J. For. Res.

(2020) 31(1):13–26
https://doi.org/10.1007/s11676-019-00935-8

ORIGINAL PAPER

Assessment of early survival and growth of planted Scots pine

(Pinus sylvestris) seedlings under extreme continental climate
conditions of northern Mongolia
Gerelbaatar Sukhbaatar1,2 • Batsaikhan Ganbaatar3 • Tsogtbaatar Jamsran3 •

Battulga Purevragchaa3 • Baatarbileg Nachin2 • Alexander Gradel4,5

Received: 19 October 2018 / Accepted: 6 January 2019 / Published online: 6 April 2019
 The Author(s) 2019

Abstract Environmental factors play vital roles in suc- region of northern Mongolia on six Scots pine plantations
cessful plantation and cultivation of tree seedlings. This ranging from 5 to 10 years. In each of the six plantations,
study focuses on problems associated with reforestation five 900 m2 permanent sample plots were established and
under extreme continental climatic conditions. The objec- survival rates and growth performance measured annually
tives were to assess relative seedling performance (survival over 7 years. Results show high variation in survival
and growth) with respect to plantation age, and to analyze among the plantations (p \ 0.001, F = 29.7). Seedling
the influence of specific climatic factors during the early survival in the first year corresponded directly to the
stages of Scots pine (Pinus sylvestris L.) plantations. The number of dry days in May. However, survival rate
study was carried out in reforested areas of the Tujyin Nars appeared to stabilize after the second year. The insignifi-
cant variation of height categories throughout the obser-
vation period indicated low competition among
Project funding: The work was supported by the Partnerships for
Enhanced Engagement in Research (PEER) Science Cycle 2 Grant
individuals. Two linear mixed-effect models show that
#296 ‘Building research and teaching capacity to aid climate change height and radial growth were best explained by relative air
and natural resource management at the National University of humidity, which we consider to be a reliable indicator of
Mongolia (NUM)’. site-specific water availability. Insufficient amounts and
uneven distribution of rainfall pose a major threat during
The online version is available at http://www.springerlink.com
the first year of plantation establishment. Humidity and
Corresponding editor: Tao Xu. water availability are decisive factors for a successful
seedling plantation. This highlights the impact of drought
& Alexander Gradel on forest plantations in northern Mongolia and the impor-
agradel@mail.de
tance of developing climate resilient reforestation
1
Department of Environment and Forest Engineering, strategies.
National University of Mongolia, Ulaanbaatar 14201,
Mongolia Keywords Reforestation  Scots pine  Pinus sylvestris L. 
2
Institute of Forest, National University of Mongolia, Survival  Increment  Growth  Climate  Mongolia
Ulaanbaatar 14201, Mongolia
3
Division of Forest Resources and Protection, Institute of
Geography and Geoecology, Mongolian Academy of Introduction
Sciences, Ulaanbaatar 15710, Mongolia
4
Faculty of Forest Sciences and Forest Ecology, Universität Forest plantations provide a range of benefits, including
Göttingen, Büsgenweg 5, 37077 Göttingen, Germany
playing a key role in ecosystem functionality (Hector and
5
Chair of Forest Policy, Forest Economy and Forest Bagchi 2007), offsetting continuing deforestation and
Management, Institute of Management and Economy of the
degradation (FAO 2010), providing carbon sequestration
Forest Sector, Saint Petersburg State Forest Technical
University, Institutskii per., 5, 194021 Saint Petersburg, (Kongsager et al. 2013) and storage (Chen et al. 2015),
Russia promoting efficient nutrient cycling (Ma et al. 2007),

123
14 G. Sukhbaatar et al.

increasing plant species diversity and community structure most recent National Forest Inventory (MPNFI 2016)
(Eycott et al. 2006), and preventing soil erosion (Lawson reported that boreal forest cover is nearly 9.1 million
and Michler 2014). Forested areas regulate streamflow and hectares (\ 6%) and is found in the transitional zone
modify the magnitude of peak flows (Buendia et al. 2015), between the Siberian boreal forest and the Central Asian
and improve soil water retention (Kahle et al. 2005). The dry steppe (Mühlenberg et al. 2012). This transitional zone
benefits of establishing plantations in specific areas of open is characterized by a highly continental climate with dry
land are therefore twofold, in that they meet increasing winters. Forests in Mongolia mainly grow on mountain
demands for timber by reducing deforestation (Pirard et al. slopes between 700 and 2500 m a.s.l. World Bank studies
2016), but also strengthen local ecosystem functions. (Crisp et al. 2004; Mühlenberg et al. 2006) have shown that
However, it can be difficult to establish forest plantations climate change may affect the distribution, natural regen-
in dry, continental regions. A number of previous studies eration and successful reforestation of Mongolian conifer
have shown that changes in seedling survival rates and forests, and that newly planted species often appear to be
growth performance may be due to limited light avail- heavily affected by human disturbances, especially unreg-
ability (Kunstler et al. 2005; Hattori et al. 2009), nutrient ulated livestock grazing. According to statistics collected
deficiencies (Oscarsson et al. 2006; Löf et al. 2007), over the last 100 years, 40% of all Mongolian forests have
interspecific competition (Osunkoya et al. 2005), and been directly affected by human activities which have
changing environmental factors (Soehartono et al. 2002; largely had adverse effects (Tsogtbaatar 2004). Overall,
Battles et al. 2008; Hattori et al. 2013). Establishing 684,000 ha of previously forested areas have not regener-
plantations on logged-over land, previously dominated by ated after fires, and another 159,000 ha have been con-
conifers, with seedlings of native species is one way to verted into non-forested ecosystems.
accelerate forest recovery mechanisms. However, Löf et al. The first reforestation trials were carried out in Mon-
(2007) reported that the mortality of seedlings planted on golia in the 1970s. However, efforts at forest rehabilitation
open sites is higher than seedlings planted under a native with native species have had relatively limited success
tree canopy. around the country. Today, most plantations are mono-
Mongolia has relatively low amount of forest resources. cultures established from 2-year-old seedlings of Larix
Forest cover represents roughly 7–8% of the territory; the sibirica Lebed. and Pinus sylvestris L. Chamshama et al.

Table 1 Description of the permanent sample plots used for this study in 2010
Sample plot Establishment Age Mean Mean Basal area Geographical location Previous
of (seedling/ height in diameter in in m2/ anthropogenic
reforestation sapling) by cm ± SD mm ± SD ha-1 in Latitude Longitude Elevation disturbance
2010 in 2010 in 2010 2010 in m impact

Bayanbulag 2003 10 145 ± 2.3 45.2 ± 0.6 3.26 50,011 106,026 720 high. i. (1999);
(BB) clear-cut
(2000–2002)
Moilt guu 2004 9 140.4 ± 1.7 35.3 ± 0.4 2.77 50,010 106,024 706 high i. (1999);
(MG) clear-cut
(2002–2003)
Bayanbulagyin 2005 8 112.9 ± 1.9 26.4 ± 0.5 1.34 50,012 106,028 698 med. i. (1997);
denj (BBD) clear-cut
(2001–2004)
Tujyin nars 2006 7 67.9 ± 1.5 16.1 ± 0.4 0.45 50,012 106,032 694 high i. (1999
(TN) and 2003);
clear-cut
(2004)
Khudgyin guu 2007 6 54.6 ± 1.4 11.8 ± 0.4 0.21 50,009 106,032 703 high i. (2005);
(KHG) clear-cut
(2005–2006)
Khudgyin 2008 5 7.6 ± 0.6 1.5 ± 0.1 0.001 50,007 106,015 700 med i. (2005);
guunyiar clear-cut
(KGA) (2006–2007)
All plots were established on previously clear-cut and burned sites: high i = high intensity fire; med i = medium intensity fire; years in brackets;
SD = standard deviation. Diameter was measured at stem base

