Ecological Modelling 251 (2013) 232–245

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Ecological Modelling
journal homepage: www.elsevier.com/locate/ecolmodel

Carbon turnover in mixed stands: Modelling possible shifts under climate change
Vladimir Shanin a,∗ , Alexander Komarov a , Yulia Khoraskina a , Sergey Bykhovets a , Tapio Linkosalo b ,
Raisa Mäkipää b
a
Institute of Physicochemical and Biological Problems in Soil Science of the Russian Academy of Sciences, 142290 Institutskaya ul., 2, Pushchino, Moscow Region, Russian Federation
b
Finnish Forest Research Institute (Metla), Jokiniemenkuja 1, BOX 18, FI-01301, Vantaa, Finland

a r t i c l e i n f o a b s t r a c t

Article history: Adaptation of forest management to changing climate may lead to remarkable changes in tree species
Received 3 September 2012 composition. Deciduous species are predicted to be favoured in a warmer climate, but the competitive
Received in revised form 7 December 2012 capacity of different tree species requires detailed analysis. The main objectives of this study were to
Accepted 10 December 2012
assess how varied initial proportions of tree species affected productivity in mixed stands, and to eval-
Available online 23 January 2013
uate the biomass production and carbon sequestration potential of trees and soil in mixed stands in a
changing climate. We used the individual-based simulation model EFIMOD combined with the model of
Keywords:
soil organic matter dynamics ROMUL to simulate spatially explicit competition between trees for light
Carbon balance
Boreal forests
and nutrients. We focused our simulations on the most common type of site (mesic site, Myrtillus type) in
Competition southern Finland. The average carbon and nitrogen pools in the forest floor and mineral soil were used as
Resource use initial soil data. The ambient climate scenario was based on measured data (1961–2007) from the Finnish
Soil nutrients Meteorological Institute. Climate change scenarios were based on runs of three general circulation mod-
els. We simulated the forest stand dynamics of two- and three-species mixtures (Silver birch, Scots pine
and Norway spruce) with different initial proportions. In mixed spruce–birch stands, spruce replaced
birch whatever the initial density, regardless of climatic scenario. In the beginning of the development
of mixed spruce–pine stands, pine grew faster and increased its proportion of biomass by 10–20%. In
pine–birch stands, the dominant species maintained its dominating position, but the proportion of pine
generally declined by the end of simulation in birch-dominated stands. In mixed stands of three tree
species, spruce tended to dominate by the end of a 100-year simulation period, independently of the
initial proportions. The highest average carbon stock in standing biomass was observed in mixed stands
with three species and in spruce-dominated stands. Climate change increased stand productivity and the
increase in coniferous stands was more remarkable than in birch-dominated stands. In addition, soil was
affected by species composition: increased proportions of pine resulted in increasing carbon stocks in
the forest floor. Climate change negatively affected accumulation of organic matter in soil, especially in
the spruce-dominated stands.
© 2013 Elsevier B.V. All rights reserved.

1. Introduction locations of climatic optima for plant species. Species having

Climate change is predicted to result in increased primary
production in boreal and temperate forests, changes in spring
phenological events and major shifts in the geographical ranges
of species (e.g. Linkosalo et al., 2009; Lousteau et al., 2005;
Nabuurs et al., 2002; Thuiller, 2007). The geographical ranges of
temperature-limited species may move northward by some 500 km
over the next 100 years due to the predicted 4 ◦ C increase in the
mean annual temperature and associated alterations in precipi-
tation patterns (Thuiller, 2007). Such environmental changes can
be considered as a slow and long-time shift of the geographical
∗ Corresponding author. Tel.: +7 9267240651; fax: +7 4967330595.
E-mail address: shaninvn@gmail.com (V. Shanin).
specific ecological niches will have differentiated responses to pre-
dicted climate change and related changes in resource availability
(Araújo and Guisan, 2006; Austin et al., 2009). In a warmer climate,
the productivity of tree species will respond differently to altered
environmental conditions, which may also lead to changes in which
tree species are dominant (e.g. Kellomäki et al., 2008; Lousteau
et al., 2005). The replacement of coniferous by broadleaved species
has been demonstrated using both paleoecological (Overpeck et al.,
1991) and simulation (Iverson and Prasad, 1998, 2001; Sykes et al.,
1996) methods. The majority of European forests are managed, and
tree species composition is to a large extent manipulated by silvi-
culture. Although management of boreal forests is often targeted
at monocultures of a desired species, forests develop as a mixture
of two or more tree species, especially during the early phases of
natural stand development. If tree species composition is driven

