Forest Ecology and Management 465 (2020) 118103

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Forest Ecology and Management

journal homepage: www.elsevier.com/locate/foreco

Relative contributions of competition, stand structure, age, and climate T

factors to tree mortality of Chinese fir plantations: Long-term spacing trials
in southern China
Xiongqing Zhanga,b, Zhen Wanga, Sophan Chhinc, Hanchen Wanga, Aiguo Duana,
⁎
Jianguo Zhanga,b,
a
State Key Laboratory of Tree Genetics and Breeding, Key Laboratory of Tree Breeding and Cultivation of the State Forestry Administration, Research Institute of Forestry,
Chinese Academy of Forestry, Beijing 100091, PR China
b
Collaborative Innovation Center of Sustainable Forestry in Southern China, Nanjing Forestry University, Nanjing 210037, PR China
c
Division of Forestry and Natural Resources, West Virginia University, 322 Percival Hall, PO Box 6125, Morgantown, WV 26506, USA

A R T I C LE I N FO A B S T R A C T

Keywords: Tree mortality is an important process during forest succession, influencing forest structure, composition, and
Tree mortality ecosystem services. Chinese fir (Cunninghamia lanceolata (Lamb.) Hook.), a native and fast growing species with
Competition high quality timber, is widely planted in southern China. Yet the mechanisms leading to tree mortality of Chinese
Age fir plantations under climate change conditions is still not clearly understood. A long-term spacing trial including
Stand structure
60 permanent plots located in Fujian, Jiangxi, Guangxi, and Sichuan provinces in southern China were used to
Climate
explore tree mortality in relation to competition, age, structural diversity, and climatic variables using a logistic
Chinese fir
mixed effects model. Results showed that tree mortality increased with increasing LnN (natural logarithm of
number of trees per ha), tree density index (TN), Gini coefficient, age (A), LnAP (natural logarithm of annual
precipitation), and mean warmest month temperature, and decreased with increasing relative density (RD),
inverse of diameter (1/D), and mean annual temperature. The relative contributions of competition, age and
structure variables to mortality were 88.82%, much larger than climate variables 11.18%. In addition, in view of
the contributions of stand factors, competition indices including RD, LnN, TN were 50.57%, much larger than
age (A, 1/D) contribution of 37.45%, and structural diversity contribution of 0.80%, which indicated that
competition was the main driver of tree mortality. Our findings highlight the mechanisms of tree mortality of
Chinese fir in southern China under climate change.

1. Introduction
Tree mortality is a fundamental process during forest succession,
affecting forest structure, composition, and ecosystem services (Lutz
and Halpern, 2006; Luo and Chen, 2011; Zhang et al., 2017). Even
small changes in tree mortality can lead to changes in the lifespan of
trees, biodiversity, and the cycling of carbon and nutrients (Dietze and
Moorcroft, 2011). The generally understood direct cause of tree mor-
tality is carbon starvation indicating that photosynthetic production
that is lower than respiratory losses can lead to failure to support the
metabolic costs of maintaining tissues (Guneralp and Gertner, 2007;
Adams et al., 2009) or defend against disturbance agents such as insect
pests (Senecal et al., 2004; Zhang et al., 2014a), and mechanical da-
mage (Ryall and Smith, 2005; Negrón-Juárez et al., 2010).
Many factors can lead to tree death and can be categorized into two
major groups: endogenous and exogenous variables (van Mantgem
et al., 2009; Lu et al., 2019). Among the endogenous variables, stand
structure can affect tree mortality since it will lead to differences in
competition intensity and in resource utilization (Kweon and Comeau,
2019). A stand structure index usually incorporates a measure of spatial
heterogeneity or tree size variation (i.e., diameter and height) alone or
together (Buongiorno et al., 1994; McElhinny et al., 2005; McRoberts
et al., 2008). Size inequality is often quantified using the Gini coeffi-
cient (Weiner and Solbrig, 1984; Sun et al., 2018), Shannon index
(Shannon and Weaver, 1949) or size variation coefficient
(Staudhammer and Lemay, 2001). Size inequality or spatial hetero-
geneity results in a size hierarchy among individual trees in a stand
(Thomas and Weiner, 1989) and affects competition processes during

⁎
Corresponding author at: State Key Laboratory of Tree Genetics and Breeding, Key Laboratory of Tree Breeding and Cultivation of the State Forestry
Administration, Research Institute of Forestry, Chinese Academy of Forestry, Beijing 100091, PR China.
E-mail address: zhangjg@caf.ac.cn (J. Zhang).

