.2–6.8 4.0–5.5 42–84
516 5.4–6.3 4.2–5.2 49–120
1900 I. Lobe et al. / Soil Biology & Biochemistry 37 (2005) 1898–1909
lengths of time under cultivation, and adjacent to native
grassland, used as grazing for either cattle or sheep. The
stocking density on the grassland was on average 0.4
large stock units per hectare at Harrismith, 0.5 at
Kroonstad, and 0.6 at Tweespruit, and the grazing period
between 1 and 3 months total per year. All three agro-
ecosystems are in the Grassland Biome (Harrismith: Moist
Cold Highveld Grassland, Kroonstad: Dry Sandy Highveld
Grassland, Tweespruit: Moist Cool Highveld Grassland;
Bredenkamp et al., 1996). The botanical composition of the
native grassland is dominated by Cymbopogon plurinodis,
Themeda triandra, Setaria sphacelata, Elionurus muticus and
Eragrostis curvula in Harrismith, Eragrostis lehmanniana,
Eragrostis obtusa, Panicum coloratum, Stipagrostis uniplu-
mis and Pentzia globosa in Kroonstad, and T. triandra in
Tweespruit. The arable land had been ploughed to a depth of
20–30 cm. The 30 and 68-year-old plots in Harrismith were
ploughed to 40 cm. The fields were cultivated with a rotation
of wheat and maize (and occasionally sunflower). Inorganic
fertilizer had been applied annually (maize: 50–70 kg N, 10–
25 kg P, 0–10 kg K haK1; wheat: 10–40 kg N, 10–25 kg P,
0–15 kg K haK1; sunflower: 20–50 kg N, 10–20 kg P, 2–
6 kg K haK1). Average grain yields varied from 2.2 to
3.75 t haK1 yearK1 for maize, 1.2–2.75 t haK1 yearK1 for
wheat, and 1.0–1.25 t haK1 yearK1 for sunflower. Within a
rotation cycle, every 1–2 years, the soils are kept clear of
vegetation for up to 6 months in the dry season so that the soil
retains its stored water. This technique might enhance the
potential loss of soil organic matter because less organic
matter is added. Hence, the risks increase the enhanced erosion
during the period without protective plant cover to stabilize the
soil.
Surface samples (0–20 cm) were taken from five subsites
on each of the native grassland and arable plots. All samples
were air-dried and sieved to !2 mm. A particle-size
fractionation was carried out on material !2 mm with 2–
6 replicates depending on the silt content. As suggested by
Amelung and Zech (1999) for minimizing artefacts, 30 g of
soil was dispersed ultrasonically in 150 ml of water (using
60 J mlK1; Heat Systems, Model W 185 F). The ultrasonic
energy output was maintained according to Amelung and
Zech (1999). The coarse sand (O250 mm) was then isolated
by wet sieving, and the remaining suspension (300 ml) with
a soil-to-water ratio of 1:10 was treated ultrasonically (using
440 J mlK1) for additional dispersion. Thereafter the clay
fraction was recovered in the supernatant after centrifuging,
and the silt (2–20 mm) and fine sand (20–250 mm) fractions
were separated by wet sieving. The silt and sand fractions
were dried at 40 8C, whereas the clay fraction was freeze-
dried. Recovery of the material after the fractionation
averaged 100G0.88% (standard error) of initial sample
weight, 96.3G4.8% of organic C and 87.8G7.1% of
total N. We pooled corresponding size separates of the
uncultivated grassland subplots for each of the three agro-
ecosystems.
2.2. Chemical analyses
Total C and total N were determined by dry combustion
with a CHNS analyzer (Elementar Analysensysteme GmbH,
Hanau, Germany). There was no detectable inorganic C,
so that we can regard the total C as organic C. Measurements
of d13C and d15N values of the samples were obtained
by continuous flow-isotope ratio mass spectrometry
(CF-IRMS), using an automated nitrogen/carbon analysis-
mass spectrometer (ANCA-MS) system (Europa 20/20,
Crewe, UK). Ground wheat flour was used as the working
standard to refer the d13C values to the international Pee Dee
Belemnite (PDB). The d13C values were analyzed in bulk soil,
clay, silt, and coarse sand fractions. Due to the low
concentrations of C and N in the fine sand fraction, d13C and
d15N could not be determined in this fraction.
2.3. Calculations
For a pure C4/C3 vegetation change, the proportion of
C3-derived carbon F was calculated as proposed by
Balesdent and Mariotti (1996)
F Z ðdarable K dgrassland Þ=ðdC3 K dgrassland Þ (1)
where darable is the d13C value of the arable land, dgrassland
that of the grassland, and dC3 that of the C3 vegetation.
The decline in the d13C and d15N values and the C and N
stocks was described by a biexponential model, the changes
in crop yields by an exponential model (Lobe et al., 2001).
2.4. Stepwise C replacement model
The calculation of the real carbon replacement to correct
the d13C values for the maize (C4) inputs requires a specific
model that describes the decline of the C stocks and the d13C
values. At each year, the C contents in our soils reflect the C
stock at the former year, S(tK1), plus the net C input by the
crops during the growing season, I(t), minus the C losses
during that year, O(t)
SðtÞ Z Sðt K 1Þ C IðtÞ K OðtÞ (2)
The actual stocks of C, S(t), were available from the C
contents (Lobe et al., 2001) and the bulk densities of the top
20 cm. The output was calculated for each agro-ecosystem.
As the yields were rather similar in the three agro-