123
Assessment of early survival and growth of planted Scots pine (Pinus sylvestris) seedlings… 15

Fig. 1 Location of the study area in the Tujyin Nars National Park (Selenge province) with the six sample sites (see Table 1 for the
abbreviations of the sites)

(2009) underline the degree to which successful plantation Materials and methods
establishment depends on planting design and the selection
of appropriate species but also relies on silvicultural, eco- Site description
logical and economic aspects. Scots pine (P. sylvestris) is
an important species for the region in terms of forest cover, The study was carried out in the Tujyin Nars National Park
timber production and reforestation management. Planting (50050 and 50120 N, 106140 and 106310 E) in the Selenge
of Scots pine is typically carried out in the first half of May province of northern Mongolia (Fig. 1). The SPA stretches
which usually coincides with a lack of rainfall, prolonged approximately 33 km from east to west and covers an area
drought and high winds. The Tujyin Nars Special Protected of 73,000 ha, of which 45,800 ha are natural pine forests
Area (SPA) is a representative area for successful refor- and 21,000 ha are Scots pine plantations (Gerelbaatar
estation in northern Mongolia. Current statistics show that 2012).
over the last two decades, some 21,000 ha of previously According to the updated world map of the Köppen–
logged land in the Tujyin Nars region have been artificially Geiger climate classification (Peel et al. 2007), the region
reforested (Gerelbaatar 2012). In spite of the importance of lies within the transition climatic zone between a cool
forest plantations in Mongolia, there is little information on continental climate (Dwc) and a cold semi-arid climate
early survival and growth performance of seedlings. The (Bsk), with small pockets exhibiting a temperate conti-
main objectives of this study were: (1) to assess seedling nental climate (Dwb). The temperature at the nearest
survival and growth with respect to plantation age; and, (2) meteorological station in Sukhbaatar, 15–20 km north-west
to analyze the effects of climate factors on seedling sur- at 660 m between 2003 and 2014 averaged 0.3 C. The
vival during the growing season in the early stages of average annual precipitation was 249.8 mm with a pre-
establishment. Following an in-depth analysis of the cipitation peak between June and August. The dry season
results, survival rate and growth of seedlings is explained extends from March to June in spring and from August to
with linear mixed-effects models (LMM). October in autumn (Regzedmaa 2008). According to
meteorological data, total annual precipitation and mean
annual temperatures have varied over previous decades.
Figure 2 provides an overview of the climate characteris-
tics of the area based on data from the Meteorological
station Sukhbaatar, 2003–2014.

123
16 G. Sukhbaatar et al.

a 80.0 30.0 b
200
60.0
20.0
40.0

Precipitation (mm)
150
Precipitation (mm)

Temperature (°C)
10.0
20.0

0.0 0.0 100

Mar

May
Jun
Jul
Apr
Jan

Aug

Nov
Feb

Sep
Oct

Dec
-20.0
-10.0
-40.0 50
-20.0
-60.0
0

Mar

Jun
Jul
Apr
May
Jan

Aug

Nov
Feb

Sep
Oct

Dec
-80.0 -30.0

Fig. 2 Overview of the climatic conditions of the study area (meteorological station Sukhbaatar, 2003–2014). a Climate diagram; b annual
precipitation; c annual temperature

The soil in the study area is mainly of haplic arenosols sample plots, (overall: 6 sites 9 5 plots = 30 plots), each
developed from sandy sediments (JICA 1998). Planting measuring 900 m2 (30 m 9 30 m) in completely random-
was carried out in the beginning of the growing season ized design were established. Annual field measurements
(first half of May) from 2003 to 2008. 2-year-old bare-root of seedlings included: diameter at stem base, total height
seedlings of Scots pine were transplanted manually using and annual height increments. The survival rate of planted
the slit planting method. The initial planting density of seedlings on each plot was assessed by counting both live
each plantation was 2500 seedlings ha-1 (4.0 m 9 1.0 m and dead individuals at the end of the annual growing
spacing). Six neighboring reforested stands were selected, season (September). The survival rate of each plantation
representing early survival and growth of Scots pine. All was estimated annually (2003–2010). Height was measured
plantations were monocultures established at different to the nearest 0.01 m with a measuring tape, and at stem
times and on clear-cut forested land following frequent base, the diameter was measured to the nearest 0.1 cm with
intense fires (Table 1). The plantations were located close calipers. Annual tree-ring width of medium-sized samples,
to each other and exhibited similar site conditions. They (5 cores from each plot), were measured at the Laboratory
were selected to represent the typical early development of Dendrochronology at the Mongolian Academy of Sci-
phase of plantations in the study area. ences (MAS).

Measurements in the plots

Field measurements and data collection were carried out

annually in September/October between 2003 and 2010.
On each of the six plantation sites five square permanent

123
Assessment of early survival and growth of planted Scots pine (Pinus sylvestris) seedlings… 17

Table 2 Survival of planted

Stand Planting year Year of measurements
seedlings over the monitoring
period (%) 2003 2004 2005 2006 2007 2008 2009 2010

BB 2003 97.92 97.60 97.60 97.52 97.52 97.48 97.48 97.48

MG 2004 92.60 92.40 92.28 92.28 92.20 92.20 92.20
BBD 2005 78.44 78.12 78.12 78.12 78.04 78.00
TN 2006 61.68 61.52 61.52 61.40 61.40
KHG 2007 76.20 75.96 75.90 75.90
KGA 2008 15.28 15.04 15.00
BB Bayanbulag, MG Moilt guu, BBD Bayanbulagyin denj, TN Tujyin nars, KHG Khudgyin guu, KGA
Kudgyin guunyiar