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 V. Shanin et al. / Ecological Modelling 251 (2013) 232–245
by changing climatic conditions, as predicted by Kellomäki et al.
(2008), relative changes in the productivity of different tree species
and the following shift in species composition will most directly
occur in mixed stands with interspecific competition. Moreover,
changes in trees productivity and species composition lead to
changes in forest soils. Thus, analysis of the development of mixed
stands is a necessary step forward in understanding the overall
effects of climate change on forest growth and carbon sequestration
in forest soils.
Studies on mixed stands have shown that growth and yield of
competing tree species, as well as yield of the entire stand, are
affected by the proportions of tree species (e.g. Hynynen et al.,
2011; Jõgiste, 1998; Mielikäinen, 1980, 1985). In general, ecological
theory predicts that a mixture of coexisting species that have com-
plementary niches will lead to a higher total use of resources and
higher biomass production (Leps, 2005; Loreau and Hector, 2001).
This seems also to be the case in forest stands with spruce–birch
mixtures (Mielikäinen, 1985). An increase in productivity with
species richness is predicted to be a general pattern in temperate
forests (Morin et al., 2011). Evidently at accounting of interspecies
competition we must include in the modelling framework key
controlling biophysical processes. Use of simple growth and yield
models that do not consider changing conditions are ill equipped to
support the needed studies even though they are based on some-
times huge amounts of empirical data.
Some attempts have been made to evaluate the competitive
effects of different species using methods based on both individual-
based modelling with competition indices (Zhao et al., 2006) and
direct measurement of intra- and interspecific competition (Coates
et al., 2009). Both of these studies evaluate the competitive effects
of several tree species and reveal such effects to be unequal and
asymmetric. Ge et al. (2011) studied the responses of two Norway
spruce-dominated stands with an admixture of pine and birch to
changing climate using a process-based model. Their results indi-
cate that the growth of mixed spruce–birch stands could be higher
than that of pure stands under water stress. However, the results
are very dependent on the mechanisms of interspecific competi-
tion, which are not clearly described in their study.
We simulated the growth of mixtures of Scots pine (Pinus
sylvestris L.), Norway spruce (Picea abies (L.) H. Karst.) and Silver
birch (Betula pendula Roth) using a process-based model EFIMOD
that describes mechanisms of competition between tree species
(Komarov et al., 2003) and accounts carbon and nitrogen turnover
in forest–soil system. In simulating mixed stands, calculation of
interspecies competition for light and nutrients is the main process
determining forest growth, soil changes, succession and dominance
of tree species. Since the predicted increase in temperature and
precipitation greatly affects the rate of soil processes and nutrient
availability, modelling of the soil processes in the analysis of the
responses of mixed stands to a changing climate is crucial.
The overall objective of this study was to evaluate the biomass
production and carbon sequestration potential of mixed stands in
both the ambient and changing climate.
Two questions to be answered by this study were: (1) how ini-
tial species composition affects the dynamics of carbon stock in
standing biomass and soil, and (2) how different species mixtures
react to climate change in terms of species composition, biomass
production and carbon dynamics.
We hypothesized that productivity will increase with increas-
ing species richness, but this will depend on the actual proportions
of competing tree species. Furthermore, we hypothesized that the
proportion will change towards dominance of coniferous species
in a changing climate scenario. The sequestration of carbon into
standing biomass is expected to increase with climate change due
to the sensitivity of productivity to the amount of available nitro-
gen. We suppose that in the changed climate, soil carbon stocks will
233
be lower due to an accelerated rate of decomposition. The overall
effect of these processes cannot be estimated without simulation
experiments.
2. Material and methods
2.1. Simulation of stand and soil dynamics
In our study, the model of the forest–soil system EFIMOD
(Komarov et al., 2003) was used to simulate the dynamics of a mixed
forest. EFIMOD describes population-level dynamics consisting of
a set of separate trees competing for light and soil nutrients with
neighbour trees. This approach makes it possible to trace the devel-
opment of forest stands, including tree mortality.
The specific features of EFIMOD are an emphasis on the impor-
tance of soil processes for ecosystem functioning and a strong
feedback mechanism between soil and vegetation. EFIMOD simu-
lates mixed forest stands consisting of forest elements (each forest
element is an even-aged and single-species cohort of trees). EFI-
MOD can simulate different types of cuttings (Mikhailov et al.,
2004; Palosuo et al., 2008). It is sensitive to climate changes and
nitrogen deposition (Kahle et al., 2008), and can simulate forest
fires and windfalls (Komarov and Kubasova, 2007).
The system of models contains four main sub-models: (1) a
model of individual tree growth; (2) a spatial stand model; (3)
the model of soil organic matter dynamics ROMUL (Chertov et al.,
2001); and (4) the statistical climate generator SCLISS (Bykhovets
and Komarov, 2002).
A detailed description of the system of models EFIMOD has been
published previously (Komarov et al., 2003). We are describing here
the mechanisms of competition in more detail, because they are
the main drivers for simulation of a mixed forest. For each tree,
two competition indices were calculated. Firstly, shading was cal-
culated as an additive sum of overlapping neighbouring crowns. A
single overlap is characterized by a coefficient of absorption, which
is related to crown opacity. For spruce, this coefficient is equal 0.22,
0.12 for birch and 0.11 for pine (Tselniker, 1978).
The second competition index is a redistribution of mineral
nitrogen which each tree collects from the area occupied by its root
system. Each tree is rooted in a homogeneous cell of size 50 × 50 cm.
During growth, a tree expands its area of nutrition (as well as its
shadowing zone) onto neighbouring cells. The area of nutrition is a
square, the side of which has an odd-numbered length (1 × 1, 3 × 3,
5 × 5 etc.). The size of the area of nutrition increases with increas-
ing tree height. An example of the relationship between tree height
and size of nutrition area is shown in Appendix A, Table A.1.
The account of the shadowing coefficient KE includes a calcula-
tion of local density for each tree. The concept of local density is
similar to the calculation of indices of competition (Pukkala, 1989;
Tomppo, 1986).
The procedure for calculating the shadowing coefficient in the
case of mixed stands takes into account that neighbours of a focal
tree may belong to different species. Models of competition are
described in more detail by Komarov et al. (2003) and in Appendix
A.
The soil sub-model ROMUL (Chertov et al., 2001) describes
biological turnover of nutrients. It simultaneously allows for cal-
culation of the rate of mineralization and humification of tree litter
and soil organic matter with the corresponding carbon dioxide and
nitrogen release for plant growth.
The theoretical background to the model is the concept of humus
types and biota functioning. The experimental basis of the ROMUL
model is laboratory experiments (Chertov and Komarov, 1997;
Chertov et al., 2001; Komarov et al., 2007) in controlled temper-
ature and moisture conditions.
234 V. Shanin et al. / Ecological Modelling 251 (2013) 232–245
The model has pools of above- and below-ground litter (decom-