ecosystems and the yields were not available for all years
at all sites, net-C input I(t) was estimated from the average
yields. In order not to bias the results by the yield of 1 year
or one farmer we calculated an average yield for maize and
wheat each, normalized on the first yield after about 2 years.
The normalized yields were re-calculated into kg mK2 using
the average first yield as basis. From the yields (Fig. 1), we
calculated the C input according to Eq. (3)
IðtÞ Z YðtÞ !a !b !c !d (3)
 I. Lobe et al. / Soil Biology & Biochemistry 37 (2005) 1898–1909
5 4.0
Maize
Wheat
4 Output
Yield (t ha–1)
3
2 1.5
1
0 0.0
0 20 40 60 80 100
Duration of cultivation (y)
Fig. 1. Yields of maize and wheat as affected by duration of cropping,
shown as means of the three agro-ecosystems. Horizontal bars representing
the span of cultivation period between the three agro-ecosystems, vertical
bars the standard error.
with Y(t) is the yield as a function of cropping time; a, the
measured average C content of the crops (41.2% for maize
and 39.4% for wheat, nZ3); b, average ratio of net
production to grain yield (3.9 for maize and 5.5 for wheat,
Buyanovsky and Wagner, 1986); c, proportion of plant
residues returned to the soil (74% for maize and 82% for
wheat, Buyanovsky and Wagner, 1986); d, proportion of the
residues that are found in the top 20 cm (90% according to
Buyanovsky and Wagner, 1986). For sunflower, we
assumed a similar input as for maize. As the inputs of
maize and wheat showed no significant difference (P!0.05,
nZ24, paired differences test; Hartung, 1989), we could
simplify the estimation of the C inputs by using the mean of
the maize and wheat inputs. For sites where the crop (or
stubble) was still on the field, we estimated that half of the
input was returned to the soil by root exudates, root litter,
and above-ground litter, as the C input of roots and root
exudates is assumed to be equal (Barber, 1979; Plénet et al.,
1993) and the ratio of the shoots to roots can be assumed as
2 (Buyanovsky and Wagner, 1986).
The C output O(t) from our system, calculated from
Eq. (2), comprises: (a) C losses via erosion, E(t); (b)
decomposition of old SOM, D(t); and (c) decomposition of
the fresh residues added during the growing season, R(t):
OðtÞ Z EðtÞ C DðtÞ C RðtÞ (4)
Van der Merwe et al. (1990) found losses of soil by
wind-erosion of 1–5 Mg haK1 yearK1 depending on the
management practices around Bloemfontein. If our sites
solely lost soil as silt, as the predominantly lost particle
size (Hennessy et al., 1986; Stahr and Herrmann, 1996;
Lobe et al., 2001), a reduction of the silt content from 10 to
5.4% within 92 years is, in a rough estimation, equivalent
to a minimum soil loss of 4.6% (SE 1.6%, nZ3) from
the top 20 cm. At given bulk density of 1.43 g cmK3,
this soil loss corresponds to an erosion of silt of
1901
Input
3.5 Stocks
3.0
2.5
C (kg m–2)
2.0
1.0
0.5
0 20 40 60 80 100
Duration of cultivation (y)
Fig. 2. Modelled C input, stocks and output with increasing duration of
cropping, shown as means of the three agro-ecosystems.
1.43 Mg haK1 yearK1 (SE 0.42 Mg haK1 yearK1, nZ3),
which fits well into the range reported by Van der Merwe
et al. (1990). With the decreasing C content of the silt
particles (Lobe et al., 2001), caused by the cultivation, we
can estimate the annual C loss due to erosion of the silt
fraction which decreases from 42.1 to 18.1 kg C
haK1 yearK1. Compared with the C stocks and the in-
and output, the effect of the estimated erosion was quite
small (Fig. 2) and therefore only the decomposition of crop
residues and SOM were used in the calculations.
The rate of decomposition losses of the crop residues
was estimated from available reports in the literature. We
used a conservative assumption for our calculations that
after 1 year 20% of the root and shoot residues remain as
Barber (1979) reported that the soils receive annually
18% of root carbon and 10% of stem carbon as input to
the SOM. At a subtropical site with comparable
precipitation, Schomberg et al. (1994) found in a
litterbag experiment that about 15–20% of the wheat
residues were incorporated into the soil, after 1 year.
This is consistent with findings from incubation studies
that about 80% of the organic inputs were lost within the
first year (Broder and Wagner, 1988).
To extend the decomposition losses of the fresh residues
also to the losses of the old SOM we assumed that in the
long-term 1% of the C input remained in the soil. There are
no long-term studies which report the proportion of plant-
derived C, which remains for the long-term in the soil, but
with assuming 1% C input remaining in the soil the model
reached the C stocks after one century and resulted in good
estimates of the d13C value of the well-documented young
sites in the model. However, this simplifying assumption
can be used only for decades but not for centuries or
thousands of years.
We used the same decomposition loss rates for wheat and
maize since Broder and Wagner (1988) stated that the rather
similar behaviour of maize and wheat during decomposition
is unlikely to cause different isotopic processes. Knowing
1902 I. Lobe et al. / Soil Biology & Biochemistry 37 (2005) 1898–1909

