Carbon accumulation in soils of forest and bog ecosystems of Southern Valdai in the Holocene

Abstract

Carbon stocks and accumulation rates in humus and peat horizons of the contiguous soil series of forest and bog ecosystems
have been studied in the Central Forest State Biosphere Reserve, Tver Region. Upland soil types (soddy podzolic, brown, and
white podzolic) have been compared to paludified (peat-enriched gley podzolic and peaty gley) and bog soils differing in trophic
status, including those of upland, transitional, and lowland bogs. The results show that carbon stocks in mineral soils are
many times smaller than in waterlogged soils and an order of magnitude smaller than in bog soils. Mineral and bog soils are
characterized by similar rates of carbon accumulation averaged over the entire period of their existence. The highest rate
of carbon accumulation has been noted for the soils of waterlogged habitats, although this process may be periodically disturbed
by fires and other stress influences.

Full text

ISSN 1062-3590, Biology Bulletin, 2008, Vol. 35, No. 5, pp. 516–524. © Pleiades Publishing, Inc., 2008.
Original Russian Text © T.Y. Minayeva, S.Ya. Trofimov, O.A. Chichagova, E.I. Dorofeyeva, A.A. Sirin, I.V. Glushkov, N.D. Mikhailov, B. Kromer, 2008, published in Izvestiya
Akademii Nauk, Seriya Biologicheskaya, 2008, No. 5, pp. 607–616.
516
INTRODUCTION
Interest in studies on soil carbon stocks and dynam-
ics has increased in view of current climate changes and
the role of atmospheric
CO
2
in them. The dynamics of
accumulation and stocks of carbon are important inte-
grated parameters characterizing biogenic processes in
the soil, including energy–mass exchange in the soils
and biogeocenosis as a whole (
Regulyatornaya rol'…,
2002).
Forest and bog ecosystems form the basis of the
patchwork landscape–biogeocenotic structure of the
taiga zone. According to the State Land Inventory, peat
bogs occupy more than 8% of Russia’s territory (“Tor-
fyanye bolota…,” 2001); with account of paludified
soils (peat layer <30cm), this proportion increases to
21.6% (Vomperskii et al., 1994). Although the area of
paludified soils is greater, the total carbon stock in their
peat horizons is significantly smaller than in bogs:
12.6

10
9
vs. 100.9

10
9
t (Vomperskii et al., 1994).
However, since these soils are unstable, the environ-
ment-forming role of their carbon stock may be dispro-
portionally more important. Of special significance
may be paludified forests, which occupy 69% of Rus-
sia’s shallow-peat lands (Vomperskii et al., 1994).
Carbon stocks in peat bogs and in non-bog soils are
often considered separately. However, when assessing
carbon deposition in taiga ecosystems, it is expedient to
consider the whole variety of soil cover components,
including mineral, paludified, and bog soils. Peat
deposits in bogs should be analyzed throughout their
depth, with regard to the biogeochemical integrity of
bog soils and underlying waterlogged layers of peat.
In this study, we made an attempt to estimate carbon
stocks and accumulation rates in humus and peat hori-
zons of the contiguous soil series of forest and bog eco-
systems. The study was performed in the Central Forest
State Biosphere Reserve (CFSBR), where a complex of
southern taiga forests and bogs differing in genesis is
well preserved. This territory is almost unaffected by
human activities, but comprehensive soil-biogeoceno-
logical studies have been performed there for several
decades (
Regulyatornaya rol'…,
2002).
MATERIALS AND METHODS
The reserve is on the southern macroslope of the
Valdai Upland (
56°26
′
–39
′
N,
32°29
′
–33°01
′
E). Its
moderately continental climate subject to seasonal oce-
anic influence (Minayeva et al., 2001), weakly dis-
sected topography, and prevalence of heavy morainic
deposits account for a wide occurrence of waterlogged
ecotopes, which occupy more than 20% of the reserve
area (
Central Forest…,
1999). Specific geomorpholog-
Carbon Accumulation in Soils of Forest and Bog Ecosystems
of Southern Valdai in the Holocene
T. Y. Minayeva
a
, S. Ya. Trofimov
b
, O. A. Chichagova
c
, E. I. Dorofeyeva
b
, A. A. Sirin
d
,
I. V. Glushkov
d
, N. D. Mikhailov
e
, and B. Kromer
f
a
Central Forest State Biosphere Reserve, p/o Zapovednik, Tver Region, 172513 Russia; e-mail: tminayeva@ecoinfo.ru
b
Faculty of Soil Science, Moscow State University, Moscow, 119991 Russia
b
Institute of Geography, Russian Academy of Sciences, Staromonetnyi per. 29, Moscow, 109017 Russia
c
Institute of Forest Science, Russian Academy of Sciences, Uspenskoe, Moscow Region, 143030 Russia
d
Institute of Geochemistry and Geophysics, National Academy of Sciences of Belarus,
ul. Kuprievicha 7, Minsk, 220141 Belarus
e
Heidelberger Akademie der Wissenschaften, Inst. f. Umweltphysik, INF 229, D-69120 Heidelberg, Germany
Received February 15, 2007
Abstract
—Carbon stocks and accumulation rates in humus and peat horizons of the contiguous soil series of
forest and bog ecosystems have been studied in the Central Forest State Biosphere Reserve, Tver Region.