https://doi.org/10.1016/j.foreco.2020.118103
Received 3 February 2020; Received in revised form 16 March 2020; Accepted 20 March 2020
0378-1127/ © 2020 Elsevier B.V. All rights reserved.
X. Zhang, et al.
Fig. 1. The location of study sites of Chinese fir plantations in southern China (altitude from low to high).
forest succession (Cordonnier and Kunstler, 2015). Therefore, stand
structure has been considered as an important predictor for predicting
future growth, productivity and mortality (Liang et al., 2005; Lei et al.,
2009; Cortini et al., 2017). However, the effects of structural diversity
such as spatial heterogeneity or size inequality on tree mortality (Lei
et al., 2009; Kweon and Comeau, 2019) are rarely explored.
Competition is another key endogenous constraint on tree growth
through competition for light, water, nutrients which in turn affects the
probability of tree mortality (Franklin et al., 2002; Zhang et al., 2017).
There are two types of competition: one is size-symmetric or one-sided
competition when belowground resources are limited; the other is size-
asymmetric or two-sided competition when growth is mainly con-
strained by light availability (Pretzsch and Biber, 2010). Asymmetric
competition is a critical mechanism influencing tree size equality
(Schwinning and Weiner, 1998; Bourdier et al., 2016). Trees with small
size are expected to suffer strong asymmetric competition from domi-
nant trees and more likely to die due to growth stagnation (Bravo-
Oviedo et al., 2006; Luo and Chen, 2011). Additionally, age also effects
tree mortality, due to senescence and reduction in growth vigor and
increases in agent-based mortality, such as insect pests, fungal patho-
gens, and drought (Franklin and Van Pelt, 2004; Bennett et al., 2015).
In addition to endogenous variables, climate factors have been
viewed as important exogenous factors influencing tree mortality
(Adams et al., 2009; Zhang et al., 2014b). Recent studies have shown
that changes in temperature and precipitation, especially drought
stress, could increase tree mortality (van Mantgem et al., 2009; Peng
et al., 2011). Allen et al. (2010) assessed global tree mortality and at-
tributed it to drought and heat stress and revealed emerging climate
change risks for global forests. However, tree mortality patterns in re-
sponse to climate change are different in stands with different struc-
tures, competition intensity, age, and site conditions (Dietze and
Moorcroft, 2011; Bell et al., 2014; Merlin et al., 2015). Luo and Chen
(2013) found that tree mortality induced by regional warming and
drought is higher in old forests as a result of a higher degree of com-
petition for resources in younger forests. Bell et al. (2014) concluded
that the effects of drought stress on tree mortality of quaking aspen
(Populus tremuloides) are mediated by stand structure, competition, and
2
Forest Ecology and Management 465 (2020) 118103
age. Since the effects of climate change on tree mortality may vary
dependent on competition, structure, forest age, site condition, and
other factors, an understanding on how these factors influence tree
mortality will assist with managing forests under future climate change.
However, a major challenge for foresters is to predict tree mortality
across different site conditions and different stages of forest succession.
Chinese fir (Cunninghamia lanceolata (Lamb.) Hook), a native and
fast-growing trees species, is widely planted in south of China. Due to
its high quality timber with straight shape and high resistance to decay,
Chinese fir is one of the most important tree species for timber pro-
duction and has more than 1000 years of planting history (Zhang et al.,
2019a). Effective and sustainable management of this vastly distributed
species needs to disentangle the effect of competition, structure, age,
site conditions, and climate change on tree mortality. Zhang et al.
(2017) modeled tree mortality in relation to climate, initial planting
density, and competition in Chinese fir plantations in Jiangxi province
using a Bayesian logistic multilevel method, and identified different
sources of variation in tree mortality from the plot- and tree-level. Lu
et al. (2019) explored the relationships between tree mortality of Chi-
nese fir and site, competition, and climate variables in Fujian province
through a Bayesian model averaging approach and found that mortality
increased with high competition and decreased with increasing tem-
perature. However, it still unclear whether patterns of tree mortality are
influenced by competition, structure, age, and climate at a larger spatial
scale and what the contributions of these variables to tree mortality of
Chinese fir plantations in southern China remains unclear. In other
words, the mechanisms leading to tree mortality of Chinese fir planta-
tions are still not well understood.
In the present study, the objectives of this study are: (1) to model
tree mortality of Chinese fir in relation to competition, structure, age,
site, and climate variables and (2) to disentangle and rank the relative
contributions of these variables to tree mortality. The findings will be
helpful for understanding the mechanisms causing tree mortality and
assist with optimization of silviculture practices for maintaining the
sustainability of Chinese fir in southern China under future climate
change.
X. Zhang, et al. Forest Ecology and Management 465 (2020) 118103

