Global Ecology and Conservation 18 (2019) e00635

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Global Ecology and Conservation

journal homepage: http://www.elsevier.com/locate/gecco

Original Research Article

Effects of soil properties on the spatial distribution of forest

vegetation across China
Ji-Zhong Wan a, b, c, Jing-Hua Yu d, Guang-Jing Yin e, Zuo-Min Song e,
Deng-Xian Wei e, Chun-Jing Wang a, b, *
a
State Key Laboratory of Plateau Ecology and Agriculture, Qinghai University, Xining, 810016, China
b
College of Agriculture and Animal Husbandry, Qinghai University, Xining, 810016, China
c
Departamento de Ecología, Facultad de Ciencias Biolo
gicas, Pontificia Universidad Cato

lica de Chile, Santiago, Chile
d
Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang, 110016, China
e
Qinghai Forest Seedling Station, Xining, 810016, China

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

Article history: The investigation of factors affecting the spatial distribution of forest vegetation on a large
Received 22 February 2019 scale is a hot topic in forestry and ecology. Numerous studies have reported that climate
Received in revised form 15 April 2019 and human activities have a considerable effect on the spatial distribution of forest
Accepted 15 April 2019
vegetation. However, few studies have focused on the effects of soil properties on the
spatial distribution of forest vegetation across China. In the present study two indicators
Keywords:
were used to explore such soil effects, namely the percentage contribution of soil prop-
China
erties to the spatial distribution of forest vegetation and the similarity of the potential
Distribution modelling
Forest vegetation
distributions based on climate variables and both climate and soil variables under distri-
Organic carbon stock bution modelling. We found that 1) soil parameters (e.g., coarse fragment volume and
Soil factors organic carbon stock) contribute to the spatial distribution of forest vegetation in China,
although this contribution may vary among different biomes and vegetation classes and 2)
the spatial distribution of forest vegetation differs among the different vegetation classes
and biomes. Furthermore, soil variables (e.g., coarse fragment volume and organic carbon
stock) could play an important role in the spatial distribution of conifer-broadleaf forest
vegetation and mountain broad-leaved and conifer-leaved forest vegetation. However,
climate variables were more important than soil properties across most vegetation types. It
is therefore suggested that 1) coarse fragment volume and organic carbon stock should be
used as indicators to monitor forest vegetation and 2) soil properties should be conserved
to facilitate reforestation programs in China. The present study provides evidence that soil
parameters affect the spatial distribution of forest vegetation in China, facilitating the
development of effective management strategies.
© 2019 The Authors. Published by Elsevier B.V. This is an open access article under the CC
BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

1. Introduction

On a global level, forests are the most important resources for humans (Hansen et al., 2010, 2013). High biodiversity,
significant carbon sequestration and effective climate regulation depend on large areas of forest vegetation across different

* Corresponding author. State Key Laboratory of Plateau Ecology and Agriculture, Qinghai University, Xining, 810016, China.
E-mail address: wangchunjing00@163.com (C.-J. Wang).

https://doi.org/10.1016/j.gecco.2019.e00635
2351-9894/© 2019 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/
licenses/by-nc-nd/4.0/).
2 J.-Z. Wan et al. / Global Ecology and Conservation 18 (2019) e00635