Upland soil types (soddy podzolic, brown, and white podzolic) have been compared to paludified (peat-
enriched gley podzolic and peaty gley) and bog soils differing in trophic status, including those of upland, tran-
sitional, and lowland bogs. The results show that carbon stocks in mineral soils are many times smaller than in
waterlogged soils and an order of magnitude smaller than in bog soils. Mineral and bog soils are characterized
by similar rates of carbon accumulation averaged over the entire period of their existence. The highest rate of
carbon accumulation has been noted for the soils of waterlogged habitats, although this process may be period-
ically disturbed by fires and other stress influences.
DOI:
10.1134/S1062359008050130
SOIL BIOLOGY

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CARBON ACCUMULATION IN SOILS OF FOREST 517
ical features and local hydrologic conditions determine
the structure of soil cover (
Genesis…,
1979)
To evaluate carbon stocks and accumulation rates,
test plots were established in forest and bog ecotopes
located on different relief elements and differing in
hydrologic conditions and dynamics of mineral nutri-
ents. Within these plots, we identified topographic ele-
ments and biogeocenoses associated with them for dat-
ing carbon accumulated in different soil types (Fig. 1).
The plots reflected the diversity of ecotopes along the
gradient from upland forest biogeocenoses on well-
drained soils to peat bogs.
Plot 1 was on a flat esker slope with soddy podzolic
soils under a nemoral wood sorrel spruce forest. The
soddy podzolic soils could be formed due to agricul-
tural land use (plowing) in this area more than
200 years ago (Karimov and Nosova, 1999). For
–1
100 200
100 300200 400 500 6000
–1
0
1
0
1
0
–2
–1
100 300200 4000
0
1
–2
–5
250 500 750 1000 15000
0
1
–6
2
100 2000
3
4
–1
1
0
–1
–2
–3
–4
898 + 400 184 + 40 106 + 40
4.1 4.2 4.3
1
2
3
4
1958 + 90
9800 + 100
8.1
8.2
Mezha River shoreline
441 +10
950 + 100
5628 + 26
10206+ 70 9959 + 24 9218 + 24
7.1 7.2 7.3 7.4 7.5 7.6
4380 + 170
6
605 + 180 192 + 40
1969 + 200 853 + 50
5.1 5.2 5.3 5.4
106 + 40
Fig. 1.
Locations of soil pits within catenae in test plots: (
1
) sites of stratigraphic core sampling (numbers of cores correspond to
those in Table 1); (
2
)
14
C
age of bottom layers, cal. BP; (
3
) day surface relief, (
4
) peat deposit bottom relief. Abscissa shows catena
length, m; ordinate shows relative elevation/depth, m.

518
BIOLOGY BULLETIN
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MINAYEVA et al.
detailed characteristics of this plot, see Shaposhnikov
et al. (1994).
Plot 2 was at a convex bend of an eastern slope
descending to a river, with brown soil under a complex
of undisturbed nemoral wood sorrel spruce forests.
Good drainage provided for a high productivity of plant
communities, which were described by Minayeva
(1988). Characteristic features of brown soil formation
under these conditions were also described previously
(Trofimov and Stroganova, 1991; Trofimov, 1998).
Plot 3 was on a flat, weakly drained watershed with
prevalence of white podzolic (peat-enriched surface-
gley podzolic) soils under true moss–bilberry spruce
forests and sphagnum–bilberry spruce forests. This plot
represented the most widespread ecotope type in the
reserve. Its characteristic features included a high water
table and a stable structure of uneven-aged tree stand
(Shaposhnikov, 1988).
Plot 4 was in a local depression on a watershed
between small rivers, with peaty gley podzolic and
peaty gley soils under sphagnum and bilberry–sphag-
num spruce forests. We studied three soil pits along the
catena representing a paludification gradient between
upland peat soils (4.1) and peaty gley soils (4.2, 4.3)
under sphagnum and bilberry–sphagnum spruce for-
ests. For more details, see the monograph (
Central For-
est…
, 1999).
Plot 5 was in a depression within a flat watershed on
a ridge top. We studied three soil pits along a catena
with peaty podzolic gley (5.2), peaty gley (5.1), and
upland peat bog soils (5.3, 5.4) formed under sphag-
num–bilberry and sphagnum spruce forests passing
into cotton grass –sphagnum pine forest. Plant commu-
nities in the plot are in a dynamic state due to climate
fluctuations and tree harvesting (Shaposhnikov et al.,
1991; Minayeva and Glushkov, 2003).
Plot 6 was a small transitional bog on a ledge of
river valley slope. The depth of peat deposit varied from
0.5 to 1.5 m. We studied the profile of upland peat soil
in the central oligotrophic area with the maximum
deposit depth.
Plot 7 was on the southern slope of a watershed
occupied by a massive upland bog with peat deposits
varying in depth. Studies were performed along a
transect passing through the highest point of the bog
area (2.5 m above the lowest point), with six soil pits in
upland bog soils with peat depth varying from 0.5 to 3.8
m.