the rate of decomposition losses and the input we were able
to calculate the amount of C3 and C4 carbon for each year
by summarizing the remains of each input at each year.
According to these calculations, the annual d13C value of
the soil organic matter, d13C(t), is given by Eq. (5)
continuous grassland to a wheat–maize-crop rotation
32 years ago, 40 years ago, 60 years ago or 90 years ago.
The data requirements are given in Table 2. No farmyard
manure was added to the soils. The parameterization for the
calculation of the C dynamics was done as follows:

d13 CðtÞ Z fd13 CGrassland !½SðtÞ K C3ðtÞ K C4ðtÞ
C C3ðtÞ !d13 CC3 C C4ðtÞ !d13 CC4 g=SðtÞ
with d13CGrassland is the d13C value of the grassland soil;
C3(t), amount of C3 carbon; C4(t), amount of C4 carbon;
d13CC3, d13C value of the C3 crop inputs; d13CC4, d13C value
of the C4 crop inputs. We used the d13C value of wheat and
sunflower for that of the C3 crop inputs, calculated for each
field from their proportion due to cropping history.
2.5. Modelled C dynamics using the Rothamsted
Carbon Model
We used the Rothamsted Carbon Model (ROTHC26-3)
(Jenkinson and Rayner, 1977; Coleman and Jenkinson,
1999) to model the C dynamics in the Tweespruit agro-
ecosystem, as required input data for the model was readily
available. The model includes the pools decomposable plant
material (DPM), resistant plant material (RPM), Cmic,
HUM and inert organic matter. This version had been tested
by Smith et al. (1997). The decay of the pools DPM, RPM,
Cmic and HUM follows first-order kinetics
Yt Z Y0 f1 K expðKabcktÞg;
(a) The amount of inert organic C of the soil from the
continuous grassland was estimated with the equation
(5) given by Falloon et al. (1998)
CI Z 0:049CT1:139 ; (7)
where CI is the amount of inert organic C, in t haK1, and
CT is the total organic C in the soil. The amount of inert
C was 0.26 kg C mK2.
(b) Then, we used the model to calculate the average annual C
input from the grass to match the measured values of soil
organic carbon of 3.23 kg C mK2 in the depth from 0 to
20 cm at Tweespruit by assuming that a steady state was
reached. We obtained a value of 0.089 kg C mK2 yearK1.
(c) Then the C inputs by maize and wheat were estimated to
reach a C storage of 1.24 kg C mK2 after 90 years of a
continuous crop rotation. As an approximation we
assumed that the C inputs were the same for wheat and
for maize. We obtained annual inputs of 0.036 kg C mK2
for wheat and an input of 0.036 kg C mK2 for maize in 2
years. In the first year of each cycle, the soil was bare from
January to November and maize was present in
December, in the second year maize was present from
January to June and the soil was bare until December
(6) (Table 2).

where Yt is the amount of carbon that decomposes in a

particular time t, Y0 is the initial amount of pool Y, a is the rate-
modifying factor for temperature, b is the rate-modifying 3. Results
factor for moisture, c is the soil cover rate-modifying factor,
and k is the decomposition rate constant for that compartment. 3.1. d13C values in bulk soil
The decomposition rate constants (in yearK1) were set to 10.0
(DPM), 0.3 (RPM), 0.66 (Cmic) and 0.02 (HUM) as suggested The d13C values of the grassland soil in the three agro-
by Coleman and Jenkinson (1999). ecosystems ranged from K13.4 to K16.1‰ (Table 3).
The model was run exemplarily for the 0–20 cm horizon Cultivation of the grassland (dominated by C4 vegetation)
at Tweespruit for the sites which were converted from a as arable land did result in a consistent decrease of the d13C
Table 2
Data requirements for the Rothamsted Carbon Model of the agro-ecosystem at Tweespruit

Variable (mean) Actual data used in the modelling

Monthly air temperature (8C)a 22.5 (J), 22.0 (F), 20.0 (M), 16.0 (A), 12.0 (M), 8.5 (J), 8.5 (J), 12.0 (A), 16.0 (S), 18.0 (O), 20.0 (N), 22.0 (D)
Monthly precipitation (mm)a 85 (J), 65 (F), 80 (M), 45 (A), 15 (M), 5 (J), 5 (J), 6 (A), 15 (S), 45 (O), 70 (N), 80 (D)
Monthly evaporation (mm)a 250 (J), 200 (F), 180 (M), 125 (A), 110 (M), 85 (J), 100 (J), 140 (A), 190 (S), 220 (O), 230 (N), 260 (D)
Soil depth (cm) 20
Soil clay content (%) 13.1
DPM/RPM ratio for wheat and maizeb 1.44
Soil cover Wheat crop production (first year): covered from May till November. Maize crop production (second and third
year): bare from January till November, covered from December till June
Monthly input of plant residues Unknown, obtained as described in the modelling section
Amount of inert organic matter Unknown, obtained as described in the modelling section
a
The weather data were taken from a nearby station.
b
The value suggested by Coleman and Jenkinson (1999) was used.
Table 3
d13C values in fine earth (!2 mm) and particle-size fractions of the surface soils (0–20 cm), d13C values of the bulk soil of the model and d13C values of the bulk soil corrected for the maize input by the model,
the minimum proportion of C3 carbon in bulk soil and particle-size fractions, and the potential proportion of new crop-derived carbon in the bulk soil of the three agro-ecosystems