spatial scales (Noss, 1999; Fang et al., 2001; Hou, 2001; Hansen et al., 2010; Alkama and Cescatti, 2016; Smith et al., 2018).
Bustamante et al. (2016) showed that tropical forests harbour a significant portion of global biodiversity and are a critical
component of the climate system. According to Fang et al. (2001), forest vegetation contributes significantly to the large
biomass of carbon stored in forests in China. However, the degradation of forest vegetation is currently advancing rapidly on a
large scale, leading to a considerable loss of suitable habitats for plants and animals and to decreased ecosystem services (e.g.,
carbon sequestration, water balance and food production) (Hansen et al., 2010; Ghazoul et al., 2015; Veldman et al., 2015).
Previous studies (e.g., Kairis et al., 2015; Salvati et al., 2015; He et al., 2016; Tsujino et al., 2016) showed that the degradation
and regrowth of forest vegetation are highly related to variations in soil conditions. Hence, for the efficient conservation of
forest resources, it is important to understand the effects of soil factors on forest vegetation.
Forests, the dominant ecosystems in China, are distributed over large areas (Hou, 2001). Hou (2001) showed that climatic
factors (e.g., temperature and precipitation) are the main drivers of the spatial distribution of forest vegetation in China.
However, numerous studies (e.g., Urbanov
a et al., 2015; Hume et al., 2016; James and Harrison, 2016; Widenfalk et al., 2016)
demonstrated significant relationships between soil variables and the coverage and species composition of forest vegetation
on a small scale. For example, soil water dynamics could shape different forest vegetation covers (Hayati et al., 2018; Molina
et al., 2019). Furthermore, some studies (e.g., Potter et al., 2003; Ter Steege et al., 2006; Wan et al., 2018a) showed that soil
factors can affect the spatial patterns of plant diversity on a large scale. However, only few studies have shown the re-
lationships between soil variables and the spatial distribution of forest vegetation on a large spatial scale. In China, as a result
of changing soil conditions, large areas are subject to forest degradation and deforestation (Wenhua, 2004; Wang, 2004; Zhai
et al., 2014; He et al., 2016; Wang et al., 2018). For example, there are obvious interactions between soil carbon, nutrient
dynamics and forest degradation (Cambardella and Elliott, 1994; Veldkamp, 1994; Feldpausch et al., 2004; Wenhua, 2004). In
this sense, it is important to investigate the effects of soil properties on the spatial distribution of forest vegetation in China,
with the aim of establishing a theoretical and practical basis for the recovery and conservation of Chinese forest resources.
Species distribution modelling (SDM) has been used to project vegetation distribution on a large scale (Tarkesh and
Jetschke, 2012; Ullerud et al., 2016; Wang et al., 2017; Wan et al., 2018b). For example, Tarkesh and Jetschke (2012)
showed that six SDMs (i.e., Bioclim, GARP, MaxEnt, MARS, NPMR, and LRT) can predict the occurrence of a given vegeta-
tion type with respect to environmental conditions. We can use occurrence records of forest vegetation and relevant envi-
ronmental variables to model the spatial distribution of forest vegetation in an SDM work environment (Tarkesh and Jetschke,
2012; Ullerud et al., 2016; Wang et al., 2017; Wan et al., 2018b). For example, Ullerud et al. (2016) established distribution
modelling of vegetation types in the boreal-alpine ecotone, using MaxEnt modelling. Wang et al. (2017) explored the response
of spatial vegetation distribution in China to climate changes since the Last Glacial Maximum (LGM), using SDMs based on
climatic variables and occurrence records of vegetation. The use of SDM, therefore, provides new insights into the projection
of spatial distribution of forest vegetation in China in consideration of soil effects.
In the present study, we tested two hypotheses to explore the effects of soil properties on the spatial distribution of forest
vegetation in China. 1) Hypothesis 1: Soil factors contribute to the spatial distribution of forest vegetation in China, based on
evidence of relationships between soil factors and forest vegetation on a small scale and between soil variables and plant
diversity on a large scale. 2) Hypothesis 2: The spatial distribution of forest vegetation differs across different vegetation
classes and biomes. Soil characteristics can vary from one forest vegetation type to another, e.g., from mountainous to
temperate to tropical vegetation. Different vegetation classes and biomes offer various soil conditions for plant diversity (Hou,
2001; Olson et al., 2001). If this hypothesis was to be supported, this would enable us to develop specific conservation and
restoration strategies for forest vegetation types, based on soil characteristics.
To test these hypotheses, we used SDM (i.e., MaxEnt) to project the spatial distribution of forest vegetation in China based,
in turn, on climate variables only and both climate and soil variables (Tarkesh and Jetschke, 2012). Subsequently, the dif-
ferences between the two sets of obtained results were compared on the basis of different vegetation types to test Hypotheses
1 and 2. Finally, we suggest management strategies for the conservation and restoration of forest vegetation in China.

2. Material and methods

2.1. Spatial distribution data on forest vegetation in China

We obtained data on the spatial distribution of forest vegetation in China from the Vegetation Atlas of China 1:1,000,000;
which contains 25,350 polygons of forest vegetation based on data from robust field investigations (Hou, 2001). Subsequently,
we transferred the polygon data into spatial distribution data (i.e., occurrence records) of forest vegetation in China at a grid
resolution of 10 arc-minutes (approx. 16
16 km2) (Hou, 2001; Wang et al., 2017; Wan et al., 2018b). We excluded vegetation
types with less than 10 occurrence records from our analysis for robust SDM (Wan et al., 2018b). In total, 189 forest vegetation
types were identified, belonging to three biomes (i.e., broad-leaved forest, conifer-broadleaf forest and conifer-leaved forest)
and 16 vegetation classes [detailed information is shown in the study of Hou (2001) and Table 1].