Plot 8 was in a cup-shaped river valley with peat
deposits (30–180 cm deep) overlying slopes and ledges
of the ancient terrace. Lowland peat soils under herba-
ceous spruce bog forest with an admixture of black
alder were studied in the plot.
Samples from humus horizons of mineral soils were
taken throughout the horizon depth. Peat samples for
determining apparent density, carbon content, and
radiocarbon age were taken from soil pits when the
deposit was shallow (no deeper than 50 cm); in other
cases, a TBG-1 auger was used. In a pit, mineral soils
and dense peat were sampled with a knife; loose peat,
with a soil cup. In both cases, the samples were no more
than 2–5 cm deep. When the auger was used or the
deposit contained liquid peat layers, total samples
including the whole test layer (10–50 cm) were ana-
lyzed. Apparent density was determined gravimetri-
cally, and ash content was determined by an ignition
method (Arinushkina, 1970).
Soil monoliths 1/16 m
2
in cross section were taken
in five to seven replications to determine carbon con-
tents in organogenic horizons of mineral soils. The
monoliths were dried and weighed, and carbon was
measured with a rapid response analyzer. In mineral
horizons, carbon content and density were determined.
We also used relevant data obtained in previous field
and laboratory studies on the CFSBR territory (Trofi-
mov et al., 1997; Trofimov, 1998). Carbon content in
peat samples was determined by dry combustion in
oxygen using an AN-7529 rapid response analyzer with
coulometric detection.
The depth of test layers for radiocarbon dating did
not exceed 2–4 cm. Samples were collected from soil
pits (with a knife) or from multiple core samples taken
within a 1-m
2
plot. The weight of a dry sample was no
more than 50 g. Before acid–base treatment, the sam-
ples were cleaned of organic debris, including plant
roots.
To prepare peat and soil samples for radiocarbon
dating, they were treated by an acid–base–acid method.
In the case of soil samples, this was followed by boiling
and washing preparations of humic acids (HAs) with
distilled water (to pH 6–7) and drying them under a
quartz lamp (Chichagova, 1985, 2005; Chichagova et
al., 2001). To obtain the substance for determining
14
C
radioactivity (benzene), the sample was subjected to
dry distillation or combustion in a reactor, and the prod-
uct was sintered with metallic lithium to obtain lithium
carbide for acetylene synthesis; acetylene was purified
and used to synthesize benzene, which was refined by
sublimation and supplemented with scintillators PPO
and POPOP (Arslanov, 1987). Measurements of
14
N
radioactivity were performed at the Institute of Geogra-
phy, Russian Academy of Sciences (IGAN), with liquid
scintillation
β
-spectrometers IGAN models 1–4 (Rus-
sia), Mark II (Nuclear Chicago, United States), and
Quantulus-1220 (LKB Wallac, Sweden), and at the
Institute of Geological Sciences, National Academy of
Sciences of Belarus (IGSB), with LKB1211, LKB1219
(Sweden), and low-background Guardian (Wallac, Fin-
land) spectrometers. Dating was also performed in
Heidelberg, where the samples were processed by the
same acid-base method but
14
N radioactivity was mea-
sured by means of
ëO
2
gas proportional counting
(Kromer and Münnich, 1992).
The dating cycle consisted in measuring specific
radioactivity of a test sample in comparison with two
reference samples, the background preparation and the

BIOLOGY BULLETIN
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CARBON ACCUMULATION IN SOILS OF FOREST 519
current standard. All the above laboratories participate
in the FIRI program (Fourth International Radiocarbon
Intercomparison), which guarantees consistency of
results. Radiocarbon ages were calibrated into calendar
years before 1950 AD (cal. yr. BP) with the Calib Rev
5.1 program (Stuiver and Reimer, 1993;
http://calib.qub.ac.uk/calib/calib.html).
The rates of organic matter decomposition were
estimated titrimetrically from
ëO
2
production at a soil
moisture of about 80% of the maximum water capacity,
at
20°ë
(Dobrovol’skii et al., 1999).
RESULTS AND DISCUSSION
The data on soil age and carbon stocks accumulation
rates (Table 1) and coefficients of organic matter min-
eralization (Table 2) characterize the main groups of
soil types including drained upland soils, peaty gley
and peaty soils of paludified forests, and true bog soils
differing in trophic capacity, including those of upland,
transitional, and lowland bogs
Drained upland soils were studied in typical habitats
on drained slopes of eskers and hills (plots 1 and 2):
these were secondary soddy podzolic soils in relict ara-
ble fields and brown soils under naturally developing
primary spruce forests of nemoral and nemoral–wood
sorrel types. The
14
C age of deep soddy podzolic soil at
the boundary of the litter and mineral horizon was esti-
mated at
108
±
30
yr. BP, and that of the humus horizon
of brown soil, at
317
±
90
yr. BP (Table 1).
The young radiocarbon age (317 yr. BP) of humic
acids (HAs), the most “mature” component of the soil
humus, in the upper Ahf horizon of brown soil may
reflect the time elapsed since the onset of the last
postwindfall succession. Alternatively, it may be
explained by continuous “renewal” of HA fragments
due to leaching of water-soluble organic compounds
and their input by soil mesofauna. The recent concept
of humic substances as a supramolecular association of
relatively low-molecular-weight compounds whose
stability is accounted for by noncovalent interactions
(Piccolo, 2002) agrees with the high rate of humus
renewal better than the classic concept that humus is an
association of high-molecular-weight macromolecules.