2. Materials and methods 2.3. Endogenous variables

2.1. Study sites and data
In this study, four Chinese fir experiments were established using
bare-root seedlings. One experiment was established in 1981 in Jiangxi
province, and the other three experiments were established in 1982 in
Fujian, Guangxi, and Sichuan provinces, in China (Fig. 1). The study
sites located in Fujian, Jiangxi, and Sichuan belong to middle-sub-
tropical climate zone. Guangxi is classified in the southern-subtropical
climate zone. The soils in Fujian, Guangxi, and Sichuan are mainly red
earth developed on granite and other parent materials, but in Jiangxi
the soil is yellow-brown earth developed on sand shale.
Each province's experiment contains five planting spacings: A
(2 m × 3 m, 1667 trees/ha), B (2 m × 1.5 m, 3333 trees/ha), C
(2 m × 1 m, 5000 trees/ha), D (1 m × 1.5 m, 6667 trees/ha), and E
(1 m × 1 m, 10,000 trees/ha). Each spacing level was replicated three
times, and a total of 15 plots were established at each site. Each plot
comprised an area of 20 m × 30 m surrounded by a 2-row buffer zone
consisting of similarly treated trees. All trees were tagged, and diameter
at breast height (D) was measured after the tree height reached 1.3 m.
In each plot, more than 50 trees were randomly selected and measured
for tree height and the stand dominant height was regarded as an
average height of the tallest six trees. Sampling was performed in the
winter every 1–3 years. Table 1 describes statistics of tree and stand
variables at each site and includes the measurement period and fre-
quency of each site location.
2.2. Climate data
To explore the effects of climate change on tree mortality of Chinese
fir plantations, we obtained climate variables of each site using
ClimateAP (Wang et al., 2012) through spatially interpolated estima-
tion according to the sites’ latitude, longitude, and elevation. Generally,
both temperature and water supply influence physiology and growth
processes of forests (Oberhuber, 2004; Ryan, 2011; Anderegg et al.,
2015). In this study, eight climate variables including mean annual
temperature (MAT, °C), annual precipitation (AP, mm), mean warmest
month temperature (MWMT, °C), mean coldest month temperature
(MCMT, °C), degree-days below 0 °C (DD0), summer mean maximum
temperature (SMMT, °C), winter mean minimum temperature (WMMT,
°C), and summer mean temperature (SMT, °C) were used to model tree
mortality. In addition, the annual heat-moisture (AHM), an inverse
form of the De Martonne aridity index (De Martonne, 1926) was also
incorporated to the model, since it integrates MAT and AP into a
combined parameter: AHM = (MAT + 10)/(AP /1000) . Larger AHM va-
lues suggest drier conditions, while lower AHM values indicate rela-
tively wetter conditions. The statistics of the climate data for each study
site are shown in Table 2.
Here, we divided the endogenous variables into four categories:
competition index, structural diversity, size, site condition. Competition
can be divided into one-sided (size-asymmetric) and two-sided (size-
symmetric) competition (Yang et al., 2003; Pretzsch and Biber, 2010).
In two-sided competition, the indexes including stand basal area (Ba),
number of trees per hectare (N), and quadratic mean diameter (Dq)
were used. In one-sided competition, here we used sum of basal areas of
trees larger than the subject tree (BAL), relative diameter (RD = D/Dq),
and tree density index (TN). BAL is commonly considered as an in-
dicator of one-sided competition and has been used to model mortality
(Monserud and Sterba, 1999; Bravo-Oviedo et al. 2006). Stage (1968)
proposed a tree-by-tree measure (here we refer to it as tree density
index, TN) of site utilization related to stand density index (SDI). Stage
(1968) indicated that SDI should be additive and calculated tree by tree
within a stand for describing the relative composition of the growing
N
stock according to sizes: SDI = ∑ TN , and TN = a + bD 2 , where
i=1
a = 0.0049Dq1.605 , b = 0.0199Dq−0.395 .
Two main indexes quantifying stand structural diversity are
Shannon index and Gini coefficient (Lei et al., 2009). Shannon index is
a widely used index as a measure of structural diversity in forest
management (Neumann and Starlinger, 2001; Vanhellemont et al.,
2018). However, Shannon index is sensitive to changes in class width.
In contrast, the Gini coefficient, the other measurement of size di-
versity, was found superior for measuring structural diversity because it
does not require arbitrarily classified diameter classes (Lexerød and Eid,
2006). Since the Gini values range from 0 to 1, the index is easy to
interpret and has been widely used for quantifying structural diversity
in a stand (Peck et al., 2014; Pach and Podlaski, 2015; Soares et al.,
2016). Larger values indicate greater size diversity. Here, we used the
Gini coefficient instead of Shannon index to measuring the structural
diversity of Chinese fir plantations. Gini coefficient was calculated
using “ineq” package in R (Zeileis, 2014; R Development Core Team,
2018) to represent the degree of diameter inequality in a stand. The
statistics of Gini coefficient of size (diameter) diversity can be found in
Table 1.
The measures of tree age (size) used in this study were the inverse of
diameter (1/D) and tree age (A). Some studies (Diéguez-Aranda et al.,
2005; Lu et al., 2019) have reported that tree mortality rates were
different under different site conditions. A common measure of site
condition is stand dominant height (Hd) (Bravo-Oviedo et al., 2008). In
addition, to account for the effect of initial spacing on tree mortality,
the planting density (IPD) was also regarded as a candidate predictor.
To avoid multicollinearity between any two predictors, the variance
inflation factor (VIF) test was performed through DAAG package in R. A
common thumb is that multicollinearity does not exist among variables
when VIF < 5.