2.2. Environmental data

We used seven climate variables [Annual Mean Temperature ( C*10); Mean Diurnal Range; Temperature Seasonality
(standard deviation *100); Annual Precipitation (mm); Precipitation during the Wettest Month (mm); Precipitation during
 J.-Z. Wan et al. / Global Ecology and Conservation 18 (2019) e00635 3

Table 1
Mean ± SE percentage contribution (PC) of soil variables to the spatial distribution of forest vegetation in China across different biomes and classes.

Vegetation type BLD CEC CLYPPT CRFVOL ORCDRC PHIHOX SLTPPT SNDPPT
Conifer 0.41 ± 0.08 0.58 ± 0.23 0.25 ± 0.05 7.19 ± 1.20 2.75 ± 0.66 1.44 ± 0.34 1.02 ± 0.31 0.30 ± 0.08
a 0.45 ± 0.20 0.48 ± 0.39 0.12 ± 0.05 4.74 ± 1.87 8.16 ± 1.89 2.32 ± 1.10 1.31 ± 0.43 0.21 ± 0.06
b 0.53 ± 0.43 0.11 ± 0.06 0.58 ± 0.39 4.49 ± 2.31 0.23 ± 0.17 0.57 ± 0.32 0.75 ± 0.63 0.18 ± 0.07
c 0.32 ± 0.07 1.22 ± 0.70 0.25 ± 0.06 2.38 ± 0.97 0.55 ± 0.25 0.56 ± 0.13 0.55 ± 0.16 0.11 ± 0.03
d 0.41 ± 0.15 0.25 ± 0.06 0.27 ± 0.09 13.55 ± 2.45 1.01 ± 0.37 1.68 ± 0.42 1.25 ± 0.80 0.54 ± 0.21
Conifer-broadleaf 0.09 ± 0.04 0.14 ± 0.05 1.75 ± 1.71 4.50 ± 2.65 9.30 ± 2.82 1.24 ± 0.46 0.46 ± 0.33 0.12 ± 0.05
e 0.12 ± 0.07 0.20 ± 0.08 0.05 ± 0.03 0.85 ± 0.27 15.47 ± 2.17 1.44 ± 0.50 0.21 ± 0.10 0.17 ± 0.07
f 0.05 ± 0.03 0.05 ± 0.04 4.30 ± 4.28 9.98 ± 5.98 0.04 ± 0.04 0.95 ± 0.95 0.83 ± 0.83 0.03 ± 0.03
Broad-leaved 0.37 ± 0.07 0.84 ± 0.16 0.34 ± 0.07 3.71 ± 0.70 1.05 ± 0.31 1.32 ± 0.27 0.80 ± 0.17 0.34 ± 0.08
g 0.25 ± 0.06 0.26 ± 0.09 0.22 ± 0.10 3.79 ± 0.90 2.75 ± 1.04 1.57 ± 0.45 1.62 ± 0.43 0.36 ± 0.16
h 0.11 ± 0.07 1.00 ± 0.47 0.07 ± 0.02 1.03 ± 0.52 0.14 ± 0.08 0.57 ± 0.43 0.44 ± 0.16 0.79 ± 0.65
i 0.21 ± 0.09 0.99 ± 0.31 0.39 ± 0.16 7.72 ± 3.17 0.99 ± 0.48 3.18 ± 2.41 0.89 ± 0.72 0.62 ± 0.42
j 0.25 ± 0.09 1.91 ± 0.90 0.66 ± 0.41 5.70 ± 3.04 0.51 ± 0.29 1.12 ± 0.40 0.83 ± 0.66 0.52 ± 0.35
k 0.68 ± 0.17 0.56 ± 0.30 0.27 ± 0.07 0.49 ± 0.15 0.23 ± 0.10 1.16 ± 0.31 0.31 ± 0.13 0.21 ± 0.08
l 0.57 ± 0.18 0.43 ± 0.17 0.28 ± 0.14 0.70 ± 0.39 0.13 ± 0.06 1.82 ± 0.77 0.45 ± 0.14 0.20 ± 0.09
m 0.10 ± 0.06 0.37 ± 0.23 0.43 ± 0.39 19.57 ± 5.62 0.12 ± 0.08 0.76 ± 0.42 0.06 ± 0.05 0.08 ± 0.07
n 0.21 ± 0.18 0.41 ± 0.35 0.87 ± 0.21 0.52 ± 0.41 0.06 ± 0.03 1.31 ± 0.39 0.29 ± 0.15 0.06 ± 0.04
o 0.28 ± 0.10 1.41 ± 0.66 0.26 ± 0.14 0.57 ± 0.32 0.26 ± 0.22 0.91 ± 0.34 0.64 ± 0.37 0.25 ± 0.12
p 0.84 ± 0.53 1.10 ± 0.37 0.34 ± 0.13 2.18 ± 1.32 0.52 ± 0.32 0.24 ± 0.10 0.07 ± 0.03 0.09 ± 0.04