The lower part of the forest litter subhorizon H in
white podzolic (peat-enriched surface-gley podzolic)
soils was dated
169
±
10
yr. BP (plot 3). The radiocar-
bon age of the lower layers of peaty gley soils at a depth
of 30-35 cm (plot 4) was only 106-184 yr. BP, which
could be attributed to “rejuvenation” of humus sub-
stances in the lower horizon, e.g., due to water-soluble
organic matter leaching out of upper horizons. In the
upland peat soil, the age of the bottom horizon of at a
depth of 55 cm reached
898
±
400
yr. BP.
In the peaty gley soil, the age of bottom layers at a
depth of 50 cm was
605
±
180
yr. BP (pit 5.1). In the
gley podzolic soil, the age of these layers was about 190
yr. BP (pit 5.2). The bottom layers of upland bog soil
were dated
1969
±
200
yr. cal BP at a depth of 95 cm
(pit 5.3) and
853
±
50
yr. BP at a depth of 60 cm (pit
5.4).
In oligotrophic sites of the transitional bog (plot 6),
the age of the deepest peat layers (125 cm) reached
4.3 kyr. BP. In the massive upland bog (plot 7), the age
of bottom layers gradually increased with depth of peat
deposit, reaching approximately 10 kyr. BP at 4 m.
In lowland peat soils of paludified forests (plot 8),
the age of bottom peat layers also increased with depth
of the deposit, reaching almost 10 kyr. BP at 160 cm. It
is noteworthy that strongly decomposed dense peat pre-
vailed in this deposit. Its decomposition could be pro-
moted by changes in the hydrologic regime of the river
and in moistening conditions that were favorable for the
mixing of peat layers due to activities of mesofauna and
small mammals.
Upland forest biogeocenoses have the minimum
carbon stocks accumulated in their mineral soils
(Fig. 2). Carbon stocks in the humus horizons of soddy
podzolic and brown soils under nemoral wood sorrel
10
8
6
4
2
0
140
120
100
80
60
40
20
0
400
350
300
250
200
150
100
50
0
1235.2
4.2
4.35.1
4.1
5.3
5.4 67.17.2
7.4
7.5
7.68.18.2
1235.24.2
4.35.1
4.15.3
5.4 67.17.2
7.47.57.68.18.2
1
2
3
5.2
4.2
4.3
5.1
4.1
5.3
5.4
6
7.1
7.2
7.4
7.5
7.6
8.1
8.2
(a)
(b)
(c)
1000 yr. BP
kg C/m
2
g C/m
2
per year
1
2
3
4
5
Fig. 2.
(a) Calibrated
14
C age, (b) carbon stocks, and
(b) average rate of carbon accumulation in soils over the
period of their existence. Abscissa shows soil pit numbers.
Designations: (
1
) mineral soils (SP, soddy podzolic; B,
brown; P, podzolic); (
2
) peaty podzolic soils; (
3
) peaty gley
soils; (
4
) upland peat soils; (
5
) lowland peat soils.

520
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MINAYEVA et al.
spruce forests are 2 and 4 kg N/m
2
, respectively. In
white podzolic (peat-enriched surface-gley podzolic)
soils under true moss–bilberry and sphagnum–bilberry
spruce forests, the carbon stock increases to or exceeds
6 kg N/m
2
. With respect to moistening conditions, these
soils are closer to paludified soils. Unlike in soddy pod-
Table 1.
Results of estimating soil age and carbon stock and accumulation rate in test plots
Plot, horizon/core Depth, cm Radiocarbon
age, nal. yr. BP
Index* and labora-
tory number
Carbon stock,
g/m
2
Average carbon accumula-
tion, g/m
2
per year
Soddy podzolic soil
Plot 1, horizon AO 6 108
±
30 IGAN-1731 2160 22
Brown soil
Plot 2, horizon Ahf 13 317
±
90 IGAN-1835 4090** 14
White podzolic soil
Plot 3, horizon H 12 169
±
10 IGAN-1733 6450 38
Peaty podzolic gley soil
Plot 5, core 5.2 40 192
±
40 IGSB-309 51500 280
Peaty gley soil
Plot 4, core 4.2 35 184
±
40 IGSB-303 15200 87
Plot 4, core 4.3 30 106
±
40 IGAN-1777 19 500 214
Plot 5, core 5.1 50 605
±
180 IGSB-308 32 200 58
Upland peat soil
Plot 4, core 4.1 55 898
±
400 IGSB–302 20900 29
Plot 5, core 5.3 95 1969
±
200 IGSB-306 24 260 12
Plot 5, core 5.4 60 853
±
50 IGSB-305 23850 28
Plot 6 42 1124
±
230 IGSB-273 27490 26
90 2496
±
330 IGSB-269 51 380 21
95 2692
±
190 IGSB-272 54 390 20
125 4380
±
170 IGSB-268 62 690*** 14
Plot 7, core 7.1 50 441 ± 10 IGSB-310 21370 48
Plot 7, core 7.2 120 950 ± 100 IGSB-311 27260 29
Plot 7, core 7.3 230 5628 ± 26 Hd-20478 No data No data
Plot 7, core 7.4 375 10206 ± 70 Hd-20479 100 700 10
Plot 7, core 7.5 390 9959 ± 24 Hd-20472 129050 13
Plot 7, core 7.6 380 9218 ± 24 Hd-20689 64 870 7
íLowland peat soil
Plot 8, core. 8.1 50 1958 ± 90 IGAN-1730 31500 16
Plot 8, core 8.2 50 1672 ± 75 IGAN-1776 18000 11
100 5589 ± 200 IGAN-1784 38 390 7
160 9800 ± 100 Hd-20480 51 100 5
Notes: * IGAN, Institute of Geography, Russian Academy of Sciences; IGSB, Institute of Geological Sciences, National Academy of Sci-
ences of Belarus; Hd, Heidelberg Academy of Sciences, Germany.