Basic analyses role, the following terms were applied: ‘accelerated

growth’ instead of ‘dominant’; relatively ‘average growth’
Anova instead of ‘codominant’; and, ‘retarded growth’ instead of
‘suppressed’.
For the analysis of survival and seedling growth, one-way
analysis of variance (ANOVA) was used to determine Comparing a negative and a positive pointer year
statistically significant differences in means among vari-
ables between the sites, and an F-test was used to deter- Based on the results on seedling survival (Table 2), the
mine equality of group means (Fisher 1925; O’Brien 1979). most positive and the most negative pointer years were
selected and the respective course of precipitation and
Correlation between increment and climatic variables temperature of the respective year compared against the
after plantation establishment average course of the climate factors during the observa-
tion period (2003–2010).
Pearson’s correlation coefficient (Edwards 1976) measured
the strength of the linear association between the sum of Quantification of influence of dryness/humidity
annual heights and radial increments of the second and on survival rate and growth of planted Scots pine
third years after plantation establishment, and the average trees in Northern Mongolia with linear mixed
values (annual and growing season) of climatic variables of models (LMM)
the respective time frame: Bayanbulag (2004/2005), Moilt
guu (2005/2006) Bayanbulagyin denj (2006/2007), Tujyin Based on the descriptive and basic results, explanatory
Nars (2007/2008), Khudgyin guu (2008/2009), Kudgyin models were tested and developed for explaining survival
guunyiar (2009/2010). The first year when the seedlings rate in the year of plantation, and growth (height growth
were planted may be considered as an adaptation phase and and radial growth) in subsequent years after establishment.
was therefore not included. The following climatic factors A planted tree population i at site j represents a sample
and indicators for the whole year and growing season unit. A linear mixed model approach (LMM) was used to
(May–August) were considered in the correlation analyses: avoid pseudoreplication and include fixed effects (e.g.,
sum of precipitation, average air temperature, average number of dry days during a certain period), and random
relative air humidity and number of dry days, the latter effects (e.g., site or plot) (Crawley 2007; Zuur et al. 2009;
refers to the average number of days below a critical level R-bloggers 2011). The models were optimized based on the
of air humidity (\ 30%). restricted maximum likelihood method (REML) (Zuur
et al. 2009; R-bloggers 2011). We also tested for interac-
Assessment of relative height categories tions between different fixed factors. Every initial model
run was evaluated by a standard procedure of regression
All seedlings were classified into three height categories diagnostics in which outliers were detected and eliminated
relevant to neighboring seedlings in each plot, which, in a based on the distribution of internally studentized residuals
similar fashion, is often applied to estimate competition in in Q–Q plots with a 95% confidence envelope (Robinson
stands (see for example, Krstic et al. 2012). Since the and Hamann 2011). The best models were selected
seedlings were in a very early competition stage during according to the following criteria: Akaike‘s Information
which direct competition played no role or a subordinate Criterion (AIC), Bayesian information criterion (BIC) and

123
18 G. Sukhbaatar et al.

the value of the log likelihood, the plausibility of the

intercept, the distribution of residuals, and the plausibility
of the respective model from an ecological point of view
(Gradel et al. 2017a). Spatial autocorrelation was accoun-
ted for within the sites in the mixed model procedure (Zuur
et al. 2009). At no time was it necessary to incorporate a
spatial dependence structure in the model. The average
survival rate (Sij) in the year of plantation establishment
was explained by the number of dry days in May (D). The
following model was selected:



Sij ¼ ðb0 ÞI þ b1;i D þ b2;j s þ eij ð1Þ

where b0, b1,i, b2,j are the parameter estimates of the

intercept (I), the number of dry days in May (D), and the
Fig. 4 Radial growth of planted trees. BB Bayanbulag, MG Moilt
site (s), respectively; eij = error term of the population i at guu, BBD Bayanbulagyin denj, TN Tujyin nars, KHG Khudgyin guu,
site j. KGA Kudgyin guunyiar
A model was developed for the description of average
growth in the subsequent years after planting. The model with an adjusted script based on a version from Gradel
explained average height growth (Hij) and average radial et al. (2017a); an initial version was provided by the Chair
growth (Rij) of the subsequent years after planting. The of Silviculture at TU Dresden.
growth model of the plantation i of site j consisted of the
following elements:


Results
Hij ¼ ðb0 ÞI þ b1;i h þ b2;j s þ eij ð2aÞ



Rij ¼ ðb0 ÞI þ b1;i h þ b2;j s þ eij ð2bÞ Survival rate
where, b0, b1,i, b2,j are the parameter estimates of the
Survival rate over the monitoring period
intercept (I), the annual relative average air humidity (h) of
the plantation and the site (s) respectively; eij = error term
There was a massive mortality of seedlings during the
of the population i at site j.
growing season the first year after plantation establishment
The following software packages/routines were used:
in 2006, 2007 and especially in 2008 (sites TN, KHG, and
R-statistics (R Development Core Team 2016) with the
KGA). There was a significant difference (p \ 0.001,
packages nls2 (Grothendieck 2013), nlme (Pinheiro et al.
F = 29.7) in the rate of seedling survival among the plan-
2018), ncf (Bjornstad 2013), car (Fox and Weisberg 2011),
tations. Table 2 provides an overview of the survival of
and lattice (Sarkar 2008). As part of the LMM (R-bloggers
planted seedlings over the monitoring period in the
2011), the test for spatial autocorrelation was conducted
plantations.

Survival rate and climate impact

We first analyzed the distribution of annual monthly air

temperatures and precipitation collected from the neigh-
boring Sukhbaatar meteorological station. The variation of
annual precipitation (p \ 0.001, F = 4.6) and air temper-
atures (p = 0.006, F = 9.1) were significantly high, and
precipitation showed an uneven distribution during grow-
ing seasons (Fig. 2). The survival of Scots pine plantations
correlated positively with monthly precipitation and air
temperatures during the growing season. In contrast,
seedling survival was more dependent on monthly precip-
itation than on air temperatures. A relatively strong cor-
relation between rainfall and seedling survival was
Fig. 3 Height growth of planted trees. BB Bayanbulag, MG Moilt
guu, BBD Bayanbulagyin denj, TN Tujyin nars, KHG Khudgyin guu, observed in June (r = 0.41; p = 0.05) and July (r = 0.46;
KGA Kudgyin guunyiar p = 0.05). However, among of all climatic factors and

123
Assessment of early survival and growth of planted Scots pine (Pinus sylvestris) seedlings… 19

Table 3 Pearson’s ‘correlation coefficient’ between the sum of (sum_prec), average air temperature (T_air), average relative air
increment (second and third year after planting) and respective humidity (H) and number of dry days (D)
values of climatic factors and indicators; sum of precipitation
sum_prec sum_prec T_air T_air H H D D
(annual) (vegetation (annual) (vegetation (annual) (vegetation (annual) (vegetation
period) period) period) period)

Height increment (sum of 0.28 0.13 0.47 0.11 0.24 0.27 0.21 0.17
second and third year)
Radial increment (sum of 0.72 0.59 0.64 0.10 0.60 0.73* 0.25 0.26
second and third year)

*With p B 0.1

100 When assessing radial growth, there was a similar ten-

dency as in height growth; the annual increment of radial
80 growth in the planting year was the lowest, which may in
Percentage (%)