Vegetation codes were following: a: Cold-temperate and temperate mountains needleleaf forests; b: Temperate needleleaf forests; c: Subtropical coniferous
forests; d: Subtropical and tropical mountains needleleaf forests; e: Temperate coniferous and deciduous broad-leaved mixed forests; f: Subtropical
mountain mixed needleleaf, broadleaf evergreen and deciduous forests; g: Deciduous broad-leaved forests; h: Temperate microphyllous deciduous
woodlands; i: Subtropical deciduous broad-leaved forests; j: Subtropical evergreen and deciduous broad-leaved mixed forests; k: Subtropical evergreen
broad-leaved forests; l: Subtropical monsoon evergreen broad-leaved forests; m: Subtropical broadleaf evergreen sclerophyllous forests; n: Tropical
monsoon rain forests; o: Tropical rain forests; p: Subtropical and tropical bamboos, forests and scrubs. Soil codes were following: BLD: Bulk density (kg/
cubic-meter); CEC: Cation exchange capacity (cmolc/kg); CLYPPT: Soil texture fraction clay (%); CRFVOL: Coarse fragments volumetric (%); ORCDRC: Soil
organic carbon stock (tonnes per ha); PHIHOX: Soil PH; SLTPPT: Soil texture fraction silt (%); SNDPPT: Soil texture fraction sand (%).

the Driest Month (mm) and Precipitation Seasonality (Coefficient of Variation); Hijmans et al., 2005], eight soil variables [Bulk
Density (kg/m2); Cation Exchange Capacity (cmolc/kg); Soil Texture Fraction Clay (%); Coarse Fragments Volumetric (%); Soil
Organic Carbon Stock (tonnes per ha); Soil PH; Soil Texture Fraction Silt (%) and Soil Texture Fraction Sand (%); Hengl et al.,
2017] and one human footprint variable (Sanderson et al., 2002) to run SDM for the spatial distribution of forest vegetation
across China. Climate data on 10 arc-minutes were downloaded from the WorldClim database (averages from 1950 to 2000;
www.worldclim.org), and soil data on 0.5 arc-minutes were downloaded from SoilGrids (www.soilgrids.org; Hengl et al.,
2017). Based on the study of Hengl et al. (2017), the spread of selected soil points could cover most areas of China, and the
translation and cleaning up of soil properties and soil classes was sufficient for robust data performance of soil variables.
Furthermore, machine-learning techniques could improve the accuracy of soil maps for China based on tests of the true
positive rate (TPR) and area under the receiver operating characteristic curve (AUC; Hengl et al., 2017). Detailed information
on soil data was shown in the study of Hengl et al. (2017). Data on the human footprint was detailed in the study of Woolmer
et al. (2008). We used ArcGIS 10.6 (https://www.esri.com/en-us/home) to transfer the resolution of soil and human footprint
data from 0.5 arc-minutes to 10.0 arc-minutes. These environmental variables were selected because they potentially in-
fluence the distribution of forest vegetation (Hou, 201; Kier et al., 2005; Chen et al., 2015; Wang et al., 2017; Wan et al., 2018a
and 2018b).