Cultivation d13C (‰) Cropping Last crop Minimum proportion of C3–C (F; Eq. (1)) (%) Crop-derived
period with C3, last Ca model,
(years) Bulk soil Coarse Silt Clay Model, Model, crop- 15 yearsb (%) Bulk soil Coarse Silt Clay bulk soil
sand bulk soil derived Ca sand
Harrismith
0 K13.4 K14.7 K12.6 K12.2
3.5 K16.1 K20.7 K14.5 K13.7 K15.6 K15.6 100 C3 24 60 16 12 20

I. Lobe et al. / Soil Biology & Biochemistry 37 (2005) 1898–1909

8 K14.3 K19.3 K13.4 K13.0 K15.0 K15.3 42 C3 9 46 7 7 17
10 K14.0 K15.0 K14.2 K12.5 K13.7 K15.3 60 C4 6 3 14 3 17
20 K16.1 K20.8 K14.9 K13.6 K15.4 K15.4 100 C3 24 60 19 11 18
30 K14.6 K18.1 K14.1 K12.5 K13.6 K15.0 41 C4c 11 34 13 3 15
45 K15.3 ND K15.1 K13.6 K15.3 K16.1 50 (C3) 17 ND 21 11 24
68 K15.1 K16.3 K15.1 K13.0 K14.1 K17.0 41 C4 15 16 21 7 32
90 K17.7 K22.9 K18.1 K15.2 K17.7 K18.0 75 C3 38 82 45 24 41
Kroonstad
0 K14.4 K16.2 K13.1 K12.9
2.5 K14.7 K17.4 K13.9 K13.1 K14.6 K18.7 55 C4 2 15 7 2 42
7.5 K17.0 K18.4 K15.6 K14.2 K15.4 K18.0 75 C4d 22 22 19 10 30
12 K14.5 K19.0 K13.8 K13.4 K14.6 K18.3 50 C4 1 34 6 5 39
20 K16.3 K23.4 K16.2 K14.1 K16.1 K18.5 50 (C3) 15 87 27 11 40
30 K16.2 K19.2 K17.4 K14.4 K15.6 K19.7 80 C4 15 30 33 12 45
40 K17.2 K25.9 K17.6 K15.2 K18.0 K20.6 18 C3 23 93 33 17 50
57 K17.9 K18.8 K19.5 K17.4 K16.8 K21.9 80 C4 28 25 47 33 61
98 K15.5 K19.3 K16.4 K14.3 K15.6 K25.4 50 C4c 9 30 25 10 89
Tweespruit
0 K16.1 K17.4 K15.4 K12.8
2 K16.2 K20.5 K15.6 K12.8 K18.6 K18.6 100 C3 1 32 1 0 22
8.5 K16.1 K20.6 K15.4 K12.0 K16.1 K18.6 78 C4 0 32 0 K6 21
12 K18.6 K22.5 K19.1 K15.8 K18.6 K18.7 92 C3 21 50 30 20 23
22 K17.0 K25.4 K17.9 K13.7 K16.3 K19.2 69 C3 8 79 20 6 27
32 K17.4 K18.4 K17.3 K15.3 K17.2 K19.2 50 C4c 12 10 15 17 27
40 K17.1 K18.5 K17.7 K14.4 K16.6 K20.2 50 (C4) 9 11 19 11 37
60 K15.5 K17.0 K17.2 K12.8 K15.8 K21.3 50 (C4) K6 K4 15 0 46
90 K16.3 K17.5 K15.9 K13.9 K16.7 K23.3 50 (C4) 2 1 4 7 65

The on of the data was 0.1‰.

a
Input of maize-C4 mathematically treated as C3 input.
b
Length of cropping when site was !15 years under cultivation.
c
Crop still on field.
d
Stubble on field.