60
part be related to transplant shock (Figs. 3, 4). For all
plantations, a gradual increase of both height and radial
40 increment was observed at the beginning of the second
year. Increasing height and radial increment of seedlings/
20 saplings was recorded each year, with the exception of
2006 and 2008, with particular focus to radial growth. The
0
KGA [3 y.] KHG [4 y.] TN [5 y.] BBD [6 y.] MG [7 y.] BB [8 y.] standard deviation of individual tree height growth was
Stand [years after establishment] often higher than for radial growth measured in each
plantation.
Fig. 5 Relative proportion of seedlings by height categories in
relation to neighborhood composition. Growth categories: white:
accelerated growth; striped: average growth; black: retarded growth. Growth performance and climate impact
BB Bayanbulag, MG Moilt guu, BBD Bayanbulagyin denj, TN Tujyin after plantation establishment
nars, KHG Khudgyin guu, KGA Kudgyin guunyiar
The initial analysis of the impact of climate variables on
indicators, the number of dry days (D) showed the stron- height and radial growth in the second and third years after
gest correlation with seedling survival: (Dmay: 0.76, Djune: planting indicated rather low values for height growth.
0.69, Dvegetation period: 0.71 with p = 0.005 and F = 3.76). With regard to effects on radial growth, precipitation, air
humidity and temperature showed higher correlation val-
Growth performance over the monitoring period ues, but only the relation between radial growth and
average annual relative air humidity, showed a significant
Height and radial increments in the first year following correlation (Table 3).
transplanting were consistently less than those measured in
the second year, and ranged from 4.8 ± 1.2 to Distribution of planted trees by height categories
7.5 ± 1.6 mm in height and 0.3 ± 0.2 to 0.9 ± 0.2 mm in
radial growth, and clearly dependent on the amount of All seedlings were divided into three height growth cate-
rainfall in the growing season. These results reveal that gories: accelerated growth, average growth and retarded
annual height increments of seedlings for the plantations growth. Overall, the distribution of trees by height cate-
exhibited similar growth trends that increased in the second gories did not significantly differ (p = 0.18; F = 2.62)
year after planting, effectively in parallel with the age of among plantations with different ages. In this study, most
the plantation (Figs. 3, 4). of the planted trees belonged to the average growth cate-
While growth trends among the plantations were simi- gory (64–81.6%), while 15.0 ± 3.2% of the remaining
lar, a significant difference in height growth was observed trees were classified as accelerated growth and
(p = 0.0004; F = 26.8). The mean annual height increment 11.8 ± 1.3% were in the retarded growth category (Fig. 5).
of seedlings differed for all plantations, of which the lowest The greatest variation in height categories were observed at
was observed in BB and highest in BBD (Fig. 3). the time of plantation establishment.

123
20 G. Sukhbaatar et al.

100
20

Precipitation (mm)
80

Temperature (°C)
10
60 0

40 -10
-20
20
-30
0 -40
Jan
Feb

May
Jun
Jul
Aug
Sep

Nov
Dec
Oct
Mar
Apr

Jan
Feb

May
Jun
Jul
Aug
Sep

Nov
Dec
Oct
Mar
Apr
120
20
Precipitation (mm)

100

Temperature (°C)
10
80
0
60
-10
40 -20
20 -30
0 -40
Jan
Feb

May
Jun
Jul
Aug
Sep

Nov
Dec
Oct
Mar
Apr

Jan
Feb

May
Jun
Jul
Aug
Sep

Nov
Dec
Oct
Mar
Apr
Fig. 6 Average course of precipitation and temperature during the period 2003–2010 shown as solid line. Actual course of precipitation shown
as dashed line. Data from meteorological station Sukhbaatar

Table 4 Overview of the linear mixed effect models for survival rate in the planting year (2003–2008 respectively) and growth in the following
years (2004–2010); only models with a level of significance of the fixed effect value with p \ 0.05 are shown
Model Variable Fixed effects Degrees of freedom Model parameter (fixed effects) AIC of the model
Intercept p value Fixed effect p value

1. Survival_rate Survival Dmay 4 106.610 0.001 - 2.559 0.034 50.386

2. Growth Height growth H 20 - 41.179 0.000 0.759 0.000 149.620
Radial growth H 17 - 3.645 0.026 0.092 0.001 39.920
Dmay = number of dry days (days with critical low value of the average relative air humidity [below 30%] in May); H = average annual air
humidity (%); AIC = Akaike‘s Information Criterion. The minimum number of dry days was Dmay = 5

a 100 b 4 c 25

80 20
Height growth (cm)
Radial growth (mm)
Survival rate (%)

60 15
2
40 10

1 5
20

0 0
0 40 60 80 100
0 5 10 15 20 25 30 40 60 80 100
Dmay Average annual air humidity (%) Average annual air humidity (%)

Fig. 7 LMM - graphics: relationship between the number of dry Scots pine trees (a). Relationship between the average annual relative
days (Dmay = days with critical low value of the average relative air air humidity and radial growth (b) or height growth (c), respectively
humidity [below 30%] in May) and the survival rate of the planted

123
Assessment of early survival and growth of planted Scots pine (Pinus sylvestris) seedlings… 21

Comparative diagrams of the most positive and most planting to re-establish appropriate root architecture sys-
negative pointer years related to planting success tems able to support the growth of apical shoots. Thus, the
first months after planting are a decisive time for planting
In terms of seedling survival in the year of planting, 2003 success. Sufficient water supplies during the initial growing
was identified as a positive pointer year (survival rate [ season, especially immediately after planting, is a major
97%) and 2008 as negative pointer year (survival rate determining factor for reforestation success. Our data
\ 16%), as shown in Table 2. The annual sum of precip- analyses indicated that a lack of humidity during and
itation of the positive pointer year (2003) was below directly after initial planting causes mass mortality of
average but above average during May. Conversely, in the seedlings (Table 2; Fig. 6). On the other hand, sufficiently
negative pointer year (2008), the precipitation in May was humid conditions during the month of May allowed for
below average, whereas the sum of precipitation was rel- fairly good survival. This became especially clear through
atively high (Fig. 6). the comparative diagrams of the pointer years and the
survival model of seedlings. Mass mortality was observed
Modeling of the influence of dryness and humidity in 2008 (KGA) (Table 2), which may be explained by a
on the survival rate and growth of planted Scots pine lack of rainfall for May. Interestingly, instead of precipi-
tation itself, it was the number of dry days in May, which
Based on the results, linear mixed models were developed we consider as a kind of dryness indicator, which gave the
to predict first-year survival and growth rates over subse- best model results for seedling survival (Fig. 7a). Critical
quent years. Sufficient humidity levels significantly influ- dry periods in June 2006 (2.5 mm) and May 2008
enced survival rates of planted pine seedlings in the first (15.4 mm) resulted in relatively low survival of first-year
year and strongly contributed to seedling growth in sub- seedlings in plantations TN (61.7%) and KGA (15.3%),
sequent years. Different indicators, however, provided the respectively (Table 2). Meanwhile, in 2003, 2004 and
best results for predicting survival and growth rates. 2007, the amount of rainfall during the growing season was
The number of dry days in late spring (Dmay) had a relatively high and resulted in better survival rates, as
particularly strong impact on first-year survival and varied recorded in BB (97.9%), MG (92.6%) and BBD (78.4%)
considerably from year to year. With regard to seedling (Table 2). These observations indicate that weather con-
growth after plantation establishment (second year ditions during spring, specifically the month of May (ver-
onwards), average annual relative air humidity (H) pro- sus the sum of precipitation measured for the whole year),
vided the most reliable results for the whole observation played a crucial role for seedling survival in this region of
period. Both explanatory approaches (height growth and Mongolia. This study provides a quantitative orientation
radial growth) resulted in highly significant p values for for projected survival rates based on the number of dry
the explanatory variable. The variation of H was relatively days (D) in May (Fig. 7a). One of the reasons why weather
low throughout all the study years; see Table 4, Fig. 7 for a conditions in May were important in the analysis of seed-
graphical representation of the models. ling survival was that the seedlings were planted during
May. Establishing plantations in autumn may lead to dif-
ferent results, in terms of both individual tree survival and
Discussion fixed-effects on forest structural parameters. The results,
however, show that environmental conditions, both during
Climate factors determine plantation success and immediately after planting, represent a key factor in
the successful survival of seedlings in Mongolia.
In this study, seedling survival and growth in the forest
steppe of northern Mongolia are largely dependent on cli- Growth performance of Scots pine
matic factors. As expected, the amount of precipitation and
its distribution throughout the growing season is crucial. Regarding growth performance after plantation establish-
The study region is characterized by highly variable rain- ment, the pattern is somewhat different. Water availability
fall regimes and relatively low air humidity levels throughout the year is of much greater importance. The
throughout the growing season. correlation of height and radial growth with total annual
precipitation was higher than for the growing season
Survival rate is related to meteorological conditions (Table 3). The LMM results for height and radial growth
modeling showed that growth of the plantations can be
First-year survival of transplanted seedlings plays a crucial directly related to air humidity throughout the year
role in the subsequent success of plantations. Brunner et al. (Fig. 7b). Air humidity may be considered as an indicator
(2015) noted that seedlings might take some time after of planting success, which incorporates both precipitation