2.3. Modelling and statistical analysis

MaxEnt modelling was applied as an SDM to analyse the effects of soil properties on the spatial distribution of forest
vegetation in China, using the same data on the spatial distribution of forest vegetation (Phillips et al., 2006; Tarkesh and
Jetschke, 2012; Wang et al., 2017; Wan et al., 2018b). Including human footprint, we built two MaxEnt models to produce
grid maps of spatial distribution probabilities of forest vegetation, based on climate variables only (Model C) and based on
both climate and soil variables (Model C þ S), according to the same sets as follows: 1) the regularization multiplier (beta) was
set to 2.0 to produce a smooth and general response shape that represents a biologically realistic behaviour (Radosavljevic
and Anderson, 2014); 2) the maximum number of background points was set to 10,000, with the same bias as the buffer
of occurrence records (Phillips et al., 2009); 3) we used a five-fold cross-validation approach to remove bias with respect to
occurrence records (Merow et al., 2013); 4) we used a complementary log-log (cloglog) transformation for the output of
MaxEnt (Phillips et al., 2017); 5) the other sets were the same as in the studies of Wang et al. (2017) and Merow et al. (2013).
We evaluated the predictive precision of the MaxEnt model using the AUC, which regards each value of the prediction result
as a possible threshold and then obtains the corresponding sensitivity and specificity values to calculate the curve. The
training omission rate showed the proportion of training grid cells of distribution areas of predicted absence distribution
areas (Phillips et al., 2006). Based on the study of Yu et al. (2014), we set grids with distribution probabilities higher than 0.5 as
4 J.-Z. Wan et al. / Global Ecology and Conservation 18 (2019) e00635

the units of spatial distribution of forest vegetation in China. We considered models with an AUC higher than 0.7 for further
analysis (Phillips et al., 2006).
To test Hypothesis 1, we used the percentage contribution (PC) from Model C þ S to evaluate the effects of climate and soil
variables on the spatial distribution of forest vegetation in China. We then quantified the PC of climate and soil variables based
on the sum of all climate or soil variables of Model C þ S. Here, the importance of climate or soil variables is high when the PC
is at least 15% of the model for each vegetation type, vegetation class and biome (Oke and Thompson, 2015).
To test Hypothesis 2, we measured the similarity of the potential distributions between Model C and Model C þ S,
exploring the effects of soil properties on the spatial distribution of forest vegetation across different vegetation types using
Schoener's D (Warren et al., 2008, 2010). To quantify Schoener's D between Model C and Model C þ S, we used ENMTools 1.4.4
(see Warren et al., 2008, 2010 for detailed information). Schoener's D ranges from 0 (vegetation with a completely discordant
distribution) to 1 (vegetation with an identical distribution; Warren et al., 2008). We compared the similarity of the potential
distributions between the two models based on vegetation types, vegetation classes and biomes. The graphing and analysis
software used was JMP 11.0 (SAS Institute, Cary, North Carolina).

3. Results

All models showed good performance for all studied vegetation types, with training and test AUC values being higher than
0.7 (Table S1). Climate was the most important variable, while the human footprint was the weakest variable to explain the
spatial distribution of forest vegetation in China across different biomes and vegetation classes (Fig. 1). The PCs of soil factors
to spatial distribution were higher than 15% for 53 forest vegetation types, and the largest contribution was 57.0% for the
spatial distribution of Abies spectabilis forests (Fig. 2a and b; Table S1). Soil variables contributed in large part to the spatial
distribution of conifer-leaved (PC: 13.9%) and conifer-broadleaf forests (PC: 17.6%; Fig. 1). Regarding vegetation classes, the
importance of soil variables was high for cold-temperate and temperate mountainous needleleaf forests (PC: 17.8%); sub-
tropical and tropical mountainous needleleaf forests (PC: 19.0%); temperate coniferous and deciduous broad-leaved mixed
forests (PC: 18.5%); subtropical mountainous mixed needleleaf, broadleaf evergreen and deciduous forests (PC: 16.2%);
subtropical deciduous broad-leaved forests (PC: 15.0%) and subtropical broadleaf evergreen sclerophyllous forests (PC: 21.5%)
(Fig. 1).
Specifically, coarse fragment volume (%) was the most important soil variable affecting the spatial distribution of conifer-
leaved forests (PC: 7.19%) (Table 1), and the variable soil organic carbon stock was the most important influence on conifer-
broadleaf forests (PC: 9.30%) (Table 1). Coarse fragment volume was the most important variable affecting the spatial distri-
bution of subtropical and tropical mountainous needleleaf forests; subtropical mountainous mixed needleleaf, broadleaf
evergreen and deciduous forests; subtropical deciduous broad-leaved forests and subtropical evergreen and deciduous broad-
leaved mixed forests (Table 1). This variable was also highly important for the spatial distribution of temperate coniferous and
deciduous broad-leaved mixed forests (PC: 19.57%), while the variable soil organic carbon stock was a significant determinant of
the spatial distribution of subtropical broadleaf evergreen sclerophyllous forests (PC: 15.47%) (Table 1).
The potential distributions between the two models shared high similarity across three biomes (conifer-leaved forests:
0.91, conifer-broadleaf forests: 0.90 and broad-leaved forests: 0.92) (Fig. 3). However, the similarity of the potential distri-
butions between Model C and Model C þ S was lower for cold-temperate and temperate mountainous needleleaf forests
(0.88), temperate coniferous and deciduous broad-leaved mixed forests (0.88), temperate microphyllous deciduous wood-
lands (0.89) and subtropical deciduous broad-leaved forests (0.89), compared with other vegetation types (Fig. 3). The
similarity was the highest for subtropical and tropical bamboo stands, forests and scrubs (0.95) (Fig. 3) and for Picea jezoensis
forests (Fig. 2c and d; Table S1).