position rates are calculated for them with different temperature
and moisture dependencies) and a cascade of soil organic mat-
ter (SOM) fractions: undecomposed litters, partly humified organic
materials (forest floor, peat and so-called ‘labile’ humus in mineral
topsoil) and humus bonded with the mineral matrix of the topsoil.
The litter input in the model can be represented by an unlimited
number of cohorts which have species-specific ash and nitrogen
contents. There are three processes of SOM humification by three
communities of microorganisms and soil fauna (without consider-
ation of their biomass) and three processes of mineralization. The
model is represented by a system of ordinary differential equations
with coefficients that depend on soil temperature and moisture, the
nitrogen and ash content in litter, and on the C/N-ratio in the min-
eral topsoil. The model also considers nitrogen release and gross
carbon dioxide emission from the soil.
The main interdependencies between the decomposition activi-
ties of the soil fauna and climatic conditions in ROMUL are the same
for different geographic zones. ROMUL has been used for wide-
scale simulation of SOM dynamics in forest ecosystem modelling
and in different vegetation zones. The model is implemented for
the investigation of the effects of climate change on SOM dynam-
ics, soil development at primary succession, plant nutrition, carbon
balance and regional evaluation of forest soil dynamics (Mäkipää
et al., 2011; Nadporozhskaya et al., 2006; Palosuo et al., 2008,
2012; Peltoniemi et al., 2007; Rantakari et al., 2012; Shaw et al.,
2006; Yurova et al., 2010). Mathematical formulations of ROMUL
are described in detail in Chertov et al. (2001), and in Appendix B.
The EFIMOD has annual time step but the ROMUL model sim-
ulates carbon and nitrogen dynamics with monthly step. It allows
taking into account seasonality and possible monthly variability
of climatic conditions in relation to decomposition rate, which
depends on temperature and forest floor or mineral soil moisture.
The tree growth sub-model has an annual time step and there-
fore it is unable to simulate phenology though it is important
(Chuine and Beaubien, 2001). Soil sub-model which makes runs
with monthly step accounts possible changes in total amount of
available nitrogen flow due to climatic changes. These changes
can give a priority to deciduous species in a case of warming or
changes in precipitation. Nevertheless we assume that in simu-
lations of mixed forest, which consists of common boreal species
(pine, spruce and birch) with wide range of tolerance to site con-
ditions in the study region, phenology is not the main determinant
of the competitiveness of tree species. We assume that regenera-
tion rules which are described further cannot switch the species
composition because regeneration in the EFIMOD depends mostly
on light availability and fertility of soil, and all three species have
a great amount of easily disseminated seeds. Possible dependence
of regeneration on soil moisture conditions (Walck et al., 2011) did
not include in the EFIMOD because of simulating forest growth in
the mild moisture conditions typical for south taiga climatic zone.
Snow influence on regeneration (Drescher and Thomas, in press) is
also not accounted in the stand model. However, the effect of snow
cover is indirectly incorporated into the climate generator, which
governs winter forest floor and mineral soil moisture.
The input data of the system of models are: (1) species compo-
sition (mixed stand is represented as a number of forest elements
which are even-aged cohorts of trees of the same species with
similar size); (2) the main dendrometric parameters (mean stand
height, diameter at breast height, number of trees/ha) of each for-
est element; (3) carbon and nitrogen pools in organic horizons (L,
F, H) and mineral soil (A1 ); (4) soil hydrology characteristics; and
(5) monthly air temperature and precipitation. The model outputs
for each annual time step are as follows: (1) dendrometric param-
eters of each forest element, including dominant height; (2) basal
area and growing stock of each forest element or each species; (3)
deadwood; and (4) pools of carbon and nitrogen for each forest
element in the stand and carbon and nitrogen pools in organic and
mineral soil.
2.2. Climate scenarios
We simulated stand development for ambient and changing
climate conditions. The ambient climate scenario, in terms of tem-
perature and precipitation, was based on the 47-year (1961–2007)
period measured from observing stations interpolated onto a
10 × 10 km grid by the Finnish Meteorological Institute (Venäläinen
et al., 2005). In this study, we used data from the grid cell clos-
est to the Hyytiälä/Smear II station. The 47-year data series was
detrended and repeated to obtain climate data for whole 100-year
simulation period in the ambient climate model.
Climate change scenarios were based on runs of three GCMs:
Max Plank Meteorological Institute (MPMI) ECHAM5 (Roeckner,
2005a,b; Wegner, 2007a,b,c,d), United Kingdom Meteorological
Office (UKMO) HadGEM1 (Lowe, 2005a,b) and National Cen-
ter for Atmospheric Research (NCAR) CCSM3 (NCAR, 2005a,b,
2007a,b,c,d), available from the Climate and Environmental
Retrieval and Archive (CERA) database of the World Data Cen-
ter for Climate, Hamburg (WDCC; http://cera-www.dkrz.de/). The
reason for choosing these models was partly that they (especially
ECHAM5) gives a relatively good result in reproducing 20th century
climate conditions for North-western regions of Russia (Zukert,
2011), the nearest to those of Finland. In addition, we compared
the results of the models runs under 20th century greenhouse
gas (GHG) conditions (20C3M scenario (Roeckner, 2005a; Wegner,
2007a,c; Lowe, 2005a; NCAR, 2005a, 2007a,c), interpolating them
to the Hyytiälä geographical coordinates 61◦ 51 N, 24◦ 17 E from
the closest models grid points) with Hyytiälä climatic data for
1961–1990. On the basis of this comparison, we concluded that
there is a good agreement between predicted monthly tempera-
tures. Larger differences in precipitation are partly explained by
the known underestimation of this interpolated precipitation data
(Venäläinen et al., 2005), and partly by the fact, that generally GCMs
still simulate precipitation not very well.
We were therefore able to use these model results as the basis of
scenarios for our simulation experiments. We used the differences
between real climatic averages and their estimates from GCM runs
for the contemporary climate, as a correction for model scenar-
ios, based on model runs for expected future conditions. This is
equivalent to the usually used approach, based on the use of the
difference between simulated results for future and ambient cli-
mate conditions instead of the absolute values for the simulated
future climate.
Scenarios of climate change for 21st century were based on
these models runs for the relatively moderate A1B SRES emission
scenario (IPCC Special Report. . ., 2001). There are available three
independent model runs of ECHAM5 (Roeckner, 2005b; Wegner,
2007b,d), three runs of CCSM3 (NCAR, 2005b, 2007b,d) and only
one run of HadGEM1 model (Lowe, 2005b) with this scenario. This
scenario predicts an increase of annual temperature of about 4.1 ◦ C
over 100 years with ECHAM 5 model, 5.3 ◦ C with HadGEM1 model
and 3.7 ◦ C with CCSM3 model (Table 1). The projected changes
of precipitation is more different for different models and model
runs: HadGEM1 predict almost no change of annual precipita-
tion, ECHAM5 predicts in average a rise of annual precipitation
on about 110 mm or 21%, CCSM3 – a rise on about 70 mm or 13%
(Table 1).
Soil temperature as well as moisture was estimated using the
SCLISS model (Bykhovets and Komarov, 2002; Komarov et al.,
2007). Soil physical properties required for soil moisture simulation
(Table 2) were based mainly on data from the Hyytiälä Experimen-
tal Catchments (Ilvesniemi et al., 2010).
 V. Shanin et al. / Ecological Modelling 251 (2013) 232–245 235