123
22 G. Sukhbaatar et al.

and temperature. In general, seedlings are typically more Forest plantations in the context of climate change
sensitive to water deficits than older trees due to their less in Mongolia
developed root systems. For example, a climate-growth
study in the Altansumber research area of the Mongolian Temperatures in the southern boreal forests of Eurasia have
mountain forest steppe showed that young birch trees were warmed rapidly in recent decades, particularly in spring,
more prone to react favorably to precipitation than older and this is projected to continue in coming decades (Schär
trees (Gradel et al. 2017b). However, these findings were et al. 2004; IPCC 2007). A number of studies have reported
observed in naturally regenerated forests. Scots pine is a that global warming, combined with increasing aridity, are
light-demanding species that is moderately drought resis- the main drivers of changes in species composition (Paw-
tant and has modest nutritional requirements which makes son et al. 2013; Benavides et al. 2015), and directly impact
it better suited to nutrient poor soils than other species on forest ecosystem rehabilitation (Hattori et al. 2013).
(Lawson and Michler 2014). The absence of shading from Consequently, forest vulnerability to climate change has
mature trees in stands might have had a favorable effect on increasingly becomes a topic of considerable interest to the
seedling growth during the early stages of plantation forestry sector, most notably in Eastern Asia (FAO 2010;
establishment. The largest differences between relative Cui et al. 2016). Recent studies have shown that semi-arid
height categories were observed in the first two years after ecosystems are particularly vulnerable to climate change
planting, which was mainly caused by individual adapta- and the increased risk of more frequent and severe droughts
tion to the new growing environment. The ever-increasing (Ma et al. 2015; Rammig and Mahecha 2015). A recent
proportion of average classified height categories and study indicated that the physiological responses of seed-
gradual reduction of accelerated height categories indicates lings to drought stress that ultimately causes them to die,
a low competition rate between individuals. Our results can differ between species (Ivanov et al. 2019). Findings
show that during the first years of plantation establishment, from this study identified carbon starvation, resulting from
variations in height among individual seedlings gradually rapid root growth at the onset of water stress, as the pri-
decreased (Fig. 5). This will most likely change as the trees mary cause of death for Scots pine seedlings. To date,
grow older and competition becomes more important. relatively few studies have addressed the effects of climate
Competition is another important factor affecting tree warming on seedling mortality and growth of forest plan-
growth (Gadow 2005; Gadow et al. 2012; Klädtke et al. tations in Mongolia (Dulamsuren et al. 2013). Forests have
2012; Gradel et al. 2017a). In the context of the current the potential to buffer climate effects, notably by protecting
developmental stage of the Scots pine plantations, drought the soil from high solar radiation and elevated summer
stress is the dominant limiting factor for seedling survival temperatures. In the context of global warming, effective
(Dulamsuren et al. 2013) and growth (Cregg and Zhang climate resilient afforestation and reforestation efforts have
2001; Turtola et al. 2003). It has also been shown that for become critically important for the future of Mongolia. The
mature Scots pine, a clear correlation exists between combined effects of accelerating climate change, ongoing
growth and water availability in the forest steppe zone desertification, and the ever-present risk of drought pro-
(Babushkina and Belokopytova 2014; Demina et al. 2017). jected to occur with higher frequency (Oyuntuya et al.
Water availability is considered the most important limit- 2015; Ubugunov et al. 2017), underline the need to develop
ing factor affecting plant productivity and is an important sustainable and effective forest planting practices to
ecosystem driver in general (Heisler-White et al. 2008; Liu respond to the unique set of conditions specific to Mon-
et al. 2013). Williams et al. (2013) highlighted the finding golia. Lower survival rates and largely ineffective refor-
that semi-arid forests experience seasonal water stress and estation efforts have limited forest plantations in Mongolia
may be particularly vulnerable to even slight increases in to only 300,000 ha. Approximately 30% of these have been
water deficit, which can inhibit tree growth and trigger established in the Selenge Aimag. Valuable insight can be
increased mortality. A study on tree distribution, growth gained from similar scenarios in other Asian countries, for
and survival within populations of Scots pine grown at sites example, by using containerized seedlings instead of bare-
in Europe and North America showed that even modest root seedlings. One option might be to incorporate the use
climate warming will likely have a positive influence on of mycorrhiza or endophytic fungi to enhance plant growth.
Scots pine survival and growth in the northern parts of the Mycorrhizal fungi are known to improve plant ability to
species range, and will negatively impact the more south- absorb both nutrients and water as they effectively increase
ern range of the species (Reich and Oleksyn 2008). the surface area of absorbing roots and, in general, lead to
improved plant health and biomass production (Dighton
and Skeffington 1987; Niemi et al. 2002; Alberton et al.
2010). For greater success in establishing plantations in
harsh environments under dry soil conditions, Cortina et al.