Table 1
Measurement years and summary statistics for stand- and tree- level factors by site.
Site Measurement year A Dq N Gini D

Mean SD Mean SD Mean SD Mean SD Mean SD

Fujian 1985–1990: every year 15 8.53 11.52 5.40 4431 2592.90 0.138 0.032 9.45 5.30
1990–2010: every 2 years
Jiangxi 1985–1989: every year 14 7.89 10.60 4.16 4615 2484.69 0.144 0.033 8.90 4.34
1989–2007: every 2 years
Guangxi 1990–1995: every year 19 6.62 12.60 3.09 3193 1837.70 0.138 0.017 10.78 4.01
1995–2012: every 2 years
Sichuan 1985–1995: every year 15 8.47 9.72 3.40 4143 2527.43 0.144 0.028 8.12 3.65
1995–2012: every 2 or 3 years

Note: A: stand age (year); Dq: stand quadratic mean diameter (cm); N: number of trees per ha; Gini: Gini coefficient of size (diameter) diversity; D: tree diameter at
breast height (cm); SD = standard deviation.

3
X. Zhang, et al. Forest Ecology and Management 465 (2020) 118103

Table 2
Summary statistics of climate factors for each site. Values in parentheses are minimum and maximum values.
Climate factors Fujian Guangxi Jiangxi Sichuan

MAT (°C) 18.91 (18.1, 19.6) 22.24(21.9, 22.6) 18.24(17.3, 19.3) 18.39(17.9, 19.1)
MWMT (°C) 28.22 (27.2, 30) 28.25 (27.5, 28.9) 28.58(27.1, 30.3) 28.2(26.9, 30.4)
MCMT (°C) 8.58(5.8, 10.2) 13.65(12.2, 15.8) 6.84(4.1, 9.1) 8.43(6.1, 10)
AP (mm) 1809.53(1453, 2350) 1466(1203, 1912) 1602.45(1334, 1899) 1187.47(1028, 1440)
AHM 16.25(12.5, 20.4) 22.29(17, 26.9) 17.83(14.6, 21.1) 24.11(19.4, 28)
DD0 1.41(1, 3) 0(0, 0) 3.19 (1, 9) 1.6(1, 3)
DD5 5050.18(4783, 5297) 6219.46(6099, 6354) 4830.63(4573, 5198) 4865.73(4696, 5111)
SMMT (°C) 32.19(31.5, 33.6) 31.8(31.1, 32.5) 32.20(30.7, 33.4) 30.72(29.8,32.6)
WMMT (°C) 4.71(2.5, 6) 12.16(11.3, 13) 4.37(2.3, 6) 7.15(5.7, 8.4)
SMT (°C) 18.39(16.9, 19.6) 22.74(22.2, 23.4) 17.68(16.7, 18.7) 18.63(17.9, 19.6)

Note: MAT: mean annual temperature; AP: annual precipitation; MWMT: mean warmest month temperature; MCMT: mean coldest month temperature; AHM: annual
heat-moisture index; DD0: degree-days below 0 °C; DD5: degree-days above 5 °C; SMMT: summer (Jun. - Aug.) mean maximum temperature; WMMT: winter (Dec
(prev. yr) - Feb.) mean minimum temperature; SMT: spring (Mar. - May) mean temperature.