Table 1 2.3. Parameterization of the model

Projected air temperature and precipitation change (differences of average annual
values for 2001–2100 and 1961–1990).
2.3.1. Soil data
GCM, run Air temperature change Precipitation change Our simulations were focused on mesic site type (Myrtillus-type,
◦
C mm % MT, according to fertility classes applied in Finland (Cajander, 1949;
Hotanen et al., 2008)), which is the most common site type repre-
ECHAM5 run1 4.1 192 37
ECHAM5 run2 4.2 92 18 senting half of the forests on mineral soils in southern Finland. In
ECHAM5 run3 4.0 48 9 addition, it is a widely spread forest type in Scandinavia and the
HadGEM1 run1 5.3 3 1 boreal zone of European Russia. All three tree species included in
CCSM3 run1 3.9 94 18 this study can be found growing together in mixed stands on this
CCSM3 run2 3.8 65 12
CCSM3 run3 3.3 50 10
site type. We used values reported in Tamminen (1991) to set initial
values (Table 4) for carbon and nitrogen pools in the forest floor and
in the mineral soil of mesic sites in southern Finland. Organic mat-
ter in mineral soil is described as the sum of stable humus and labile
Table 2 humus originating from root litter. We calculated labile humus to
Soil (and forest floor) physical properties. be 15% of total organic matter measured for mineral soil, and in
assigning the nitrogen amount assumed that the C/N-ratio is similar
Physical properties Forest floor Mineral soil
to the C/N-ratio of the forest floor.
−3
Bulk density, g cm 0.14 1.35 Soil variability was taken into account by calculating Monte-
Moisture at permanent 7.1 7.1
wilting point, vol.%
Carlo runs with a 20% standard deviation from the average. The
Moisture at field capacity, 33.0 29.0 proportion of labile humus was controlled from the model output,
vol.% being approximately constant for the simulation period.
Total porosity (or moisture 62.8 47.1
at full saturation), vol.%
2.3.2. Species-specific data
The main model calibration parameters were allocation of
annual tree increments to different tree compartments: stem,
foliage, coarse roots and fine roots in relation to the age status of the
tree, similar to Komarov et al. (2003). Age status is represented as
Simulated litter temperature (Table 3) at the end of 21st cen- three ontogenetic periods: young, mature and over-mature trees.
tury as compared to contemporary climate conditions increases Each group has species-specific similar coefficients of allocation
on about 1–2 ◦ C in winter, 2–4 ◦ C throughout the rest of year, and of annual tree increments. These coefficients play a crucial role in
2–3 ◦ C for annual mean (results of HadGEM1 is about 1 ◦ C higher). the competitive abilities of tree species. They define growth rate
In winter soil temperature is less sensitive to the change of air in terms of dependence on age, height and crown size (competi-
temperature than in warm seasons. For simulated litter moisture tion for light) and biomass of fine roots (competition for nitrogen).
(Table 3) we have more significant increase in winter and relatively For the description of mixed forests, we used growth calibrated in
small change in summer (small increase or small decrease). And, in correspondence with growth tables for pure stands. The nitrogen
this case, differences between results of different runs of the same and ash contents of the compartments of different trees are also of
model are comparable with differences between models. great importance (Appendix A, Table A.2). They point to biological

Table 3
Projected monthly average temperature and moisture of litter for 2071–2100 vs. those calculated from the actual weather for 1961–1990 (±standard deviations).