4. Discussion

We tested the effects of soil properties on the spatial distribution of forest vegetation across China, using MaxEnt
modelling. Our results indicate that 1) soil factors potentially contribute to the spatial distribution of forest vegetation, but
such contributions may vary among biomes and vegetation classes, and 2) the spatial distribution of forest vegetation varies
among vegetation classes and biomes. Hence, climate variables are the main drivers of the spatial distribution of forest
vegetation, although soil variables play an important role in shaping the distribution of forest vegetation on a large spatial
scale. The results of our study provide a scientific basis for the development of specific management strategies tailored to the
requirements of forest vegetation across China.

4.1. Effects of soil properties on the spatial distribution of broad-leaved forest vegetation

The contribution of soil variables to spatial forest vegetation distribution and the similarity of the potential distributions
between the two models were relatively low compared with the other two biomes (i.e., conifer-broadleaf and conifer-leaved
forests), indicating that soil parameters have only minor effects on the spatial distribution of broad-leaved forest vegetation,
while climate variables determine, in large part, the spatial distribution of broad-leaved forest vegetation in China. However,
such effects may depend strongly on the vegetation types and classes (Figs. 1 and 3). Particularly, different soil factors could
contribute to the spatial distribution of different broad-leaved forest vegetation types.
 J.-Z. Wan et al. / Global Ecology and Conservation 18 (2019) e00635 5

Fig. 1. Mean ± SE percentage contribution (PC) of climate, soil and human footprint variables to the spatial distribution of forest vegetation in China across
different biomes and classes. Vegetation codes were following: a: Cold-temperate and temperate mountains needleleaf forests; b: Temperate needleleaf forests;
c: Subtropical coniferous forests; d: Subtropical and tropical mountains needleleaf forests; e: Temperate coniferous and deciduous broad-leaved mixed forests; f:
Subtropical mountain mixed needleleaf, broadleaf evergreen and deciduous forests; g: Deciduous broad-leaved forests; h: Temperate microphyllous deciduous
woodlands; i: Subtropical deciduous broad-leaved forests; j: Subtropical evergreen and deciduous broad-leaved mixed forests; k: Subtropical evergreen broad-
leaved forests; l: Subtropical monsoon evergreen broad-leaved forests; m: Subtropical broadleaf evergreen sclerophyllous forests; n: Tropical monsoon rain
forests; o: Tropical rain forests; p: Subtropical and tropical bamboos, forests and scrubs.
6 J.-Z. Wan et al. / Global Ecology and Conservation 18 (2019) e00635

Fig. 2. Spatial distribution of Abies spectabilis forests (a and b) and Picea jezoensis forests (c and d) across China based on climate variables only (Model C) and
based on both climate and soil variables (Model C þ S). The colour on the maps ranging from brown to blue represents the distribution probability ranges of Abies
spectabilis forests and Picea jezoensis forests ranging from 0.5 to 1.0. (For interpretation of the references to colour in this figure legend, the reader is referred to
the Web version of this article.)