2.4. Model development
Logistic regression model has been regarded as a preferred choice
and has been widely used for modelling tree mortality (Hamilton, 1974;
Zhang et al., 2017; Zhang et al., 2019b):
Ps = {1 + exp[−(α1 + βX )]}−q
where Ps is survival probability, X is a vector of predictors, including
competition, structure (Gini coefficient), size, site condition, initial
planting density (IPD) and climate variables. α1 is the intercept, β is a
vector of parameters including intercept, and q is the measurement
interval.
Since data collected from four sites and 15 plots in each site, the
random effects of site and plot were added to the intercept of the tree
mortality model. In addition, we used the unstructured covariance
structure (Littell et al., 1996) for describing the variance-covariance
structure of random effects. The parameter estimation was im-
plemented using the procedure NLMIXED in SAS (SAS Institute, 2011).
2.5. Model evaluation
increase in Fujian (p = 0.004), and Jiangxi (p = 0.001), but no sig-
nificant change in Guangxi and Sichuan (p = 0.157 and 0.500, re-
spectively). AP increased significantly in Fujian (p = 0.042), but there
were no significant changes in Guangxi, Jiangxi, and Sichuan (0.362,
0.89, and 0.384, respectively) (Fig. 3).
(1)
3.2. Tree survival model
In order to reduce the heteroskedasticity of the tree survival model,
we transformed N and AP into their natural logarithms. Accounting for
the random effects of site and plot on tree mortality, the site- and plot-
level random-effects parameter were incorporated into the intercept.
However, based on our preliminary analysis, it was difficult to achieve
convergence in the analysis if we considered the two levels together.
Therefore, only the site- or plot-level random effects parameter was
incorporated to the model, which was defined as: μÑ(0, v ) , where v is
the variance.
Ps = {1 + exp[−(μ + α1 + β1 LnN + β2 RD + β3 TN + β4 Gini + β5 A
+ β6/ D + β7 MAT + β8 LnAP + β9 WMMT )]}−q (2)

ROC (receiver operating characteristic) curve has been widely re-
garded as a criterion to evaluate tree mortality models (Cortini et al.,
2017; Kweon and Comeau, 2019). The AUC (the area under the ROC)
value reflects the performance of the mortality model: the larger the
value of AUC, the better the model performs (Hosmer and Lemeshow,
2000). A common rule for AUC is: 0.9–1 = excellent; 0.8–0.9 = good;
0.7–0.8 = fair; 0.6–0.7 = poor; and 0.5–0.6 = fail (Zhang et al., 2011).
For disentangling the relative contributions of competition, struc-
ture, age, site, and climate variables in the tree mortality model, hier-
archical partitioning (HP) analysis was used, which was implemented
through the hier.part package in R (Mac Nally and Walsh, 2004). HP
analysis compares the influence of each independent variable on a re-
sponse variable over the entire hierarchy of all models (Chevan and
Sutherland, 1991).
3. Results
3.1. Changes of Gini coefficient and climate variables
Based on the VIF test, there was no multicollinearity among N, RD,
TN, Gini, A, 1/D, MAT, AP, and WMMT (VIF < 5). The Gini value in
the four sites were nearly equal, with a Gini value of 0.138 in Fujian
and Guangxi, and a Gini value of 0.144 in Jiangxi and Sichuan
(Table 1). The Gini value decreased until about age 10, then increased
moderately up to age 25, and decreased after age 25 (Fig. 2). We also
found that the Gini value increased with an increase in stand density
(trees per ha) (Fig. 2).
During the time period of the study, MAT showed a significant
All of the variables selected by the VIF test were significant
(p < 0.05) (Table 3). Comparing the site-level random effects model
with plot-level model, we found that plot-level random effects model
(AIC = 64700) performed better than the site-level model
(AIC = 64881). AUC value from the plot-level random effects model
was 0.8965 (Fig. 4) showed that the model fitted the data and predicted
tree survival morality correctly. The subsequent results were all derived
from the plot-level random effects model.
3.3. Contributions of competition, structure, age, and climate factors to tree
mortality
Trees were likely to survive with increasing RD and 1/D (Table 3).
However, the effects of LnN, TN, Gini, and age had negative effects
(Table 3, Fig. 5). In view of the effects of climate on survival, trees were
likely to be alive with increasing MAT. However, survival decreased
with increasing LnAP and WMMT (Table 3). According to the HP
analysis, the relative contribution of RD to tree survival was the most,
followed by A, TN, MWMT, 1/D, MAT, LnN, lnAP, and Gini coefficient
(Table 4). The relative contributions of competition, age and structure
variables to survival were 88.82%, much larger than climate variables
(11.18%) (Table 4, Fig. 6). Furthermore, the relative contributions of
competition indices including RD, LnN, TN were 50.57%, age (age, 1/
D) was 37.45%, and structural diversity was 0.80% (Table 4, Fig. 6).
4. Discussion
In this study, the results showed that tree mortality of Chinese fir