Month Calculated from actual ECHAM5 run 1 ECHAM5 run 2 ECHAM5 run 3 HadGEM1 run 1 CCSM3 run 1 CCSM3 run 2 CCSM3 run 3
weather

Temperature, ◦ C
Jan −2.3 ± 1.1 −0.5 ± 0.9 −0.5 ± 0.8 −0.5 ± 0.7 −0.5 ± 0.9 −0.4 ± 0.7 −0.8 ± 0.7 −0.8 ± 0.8
Feb −1.9 ± 0.6 −0.7 ± 0.6 −0.6 ± 0.6 −0.8 ± 0.5 −0.6 ± 0.7 −0.7 ± 0.4 −0.9 ± 0.5 −1.0 ± 0.5
Mar −0.9 ± 0.6 0.4 ± 0.9 1.0 ± 1.5 1.1 ± 1.6 1.7 ± 2.0 0.9 ± 1.1 0.6 ± 1.1 0.1 ± 0.8
Apr 1.4 ± 1.3 4.8 ± 1.7 5.3 ± 1.3 5.3 ± 1.7 8.1 ± 1.9 4.2 ± 1.7 5.0 ± 1.6 3.7 ± 1.1
May 8.7 ± 1.7 11.9 ± 2.0 12.3 ± 1.5 11.3 ± 1.8 13.6 ± 2.0 11.8 ± 1.9 11.9 ± 2.1 10.6 ± 1.6
Jun 14.2 ± 1.9 16.7 ± 1.6 17.1 ± 1.7 16.8 ± 2.0 19.2 ± 2.0 16.4 ± 1.8 17.2 ± 2.3 16.8 ± 1.6
Jul 16.0 ± 1.5 18.9 ± 2.3 18.8 ± 1.8 18.4 ± 1.7 20.3 ± 1.7 19.3 ± 2.2 19.8 ± 2.5 19.9 ± 2.8
Aug 13.8 ± 1.2 16.5 ± 1.8 16.4 ± 1.4 16.7 ± 1.5 18.1 ± 1.3 16.7 ± 1.7 17.5 ± 1.6 17.5 ± 2.1
Sep 8.5 ± 1.6 12.2 ± 1.4 12.0 ± 1.7 12.0 ± 1.5 12.9 ± 1.5 11.1 ± 0.9 11.2 ± 1.4 11.3 ± 1.3
Oct 3.3 ± 2.0 7.4 ± 1.4 7.6 ± 1.5 7.8 ± 1.6 7.9 ± 1.7 7.0 ± 1.2 6.0 ± 1.1 6.1 ± 1.5
Nov −0.8 ± 1.1 2.8 ± 2.1 2.1 ± 2.0 2.0 ± 1.9 3.4 ± 2.6 1.8 ± 1.4 1.2 ± 1.3 1.2 ± 1.2
Dec −2.8 ± 1.4 −0.1 ± 1.5 −0.1 ± 1.2 −0.2 ± 1.1 0.3 ± 1.9 −0.7 ± 1.1 −0.9 ± 1.1 −0.8 ± 1.3

Moisture, mass%
Jan 88.7 ± 8.2 109.8 ± 8.4 101.4 ± 12.0 92.8 ± 11.0 87.3 ± 10.3 101.2 ± 8.6 85.2 ± 7.3 94.5 ± 10.0
Feb 88.7 ± 8.2 110.9 ± 8.8 102.9 ± 11.6 93.5 ± 11.9 88.5 ± 10.4 101.9 ± 9.2 85.3 ± 7.3 94.5 ± 10.0
Mar 89.2 ± 8.1 113.4 ± 8.5 107.9 ± 9.5 99.5 ± 11.7 94.1 ± 10.1 106.1 ± 9.3 91.1 ± 8.1 97.2 ± 11.2
Apr 96.4 ± 7.7 115.4 ± 8.6 110.1 ± 8.0 103.0 ± 10.4 95.7 ± 8.6 107.8 ± 7.3 99.9 ± 8.0 103.2 ± 9.6
May 98.1 ± 6.7 106.6 ± 9.5 97.4 ± 7.5 93.5 ± 8.0 84.4 ± 6.7 96.4 ± 6.4 97.0 ± 6.4 98.9 ± 7.9
Jun 86.7 ± 6.6 93.6 ± 7.4 81.2 ± 6.6 80.4 ± 7.1 72.1 ± 6.2 82.8 ± 6.0 85.9 ± 6.9 84.8 ± 6.7
Jul 76.1 ± 6.3 83.8 ± 8.6 72.3 ± 8.3 71.1 ± 7.4 68.0 ± 8.2 74.9 ± 6.6 76.6 ± 9.3 72.4 ± 6.8
Aug 73.3 ± 5.6 82.3 ± 9.4 69.1 ± 8.4 70.0 ± 8.2 67.6 ± 9.7 74.8 ± 9.7 69.4 ± 8.7 67.9 ± 9.3
Sep 76.6 ± 6.0 86.8 ± 10.0 74.2 ± 8.5 72.8 ± 9.6 67.6 ± 8.6 79.9 ± 9.9 67.2 ± 7.5 72.1 ± 11.4
Oct 82.2 ± 7.0 92.8 ± 9.8 84.7 ± 9.3 78.5 ± 10.5 73.1 ± 8.2 87.0 ± 7.8 73.9 ± 6.7 80.2 ± 11.5
Nov 87.0 ± 7.9 102.0 ± 9.1 94.2 ± 9.5 87.4 ± 10.4 81.1 ± 8.3 95.8 ± 7.4 82.4 ± 6.3 89.5 ± 10.0
Dec 87.7 ± 8.1 107.3 ± 8.2 99.8 ± 10.5 91.7 ± 10.7 85.5 ± 9.4 100.2 ± 8.0 85.3 ± 7.1 93.9 ± 9.7
236 V. Shanin et al. / Ecological Modelling 251 (2013) 232–245

Table 4 Table 5
Initial soil parameters for MT forest types on mineral soils (kg m−2 ) used to run soil Parameters for saplings of different species for simulation of natural regeneration.
sub-model. S.D. – standard deviation.

