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A Bayesian Network With Fuzzy Mathematics For Species Habitat Suitability Analysis
A Bayesian Network With Fuzzy Mathematics For Species Habitat Suitability Analysis
Ecological Modelling
journal homepage: www.elsevier.com/locate/ecolmodel
A R T I C L E I N F O A B S T R A C T
Keywords: There are many different types of species distribution models (SDMs) that are widely used in the field of ecology.
Bayesian network (BN) In this research, we explored a new advanced mechanism for predicting the distribution of species based on fuzzy
Fuzzy matter element model (FME) membership function, principle of maximum entropy, fuzzy mathematics comprehensive evaluation, and the
Fuzzy mathematics and Bayesian network
framework of Bayesian networks. We use fuzzy mathematics and Bayesian network model (FBM) to simulate
model (FBM)
Species habitat suitability
relationships between species’ habitats and environmental variables, and the relationship may be difficult to
Angelica sinensis (Oliv.) Diels quantify effectively. FBM, which combines species data, environmental data, expert experience, and machine
learning, could reduce the data and system error. In the case of medicinal plant, Angelica sinensis (Oliv.) Diels,
many approaches have been applied, including nine learning sequence of sampling sites, three FBM models, two
types of information classification by fuzzy mathematical classification (FMC) and equal interval classification
(EIC), and the evaluation of AIC and log-likelihood. Through the comparison of reasoning results between FBM
and fuzzy matter element model (FME) in testing sites, the result shows that the combination of objective data
and empirical model structure makes FBM have better result output. Besides, FBM sensitivity analysis helps
researchers explore in detail the impact of environmental factors on each level of species habitat suitability. The
temperature factor has an important influence on the highly suitable, moderately suitable, and lowly suitable
habitats of A. sinensis. Through FMC and sensitivity analysis, annual mean temperature (Bio1) in 5.92 ◦ C-9.05 ◦ C
and mean temperature of warmest quarter (Bio10) in 14.80 ◦ C-18.60 ◦ C are the highly suitable habitat tem
perature range of A. sinensis.
1. Introduction up so far.
The Generalized linear model (GLM) proposed by Nelder and Wed
Many researchers are interested in species distribution models derburn (1972) appeared very early and has been applied to the suit
(SDMs). On this basis, the relationship model between habitat occur ability research of species. As a habitat suitability model, GLM performs
rence localities and environmental requirements was constructed a good analysis when there are a large number of species or lack of data
(Phillips et al., 2006). Since the late twentieth century, scholars have (Hirzel et al., 2001). The Generalized additive model (GAM) is a
been using mathematical methods to construct species environmental non-parametric extension of GLM and is used to deal with complex
suitability models to help people understand the living conditions of geographic data to predict spatial distributions of species (Zaniewski
species (Friedman, 1991; Manel et al., 1999; Breiman, 2001; Elith et al., et al., 2002). Unlike GLM or GAM, the genetic algorithm for rule-set
2006). Each model has its own theoretical framework and is different production (GARP) could predict the distribution of plants and ani
from the concept definition with species suitability. Due to the system mals through automated spatial modeling and filter out potential sour
atic error of different SDMs, the final prediction results will also be ces of errors (Stockwell, 1999). Subsequently, the Maxent model
distinguished. After long-term application and screening, a few SDMs appeared at the historic moment, which used maximum-entropy tech
coped well with environmental conditions, and have been widely used niques to model the distribution of species. And its sequential update
* Corresponding author.
E-mail addresses: weihy@snnu.edu.cn (H. Wei), weigu@snnu.edu.cn (W. Gu).
https://doi.org/10.1016/j.ecolmodel.2021.109560
Received 26 June 2020; Received in revised form 2 February 2021; Accepted 3 April 2021
Available online 22 April 2021
0304-3800/© 2021 Elsevier B.V. All rights reserved.
Q. Zhang et al. Ecological Modelling 450 (2021) 109560
Table 1
Our team uses FME to study the potential distribution of medicinal plants.