Chinese broad-leaved forests are distributed throughout tropical and subtropical regions and are characterized by a high
level of plant diversity (Hou, 2001; Liu et al., 2015; Ouyang et al., 2016; Ding et al., 2017). Furthermore, plant diversity shows a
narrow thermal tolerance to climatic changes and low dispersal on a large spatial scale (García-Robledo et al., 2016; Bush
et al., 2018; Polato et al., 2018). The features of climatic tolerance and dispersal ability are extremely vulnerable to climate
change (Bush et al., 2018; Polato et al., 2018; Wan et al., 2018a). For example, Zhou et al. (2013) showed that climate variance
(e.g., global warming and drought stress) could lead to predicted reductions in rainfall and forest productivity, increased tree
mortality and declining forest biomass carbon sinks in a subtropical, monsoon, evergreen broad-leaved forest in Southern
China. This leads us to infer that climate variables largely determine the spatial distribution of broad-leaved forest vegetation
in China (Hou, 2001; Zhou et al., 2013; Bush et al., 2018; Polato et al., 2018). Although the effects of soil parameters were not
that high, we should pay attention to the effects of soil on spatial distribution across different vegetation types and classes.
Soil parameters, particularly coarse fragment volume, could affect the spatial distribution of subtropical, deciduous broad-
leaved forests in China, according to the results of PC and Schoener's D (Figs. 1 and 3). Previous studies have shown significant
relationships between soil parameters and plant diversity in deciduous broad-leaved forests (Cerný  et al., 2013; Kooch et al.,
2017; Wan et al., 2018b). On a relatively large spatial scale, the resources available to plants could be altered in subtropical
deciduous broad-leaved forests due to frequent changes in water availability (Forrester, 2014; Chen et al., 2015). Coarse
fragment volume may be largely related to soil water retention (Perie and Ouimet, 2008). In this sense, the soil resources may
shape the spatial distribution of subtropical, deciduous broad-leaved forests in China due to water availability (Hanson and
Blevins, 1979; Hudson, 1994).

4.2. Effects of soil properties on the spatial distribution of conifer-broadleaf forest vegetation

The values of PC and Schoener's D were the highest across all three biomes, indicating that soil variables contributed
significantly to the spatial distribution of conifer-broadleaf forest vegetation across China. Furthermore, temperate coniferous
and deciduous broad-leaved mixed forests; subtropical mountain mixed needleleaf, broadleaf evergreen and deciduous
forests; subtropical evergreen and deciduous broad-leaved mixed forests and temperate coniferous and deciduous broad-
leaved mixed forests could be severely affected by changes in soil parameters. Our study, therefore, provides important
references for the mechanism of spatial distribution of conifer-broadleaf forest vegetation in China.
 J.-Z. Wan et al. / Global Ecology and Conservation 18 (2019) e00635 7

Fig. 3. Similarity of the potential distributions of forest vegetation between Model C and Model C þ S, using Schoener's D. Range: the range of similarity of the
potential distributions between Model C and Model C þ S; the transverse line of the boxes: the average values of similarity of the potential distributions between
Model C and Model C þ S. Vegetation codes were following: a: Cold-temperate and temperate mountains needleleaf forests; b: Temperate needleleaf forests; c:
Subtropical coniferous forests; d: Subtropical and tropical mountains needleleaf forests; e: Temperate coniferous and deciduous broad-leaved mixed forests; f:
Subtropical mountain mixed needleleaf, broadleaf evergreen and deciduous forests; g: Deciduous broad-leaved forests; h: Temperate microphyllous deciduous
woodlands; i: Subtropical deciduous broad-leaved forests; j: Subtropical evergreen and deciduous broad-leaved mixed forests; k: Subtropical evergreen broad-
leaved forests; l: Subtropical monsoon evergreen broad-leaved forests; m: Subtropical broadleaf evergreen sclerophyllous forests; n: Tropical monsoon rain
forests; o: Tropical rain forests; p: Subtropical and tropical bamboos, forests and scrubs.

Previous studies have shown a significant interaction between soils (mainly coarse fragment volume and organic carbon
stock) and plant species composition for conifer-broadleaf forest vegetation (Quideau et al., 2001; Sun et al., 2004; De Deyn
et al., 2008; Lange et al., 2015). Changes in the plant species composition may result in different types of conifer-broadleaf
forest vegetation (Hou, 2001; Sollins, 1998). The levels of both coarse fragment volume and organic carbon stock could
lead to variation in plant species composition (Hou, 2001; Sollins, 1998; Saha et al., 2009; Zhang et al., 2011). The rate of
change in soil carbon stocks could also alter the composition of tree species in conifer-broadleaf forests (Saha et al., 2009;
Zhang et al., 2011). Soil carbon stocks can enhance aboveground net primary productivity and belowground biomass, thereby
increasing plant species diversity, which can in turn drive changes in plant species composition (Clark et al., 2001; Chase and
Leibold, 2002; Tilman et al., 2012; Liang et al., 2016). Climate variables could drive changes in organic carbon stock in Chinese
conifer-broadleaf forests (Sun et al., 2019). Combined with our results that climate variables are the main drivers of the spatial
distribution of Chinese conifer-broadleaf forest vegetation, the spatial distribution of conifer-broadleaf forest vegetation
could be affected by a changing climate and changes in plant species composition (Thapa et al., 2016; Wan et al., 2017; Sun
et al., 2019).
8 J.-Z. Wan et al. / Global Ecology and Conservation 18 (2019) e00635