1903
1904 I. Lobe et al. / Soil Biology & Biochemistry 37 (2005) 1898–1909
values with increasing length of cultivation (Fig. 5). This
was caused by the input of the C3 crops wheat and sunflower
with more negative d13C values than the grassland
vegetation. However, the irregular input of maize (C4)
increased the d13C values with increasing duration of
cultivation. Larger proportions of C4 crops during the last
15 years increased the d13C values, in comparison with the
average course of the decreasing d13C values (e.g. 8-, 30-,
and 68-year-old Harrismith site; Table 3), while larger
proportions of C3 crops decreased the d13C values (e.g. 7.5-
year-old Kroonstad site, and 12-year-old Tweespruit site;
Table 3). In addition, the last crop at the sites had a strong
influence on the d13C values, e.g. 3.5-, 20-, and 90-year-old
Harrismith site, 40-year-old Kroonstad, and 22-year-old
Tweespruit (Table 3).
3.2. Estimations of C3 crop-derived organic matter
and turnover rates
At our sites, the original vegetation was dominated by C4
plants. Disregarding the effect of maize inputs gives
therefore a first estimation of the minimum amount of C3
organic matter that has replaced C4 organic material. The
minimum proportion of the newly formed C3-derived
organic matter (Eq. (1)) accounted for up to 21–38% of
the total C in the three different agro-ecosystems (Table 3).
The largest proportion of C3-derived organic matter was
found in the 90-year-old field at Harrismith that was grown
for most of the cultivation period with wheat.
With increasing duration of cultivation the yields
declined exponentially (Fig. 1). The results revealed that
due to the declining yields, the annual C input rates also
declined rapidly after conversion of the grassland to
cropland, resulting in rapid annual C output rates that
approached steady-state equilibrium after ca. 20 years of
cropping (Fig. 2). Further decline in C stocks after this time
reflects continuous C output at low input rates as the output
was still slightly higher than the input.
The calculated C stocks of the stepwise C replacement
model describe the measured values very well. We found
differences between measured and modelled values of 0.1–
6.9 Mg C haK1 and on average 0.2G2.2 (SD) Mg C haK1
(nZ24). Running the model with the data of the cropping
history resulted in differences between measured and
modelled (Eq. (5)) values of 0.0–2.5‰ and on average
0.21G0.75‰ (SD, nZ24) (Table 3). The value of the
2-year-old site in Tweespruit, which was 2.5‰ too positive,
might be due to the irrigation of this site resulting in
increased d13C values (Boutton, 1996).
When we replace all C4 maize inputs mathematically by
C3 inputs, we receive the proportion of the newly formed
soil organic matter. After 90 years of cropping, 41–89% of
the total C in the three different agro-ecosystems was
derived from the new crop-derived organic matter (Table 3).
The results of our stepwise C replacement model
revealed no linear increase of the amount of crop-derived
0.8 0.8
Crop-derived C (kg m–2)
Crop-derived C / total C
0.6 0.6
0.4 0.4
0.2 Crop-derived C / total C
0.2
Crop-derived C
0.0 0.0
0 20 40 60 80 100
Duration of cultivation (y)
Fig. 3. Proportion of crop-derived C in total C in the bulk soil, as well as the
total amount of crop-derived C as affected by duration of cropping. Shown
as the mean of the three agro-ecosystems with horizontal bars representing
the span of the cultivation period and vertical bars the standard error.
C with the duration of cultivation (Fig. 3). Both the absolute
amount as well as the relative proportions of newly
incorporated crop-derived C increased rapidly within
the first 2–3 years of arable land-use. After this initial
period, about one-third of the crop-derived C was lost again,
however, after 20 years of cropping, the proportions of crop-
derived C escalated again. To reach the level of the 2–3-
year-old fields about 60 years were necessary after the
conversion of grassland to arable land.
The modelling of the C dynamics with the Rothamsted
Carbon Model found that this model described the stock of C
in the surface soil (0–20 cm) well after 90 years of the
wheat/maize crop rotation (Fig. 4). The same was true for
the 60-year old site, whereas 32 and 40 years after conversion
the measured values were much lower than predicted (Fig. 4).
3.3. d13C values in particle-size separates
The d13C values of the grassland increased from coarse
sand to clay (Fig. 5). With increasing duration of cultivation,
the d13C values of the particle-size fractions declined in
accordance with the bulk soil. This decrease in the d13C
values during cultivation was more expressed in the order
clay!silt!coarse sand indicating that more new C3
material was incorporated in the coarse sand than in the
clay-size separates (Fig. 5). The proportion of the new C3
organic matter increases therefore from the clay to the
coarse sand fraction (Table 3).
In the grassland, the d13C values in the silt fraction
showed an enrichment of 13C relative to the d13C value in
the bulk soil showing the higher decomposition stage of the
organic matter in that fraction. With increasing duration of
cultivation, however, the d13C values became more negative
in comparison with the bulk soil, indicating a higher
proportion of younger C3 organic matter in the silt fraction
after prolonged cultivation.
 I. Lobe et al. / Soil Biology & Biochemistry 37 (2005) 1898–1909 1905

3.5 12

C3 - derived 10
C4 - derived
3.0 Total 8

δ15N (o/oo)
6
2.5
4

Bulk soil
2 Clay
Soil C (kg C m–2)

2.0 Silt
Coarse sand
0
0 20 40 60 80 100
Duration of cultivation (y)
1.5
Fig. 6. Effect of duration of cropping on the d15N values of the bulk soil and
the clay, silt and coarse sand fractions, shown as means of the three agro-
ecosystems. Vertical bars represent the standard error between the three
1.0 agro-ecosystems.

the d13C values which decreased. With increasing duration

of cultivation, the d15N values of the clay fraction did
0.5 increase similar to those of the bulk soil, while the signature
of the silt fraction remained unchanged. In the coarse sand
fraction the d15N values decreased with prolonged cropping,
0.0 suggesting different N dynamics in the size separates.
0 10 20 30 40 50 60 70 80 90
Duration of cultivation (y)

Fig. 4. Modelled total, C3- and C4-derived soil C stocks using the Roth-C 4. Discussion
Model for Tweespruit agro-ecosystem.
4.1. d13C values in bulk soil
3.4. d15N values in bulk soil and soil separates
The d13C values of the grassland soil in the three agro-
15
The d N values of the grassland ranged from 7.6 to ecosystems ranged within 2.7‰ (Table 3). A slight
8.2‰ revealing a much lesser variation than the d13C difference in composition of the native vegetation (some
values. With increasing duration of cultivation the d15N minor contribution of C3 vegetation, e.g. C3 shrubs, to the
values increased logarithmically (Fig. 6), in contrast to C4 grasses) might have caused these differences. We do not
attribute this difference in d13C to increasing precipitation
from Tweespruit to Harrismith, resulting in a lower
–12 discrimination of 13C and therefore increasing d13C values
–14 (Boutton, 1996; Henderson et al., 1992), because the
d-values of the grasses were rather similar, as indicated by
–16
δ13C (o/oo)