** Including 1940 g N/m2 in organogenic horizons and 2150 g N/m2 in the Ahf horizon.
*** For a 110-cm layer.

BIOLOGY BULLETIN Vol. 35 No. 5 2008
CARBON ACCUMULATION IN SOILS OF FOREST 521
zolic and brown soils, the coefficients of organic matter
mineralization in white podzolic soils are lower and
decrease more abruptly down the soil profile. Their car-
bon stock increases along the gradient of soil moisten-
ing. In peaty gley podzolic soils with better developed
organogenic horizons (30 cm or deeper), the carbon
stock increases by an order of magnitude and exceeds
50 kg C/m2. This value is even greater than those
recorded in peaty gley and even upland peat bog soils
with peat horizons deeper than 30 cm.
The carbon stock of bog biogeocenoses progres-
sively increases along with increase in peat deposit
depth. In most sites of upland, transitional, and lowland
bogs, the carbon stock of paludified soils was 20-30 kg
C/m2; only in single cases, when the peat layer was
deeper than 1 m, it exceeded 50 kg C/m2. Thus,
paludified and surface-peat forest–bog biogeocenoses
have similar carbon stocks. With peat deposit depth
being the same, the carbon stock in lowland bogs is sig-
nificantly greater than in upland bogs. Dense carbon
“packing” in well-decomposed and structurally homo-
geneous lowland deposits accounts for its considerable
amounts stored per unit area. Carbon stocks increase
severalfold in bogs where the depth of peat deposits
reaches several meters. In the massive upland bog with
a peat depth about 4 m, soil carbon stocks in some sites
exceeded 100 kg C/m2.
Our data on carbon stocks in humus and peat hori-
zons in the contiguous series of mineral, peat-enriched,
and peat bog soils indicate that paludified and bog bio-
geocenoses play a major role in the formation of soil
carbon stocks. They confirm the general conclusion
that bogs and paludified habitats of the southern taiga
subzone are rich in carbon due to the presence of peat
deposits, which function as a long-term accumulator of
atmospheric carbon.
An explanation for differences between the rates of
carbon accumulation in mineral, paludified, and bog
soils is that these systems differ in the degree of open-
ness and, therefore, in the intensity of carbon exchange
over the period of existence of their humus and peat
horizons. No less significant are differences in bio-
geochemical processes developing in the lower hori-
zons (permanently saturated with water) and the upper,
periodically undersaturated horizons of bog soils. The
rate of organic matter decomposition in the latter, under
aerobic conditions, is approximately an order of magni-
tude higher than that in permanently moistened layers
of peat deposits, where conditions are almost anaero-
bic. This is why we limited ourselves to a general com-
parison of carbon accumulation in soils of different
habitats, relying on integrated estimations over the
entire period of their existence (calculated from the age
of bottom layers) and assuming that the dynamics of
organic matter destruction in the soil profile are linear.
Our calculations have shown that the average rates
of carbon accumulation in humus horizons of soddy
podzolic and brown soils are 22 and 14 g N/m2 per year,
respectively. These rates are markedly lower than those
for the forest litter at windfall sites, which may reach 50
g N/m2 per year (Bobrov et al., 1997; Dobrovol’skii et
al., 1999). Assuming that carbon accumulation rates
tend to slow down with time, tending to zero under
steady-state conditions of soil functioning, the result of
these calculations may be regarded as an integrated
index of peat accumulation, with the carbon accumula-
tion rate being high in the first decades of organic pro-
file formation and relatively low at subsequent stages.
It may well be that this is a normal process of humus
horizon restoration in the course of normal forest
regeneration, with the calculated rates of carbon accu-
Table 2. Coefficients of organic matter mineralization in
soils of test plots
Object Coefficient of mineralization
(Cm)
Horizon Depth, cm % Norg
per day
Norg
per month
Soddy podzolic soil, plot 1
L 0–1.5 0.817 24.51
F 1.5–4 0.444 13.32
H 4–6 0.138 4.14
Brown soil, plot 2
L 0–1 0.562 16.86
F 1–3 0.221 6.63
AO 3–5 0.203 6.09
Ahf 5–12 <0.008 <0.25
Peaty white podzolic soil, plot 3
L 0–2 0.427 12.81
F 2–5 0.116 3.48
H 5–11 0.067 2.01
A2h 11–17 <0.008 < 0.25
Upland peat soil, plot 5
T1 0–6 0.006 0.18
T2 12–16 0.007 0.21
T3 35–40 0.009 0.27
Lowland peat soil, plot 8
L 0–0.5 1.085 32.55
F 0.5–3 0.718 21.54
T1 3–5 0.048 1.44
T2 70–80 0.023 0.69
T3 140–150 0.086 2.58

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MINAYEVA et al.
mulation reflecting the actual processes taking place
during this period.