Pools SOM pool Nitrogen pool Species Age, years Height, m DBH, cm

Forest floor 2.9 0.052 Mean S.D. Mean S.D.

Stable humus in mineral soil 9.3 0.25
Labile humus in mineral soil 1.5 0.027 Initial planting:
Birch 20 7.3 3.5 5.2 2.5
Pine 20 5.9 2.5 7.2 3.5
Spruce 20 3.3 1.5 3.5 1.7
turnover of elements due to controlling the full balance of nitrogen Natural regeneration:
in the model. Birch 5 1.50 0.75 0.95 0.48
Pine 5 1.25 0.62 1.10 0.55
Spruce 5 0.46 0.23 0.21 0.10
2.4. Model verification

Verification of the EFIMOD model has previously been under-

taken for different climatic and environmental conditions (Chertov
et al., 2009; Palosuo et al., 2008; Shanin et al., 2011, 2012; Shaw
et al., 2006; Verkerk, 2004). Many examples of EFIMOD applications
are collected in a monograph (Komarov et al., 2007).
Verification was carried out by data from regional yield tables
(Shvidenko et al., 2006). We performed Monte-Carlo runs of the
model using the initial soil data distributed with a 20% standard
deviation from average. These simulations provided predicted
dynamics for some key characteristics (average height and diame-
ter, stand density and basal area) of pure stands for all three species.
EFIMOD fitted well with data from yield tables and measure-
ments reported by Ilvessalo and Ilvessalo (1975) (Fig. 1). We also
Fig. 1. Model verification against yield tables and field data for stand density (a) and
mean height (b).
compared model outputs with the growth tables given by Koivisto
(1959), but they looked unrealistic for young spruce and pine and
did not fit well with the model outputs.
2.5. Initial parameters and scenarios of simulation
Initial datasets with different species composition and their ini-
tial shares were constructed for the simulations. The first group
of initial sets consisted of two-species mixtures (birch and pine,
birch and spruce, pine and spruce) with different initial propor-
tions (9:1, 7:3, 5:5, 3:7, 1:9). These shares represented different
kinds of mixed forests: ‘almost pure’ stands with a small admix-
ture of another species, equal share, and an intermediate case.
We used proportions based on stem numbers rather than biomass
to simplify calculations of initial density for each species. The
second group of initial sets were mixed stands with all three
species in various proportions (2:2:4, 2:4:2, 4:2:2). We constructed
a total of 18 sets of initial conditions. Initial stand density was
set to 10,000 trees ha−1 , for all cases. The short codes of initial
species composition were used for representation on figures. Such
codes consist of first letter of the names of species in mixture
and theirs proportional factors. For example, code 7P3S denotes
stand with initial portions of pine and spruce equal to 70 and 30%,
correspondingly.
Since the model has no specialized regeneration module, the
regeneration was simulated as the planting of new trees every 10
years with a total density of 2000 trees ha−1 . Shares of species
were the same as existed on the site at the moment of regeneration
(basal area was taken as the basis for calculation of the share); the
quantity of birch saplings was taken to be two times higher than
calculated on the basis of existing species proportions because of
the higher seed productivity of this species. We used approximate
rather than exact estimates of the quantity of seedlings because we
applied spatially explicit models of crown and root competition,
which simulates self-thinning due to competition and determines
the number of trees of each species, ‘smoothing out’ errors in deter-
mining their initial amounts. Both at initial planting and simulation
of natural regeneration, trees were populated on a square grid
25 × 25 m in size divided into cells 0.5 × 0.5 m in size. Initial dis-
placement was described by contagious distribution (Grabarnik
and Sarkka, 2009).
The age of all saplings at regeneration was 5 years; the values
for height and diameter of stem (Table 5) were taken from the
corresponding yield tables (Shvidenko et al., 2006).
The scenarios, which we used assumed no management options.
In order to analyse the uncertainties we ran the model in Monte-
Carlo mode with initial data on stands and soil distributed with a
20% standard deviation. The model outputs were the average values
and standard deviations.
 V. Shanin et al. / Ecological Modelling 251 (2013) 232–245 237

Fig. 2. Changes in portions of pine at simulation steps 30, 70 and 100 (final-to-initial
species portion ratio) and standard deviations obtained from Monte-Carlo runs in
mixed birch–pine stands at different initial proportions of species. Logarithmic scale
is used. Labels on X-axis denote initial species composition where letters are the first
letters of species’ names and numbers are theirs proportional factors.
Fig. 4. Changes in portions of spruce at simulation steps 30, 70 and 100 (final-to-
initial species portion ratio) and standard deviations obtained from Monte-Carlo
runs in mixed pine–spruce stands at different initial proportions of species. Log-
arithmic scale is used. Labels on X-axis denote initial species composition where
letters are the first letters of species’ names and numbers are theirs proportional
factors.

3. Results

share, but in other two cases its proportion remarkably decreased.

3.1. Dynamics of tree species proportions
Birch was replaced by other species after 30–50 years (Fig. 5).
In general, variation in dynamics of species composition due to
In the ambient climate scenario, a dominant species in mixed
dispersion of stand and soil characteristics (Monte-Carlo runs) was
pine–birch stands was able to maintain its dominant position. In
not proportional to the degree of changes in species proportions,
stands with initial domination of birch, the proportion of pine (in
and was similar both for remarkable changes and for the cases with
terms of biomass) had greatly declined by the end of the simulation
almost no change.
while in initially pine-dominated stands there were no remark-
In climate change scenarios, growth of pine in mixed
able changes in species composition (Fig. 2). To describe changes
birch–pine stands was generally favoured by changing conditions
in species performance during development of stands, we used the
in comparison to birch (Fig. 6a). Additionally, the change from
ratio between the proportion of given species at simulation steps
birch-dominated to spruce-dominated stands was generally faster
30, 70 and 100, and its initial proportion. For the better representa-
than in the ambient climate scenario, with the most remarkable
tion of results, the logarithmic scale was used. With such scales, bars
increase in the case with initial 10% spruce portion (Fig. 6b). Mixed
above the horizontal axis (ratio > 1) represent increase in species
stands of pine and spruce showed no clear response to climate
proportion, and bars below scale (ratio < 1) represent decrease in
change (Fig. 6c). In the case of the three-species mixture, cli-
this parameter.
mate change positively affected pine in initially pine-dominated
In spruce–birch mixed stands, spruce replaced birch at any ini-
tial density. When the proportion of birch biomass was low at the
beginning of the simulation, birch was able to increase its propor-
tion in the young stands but thereafter decreased (Fig. 3).
In spruce–pine mixed stands, at the beginning of stand devel-
opment pine grew faster and increased its proportion of biomass. If
the initial proportion of spruce was 50% or higher, the stand turned
to being dominated by spruce within a 100-year simulation period
(Fig. 4).
In mixed stands of three tree species (spruce–pine–birch),
spruce tended to be dominant by the end of the 100-year simulation
period, independently of the initial proportions. If pine was dom-
inant at the beginning of the simulation it retained its proportion