4
X. Zhang, et al.
Fig. 2. Relationship of Gini coefficient of size diversity against age and number of trees per ha.
Fig. 3. Trends in annual mean temperature (MAT) and precipitation (AP) in relation to age for each site.
were influenced by competition, structural diversity, age, and climate
variables (Table 3). The relative contributions of competition, age, and
structure variables to survival were 88.82%, which was much larger
than climate variables 11.18% (Table 4). Competition was the main
driver (50.57%), followed by age (37.46%), climate (11.18%), and
structural diversity (0.80%) (Fig. 6).
4.1. Contributions of competition, structure and age to mortality
The results suggested that inter-tree competition for light, water,
and nutrients was the main agent influencing tree mortality (Franklin
et al., 2002; Yang et al., 2003). In general, larger trees in a stand have
advantages in comparison with smaller trees but not vice versa (Cannel
et al., 1984). As an asymmetric measure of competition, the positive
effect of RD on tree survival indicated that trees were more likely to be
alive when trees held a higher competitive status (Table 3), which was
consistent with the results of other studies (Laarmann et al., 2009;
Zhang et al., 2017). Stand density index is a useful measure of stand
stocking, but it cannot describe the contribution of trees to the total
stand index. Tree density index (TN), a tree-by-tree measure of site
5
Forest Ecology and Management 465 (2020) 118103
utilization related to stand density index proposed by Stage (1968)
could be applied in predicting tree growth and mortality in even-aged
forest, as well as uneven-age forest and mixed-species forest. In the
current study, tree density index (TN) was found to be negatively as-
sociated with tree survival (Table 3), which indicates that trees are
more prone to die in the context of higher site utilization. Zhang et al.
(2020) incorporated the tree density index into the height/diameter
ratio model, and found TN was negatively associated with height/dia-
meter ratio and the relative importance to the overall model was about
13.65%. In a dense stand, trees experience intense competition from
neighboring trees, which would lead to higher rates of mortality. This
was reflected in our finding that tree mortality increased with in-
creasing number of trees per ha (Table 3). Yang et al. (2003) reported
that stand basal area was a good index measuring stand crowdedness
and capturing asymmetric competition. However, stand basal area was
removed in the model because it could not pass the VIF test.
Tree mortality increased with increasing age (A) (Table 3). As an
endogenous variable, age also is a critical variable influencing tree
mortality. Physiological functions such as photosynthetic efficiency
decline with increasing tree age (Barnes et al., 1998), which leads to
X. Zhang, et al. Forest Ecology and Management 465 (2020) 118103