The calculated average rates of carbon accumula-
tion in paludified habitats proved to the highest, varying
from 60 to 210 g C/m2 per year for peaty gley soil and
reaching 280 g N/m2 per year for peaty gley podzolic
soils. High variation in the group of peaty soils may be
explained by the impact of fires, which are frequent in
paludified forests. In particular, this follows from the
presence of soil layers with increased ash contents or,
in some cases, charcoal layers immediately overlying
mineral strata.
The calculated average rates of carbon accumula-
tion by peat in bogs proved to be very low, especially in
the lowland bog. This is not surprising, since this bog is
abundantly supplied with ground waters rich in mineral
substances and its location of the slope provides for a
certain degree of drainage and, consequently, surface
water saturation with oxygen. As a result, mineraliza-
tion proceeds more actively, a large proportion of the
aboveground phytomass is decomposed, and the coeffi-
cient of mineralization (Cm) is higher than in other soil
types studied (Table 2).
Comparing the data on soil pits in different parts of
the massive upland bog, is can be seen that the calcu-
lated average rate of carbon accumulation increases
from 7 g C/m2 per year at the margin to 48 g C/m2 per
year in the center of the bog. A probable reason is that
the bottom peat layers of differing age and depth per-
tain to paleoclimatic intervals with different conditions
for peat accumulation, which is true both of Eurasia as
a whole (Klimanov and Sirin, 1997) and of the study
region (Klimenko et al., 2003; Kozharinov et al., 2003;
Minayeva et al., 2006). It is also possible that the mar-
ginal areas of bogs accumulate carbon at higher rates
because of accelerated accumulation of peat. In this
respect, bog margins are somewhat similar to paludified
and surface-peat habitats, where the rates of peat and
carbon accumulation also reach the highest values. In
paludified forests and surface-peat forest bogs, peat
deposits are often affected by forest fires and other
stress factors usually associated with cyclic changes in
climate and, therefore, in the water regime (Minayeva
et al., 2004). In favorable periods, this is counterbal-
anced by more intensive peat accumulation, and it is not
accidental that the highest increments of sphagnum
mosses are observed in such areas (Minayeva and Star-
odubtseva, 1997).
Mineral and bog soils have similar average rated of
carbon accumulation over the periods of their exist-
ence, which means that corresponding ecosystems have
reached a certain steady state in terms of structure and
functioning. Exceptions are marginal areas of bogs,
paludified forests, and, probably, areas disturbed by
windfall or in other ways. Thus, throughout the catena,
in both hydromorphic and automorphic habitats, accu-
mulation of carbon has prevailed over its loss during the
past several centuries.
CONCLUSIONS
The results of this study show that carbon stocks in
mineral soils are many times lower than in paludified
soils and about an order of magnitude lower than in bog
soils. The carbon stock of soddy podzolic soils and
brown soils under nemoral wood sorrel spruce forests
does not exceed 2–4 kg N/m2; that of peat-enriched
white podzolic soils under true moss and sphagnum–
bilberry spruce forests is 6 kg N/m2. Soil carbon stocks
in habitats with more abundant moisture supply sharply
increase, reaching 50 kg N/m2 in paludified forests and
more than 100 kg N/m2 in bog biogeocenoses.
Assuming that peat accumulation is a linear process,
its average rates in mineral and bog soils over the
period of their existence appear to be fairly similar. Car-
bon is accumulated most actively in soils of paludified
habitats, but this process is periodically disturbed by
fires and other stress influences. Bogged soils counter-
balance these disturbances by the maximum rates of
peat and carbon accumulation in favorable periods. In
massive bogs, the highest parameters of carbon accu-
mulation are characteristic of marginal areas account-
ing for the lateral growth of the bog.
These results provide evidence for differences in the
functioning of forest–bog soils as carbon accumulators.
Analyzing carbon deposition in taiga ecosystems, it is
expedient to consider the whole range of constituent
soil types, including mineral, paludified, and bog soils.
such a comprehensive analysis appears promising, as it
can reveal general and specific trends in organization of
these soils and provide a new outlook on their role in
carbon turnover.
ACKNOWLEDGMENTS
The authors are grateful to O.A. Starodubtseva
(Institute of Forest Science, Russian Academy of Sci-
ences) and to T.B. Men’shikh (Moscow State Univer-
sity) for their help in the collection and processing of
field material.
This study was supported by the program “Biodiver-
sity and Gene Pool Dynamics” of the Presidium of the
Russian Academy of Sciences, subprogram “State and
Resource-Ecological Potential of Terrestrial Ecosys-
tems under Conditions of Global Changes in Northern
Eurasia.”
REFERENCES
Arinushkina, E.V., Rukovodstvo po khimicheskomu analizu
pochv (A Manual of Chemical Soil Analysis), Moscow:
Mosk. Gos. Univ., 1970.