Fig. 3. Changes in portions of spruce at simulation steps 30, 70 and 100 (final-to-
initial species portion ratio) and standard deviations obtained from Monte-Carlo
runs in mixed birch–spruce stands at different initial proportions of species. Log-
arithmic scale is used. Labels on X-axis denote initial species composition where
letters are the first letters of species’ names and numbers are theirs proportional
factors.
Fig. 5. Changes in portions of pine (a) and spruce (b) at simulation steps 30, 70 and
100 (final-to-initial species portion ratio) and standard deviations obtained from
Monte-Carlo runs in mixed birch–pine–spruce stands at different initial propor-
tions of species. Logarithmic scale is used. Labels on X-axis denote initial species
composition where letters are the first letters of species’ names and numbers are
theirs proportional factors.
238 V. Shanin et al. / Ecological Modelling 251 (2013) 232–245

Fig. 6. Differences (%) between proportions of pine (white) and spruce (grey) in total stand biomass in comparison to ambient climate (average for different climate change
scenarios and standard deviations obtained from dispersion of climate characteristics). a – birch–pine stands, b – birch–spruce stands, c – pine–spruce stands, d – three-species
mixtures. Labels on X-axis denote initial species composition where letters are the first letters of species’ names and numbers are theirs proportional factors.

stands (Fig. 6d) while in other two cases both conifer species were
favoured by climate change.
3.2. Dynamics of basic carbon pools
The average carbon stock of trees over the whole simulation
period was highest in the three-species mixtures, especially those
with initial domination of spruce (12–15 kg m−2 , Fig. 7). Carbon
stock in standing biomass was also high in spruce-dominated
stands because of higher productivity and capacity to accumulate
carbon in living biomass of this species. In mixed stands without
spruce, the maximum values of carbon stock in standing biomass
were found in stands with similar proportions of pine and birch
(8 kg m−2 ).
In general, the SOM on the forest floor increased with stand age,
but decreased in the short-term in young stands and at times of
shifts in the dominant species. This could be explained by a relative
deficit of litter in the first half of stand development. SOM in mineral
soil reflected these changes, but to a much smaller extent, almost
without any remarkable differences between cases.
In the ambient climate scenario, the higher the proportion of
pine the higher the SOM stock of the forest floor in pine–birch
and pine–spruce mixed stands, as well as when all tree species
were presented (Fig. 7). In spruce–birch mixed stands, there were
no remarkable differences in the SOM stock of the forest floor at
different initial proportions of tree species (Fig. 7).
Climate change led to a 10–25% increase (in comparison to the
ambient climate scenario) of carbon stock in standing biomass as
a result of increased productivity (Fig. 8). Carbon stocks of conifer-
ous stands increased more than in birch-dominated stands while
birch and pine mixtures with initial domination of birch showed
remarkably lower increase in standing biomass.
In the changing climate scenario, the average SOM stock of the
forest floor over the whole simulation period was generally 5–10%
lower than in the ambient climate scenario (Fig. 8) but it showed
the highest variation among all three carbon pools. The decrease
of the SOM stock in mineral soil due to climate change was more
remarkable than on the forest floor. In conclusion, climate change
negatively affected the accumulation of organic matter in soil.
In general, changes in carbon stocks in mineral soil and standing
biomass were quite similar for all cases. Additionally, variation in
results of simulations due to dispersion of climatic characteristics
(Fig. 8) was higher than this one due to dispersion of initial soil
conditions and characteristics of stands (Fig. 7).
3.3. Carbon balance in the ecosystem
Climate change positively affected carbon accumulation in
standing biomass, but had a negative influence on soil carbon stock.
To reveal the overall effect of these opposite processes, the val-
ues of the net primary production of stands and heterotrophic
respiration from the soil were analysed and the carbon balance
was calculated as the difference between net primary productiv-
ity (NPP) and emission of CO2 (Fig. 9). Since simulations started
with young stands, there was an accumulation of carbon during
stand development, which resulted in a positive carbon balance in
all cases (0.07–0.22 kg m−2 year−1 ). NPP generally varied between
0.4 and 0.5 kg m−2 year−1 , and heterotrophic respiration generally
varied between 0.3 and 0.4 kg m−2 year−1 ; both values being higher
in pine-dominated stands. Climate change positively affected both
increases in NPP and increase of decomposition rates (by ∼20%).
However, productivity gain was higher, so that carbon balance
under conditions of climate change not only remained positive
but even increased in such cases (Fig. 10). The results obtained
for single-species stands are also described in Figs. 9 and 10, for
 V. Shanin et al. / Ecological Modelling 251 (2013) 232–245 239

Fig. 7. Average values and standard deviations obtained from Monte-Carlo runs for carbon stocks in standing biomass, forest floor and mineral soil at ambient climate over
the whole period of simulation. Labels on Y-axis denote initial species composition where letters are the first letters of species’ names and numbers are theirs proportional
factors.

comparison. The NPP value was the highest for spruce stands and
the lowest for pine, but the differences were small. Notably, the
variation in heterotrophic respiration due to dispersion of initial
soil characteristics (Fig. 9) is higher than this one due to dispersion
of climatic variables (Fig. 10).
4. Discussion
4.1. Competition driven stand development
Our results showed that replacement of pioneering by late-
seral species was accelerated by climate change and the proportion
of coniferous species increased. The reason for this is that cli-
mate change led to an increased rate of decomposition of plant
residues in soil, which increased the amount of nitrogen occurring
in forms available for plant nutrition. Therefore spruce, which is
more dependent on nitrogen uptake, gained a competitive advan-
tage. This is more significant for spruce with its lateral root system,
which covers a large area of possible nutritional uptake (Schmid
and Kazda, 2002).
In stands formed by shade-tolerant spruce together with other
species, there were two clearly distinct tree layers; the upper
layer, formed by fast-growing tree species (pine or birch), and
the lower one, consisting of spruce trees. After the breakdown
of the birch canopy, spruce occupied the overstorey and formed
a close canopy, preventing regeneration of other tree species.
Thus, spruce outcompeted other tree species after early develop-
ment of a stand. According to our results, spruce grew better in
cases where its initial density was low (10 or 30%), which can be
explained by a weakening of intraspecific competition. In mixed
birch–spruce stands, spruce grew slowly in the early stages of
stand development and faster in the later stages of stand devel-
opment. This result is in a good agreement with that reported
by Cavard et al. (2011). One more factor negatively affecting the
growth of pine and birch in spruce-dominated stands is, prob-
ably, a decrease in soil fertility due to incoming spruce litter
(Cavard et al., 2011).
In the case of mixed stands of two species with similar life
strategies (pine and birch), the succession was dependent on ini-
tial species proportions. In birch-dominated stands (initial share of
birch ≥50%), birch tended to increase its proportion, possibly due
to its more effective use of available resources (for example, solar
radiation), which resulted in a higher growth rate than that of pine.
We also observed that pine competed with spruce more effectively
than it did with birch, because its nitrogen uptake for biomass is
lower than for pine.
240 V. Shanin et al. / Ecological Modelling 251 (2013) 232–245

Fig. 8. Changes (%) in carbon stocks in standing biomass, forest floor and mineral soil in comparison to ambient climate (average values over the whole period of simulation
for different climate change scenarios and standard deviations obtained from dispersion of climate characteristics). Labels on Y-axis denote initial species composition where
letters are the first letters of species’ names and numbers are theirs proportional factors.