Predicted Species of medicinal plant Number of environment Characteristic values of Study area in China Literature
period variables species
Current Schisandra sphenanthera Rehd. et Wils. 15 schisantherin A Qinling Mountains Lu et al., 2012
Schisandra sphenanthera Rehd. et Wils. 15 deoxyschizandrin Qinling Mountains Guo et al., 2013
γ-schizandrin
Scutellaria baicalensis Georgi 15 baicalin The whole of China Zhu et al., 2015
Cornus officinalis 21 loganin Qinling Mountains Sang et al., 2015
Sinopodophyllum hexandrum (Royle) 12 fruit yield 7 provinces in western of Guo et al., 2015
Ying China
Angelica sinensis (Oliv.) Diels 14 ferulic acid 10 provinces in western of Shang et al.,
China 2015
Schisandra chinensis (Turcz.) Baill. 18 lignan 4 provinces of China Mao et al., 2016
Gynostemma pentaphyllum (Thunb.) 19 gypenosides The whole of China Zhao et al., 2017
Makino
Future Schisandra sphenanthera Rehd. et Wils. 19 schisantherin A Qinling Mountains Guo et al., 2016
algorithms could handle a very large number of features (Phillips et al., European hake (Merluccius merluccius), anglerfishes (Lophius piscatorius
2006). By contrast with GARP, Maxent predicted the distribution more and L. budegassa), and red mullets (Mullus barbatus and M. surmuletus) in
accurately when it considered presence-only data (Ray et al., 2018). an exploited marine ecosystem of the Northwestern Mediterranean Sea.
With the advancement of machine learning techniques (Džeroski et al., In addition to animals, few studies have used Bayesian theory to focus on
1997; Chon et al., 2001), several models have been developed using a the suitability distribution of terrestrial plants. Meineri et al. (2015)
variety of algorithms, such as classification trees (Debeljak et al., 2001), disentangled direct and indirect associations between landscape physi
Random forest (RF) (Freeman et al., 2012), and Generalized boosting ography, environmental variables, and species distribution of four
model which is usually called Boosted regression trees (GBM/BRT) common plants at their northern range limit in Sweden by Gaussian BN.
(Hallstan et al., 2013). However, due to the difficulty of data collection and experience acqui
The fuzzy matter element model (FME), which is established on the sition, no scholars have directly used BN to predict the distribution of
basis of fuzzy mathematics and GIS could provide and quantify an plant species suitability. This inspired us to make a species suitability
effective method for the evaluation of plant habitat suitability. FME fits prediction of BN supported by fuzzy mathematics.
the relationship between environmental variables data at sampling sites Our approach closely links to ecological niche theory (Patten and
and species characteristics, such as the content of marker compounds Auble, 1981). Chinese mathematician Cai founded matter element the
extracted from the species (Nino et al., 2017). In 2012, our team firstly ory (Cai, 1983). In this research, we use matter element theory, fuzzy
used the FME to predict the suitability distribution of Schisandra sphe membership function (Zadeh, 1965), the principle of maximum entropy
nanthera Rehd. et Wils. in Qinling Mountains (Lu et al., 2012). After our (Parkash et al., 2008), and fuzzy mathematics comprehensive evaluation
effort, we have applied FME to study the suitability distribution of many (Lu et al., 2012) to assist BN modeling. Fuzzy membership functions
medicinal plants using different number of environment variables could fit the membership relations between fuzzy features, such as
(Table 1). Hence, we have accumulated valuable experience in opti medicinal effective contents of species, and different numerical envi
mizing and improving FME. ronmental variables. In general, the scope of fuzzy membership degree is
Bayesian theories or methods are widely used in various fields of [0,1] (Zadeh, 1965). The principle of maximum entropy of information
research, such as financial econometrics (Wang et al., 2017a), risk could determine the weighting coefficients associated with the observed
management (Xin et al., 2017), environmental assessment and man data, avoiding the subjective bias with only using experience. Fuzzy
agement (Marcot and Penman, 2018), road infrastructure (Sierra et al., mathematics comprehensive evaluation could achieve the original data
2018), mechanical engineering (Wang et al., 2017b; Wang and Matth standardization according to the membership relations (Lu et al., 2012;
ies, 2019), biotechnology (Gendelman et al., 2017), and ecology (Sal Zhao et al., 2017). Fuzzy mathematics could help us build habitat suit
liou et al., 2017). There are some studies focused on species habitat ability models in the absence of expert experience. In this research, the
research based on Bayesian theory (Feki-Sahnoun et al., 2018; Wiest role of fuzzy mathematics mainly reflects in the value classification of
et al., 2019; Thompson et al., 2020). In terms of ecological suitability, states of single environmental variables (SEVs), the determination of
some researchers chose Bayesian methods to study the habitat distri states of integrated environmental variables (IEVs), and habitat suit
bution of animals. For vertebrates, they have proved that the Bayesian ability node. As for BN, it mainly reflects in the construction of model’s
network (BN) based on abundant prior knowledge is sufficient to frame, the calculation of the probability of species habitat suitability,
establish a model with reasonable sensitivity (Tantipisanuh et al., 2014). and sensitivity analysis. The results of fuzzy mathematics will process
Gieder et al. (2014) used BN to study the response of habitat to sea level into the BN model.