123
Assessment of early survival and growth of planted Scots pine (Pinus sylvestris) seedlings… 23

(2013) suggested planting seedlings previously cultivated Conclusions

in the open air and under nutrient-limited conditions
favorable for root development. An evaluation by Cao et al. The Tujyin Nars region is a representative site of suc-
(2011) concluded that four key factors should systemati- cessful tree planting in Mongolia, and provides ample
cally be considered during the early stages of afforestation opportunity to derive general conclusions and recommen-
projects: climate, pedology, hydrology and landscape dations for future planting programs. Our study showed
characteristics. In certain cases, integrating protective high survival for Scots pine after the first growing season.
coverage of native herb-layer vegetation is appropriate for Observations of steadily increasing height and radial
generating revenue for the local communities (e.g., medical increments, combined with a gradual reduction of growth
herbs) (Jiang et al. 2016). With regard to species adapt- variations, indicate considerable potential for plantation
ability to new environments with drier conditions, several establishment with this well-adapted native species. Sur-
studies have proposed provenance tests of different regio- vival of Scots pine seedlings are directly related to mete-
nal Scots pine (e.g., Gülcü and Bilir 2017). orological conditions in May (see Fig. 7a). This planting
period, however, coincides with unfavorable environmental
Ecosystem-based adaptation and climate-smart conditions such as prolonged drought and frequent wind-
forestry in Mongolia storms, both which effectively create conditions for mass
loss of seedlings. This study provides threshold limits for
A targeted approach, such as climate-smart forestry, to dry days in May that relate to plantation success. As cli-
build resilient forests in the mountain steppes may allow matic factors and time of planting have a major effect on
for a mixture of two or three species, including broad- plantation success, it is critical to rethink current planting
leaved pioneer species (e.g., birch or aspen). In reforesta- practices and research alternatives. Growth performance
tion efforts with mixed plantations, preference should be during subsequent years was best explained by relative air
given to native species and, where possible (depending on humidity levels throughout the entire growing season
specific site conditions and seedling availability), to more (Fig. 7b), which may be considered as an indicator of
than one native species. Reforestation with multiple native general water availability. Climate-resilient reforestation
species may help to ensure plantation success. In more guidelines should be developed for different regions
forested regions (e.g., Khentii Mountains), enrichment throughout Mongolia. Historical data on previous forest
planting might be considered as a means of protecting cover, soil characteristics, planting schemes, and optimal
existing stands and associated ecosystem services by fur- planting seasons should be considered. If the site was
ther adding more valuable species, e.g. Siberian larch previously forested, species selection should focus on the
(L. sibirica Ledeb.) or Siberian stone pine (Pinus sibirica natural dominant species.
Du Tour). Wang et al. (2012) recommended that forests
should be planted with an optimal structure and spatial Acknowledgements The authors would like to acknowledge col-
leagues at the Tujyin Nars SPA Administration for providing human
distribution adapted to the water-carrying capacity of local
resources necessary for field data collection. Our sincere thanks to
soils and groundwater conditions. It may therefore also be Matthias Meyer (TU Dresden) for valuable comments and to Aimee
interesting to research the effect of different planting Orsini (Berkeley, California) for the careful language editing of our
schemes, such as planting in groups and troops, on survival manuscript. We are thankful for the useful comments and recom-
mendations of two anonymous reviewers. The R-script for the test for
rate and growth. For example, a comparative study of
spatial autocorrelation is based on an earlier version from the Chair of
different planting schemes for oak (Quercus spp.) in Ger- Silviculture at TU Dresden (see Gradel et al. 2017a). Results of this
many, Austria and Switzerland found that troop planting study were presented at the GMIT Symposium on Environmental
resulted in better stem shape. Troop sizes with more than Science and Engineering held in Ulaanbaatar, Mongolia in August,
2018 (Gerelbaatar et al. 2018).
twelve individuals showed higher survival rates than
planting in rows (Saha et al. 2013). There is also an indi- Open Access This article is distributed under the terms of the
cation that, for different reasons, clumped tree distribution, Creative Commons Attribution 4.0 International License (http://crea
(potential effects of environmental disturbance and tivecommons.org/licenses/by/4.0/), which permits unrestricted use,
distribution, and reproduction in any medium, provided you give
microclimatic conditions), is a common and, in part, nat-
appropriate credit to the original author(s) and the source, provide a
ural feature of forest types in the Mongolian mountain link to the Creative Commons license, and indicate if changes were
forest steppes (e.g., larch and birch forests) (Gradel et al. made.
2015, 2017b). Landscape features should be considered,
e.g., on drier slopes to concentrate planting on the lower
parts between humps and small valleys and avoid planting
on dry humps.

123
24 G. Sukhbaatar et al.

References 107(1):191–202. https://doi.org/10.1111/j.1469-8137.1987.