Arslanov, Kh.A., Radiouglerod: geokhimiya i geokhro-
nologiya (Radiocarbon: Geochemistry and Geochronology),
Leningrad: Leningr. Gos. Univ., 1987.
Bobrov, A.A., Korobov, E.D., Trofimov, S.Ya., et al,. Effect
of Windfall Damage on the Structure of Soil Meso- and
Nanofauna: An Example of Spruce Forests in the Central

BIOLOGY BULLETIN Vol. 35 No. 5 2008
CARBON ACCUMULATION IN SOILS OF FOREST 523
Forest State Biosphere Reserve, Vestn. Mosk. Gos. Univ., Ser.
17: Pochvoved.,, 1997, no. 3, pp. 41–48.
Central Forest Biosphere Reserve, in Suktsessionnye prot-
sessy v zapovednikakh Rossii i problemy sokhraneniya bio-
logicheskogo raznoobraziya (Successional Processes and
Problems of Biological Diversity Conservation in Russian
Nature Reserves), Smirnova, O.V. and Shaposnikov, E.S.,
Eds., St. Petersburg: RBO, 1999, pp. 296–419.
Chichagova, O.A., Radiouglerodnoe datirovanie gumusa
pochv (Radiocarbon Dating of Soil Humus), Moscow,
Nauka, 1985.
Chichagova, O.A., Development of I.P. Gerasimov’s Ideals
in the Field of Radiocarbon Research, in: Mnogolikaya
geografiya. Razvitie idei Innokentiya Petrovicha Gerasi-
mova (K 100-letiyu so dnya rozhdeniya) [Multifaced Geog-
raphy: Development of I.P. Gerasimov’s Ideas (on the 100th
Anniversary of His Birth)], Moscow: KMK, 2005, pp. 163–
180.
Chichagova, O.A., Goryachkin, S.V., Aleksandrovskii, A.L.,
and Kovda, I.V., Radiocarbon Research As a Basis for
Assessing Carbon Flows in the Soil–Atmosphere System,
Izv. Akad. Nauk, Ser. Goeor., 2001, no. 4, pp. 1–7.
Dobrovol’skii, G.V., Trofimov, S.Ya., Dorofeeva, E.I., et al.,
The Rate of Litter Decomposition in Southern Taiga Spruce
Forests, Lesovedenie, 1999, no. 1, pp. 3–9.
Genezis i ekologiya pochv Tsentral’no-lesnogo zapovednika
(Genesis and Ecology of Soils in the Central Forest Nature
Reserve), Moscow, Nauka, 1979.
Karimov, A.E. and Nosova, M.B., Land Use and Impact on
Nature in the Central Forest Nature Reserve between the late
16th and early 20th centuries, in Suktsessionnye protsessy v
zapovednikakh Rossii i problemy sokhraneniya biolog-
icheskogo raznoobraziya (Successional Processes and Prob-
lems of Biological Diversity Conservation in Russian Nature
Reserves), St. Petersburg: RBO, 1999, pp. 299–310.
Klimanov, V.A., Sirin, A.A., Peat Accumulation Dynamics in
Bogs of Northern Eurasia over the Past 3000 Years, Dokl.
Akad. Nauk,. 1997, vol. 354, no. 5, pp. 683–686.
Klimenko, V.V., Klimanov, V.A., Sirin, A.A., et al., Climatic
Changes in the West of European Russia during the Late
Holocene, Dokl. Akad. Nauk, 2001, vol. 376, no. 5, pp. 679–
683.
Kozharinov, A.V., Sirin, A.A., Klimenko, V.V. et al.,
Dynamic of Vegetation and Climate in the Western Dvina
Lowland (Tver Region) over the Past 5000 Years, Bot. Zh.,
2003, vol. 88, no. 3, pp. 90–102.
Kromer, B., Münnich, K.-O., CO2 Gas Proportional Count-
ing in Radiocarbon Dating: Review and Perspective, in
Radiocarbon after Four Decades, Taylor, R.E., Long, A., and
Kra, R.S., Eds., New York: Springer, 1992, pp. 184–197.
Minayeva, T.Yu., Experience in Studies on Forest Phyto-
cenoses in Permanent Test Plots in the Central Forest Nature
Reserve, in Opyt issledovaniya rastitel’nykh soobstchestv v
zapovednikakh (Experience in Studies on Plant Communities
in Nature Reserves), Moscow, Glavokhota, 1988, pp. 130–
146.
Minayeva T. and Glushkov, I., Ground Water Regime in
Paludified Forests in Central European Russia: Application
to Ecosystem Development and Dynamics, in Ecohydrologi-
cal Processes in Northern Wetlands: Selected Papers, Tartu,
2003, pp. 166–171.
Minayeva, T.Yu. and Starodubtseva, O.A., Evaluation of
Sphagnum Growth in Spruce Forest Communities of
Paludified Series, Tezisy VI Molodezhnoy konferentsii botan-
ikov v S.-Peterburge 12–16 maya 1997 g. (Abstr. VI Young
Botanists Conf., St. Petersburg, May 12–16, 1997), St.
Petersburg, 1997, p. 83.