Table 4
–18 Average isotopic signature of the shoots and roots of the plants of the sites
(standard errors given in parentheses, nZ3)
–20
Plants C (g kgK1) N (g kgK1) d13C (‰) d15N (‰)
–22 Bulk soil
Clay Shoots of
Silt
–24 Grass 433G7 9.0G1.5 K13.3G0.1 2.8G0.9
Coarse sand
Maize 435G1 6.6G1.6 K12.4G0.1 2.9G1.0
–26 Wheat 439G12 7.2G2.4 K25.4G1.0 4.3G1.3
0 20 40 60 80 100 Sunflower 444G6 4.6G0.4 K28.6G0.2 2.5G0.3
Duration of cultivation (y) Roots of
Grass 422G9 8.8G0.6 K13.3G0.2 4.7G1.8
Fig. 5. Effect of duration of cropping on the d13C values of the bulk soil and Maize 389G26 9.1G2.1 K12.4G0.2 0.6G1.5
the clay, silt and coarse sand fractions, shown as means of the three agro- Wheat 348G31 7.9G1.7 K26.2G1.1 5.9G0.2
ecosystems. Vertical bars represent the standard error between the three Sunflower 459G1 3.8G0.1 K28.1G0.8 1.9G0.2
agro-ecosystems.
1906 I. Lobe et al. / Soil Biology & Biochemistry 37 (2005) 1898–1909
the small standard error (SE, nZ3) in Table 4. The d13C
changes in C4 plants in response to environmental factors is
generally low, ca. 1‰ (Boutton, 1996) and not high enough
to explain the differences among the grassland of the three
ecosystems.
The cultivation of the grasslands (dominated by C4
vegetation) to arable land did result in a consistent decrease
of the d13C values with increasing length of cultivation
(Fig. 5). This was caused by the input of the C3 crops wheat
and sunflower with more negative d13C values than the
grassland vegetation. This exponential decrease of the d13C
signal can be attributed to one or more of the following
reasons:
(a) Advanced decomposition of the organic matter due to
the longer duration of C3 crop cultivation leading to
higher soil 13C values, which would reduce the
‘observed’ change in soil 13C values of the SOM on
the longer cropped sites. However, usually less than
0.5‰ are attributed to the effect of decomposition in the
surface soil (O’Brien and Stout, 1978; Monson and
Hayes, 1982; Balesdent et al., 1990).
(b) Enhanced mineralization as a result of the breaking of
aggregates after the cultivation of grassland and thus the
access to formerly physically protected organic matter,
derived from C4 plants, might also explain the higher
decline in the d13C values in the beginning of
cultivation.
(c) Effects of the decrease of d13C values in the
atmospheric CO2, the so-called Suess-effect, was
considered to account for 0.2 ‰ for cultivated
ecosystems in carbon cycle models (Fung et al.,
1997). But in our case, this effect can be neglected as
we use the grassland as the reference for the initial C4
soil organic matter.
(d) Irregular maize inputs (C4) besides C3 inputs of wheat
and sunflower should be the major reason for the non-
linear decrease of the d13C values. The maize inputs
comprise organic carbon that has even more positive
d13C values (12.4‰) than the organic material derived
from the grassland vegetation (13.3‰). As a result,
these maize inputs blur the input of the C3 plants.
(e) Furthermore, the shorter cultivated fields were grown to
a higher proportion with C3 crops than the longer
cultivated ones, resulting in a faster response to C3
signature in the younger fields, and therefore a faster
decline of the d13C values in the beginning of the
cultivation than later on (Table 3).
The irregular input of maize (C4) should have the largest
impact on the increase of the d13C values with increasing
duration of cultivation. This is supported by two obser-
vations. On the one hand, d13C values, which were obviously
higher or lower than the average exponential decrease, could
be well explained by the different proportions of C3 crops
during the last 15 years, e.g. 8-, 30-, and 68-year-old
Harrismith site, 7.5-year-old Kroonstad site, and 12-year-
old Tweespruit site (Table 3). On the other hand, the last crop
at the sites had a strong influence on the d13C values, e.g. 3.5-,
20-, and 90-year-old Harrismith site, 40-year-old Kroonstad,
and 22-year-old Tweespruit (Table 3).
Some of the sites have been ploughed to more than
20 cm. Taking into account that ploughing deeper than
20 cm results in a dilution of the fresh organic matter in the
upper 20 cm the measured values on these sites show higher
d13C values and therefore smaller proportions of replaced
C4 organic matter. As a consequence, the values on these
sites represent the minimal input of C3 organic matter that
occurred on these fields with respect to the top 20 cm.
For the determination of the carbon dynamics the decline
in the d13C values has to be corrected for the maize inputs;
otherwise, the loss rates of the savanna C will be
underestimated.
4.2. Estimations of C3 crop-derived organic matter
and turnover rates
At our sites, the original vegetation was dominated by C4
plants. Hence, all effects changing the d13C values that are
not associated with the change of the vegetation, namely
decomposition, increase the d13C values. Disregarding the
effect of maize inputs gives therefore an estimation of the
minimum amount of C3 organic matter that has replaced C4
organic material. At least 21–38% of the original organic
matter was replaced by wheat-derived C within 90 years of
cultivation in this subtropical agro-ecosystem. The esti-
mation of the whole newly formed soil organic matter
suggested that up to 41–89% of the total C in the three
different agro-ecosystems was derived from the new crop-
derived organic matter (Table 3). On average, 65% of the
original organic matter was replaced by wheat and maize
crop debris within 90 years of cultivation in this subtropical
agro-ecosystem. Compared with findings of Paul et al.
(1997) that 70% of the total C was prairie-derived after
85 years of cultivation with wheat in the Great Plains,
whose texture is silt dominated, partly a larger amount of
SOM was substituted by crop-derived organic matter in the
sandy soils of the South African Highveld