Table 3 Shannon index, and generalized variance (Staudhammer and LeMay,

Parameter estimates (including mean and standard deviation error, Std. error) 2001; Lexerød and Eid, 2006) which has been widely used for quanti-
and AIC values for the site-level and plot-level models of tree survival. fying size equality (Lei et al., 2009; Sun et al., 2018). During early stand
Variable Site-level Plot-level development, the Gini coefficient decreased until about age 10, and
increased slightly up to age 25, and then decreased again (Fig. 2).
Mean Std. error Mean Std. error During early stand development, tree growth variability has been
mainly attributed to environmental heterogeneity (Schume et al., 2004)
Intercept 27.775 1.082 20.4334 0.926
LnN −1.50 0.041 −0.763 0.049 or genetic variability between individuals (Boyden et al., 2008). With
RD 7.129 0.116 7.062 0.117 increasing tree size and canopy closer, smaller trees will eventually die
TN −0.049 0.003 −0.047 0.003 which leads to size equality and correspondingly smaller Gini coeffi-
Gini −5.173 0.445 −2.809 0.555
cient. Other studies (Macdonald and Yin, 1999; van Kuijk et al., 2008;
A −0.140 0.003 −0.118 0.003
1/D 3.276 0.203 3.045 0.199 Soares et al., 2016) have indicated that structural diversity increased
MAT 0.159 0.042 0.368 0.020 with age under size-asymmetric growth conditions or decreased with
LnAP −2.168 0.091 −1.286 0.074 age under size-symmetric conditions. The Gini value increased with
WMMT −0.083 0.019 −0.356 0.011 increasing number of trees per ha (Fig. 2), which is consistent with
AIC 64,881 64,700
previous studies (Soares et al., 2016; Sun et al., 2018).
Note: LnN: natural logarithm of number of trees per ha; D: diameter at breast We found tree survival decreased with increasing value of the Gini
height/cm; RD: relative diameter (D/Dq); Dq: stand quadratic mean diameter/ coefficient (Table 3). The results were consistent with Liang et al.
cm; A: stand age/year; TN: Tree stand density; MAT: mean annual temperature/ (2007), who found tree mortality in mixed forests was higher in plots of
°C; LnN: natural logarithm of annual precipitation; WMMT: winter mean larger structural diversity. However, Lei et al. (2009) reported no sig-
minimum temperature/°C. nificant effects of structural diversity on mortality in spruce-dominated
forests. Larger Gini values indicate asymmetric competition conditions
in the stand, which accelerates the mortality of smaller trees. In addi-
tion, a considerable amount of attention has been devoted to examining
relationships between structural diversity and stand growth. In such
studies, the effects of structural diversity were often negative (Liang
et al., 2005, 2007; Bourdier et al., 2016; Soares et al., 2016), sometimes
were positive (Lei et al., 2009; Zhang and Chen, 2015; Dănescu et al.,
2016), and sometimes were non-significant (Long and Shaw, 2010).
Forrester (2019) noted that structural diversity is positively or nega-
tively correlated with growth because of correlation with other vari-
ables, such as size-growth relationships. While structural diversity af-
fected tree mortality of Chinese fir, the contribution was small 0.80%
(Fig. 6).
Some studies have reported higher mortality rates at better sites
(Yao et al., 2001; Zhang et al., 2015; Zhang et al., 2017). In contrast,
some reports found tree mortality increased with decreasing site index
(Woollons, 1998). Therefore, site quality should be considered with
caution before their inclusion in mortality models. In the current study,
stand dominant height, a proxy of site index, was not incorporated in
our model because of failure to pass the VIF test. This result could be
due to high correlation and multicollinearity between Hd and age
(Castedo-Dorado et al., 2006). Consequently, the Hd of the stands was
implicitly included in the model.

Fig. 4. ROC (receiver operating characteristic) curve for the plot-level random 4.2. Contributions of climate variables to mortality
effects model of tree survival of Chinese fir plantations.
The impact of climate change on tree mortality has received wide
attention and modelling efforts (Peng et al., 2011; Zhang et al., 2017;
tree mortality induced by carbon starvation (Lugo and Scatena, 1996)
Lu et al., 2019; Brandl et al., 2020). Generally, when water availability
or reduced hydraulic conductivity (Smith and Long, 2001). In addition,
is not limited, tree survival probability was higher in warmer en-
older trees may have a higher risk of mortality because of low vigor
vironments. In this study, tree survival probability increased with in-
indicated by low diameter and height growth rates (Wunder et al.,
creasing MAT (Table 3), which was different with previous studies
2006; Reyes-Hernández and Comeau, 2014). Domec et al. (2008) re-
showing that annual temperature accelerates tree mortality (Peng et al.,
ported that larger trees may have a higher risk of death from size
2011; Zhang et al., 2014b). Chinese fir is a shade intolerant species and
constraints imposed on water and nutrient transport to their crowns.
its growth tends to increase with warmer condition, which in turn de-
Here, we found tree survival increased with decreasing diameter
creases the probability of mortality (Wu, 1984). Warm temperature
(Table 3), because the stands in this study were not old enough. Cortini
likely contributed to more rapid increases in photosynthetic activity
and Comeau (2020) indicated a decreased risk of mortality associated
than respiration rates, leading to a higher carbon assimilation rate and
with increasing diameter size, but in older forests the opposite was true.
increased tree growth (Way and Oren, 2010; Lines et al., 2012).
Structural diversity leads to a size hierarchy among individual trees
Therefore, increased growth translates into increased tree vigor which
and has a large impact on tree growth in a forest stand (Cordonnier and
is helpful for trees resisting insect pests or mechanical damage leading
Kunstler, 2015), as well as other processes, such as water-use efficiency,
to a net reduction in mortality rates. Schmidt-Vogt (1989) reported tree
carbon partitioning, nutrient cycling, and light absorption (Forrester,
death was mainly induced by overly-wet or dry years. If the environ-
2019). In forestry, the Gini coefficient is an efficient index for char-
ment gets excessive rainfall, it would cause death of fine roots due to
acterizing structural diversity compared to other indices, such as the
waterlogging and soil anoxia (Vygodskaya et al., 2002), which in turn

6
X. Zhang, et al. Forest Ecology and Management 465 (2020) 118103

Fig. 5. Predicted survival probability against 1/D (where D is diameter at breast height), RD (relative diameter), and Gini coefficient of size (diameter) diversity.