In this context, we should pay attention to the significant effects of soil parameters on subtropical mountainous mixed
needleleaf, broadleaf evergreen and deciduous forests. Previous studies have shown that forest vegetation and plant species
composition are sensitive to climatic changes (Bonan, 2008; Arasa-Gisbert et al., 2018; Sun et al., 2019). Our study provides
evidence that soil parameters shape the spatial distribution of conifer-broadleaf forest vegetation. Soil microbial communities
of subtropical mountainous forest ecosystems can affect the distribution of plant species by changing the soil carbon and
nutrient dynamics along the elevational gradient (Jiang et al., 2012; Li et al., 2014; Fang et al., 2017). Thus, variations in coarse
fragment volume and organic carbon stocks could play an important role in changes in microbial communities in subtropical
mountainous forest ecosystems (Rhoades et al., 2000; Perie and Ouimet, 2008; Gupta and Sharma, 2013; Prietzel and
Christophel, 2014). Hence, soils could affect the spatial distribution of subtropical mountainous mixed needleleaf, broad-
leaf evergreen and deciduous forests across China.

4.3. Effects of soil properties on the spatial distribution of conifer-leaved forest vegetation

The values of PC and Schoener's D were relatively high, indicating significant effects of soil parameters on the spatial
distribution of conifer-leaved forest vegetation. Furthermore, such soil effects could be enhanced following changes in
vegetation types and classes in conifer-leaved forest vegetation. Cold-temperate and temperate mountainous needleleaf
forests (e.g., Abies spectabilis forests) could be affected significantly by soil parameters (He et al., 2016). Hence, there are large
effects of soils on the spatial distribution of conifer-leaved forest vegetation in mountainous areas of China.
Conifer-leaved forest vegetation is widely distributed in mountainous areas of southwestern China (Hou, 2001). Such areas
are characterized by relatively poor soils compared with other regions in China (Hou, 2001). In other words, the conifer-
leaved forest vegetation in mountainous areas requires specific soil conditions suitable for regeneration and tree growth
(Hou, 2001). For example, Picea jezoensis forests and Abies spectabilis forests need certain levels of soil organic carbon, related
to their spatial distribution (Hou, 2001; Lei et al., 2007; He et al., 2016). The effects of soil organic carbon on conifer-leaved
forest vegetation may be due to the requirements of conifer-leaved forests in terms of soil nutrient availability and terrestrial
carbon cycling (Carleton and Read, 1991; Vogt, 1991; Macinnis-Ng and Schwendenmann, 2015). Previous studies have shown
that mountainous tree species inhabit a relatively narrow ecological niche and have a low ability to adapt to rapid climatic
changes (Hou, 2001; Bush et al., 2018; Polato et al., 2018). Our study implies not only that climate variables are the main driver
of the spatial distribution of conifer-leaved forest vegetation but also that soils could play a key role in driving the spatial
distribution of conifer-leaved forest vegetation in China.

5. Conclusions and management recommendations

Our study developed two indicators (i.e., the contribution of soil variables to spatial distribution and the similarity of the
potential distributions based on climate variables and on both climate and soil variables) to investigate the effects of soil
parameters on the spatial distribution of forest vegetation in China. Our results lead us to infer that soil parameters (e.g.,
coarse fragment volume and organic carbon stock) mostly contribute to the spatial distribution of conifer-broadleaf forest
vegetation as well as mountainous broad-leaved and conifer-leaved forest vegetation. In this context, we suggest that 1) the
parameters coarse fragment volume and organic carbon stock should be used as indicators to monitor forest vegetation and 2)
soil conservation should be considered for effective management of forest resources and to support afforestation efforts.
Future studies should, therefore, develop vegetation distribution models based on soil variables and explore the effects on
forest species composition on a global scale.

Acknowledgment

We thank the editor and two reviewers for the useful comments on the improvement of our early manuscript. This work
has been supported by the National Natural Science Foundation of China (Nos. 31800449 and 31800464), the 13th Five-year
Informatization Plan of Chinese Academy of Sciences (No. XXH13506), and the Basic Research Project of Qinghai Province,
China (No. 2019-ZJ-936Q).

Appendix A. Supplementary data

Supplementary data to this article can be found online at https://doi.org/10.1016/j.gecco.2019.e00635.

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