Minayeva. T.Yu., Istomin. A.V., Abrazhko. V.I.. et al., On the
Response of Biota to Climate Change in the Central Forest
Nature Reserve, in Vliyanie izmeneniya klimata na eko-
sistemy. Seriya publikatsiy Departamenta Prirodookhrannoy
politiki i ekspertizy Vsemirnogo fonda dikoi prirody (Effect
of Climate Change on Ecosystems. Publications of the World
Wildlife Foundation Department of Nature Conservation
Policy and Expert Assessment), Moscow: Russkii Univer-
sitet, 2001, issue 4, pp. 87–100.
Minayeva, T., Glushkov, I., Sulerzhicky, L., et al., On Tem-
poral Aspects of Shallow Peat Accumulation in Boreal
Paludified Forests: Data from Case Studies in Central Euro-
pean Russia, Wise Use of Peatlands: Proc. 12th Int. Peat
Congr., Saarijarvi: Saarijarven Offset Oy, 2004, vol. 1,
pp. 150–155.
Minayeva T., Bleuten, W., Sirin, A. et al., Eurasian Mires of
the Southern Taiga Belt: Modern Features and Response to
Holocene Paleoclimate, in Wetlands and Natural Resource
Management, Berlin: Springer, 2006, vol. 190, pp. 315–341.
Piccolo, A., The Supramolecular Structure of Humic Sub-
stances: A Novel Understanding of Humus Chemistry and
Implications in Soil Science, Adv. Agron., 2002, vol. 75, pp.
57–134.
Regulyatornaya rol' pochvy v funktsionirovanii taezhnykh
ekosistem (Regulatory Role of Soils in the Functioning of
Taiga Ecosystems), Dobrovol’skii, G.V., Ed., Moscow:
Nauka, 2002.
Shaposhnikov, E.S., Associations of Spruce Forests in the
Central Forest Nature Reserve, Extended Abstract of Cand.
Sci. (Biol.) Dissertation, Leningrad: Bot. Inst., USSR Acad.
Sci., 1988.
Shaposhnikov, E.S., Yushchenkova, L.N., and Minayeva,
T.Yu., Changes in the Community of Cotton Grass–Sedge–
Sphagnum Spruce Forest Resulting from Progressing
Paludification, Bolota okhranyaemykh territorii. Tezisy
dokladov XI seminara-ekskursii po bolotovedeniyu @ (Bogs
in Protected Areas. Proc. XI Field Seminar on Peatland Sci-
ence), Boch, M.S., Ed., Leningrad: Ross. Bot. Obshch.,
1991, pp. 73–77.
Shaposhnikov, E.S., Kennel, M., and Minajewa, T.J., Die
Gemeinsambeschreibung der Struktur des Pflanzenreiches in
der Bestandesmessflaeche der Waldklimastation: Demon-
strationsvorhaben Waldklimastation im staatlichen Russis-
chen Biosphaerenschutzgebiet zentrales Waldreservat, Freis-
ing, 1994, pp. 82–98.
Stuiver, M. and Reimer, P.J., Extended 14C Database and
Revised CALIB Radio-Carbon Calibration Program, Radio-
carbon, 1993, vol. 35, pp. 215–230.
Torfyanye bolota Rossii: k analizu otraslevoi informatsii
(Peat Bogs of Russia: On Analysis of Departmental Informa-
tion), Sirin, A.A. and Minayeva, T.Yu., Eds., Moscow, Geos,
2001. 190 p.
Trofimov, S.Ya., Soil Functioning in Undisturbed Southern
Taiga Biogeocenoses, Extended Abstract of Doctoral (Biol.)
Dissertation, Moscow, Moscow State Univ., 1998.

524
BIOLOGY BULLETIN Vol. 35 No. 5 2008
MINAYEVA et al.
Trofimov, S.Ya. and Stroganova, M.N., Specific Features of
Soil Formation on Drained Slopes in Undisturbed Southern
Taiga Biogeocenoses, Vestn. Mosk. Gos. Univ., Ser. 17:
Pochvoved., 1991, no. 3, pp. 8–16.
Trofimov, S.Ya., Goncharuk, N.Yu., and Dorofeeva, E.I., Soil
Carbon Stocks in Undisturbed Southern Taiga Biogeo-
cenoses: An Example of Central Forest Nature Reserve,
Pochvovedenie, 1997, no. 10, pp. 1211–1216.
Trofimov, S.Ya., Men’shikh, T.B., Dorofeeva, E.I., Gon-
charuk, N.Yu., Stocks of Organic Matter and Rates of Its
Mineralization in Soils of Paludified Spruce Forests in the
Central Forest State Biosphere Reserve, Pochvovedenie,
1998, no. 4, pp. 416–422.
Vomperskii, S.E., Ivanov, A.I., Tsyganova, O.P., et al., Rus-
sia’s Paludified Organogenic Soils and Bogs and Their Peat
Carbon Stocks, Pochvovedenie, 1994, no. 12, pp. 17–25.
Vomperskii, S.E., Sirin, A.A., Tsyganova, O.P., et al., Bogs
and Paludified Lands in Russia: An Attempt at Analysis of
Spatial Distribution and Diversity, Izv. Akad. Nauk, Ser.
Geogr., 2005, no. 5, pp. 21–33.
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