According to our stepwise C replacement model, the
amount of crop-derived C did not increase linearly with the
duration of cultivation (Fig. 3). Within the first 2–3 years of
arable land-use, both the absolute amount as well as the
relative proportions of newly incorporated crop-derived C
increased rapidly, because high crop yields could still be
maintained (Fig. 2) while the loss of savanna C occurred
rapidly from the labile C pools (Lobe et al., 2001). After this
initial period, about one-third of the crop-derived C was lost
again, probably reflecting rapid decomposition of the newly
formed humus as well as the decrease in crop yields as
cultivation was prolonged. After 20 years of cropping,
however, the proportions of crop-derived C escalated again.
It took about 60 years after the conversion of grassland to
 I. Lobe et al. / Soil Biology & Biochemistry 37 (2005) 1898–1909
arable land to reach the level of the 2–3-year-old fields. The
results emphasize the high contribution of fresh crop
residues, entering the soil, to the SOM. The input of the
new organic matter could not substitute the losses of SOM
in the first 3–20 years, as newly formed crop-derived SOM
was not stable against decay. It seems reasonable to assume,
therefore, that without additional C input by interrupting the
plough periods, a sustainable management for organic
matter stocks and associated soil fertility will thus hardly be
possible. It has to be taken into consideration that our model
was established to simulate the d13C values of the
investigated agro-ecosystems and their duration of cultiva-
tion. As the model lacks a term to take the slow decay of the
stable humus pool into account, it cannot be accurately used
for predicting longer cultivation periods.
However, the data could be successfully described using
the Roth-C Model (Fig. 4); here it has to be noted that the
modelled C inputs were unrealistically low compared with
those in Fig. 2, in order to match the measured C stocks
90 years after conversion. In Tweespruit, the straw was not
removed from the fields and the grain yields were low, 1.5 t
dry matter haK1 for wheat and 3 t dry matter haK1 for
maize. But even when assuming a lower ratio of straw to
grain of 1:1 and a lower ratio of under-ground/above-ground
biomass of 0.2, therefore resulting in a lower ratio of net
production to grain yield like for the calculation of the input
in Fig.2, the total C inputs into the soil are with about 0.1
Kg C mK2 yearK1 much higher than the modelled C inputs.
The main reason for that underestimation of the C inputs
might be that the model may not work well under semi-arid
conditions, because the evaporation is so high, the rate-
modifying factor for moisture is always 0.2, so decompo-
sition may be less in the model than it is in reality (K.
Coleman, pers. comm). Thus, model modifications of the
estimation of the rate-modifying factor might be required to
obtain the same C stocks as shown in Fig. 4, but by using
more realistic C inputs in the model.
4.3. d13C values in particle-size separates
The d13C values of the grassland increased from coarse
sand to clay (Fig. 5). This is the result of the different stage
of decomposition of SOM in the fractions. The more
decomposed the organic matter is in a particle-size fraction
the more enriched this fraction is in 13C.
With increasing duration of cultivation, the d13C values
of the particle-size fractions declined, reflecting the
different velocity of the incorporation of the C3 inputs to
the particle-size fractions and their turnover. The more
expressed decrease in the d13C values during cultivation
from the clay to the coarse sand fraction indicated that more
new C3 material was incorporated in the coarse sand than in
the clay-size separates (Fig. 5). Therefore, the proportion of
the new C3 organic matter increased from the clay to the
coarse sand fraction (Table 3).
1907
The d13C values of the silt and the clay fraction were well
correlated with the d13C values of the bulk soil, i.e. when the
d13C values were high in the bulk soil, due to a higher
proportion of C4 inputs, the ones of the two fractions were
higher, too, and vice versa. In contrast, the d13C values of
the coarse sand did not reflect the course of the bulk soil
(e.g. 68-year-old Harrismith site, 12-year-old Kroonstad site
and 57-year-old Kroonstad site, 22-year-old Tweespruit
site, Table 3). This indicated that the 13C signature of the
coarse sand was predominantly determined by the crop that
was grown in the last year. Nevertheless, the cropping
history of the soil, i.e. the percentage of C3 crops, can also
be seen in the 13C signature of the coarse sand but to a minor
extent (especially in the first years with the same last crop,
Table 3). When the crop was still on the field (in all cases
maize), the d13C value of the coarse sand was lower than on
other sites with maize as the last crop, thereby indicating the
former C3 inputs, and that only root residues and root
exudates contributed to the new C4 input.
With increasing duration of cultivation the d13C values in
the silt fraction became more negative than in the bulk soil,
though in the grassland, the higher decomposition stage of
the organic matter in the silt fraction resulted in higher d13C
values relative to those in the bulk soil. This indicated a
higher proportion of younger C3 organic matter in the silt
fraction after prolonged cultivation. In addition, the d13C
values became more negative in the silt than in the clay
fraction with prolonged arable cropping in comparison with
the d13C values of the grassland. In contrast, Balesdent et al.
(1998) found a similar change in the d13C values of the silt
and the clay fraction due to cultivation. Taking into account
that arable cropping on the investigated sites resulted in
linear losses of the silt fraction itself (Lobe et al., 2001), the
more negative d13C values in the silt fraction may be