tb04893.x
Alberton O, Kuyper TW, Summerbell RC (2010) Dark septate root Dulamsuren Ch, Hauk M, Leuschner C (2013) Seedling emergence
endophytic fungi increase growth of Scots pine seedlings under and establishment of Pinus sylvestris in the Mongolian forest-
steppe ecotone. Plant Ecol 214(1):139–152. https://doi.org/10.
elevated CO2 through enhanced nitrogen use efficiency. Plant
Soil 328(1–2):459–470 1007/s11258-012-0152-z
Babushkina EA, Belokopytova LV (2014) Climatic signal in radial Edwards AL (1976) The correlation coefficient. An introduction to
increment of conifers in forest-steppe of southern Sibiria and its Linear regression and Correlation. W. H. Freeman, San
Francisco, pp 33–46
dependence on local growing conditions. Russ J Ecol
45(5):325–332 Eycott AE, Watkinson AR, Dolman PM (2006) Ecological patterns of
Battles JJ, Robards T, Das A, Waring K, Keith Gilles J, Biging G, plant diversity in a plantation forest managed by clearfelling.
Schurr F (2008) Climate change impact on forest growth and tree J Appl Ecol 43(6):1160–1171
mortality: a data-driven modeling study in the mixed conifer FAO (2010) Forests and climate change in the Asia–Pacific region.
forest of the Sierra Nevada, California. Clim Change Forest and Climate Change Working Paper 7, Food and
87(1):193–213 Agriculture Organization of the United Nations, Rome, Italy,
Benavides R, Escudero A, Coll L, Ferrandis P, Gouriveau F, Hodar p 126
JA, Ogaya R, Rabasa SG, Granda E, Santamaria BP, Martinez- Fisher RA (1925) Statistical methods for research workers. Oliver and
Vilalta J, Zamaro R, Espelta JM, Penueles J, Valladares F (2015) Boyd (Edinburgh), p 336. http://www.haghish.com/resources/
Survival vs. growth trade-off in early recruitment challenges materials/Statistical_Methods_for_Research_Workers.pdf. Acces-
sed 20 Nov 2016
global warming impacts on Mediterranean mountain trees.
Perspect Plant Ecol Evol Syst 17(5):369–378 Fox J, Weisberg S (2011) An R companion to applied regression, 2nd
Bjornstad ON (2013) NCF: spatial nonparametric covariance func- edn. Sage, Thousand Oaks. http://socserv.socsci.mcmaster.ca/
tions. R Package Version 1.1-5. http://CRAN.R-project.org/ jfox/Books/Companion. Accessed 15 Sept 2017
package=ncf. Accessed 29 May 2016 Gadow KV (2005) Forsteinrichtung. Analyse und Entwurf der
Brunner I, Herzog C, Dawes MA, Arend M, Sperisen C (2015) How Waldentwicklung. Universitätsverlag Göttingen, Reihe Univer-
tree roots respond to drought. Front Plant Sci 6:1–16 sitätsdrucke, Göttingen, p 342
Buendia C, Bussi G, Tuset J, Vericat D, Sabater S, Palau A, Batalla Gadow KV, Zhang CY, Wehenkel C, Pommerening A, Corral-Rivas
RJ (2015) Effects of afforestation on runoff and sediment load in J, Korol M, Myklush S, Hui GY, Kiviste A, Zhao XH (2012)
an upland Mediterranean catchment. Sci Total Environ Forest structure and diversity. In: Pukkula T, Gadow KV (eds)
540(1):144–157 Continuous cover forestry. Managing forest ecosystems.
Springer, Dordrecht, pp 29–83
Cao SX, Chen L, Shankman D, Wang CM, Wang XB, Zhang H
(2011) Excessive reliance on afforestation in China’s arid and Gerelbaatar S (2012) Specifics of the formation of Scots pine (Pinus
semi-arid regions: lessons in ecological restoration. Earth Sci sylvestris L.) plantations. Ph.D. Thesis, National University of
Rev 104:240–245 Mongolia, Ulaanbaatar, NUM Press, Ulaanbaatar, p 117 (in
Mongolian)
Chamshama SAO, Nwonwu FOC, Lundgren B, Kowero GS (2009)
Plantation forestry in Sub Saharan Africa: silvicultural, ecolog- Gerelbaatar S, Batsaikhan G, Tsogtbaatar J, Battulga P, Batarbileg N,
ical and economic aspects. Discov Innov 21:42–49 Gradel A (2018) Early survival and growth of planted Scots pine
Chen YQ, Liu ZF, Rao XQ, Wang XL, Liang CF, Lin YB, Zhou LX, (Pinus sylvestris L.) seedlings in northern Mongolia. In: Vossen
Cai XA, Fu SL (2015) Carbon storage and allocation pattern in P, Karthe D, Gunsmaa B, Enkhjargal S (eds) Book of abstracts-
plant biomass among different forest plantation stands in GMIT symposium on environmental science and engineering,
Guangdong, China. Forests 6:794–808. https://doi.org/10.3390/ Nalaikh, pp 8–9. http://gmit.edu.mn/site/files/downloads/gmit_
f6030794 sese-2018_book-of-abstracts.pdf. Accessed 06 Dec 2018
Cortina J, Vilagrosa A, Trubat R (2013) The role of nutrients for Gradel A, Ochirragchaa N, Altaev AA, Voinkov AA, Enkhtuya B
improving seedling quality in drylands. New For 44:719–732 (2015) Spatial distribution of trees on light taiga plots before
Crawley MJ (2007) The R-book. Wiley, New York, p 942 selective thinning. Mong J Agric Sci 15(2):91–99
Gradel A, Ammer C, Batsaikhan G, Ochirragchaa N, Batdorj D,
Cregg BM, Zhang JW (2001) Physiology and morphology of Pinus
sylvestris seedlings from diverse sources under cyclic drought Wagner S (2017a) On the effect of thinning on tree growth and
stress. For Ecol Manag 154(1/2):131–139. https://doi.org/10. stand structure of white birch (Betula platyphylla Sukaczev) and
1016/S0378-1127(00)00626-5 Siberian larch (Larix sibirica Ledeb.) in Mongolia. Forests
Crisp N, Dick J, Mullins M (2004) Mongolia forestry sector review 8(4):105. https://doi.org/10.3390/f8040105
(English, Mongolian). World Bank, Washington, p 146. http:// Gradel A, Haensch C, Ganbaatar B, Batdorj D, Ochirragchaa N,
documents.worldbank.org/curated/en/805041468773956869/ Günther B (2017b) Response of white birch (Betula platyphylla
Mongolia-forestry-sector-review. Accessed 20 Nov 2016 Sukaczev) to temperature and precipitation in the mountain
Cui GS, Kwak H, Choi S, Kim M, Lim CH, Lee WK, Kim JS, Chae Y forest steppe and taiga of northern Mongolia. Dendrochronologia
(2016) Assessing vulnerability of forests to climate change in 4:24–33. https://doi.org/10.1016/j.dendro.2016.03.005
South Korea. J For Res 27(3):489–503. https://doi.org/10.1007/ Grothendieck G (2013) nls2: Non-linear regression with brute force.
R Package Version 0.2. http://CRAN.R-project.org/package=
s11676-015-0201-2
Demina AV, Belokopytova LV, Andreev SG, Kostyakova TV, nls2. Accessed 28 Nov 2017
Babushkina EA (2017) Radial increment dynamics of Scots Gülcü S, Bilir N (2017) Growth and survival variation among Scots
pine (Pinus sylvestris L.) as an indicator of hydrothermal regime pine (Pinus sylvestris L.) provenances. Int J Genom
2017:1904623. https://doi.org/10.1155/2017/1904623
of the Western Transbaikalia forest steppe. Contemp Probl Ecol
10(5):476–487. https://link.springer.com/article/10.1134% Hattori D, Kenzo T, Kendawang JJ, Irino KO, Tanaka S, Ichie T,
2FS1995425517050031. Accessed 24 March 2018 Ninomiya I, Sakurai K (2009) Effect of light intensity and soil
Dighton J, Skeffington RA (1987) Effects of artificial acid precipi- physic-chemical properties on seedling mortality and growth of
tation on the mycorrhizas of Scots pine seedlings. New Phytol six dipterocarp species planted for rehabilitation of degraded
grassland, secondary forest and logged forests in Sarawak.