Fig. 9. Net primary production (NPP), heterotrophic respiration (in terms of carbon) and carbon balance (NPP minus CO2 emission) at different initial proportions of species.
Average values over the whole period of simulation and standard deviations obtained from Monte-Carlo runs. Labels on X-axis denote initial species composition where
letters are the first letters of species’ names and numbers are theirs proportional factors.
 V. Shanin et al. / Ecological Modelling 251 (2013) 232–245
Fig. 10. Net primary production (NPP), heterotrophic respiration (in terms of carbon) and carbon balance (NPP minus CO2 emission) at different climatic scenarios. Average
values over the whole period of simulation and standard deviations obtained from dispersion of climate characteristics. Labels on X-axis denote initial species composition
where letters are the first letters of species’ names and numbers are theirs proportional factors.
4.2. Total productivity of mixed stands
Productivity increased with increasing share of more fast-
growing pioneering species. However, in the case of spruce–birch
mixtures, the total biomass production was highest when there
was an equal initial share of these species. This can be explained by
strongly decreased intraspecific competition at such initial propor-
tions, whereas interspecific competition was low due to differences
in their life strategies. This resulted in a considerable productiv-
ity gain for both species. In general, mixed stands showed higher
biomass production than pure stands, since species have differ-
ent resource requirements and use available resources effectively.
It is widely known that the growth of stands comprising a mix-
ture of species with different life strategies is better than that of
pure stands (Kelty, 1992; Mielikäinen, 1980, 1985; Morin et al.,
2011). Some explanations for this are listed in Cavard et al. (2011)
and Sterba et al. (2002). Firstly, trees of species with different
shade tolerances in mixed stands may use solar radiation more
efficiently. For example, shade-tolerant species growing under a
canopy formed by a shade intolerant species can use light that is
not intercepted by the latter. A second way to decrease competition
is the occupation of different soil horizons by roots of different tree
species, and our model is able to account for the different root strat-
ification of different tree species. Some tree species, mostly those
with taproot systems, are able to penetrate the deeper soil hori-
zons (Brassard et al., 2011; Hendricks and Bianchi, 1995; Jose et al.,
2006; Strong and La Roi, 1983). For example, competition in mixed
stands with broadleaves may result in a shallower root system for
Norway spruce (Bolte and Villanueva, 2006; Schmid and Kazda,
2002). In other words, there is ‘niche segregation’ between differ-
ent tree species in both above- and below-ground competition. A
third explanation is the ‘facilitation effect’ of the litter produced
by broadleaved trees, which is usually richer in fast-decomposing
compounds (Côté et al., 2000; Fenton et al., 2005; Laganière et al.,
2009, 2010; Légaré et al., 2005a,b; Prescott et al., 2000; Vesterdal
et al., 2012). The input of such litter may result in an increased
turnover rate of nutrients. Another mechanism proposed to explain
this effect (Jose et al., 2006) is the additional enrichment of the
forest floor by nitrogen when one of the species in the mixture is
nitrogen-fixing, such as birch, for example (Nohrstedt, 1988). In
addition, it is suggested that spruce grows better in pure stands
241
in comparison with spruce–beech mixed stands, because young
spruce trees in mixtures have a shallower root system and, there-
fore, suffer from drought stress (Sterba et al., 2002). However, in
boreal spruce–birch stands, growth of spruce is higher than in pure
stands (Mielikäinen, 1985).
In our simulations, climate change resulted in an increase of pro-
ductivity due to the additional amount of nitrogen resulting from
the increased rate of decomposition of soil organic matter, which is
noted to cause a productivity gain in many studies (Callahan et al.,
2008; Högberg, 2012; Vetter et al., 2005). A slightly more remark-
able increment was observed in spruce-dominated stands, which
turned out to be the species that was the most sensitive to the
amount of available nitrogen. Ge et al. (2011) did not find a similar
response for Norway spruce in their simulation of the develop-
ment of mixed stands. Their model accounts for water deficiencies
in more detail, but does not describe mechanisms for competition
between tree species for nutrients. These results indicate that both
nutrient dynamics and below-ground competition for water and
nutrients have to be modelled in more detail in order to predict
species responses to changing environmental conditions.
4.3. Soil carbon dynamics affected by tree species composition
The highest accumulation of organic matter in the organic layer
was observed in pine-dominated stands. This can be explained by
the relatively low nitrogen content in litter produced by coniferous
species, making them resistant to fast decomposition (Kelty, 2006;
Vesterdal et al., 2012). Spruce litter is also slow-decomposing, but
the amount of litter produced was considerably lower than that
produced by pine, due to the slower growth rate of spruce and
its much lower mortality. In contrast, litter produced by birch is
influenced by relatively fast decomposition.
The range of heterotrophic respiration we predicted
(0.3–04 kg m−2 year−1 ) was about half of the total respira-
tion measured in study sites that represent similar conditions
(0.6–0.7 kg m−2 year−1 ; Kolari et al., 2004; Valentini et al., 2000).
It is estimated that respiration of roots in forest soils is about
30–35% of the total soil respiration (Larionova et al., 2006). It is
inexplicitly accounted in our model and, therefore, we are able to
obtain the comparable balancing values for different variations of
initial species composition.
242 V. Shanin et al. / Ecological Modelling 251 (2013) 232–245
The change of carbon stock in the organic layer varied in
different cases from −0.3 to 1.1 kg m−2 for the whole simula-
tion period of 100 years with an average of 0.0032 kg m−2 year−1 .
These values were in general lower than reported for middle-aged
stands by Häkkinen et al. (2011) (0.023 ± 0.002 kg m−2 year−1 ),
but lay within the range reported in that paper (from −0.015 to
0.072 kg m−2 year−1 ). The predicted rate of soil carbon accumu-
lation is consistent with the results of chronosequence studies
(0.0042–0.008 kg m−2 year−1 ; Bormann et al., 1995; Peltoniemi
et al., 2004; Wardle et al., 2003). We simulated a long-term devel-
opment of mixed forests from young to mature where we observed
the ageing and breakdown of the canopy formed by the first gen-
eration of trees and the following transformation to uneven-aged
stands.
4.4. The effect of climate change on total carbon balance
According to our simulations, the biomass production and car-
bon stock of trees increased due to climate change, but the soil
carbon stock decreased as a result of the accelerated decomposi-
tion of soil organic matter. Such positive effects of climate change
on forest productivity, particularly for boreal forests of the north-
ern hemisphere, are reported in many studies (Lopatin et al., 2006;
Myneni et al., 1997; Shanin et al., 2011, 2012; Spiecker, 1999). In
addition, a decrease in the soil carbon stock as a result of increased
temperatures and precipitation was earlier predicted by a gap-type
ecosystem model (Mäkipää et al., 1999), and a dynamic soil car-
bon model combined with a forest scenario model (Karjalainen
et al., 2002). However, some studies have reported an opposite
trend (Cao and Woodward, 1998; Karjalainen et al., 2003; Mäkipää
et al., 2011). An increase in soil respiration and nitrogen mineral-
ization rates is also noted in many studies (e.g. Rustad et al., 2001;
Vetter et al., 2005) and a positive response to warming was gen-
erally found to be larger in colder forest ecosystems (Strömgren
and Linder, 2002). This, in turn, leads to an increase in stand pro-
ductivity because nitrogen stock is the limiting factor in most
boreal ecosystems (Bobbink et al., 2010). Summing up, there are
two simultaneous processes in ecosystems. On one hand, climate
change could result in increase of productivity and, consequently,
litter income. On the other hand, changes in climate variables lead
to increased rate of decomposition processes in soil. The relation-
ship between opposite processes and overall carbon balance is still
a question of debate and apparently depends both on climatic zone
and forest type.
The effect of occurrences of drought stress on the activity of soil
decomposers can be one more source of uncertainty in predicting
changes of carbon stocks in soil at climate change. In general, ignor-
ing this fact could potentially result in overestimation of the rates
of SOM transformation. However, climate change scenarios applied
in our study showed neither remarkable increase nor decrease in
moisture of forest floor (see Table 3) so it can be assumed that there
was no drought stress.
4.5. Possible uncertainties: evaluation of the model performance
The model that we applied had some limitations related to the
rather simple sub-model of productivity used in our system. Such a
sub-model requires a small number of parameters for initialization
but has a high operational speed. EFIMOD is not able to simulate the
effect of CO2 fertilization on tree productivity, whereas increased
carbon dioxide concentration is considered as a possible source
of increased biomass production (Boisvenue and Running, 2006;
Cole et al., 2010; Coops and Waring, 2001; Ellsworth et al., 2012;
Kozovits et al., 2005; Moore et al., 2006). However, the increase in
the NPP of boreal forests due to this fertilization effect remains
a subject of debate (Kurz et al., 2007), because no increase in
productivity has been observed in other studies (Bond-Lamberty
et al., 2007; Percy et al., 2002). Other factors negatively affecting
productivity can be drought stress (Hartmann, 2011; Huang et al.,
2010; Martínez-Vilalta et al., 2008) or a reduction in the amount
of available solar radiation due to increased cloudiness (Breshears
et al., 2009; Johnston et al., 2009). We believe that introducing a
more detailed ecophysiological model to our simulation experi-
ments will deliver more clarity to this issue. Detailed modelling
of belowground competition is also important for descriptions of
competitive mechanisms in mixed forests.
We simulated mixed stands of competing tree species in the
changing climate with a model that accounted the accelerated rate
of soil processes and the higher nitrogen availability in the warmer
temperatures. In our simulations, the responses of the studied tree
species to climate change reflected the differences in the nutri-
ent requirements. Tree species may also have differences in their
responses to direct climate impacts. Widely applied climate enve-
lope approach summarize climate niche of the tree species and
predict changes in the tree species performance particularly at the
margins of their current range (e.g. Elith et al., 2006; Zimmermann
et al., 2009). We studied tree species that have large distributional
ranges and wide acclimation capacity on region that is central in
their biogeographical range. Thus, we cannot assume remarkable
differences in the competitive status of tree species in changing
climate based on species climate niches. We have assumed that all
studied species can respond positively to direct climate impacts,
since current populations of many tree species are shown to inhabit
suboptimal environments (Leites et al., 2012; Tchebakova et al.,
2005; Rehfeldt et al., 2002) and the studied species have shown
positive responses to warmer climate (e.g. Rousi et al., 2012).
A description of the processes of SOM dynamics in different bio-
geochemical cycles relating to elements of plant nutrition under
climate change scenarios is also of great importance. It could be
improved by including more detailed SOM dynamics models, for
example by including dynamics of soil biota (Kuzyakov et al., 2000;
Talbot et al., 2008). However, we must maintain a balance between
providing a sufficiently detailed model and the number of parame-
ters in the context of experimental accuracy, because complication
of the model’s structure can lead to uncertainties resulting from the
joint action of uncertainties for individual parameters (Larocque
et al., 2008).
Nevertheless, we believe that our results reflect the main ten-
dencies of changes in tree species composition due to climate
change and corresponding changes in carbon pools for boreal
forests. Uncertainties in the model structure mentioned before
could be significant in more complicated temperate forests, where
differences in water support, root competition and distribution of
light in the canopy are more significant, in comparison to boreal
zone.
5. Conclusion
The development of mixed stands of competing tree species
under conditions of the ambient and changing climate conditions
can be evaluated with a stand simulator that accounts for competi-
tion of both light and soil resources. We simulated development of
mixed stands with a dynamic modelling of nitrogen cycling, which
provides feedback in relation to nutrient availability. The climate
change increased the rates of decomposition and correspondingly
increased the amount of available nitrogen for tree growth, which
has to be accounted for when evaluating competition between tree
species in a changing climate. Our simulations showed that com-
petition for nutrients under conditions of climate change shifts
species composition, due to changes in the allocation of carbon
and nitrogen to more competitive species, which require more
 V. Shanin et al. / Ecological Modelling 251 (2013) 232–245
nutrients and/or have a faster growth rate. Thus, we conclude that,
in a changing climate, deciduous birch will be outcompeted by
coniferous species in mixed stands.
According to our analysis, changes in the climatic conditions
affected standing biomass positively and soil organic matter neg-
atively, having an overall positive influence on forest carbon
sequestration. We also showed that tree species composition has
a remarkable influence on the carbon stock of both vegetation and
the forest floor. Earlier studies have demonstrated that forest car-
bon sequestration and its potential to mitigate climate change can
be controlled by thinning regime and length of the rotation period.
We conclude that management decisions that control tree species
dominance can greatly affect forest carbon sequestration capac-
ity. According to our simulations, mixed stands of all three tree
species with a dominance of spruce is the most favourable option
to increase forest carbon stock and mitigate climate change.
Our results show the potential opportunities and risks of for-
est management in a scenario of a changing climate, and further
studies on different site types are needed to provide guidance on
management practices and selection of tree species in changing
conditions.
Acknowledgements
This study was supported by the Academy of Finland (project
numbers 140766 and 257845), Programme 4 of the Presidium of the
Russian Academy of Sciences and collaborative project of the Rus-
sian Academy of Sciences and the Academy of Finland ‘Adaptation
of boreal forest management in the changing climate: modelling
forest stand and soil development for main boreal tree species’.
This work was supported by the Russian Foundation for Basic
Research, grant number 12-04-31635. We also thank three anony-
mous reviewers for their insightful comments.
Appendix A. Supplementary data
Supplementary data associated with this article can be found,
in the online version, at http://dx.doi.org/10.1016/j.ecolmodel.
2012.12.015.
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