explained by the relatively higher input of fresh C3-derived
organic matter to a lower amount of the remaining silt. The
results of the d13C values therewith support the findings
from previous lignin analyses (Lobe et al., 2002), suggesting
that old SOM of the silt is rejuvenated upon silt losses.
It was not possible to simulate the crop-derived C in the
particle-size fractions as no information was available as to
how much of the input was transferred to the different
particle-size pools. Therefore, our model was restricted to
the C dynamics in the bulk soil.
4.4. d15N values in bulk soil and soil separates
The logarithmic increase of the d15N values with
increasing duration of cultivation (Fig. 6) can be explained
by two reasons:
(a) Decomposition of plant residues increases the values of
d15N (Turner et al., 1983; Nadelhoffer and Fry, 1988;
Yoneyama, 1996). An increased mineralization of
easily mineralizable organic N in the beginning of the
cultivation results therefore in a loss of 15N-depleted
1908 I. Lobe et al. / Soil Biology & Biochemistry 37 (2005) 1898–1909
compounds and an enrichment of remaining 15N-
enriched compounds of the soil.
(b) The air nitrogen (0‰) which is taken up by plants by
N2-fixation turns the d15N values of the vegetation
towards 0‰ and as a consequence the ones of the soil
(Yoneyama, 1996). A higher contribution of N2-fixation
of the grassland vegetation compared with the crops
thus causes an increase of the d15N values with
increasing duration of cultivation.
The small differences in the soil pH that might affect N
mineralization processes seem to have no influence on the
d15N values, as there is no correlation between the
oscillating soil pH and the d15N values (P!0.05).
In the particle-size fractions, the d15N signature differed
due to the stage of decomposition of the organic N material.
As a consequence, the d15N values in the grassland
increased from the coarse sand to the clay fraction. In the
coarse sand fraction, the d15N values were nearest to the
values of the plants as a result of the highest content of
particulate plant residues (Table 4).
With increasing duration of cultivation, the d15N values
of the clay fraction did increase similar to the bulk soil. This
suggested that the effect of the faster mineralization during
the initial cultivation stages mainly affected the clay
fraction, probably as the microbial N-products predomi-
nantly located there. Amelung et al. (2002) indicated that
the clay fraction contains the majority of the amino sugars
representing microbial residues. Decreasing d15N values in
the coarse sand fraction with increasing duration of
cultivation could be attributed to lower d15N values in the
C3 crop plants than in the C4 grassland, especially of the
roots (Table 4). However, we should then expect to observe
an exponential decrease of the d15N values as seen in the
d13C values. Therefore, a second process seems to be
responsible for or to contribute to the linear decline of the
d15N values. The uptake of fertilizer-N, which has d15N
values around 0G3‰ (Yoneyama, 1996), would lead to a
decrease of the d15N values in the plants and consequently in
the coarse sand fraction, which mainly contains decompos-
ing plant debris. This effect is probably enhanced as, with
increasing duration of cultivation, the application of
fertilizers increased due to decreasing yields. We thus
attribute the decreasing d15N values of the coarse sand
fraction to an increased contribution of fertilizers to the 15N
signature. As a result, the proportion of fertilizer-derived N
in the soil organic matter will increase. Yet, this effect was
not detectable by d15N natural abundance measurements for
other fractions than coarse sand.
5. Conclusions
Sandy soils of the South African Highveld have little
potential for stabilization of organic matter. Our results
confirm that after 90 years of arable cropping 41–89%
(65% on average) of the remaining 1.3 kg mK2 soil C was
crop-derived in the top 20 cm. By means of the 13C natural
abundance tracer technique it was shown that replacement
rates of organic matter increased as particle-size increased;
the isotopic signature of the coarse sand fractions exhibited
the most recent plant input.
Using our newly developed stepwise C replacement
model, it was possible to correct soil d13C data for the
irregular inputs of maize (C4) or wheat (C3) C inputs. Soil C
stocks after 90 years of the wheat/maize crop rotation were
also accurately described with the Rothamsted Carbon
Model, although the modelled C inputs to the soil were very
low.
Furthermore, at least up to 33% of the carbon in the clay
fraction was formed by the crop inputs indicating that the
old SOM reserves were used up due to decreasing yields and
enhanced mineralization. The d15N values were not related
to the origin or the age of the N, as was the case for 13C.
However, it traced the origin of fertilizer N within the recent
plant debris, to the coarse sand fraction. We conclude that
the natural abundance tracer technique is not restricted to
soils with an ideal C4/C3 vegetation change, but it can be
applied profitably to sites with mixed cropping history,
provided that land-use history and yields are well reported.
Acknowledgements
This study was financially supported by the German
Research Foundation. IGER is grant aided by the Biotech-
nology and Biological Sciences Research Council. We
thank all the South African farmers who allowed us to
sample their fields, and gave us the important information
about the cropping history of the sites, and De Villiers
Bosman, David Fouche, Donny Holmes, Deon Maree, Leon
Strachan, and Frikkie Swanepoel for organizing the contacts
with the farmers. We thank Prof. D. Jenkinson and Mr
K. Coleman for their help and discussion with regards to the
Roth-C Model.
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