123
Assessment of early survival and growth of planted Scots pine (Pinus sylvestris) seedlings… 25

Malaysia. Jpn J For Environ 51(2):105–115. https://doi.org/10. Mühlenberg M, Batkhishig T, Dashzeveg TS, Drößler L, Neusel B,
18922/jjfe.51.2_105 Tsogtbaatar J (2006) Lessons from tree planting initatives in
Hattori D, Kenzo T, Kendawang JJ, Irino K, Tanaka S, Tomoaki I, Mongolia. Mongolia Discussion papers, East Asia and Pacific
Ninomiya I, Sakurai K (2013) Effect of environmental factors on Environment and Social Development Department. World Bank,
growth and mortality of Parashoreea macrophylla (Diptero- Washington, p 38. http://documents.worldbank.org/curated/en/
carpaceae) planted on slopes and valleys in a degraded tropical 143891468059947579/pdf/377950MOG0less1ing0P09260901
secondary forest in Sarawak, Malaysia. Soil Sci Plant Nutr PUBLIC1.pdf. Accessed 28 Nov 2017
59:218–228. https://doi.org/10.1080/00380768.2012.762895 Mühlenberg M, Appelfelder H, Hoffman H, Ayush E, Wilson KJ
Hector A, Bagchi R (2007) Biodiversity and ecosystem multifunc- (2012) Structure of the montane taiga forests of West Khentii,
tionality. Nature 448:188–190 Northern Mongolia. J For Sci 58:45–56
Heisler-White JL, Knapp AK, Kelly EF (2008) Increasing precipi- Niemi K, Vuorinen T, Ernstsen A (2002) Ectomycorrhizal fungi and
tation event size increases aboveground net primary productivity exogenous auxins influence root and mycorrhiza formation of
in a semi-arid grassland. Oecologia 158(1):129–140 Scots pine hypocotyl cuttings in vitro. Tree Physiol
IPCC (2007) Climate change 2007: The physical science basis. IPCC 22(17):1231–1239. https://doi.org/10.1093/treephys/22.17.1231
Fourth Assessment Report. Contribution of working group I to O’Brien RG (1979) A general ANOVA method for robust tests of
the Fourth Assessment Report of the intergovernmental panel on additive models for variances. J Am Stat Assoc 74(368):877–880
climate change. IPCC, Geneva. Switzerland. Cambridge Univer- Oscarsson H, Sigurgeirsson A, Raulund-Rasmussen K (2006) Sur-
sity Press, Cambridge, p 996 vival, growth, and nutrition of tree seedlings fertilized at planting
Ivanov YV, Kartashov AV, Zlobin IE, Sarvin B, Stavrianidi AN, on Andisol soils in Iceland: six-year results. For Ecol Manag
Kuznetsov VV (2019) Water deficit-dependent changes in non- 229:88–97
structural carbohydrate profiles, growth and mortality of pine Osunkoya OO, Othman FE, Kahar RS (2005) Growth and compe-
and spruce seedlings in hydroculture. Environ Exp Bot tition between seedlings of an invasive plantation tree, Acacia
157:151–160. https://doi.org/10.1016/j.envexpbot.2018.10.016 mangium, and those of a native Borneo heath-forest species,
Jiang WY, Yang SL, Yang XX, Gu N (2016) Negative impacts of Melastoma beccarianum. Ecol Res 20(2):205–214
afforestation and economic forestry on the Chinese Loess Oyuntuya S, Dorj B, Shurentsetseg B, Bayarjargal E (2015)
Plateau and proposed solutions. Quatern Int 399:165–173 Agrometeorological information for the adaptation to climate
JICA (1998) The forest resources management study in Selenge change. In: Badmaev NB, Khutakova CB (eds) Soils of steppe
province. Final report. Asia Air Survey Co., Ltd, Tokyo, p 116 and forest steppe ecosystems of inner Asia and problems of their
Kahle P, Baum C, Boelcke B (2005) Effect of afforestation on soil sustainable utilization: international scientific conference. Buryat
properties and mycorrhizal formation. Pedosphere State Academy of Agriculture named after V.R. Philipov, Ulan-
15(6):754–760 Ude, pp 135–140
Klädtke J, Kohnle U, Kublin E, Ehring A, Pretzsch H, Uhl E, Pawson SM, Brin A, Brockerhoff EG, Lamb D, Payn A, Paquette A,
Spellmann H, Weller A (2012) Wachstum und wertleistung der Parrotta JA (2013) Plantation forests, climate change and
douglasie in Abhängigkeit von der Standraumgestaltung. Sch- biodiversity. Biodivers Conserv 22(5):1203–1227
weiz Z Forstwes 163:96–104 (in German) Peel MC, Finlayson BL, McMahon TA (2007) Updated world map of
Kongsager R, Napier J, Mertz O (2013) The carbon sequestration the Köppen–Geiger climate classification. Hydrol Earth Syst Sci
potential of tree crop plantations. Mitig Adapt Strat Glob Change Eur Geosci Union 11(5):1633–1644. https://doi.org/10.5194/
18:1197–1213 hess-11-1633-2007
Krstic M, Stavretovic N, Isajev V, Bjelanovic I (2012) Crown Pinheiro J, Bates D, DebRoy S, Sarkar D, R Core Team (2018) nlme:
structure of Picea omorika trees in the plantation. Arch Biol Sci linear and nonlinear mixed effects models. R Package version
64(2):605–611 3.1-137. https://CRAN.R-project.org/package=nlme. Accessed
Kunstler G, Curt T, Bouchaud M, Lepart J (2005) Growth, mortality 09 Nov 2017
and morphological response of European beech and downy oak Pirard R, Secco LD, Russell R (2016) Do timber plantations
along a light gradient in sub-Mediterranean forest. Can J For Res contribute to forest conservation? Environ Sci Policy
35(7):1657–1668 57:122–130
Lawson SS, Michler CH (2014) Afforestation, restoration and R Development Core Team (2016) R: a language and environment for
regeneration—not all trees are created equal. J For Res statistical computing; Version 3.0.2. http://packages.renjin.org/
25(1):3–20 package/org.renjin.cran/nlme/3.1-113. Accessed 28 Nov 2016
Liu HY, Williams AP, Allen CD, Guo DL, Wu XC, Anenkhonov OA, Rammig A, Mahecha MD (2015) Ecosystem responses to climate
Liang EY, Sandanov DV, Yin Y, Qi ZH, Badmaeva NK (2013) extremes. Nature 527:315–316
Rapid warming accelerates tree growth decline in semi-arid R-bloggers (2011) Linear mixed models. Post by Luis. https://www.r-
forests of Inner Asia. Glob Change Biol 19(8):2500–2510 bloggers.com/linear-mixed-models-in-r/. Accessed 24 Nov 2017
Löf M, Karlsson M, Sonesson K, Welander T (2007) Growth and Regzedmaa M (2008) Climate resources and changes of meteorolog-
mortality in underplanted tree seedlings in response to variations ical data of Selenge Province. Selenge Press, Sukhbaatar,
in canopy closure of Norway spruce stands. Forestry pp 11–25 (in Mongolian)
80(4):371–383 Reich PB, Oleksyn J (2008) Climate warming will reduce growth and
Ma XQ, Heal KV, Liu AQ, Jarvis PG (2007) Nutrient cycling and survival of Scots pine except in the far north. Ecol Lett
distribution in different-aged plantations of Chinese fir in 11(6):588–597. https://doi.org/10.1111/j.1461-0248.2008.01172.
southern China. For Ecol Manag 243:61–74 x
Ma XL, Huete A, Moran S, Ponce-Campos G, Eamus D (2015) Robinson AP, Hamann JD (2011) Forest analytics with R—an
Abrupt shifts in phenology and vegetation productivity under introduction. Springer, New York, p 354
climate extremes. JGR Biogeosci 120:2036–2052. https://doi. Saha S, Kühne C, Kohnle U, Bauhus J (2013) Eignung von Nester-
org/10.1002/2015JG003144 und Trupppflanzungen für die Begründung von Eichenbestän-
MPNFI (2016) Mongolian multipurpose national forest inventory den. AFZ-Der Wald 2(2013):37–39 (in German)
report (2014–2016). Ministry of Nature and Environment, Sarkar D (2008) Lattice: multivariate data visualization with R.
Ulaanbaatar, p 126 Springer, New York, p 268

123
26 G. Sukhbaatar et al.

Schär C, Vidale PL, Lüthi D, Frei C, Häberli C, Liniger MA, steppe ecosystems in the Barguzin depression. Arid Ecosyst
Appenzeller C (2004) The role of increasing temperature 7(3):142–154
variability in European summer heatwaves. Nature Wang YH, Bonell M, Feger KH, Yu PT, Xiong W, Xu LH (2012)
427:332–336. https://doi.org/10.1038/nature0230010.1038/ Changing forestry policy by integrating water aspects into forest/
nature02300 vegetation restoration in dryland areas in China. Bull Chin Acad
Soehartono T, Newton A, Mardiastuti A (2002) Factors influencing Sci 26:59–67. http://english.cas.cn/bcas/2012_1/201411/
the survival and growth of Aquilaria Malaccensis seedling in P020141121533282744018.pdf
Indonesia. J Trop For Sci 14(3):364–378 Williams AP, Allen CD, Macalady AK (2013) Temperature as a
Tsogtbaatar J (2004) Deforestation and reforestation needs in potent driver of regional forest drought stress and tree mortality.
Mongolia. For Ecol Manag 201(1):57–63 Nat Clim Change 3:292–297
Turtola S, Manninen AM, Rikala R, Kainulainen P (2003) Drought Zuur AF, Ieno EN, Walker NJ, Saveliev AA, Smith GM (2009)
stress alters the concentration on wood terpenoids in Scots pine Mixed effects models and extensions in ecology with R.
and Norway Spruce seedlings. J Chem Ecol 29(9):1981–1995. Springer, New York, p 574
https://doi.org/10.1023/A:1025674116183
Ubugunov VL, Gunin PD, Bazha SN, Drobyshev YuI, Ubugunova VI Publisher’s Note Springer Nature remains neutral with regard to
(2017) Soil desiccation as indicator of desertification of forest- jurisdictional claims in published maps and institutional affiliations.

123