Table 4 precipitation conditions. In line with these results, we found in the

Relative contribution of each variable to tree mor- current study that tree mortality increased with increasing annual
tality of Chinese fir. precipitation (Table 3). However, in arid regions, drought was the
Variable Contribution/% primary driver of inducing tree death (Bigler et al., 2006; Peng et al.,
2011; Zhang et al., 2014b). We also found that tree mortality increased
LnN 3.67 with rising winter mean minimum temperature, WMMT (Table 3).
RD 37.18
Kliejunas et al. (2009) showed that tree mortality may increase with
TN 9.72
Gini 0.80 warmer winter temperature if pests increase their rates of over-win-
A 32.64 tering survival
1/D 4.81
MAT 3.80
LnAP 1.14
4.3. Interactive effects of endogenous and climate variables on tree
MWMT 6.24
mortality

Since tree mortality is one of the main important processes during

Fig. 6. Relative Contributions of competition, age, structure, and climate
variables to tree mortality of Chinese fir in southern China.
can increase rates of soil erosion, slope failure, and fungal (i.e., phy-
tophthora) infection. Vizcaíno-Palomar et al. (2016) reported that
higher rainfall in some sites might induce poorer soil quality, due to
increasing runoff and nutrient leaching. Zhang et al. (2018) reported
that the self-thinning slope of Chinese fir was much steeper in higher
forest succession, identifying the contributions of competition, struc-
ture, age, and climate variables to tree mortality is critical for devel-
oping climate adaptation strategies to maintain biological diversity in
forest ecosystems. Competition, as a main constraint to forest growth
through limited resources and growing space, may influence tree
mortality in response to climate change. Recent studies have shown that
competition can mediate climate-tree growth relationships (Clark et al.,
2014; Ford et al., 2016), and can thus influence different tree mortality.
Dietze and Moorcroft (2011) found that tree mortality in the eastern
and central United States was mainly driven by stand characteristics.
Zhang et al. (2015) used long-term observations in western Canada and
found that competition was the primary factor causing dynamic
changes in tree mortality. Luo and Chen (2015) indicated that the ef-
fects of intra-specific competition have intensified temporally as a
mechanism for the increased mortality of shade-intolerant tree species.
Therefore, as a shade-intolerant species, we should pay more attention
to these variables when controlling stocking levels of Chinese fir
plantations through either plantation spacing or thinning to maintain
stand vigor.

7
X. Zhang, et al.
5. Conclusions
This study modeled annual tree mortality of Chinese fir plantations
using long-term spacing trials at four sites in southern China in relation
to competition, age, structural diversity, and climatic variables through
logistic regression and a mixed-effects approach. Results showed that
tree mortality increased with increasing LnN, TN, Gini coefficient, A,
LnAP, and WMMT, and decreased with increasing RD, 1/D, and MAT.
The relative contributions of competition, age and structure factors to
mortality were 88.82%, which was much larger than climate variables
(11.18%). In addition, the relative contributions of competition indices
including RD, LnN, TN were 50.57%, which was much larger than the
contribution of age related variables (A, 1/D) (37.45%), and structural
diversity (0.80%). Our findings will be helpful for understanding the
mechanisms of tree mortality and optimization of silviculture practices
for promoting sustainable management of Chinese fir plantations in
southern China under climate change.
CRediT authorship contribution statement
Xiongqing Zhang: Data curation, Formal analysis, Writing -
Original draft preparation. Zhen Wang: Investigation, Analysis.
Sophan Chhin: Review and edit. Hanchen Wang: Investigation. Aiguo
Duan: Investigation. Jianguo Zhang: Supervision.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influ-
ence the work reported in this paper.
Acknowledgments
The study was supported by the National Natural Science
Foundation of China (No. 31670634) and State Administration of
Forestry and Grassland of China (2019132605).
Appendix A. Supplementary material
Supplementary data to this article can be found online at https://
doi.org/10.1016/j.foreco.2020.118103.
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