rise, and successfully predicted the existence of bird nests on the barrier Angelica sinensis (Oliv.) Diels as an example of model construction in
islands. Havron et al. (2017) applied BN to a small and marine dataset to this article. A. sinensis is a perennial herb, mainly distributed in the
model the probability of occurrence of three macrofauna species. They provinces of Gansu, Sichuan, Shaanxi, Hubei, and Yunnan (Committee
were the first to show that BN can effectively model habitat suitability of of Flora of China, 1992). The roots use as Chinese herbal medicine
benthic macrofauna. Hradsky et al. (2017) showed that although only (CHM), and marker compound of ferulic acid content should be no lower
remote-sensed or mapped variables are provided, BN can effectively than 0.05% (Committee of National Pharmacopoeia, 2020). The roots of
distinguish the existence of most native mammal species in the Otway A. sinensis have been used historically as a tonic, hematopoietic, and
Ranges in southeastern Australia. Using Bayesian hierarchical species anti-inflammatory agent for the treatment of gynecological diseases
distribution model, Coll et al. (2019) predicted the distribution of such as menstrual disorders, amenorrhea and dysmenorrheal for
2
Q. Zhang et al. Ecological Modelling 450 (2021) 109560
Fig. 1. 90 sampling sites (orange sites) and 50 predicted sites (black sites) of A. sinensis in China. Sampling sites are obtained mainly from the provinces of Gansu,
Hubei, Sichuan, and Yunnan. The distribution of the predicted sites is symmetrical, and the habitat suitability of them will be predicted by FBM after a series of
machine learning and reasoning.
thousands of years in Chinese herbal medicine prescriptions (Lü et al., this paper attempts to prove that FBM has great potential in predicting
2009; Cao et al., 2010). the suitability of medicinal plant habitats.
Our modeling objective is to build fuzzy mathematics and Bayesian
network model (FBM), to reconstruct the real relationship between 2. Materials and methods
environmental variables and A. sinensis’ habitats, to predict the habitat
suitability of A. sinensis, and to show the optimal results of the predicted 2.1. Study area
sites given by FBM. The original data will discretize by fuzzy mathe
matics classification (FMC) and equal interval classification (EIC). We We obtain the location information of distribution sites of A. sinensis
also use additional testing sites to compare the predicted results of the by consulting Flora of China, investigation resource reports, literature,
FBM and FME to verify the reliability of the FBM. Using general data, and digital herbarium. The sites with ferulic acid content acquired from
Fig. 2. The three kinds of FBM networks. (a) Model A, which is a simple network structure by learning with fuzzy mathematics classification (FMC); (b) Model B,
which increases three IEVs nodes and directed arrows (green), reflecting the relationship between the different IEVs; (c) Model C, which based on model B, increased
two arrows (red) which point directly to the habitat suitability node to show the importance of temperature and precipitation factors.
3
Q. Zhang et al. Ecological Modelling 450 (2021) 109560
Table 2
The eight SEVs’ states divided by fuzzy mathematics classification (FMC) method.
SEVs (unit) Four states of SEVs node in FBM network
1 2 3 4
Shang et al. (2015). In order to remove the errors caused by the exces Hence, the state of the parent nodes could affect the state of the child
sive concentration of sampling sites, we select samples by eliminating nodes by the arrows (Fuster-Parra et al., 2015). As a part of FBM con
adjacent sampling sites. Finally, we obtain 90 sampling sites of struction, BN is an important assumption that there is no loop in the
A. sinensis with complete information including ferulic acid content network. Formally, the network is a directed acyclic graph.
(Fig. 1). Here, the theoretical framework of BN proposed by FBM includes
three parts, including eight SEVs nodes (Bio1, Bio10, Bio12, Bio15,
2.2. Collection of environmental variables Bio18, SOM, Totalrad13 and EL), three IEVs nodes (temperature suit
ability, precipitation suitability, and soil suitability), and habitat suit
We collect bio-climatic variables, soil variables, topographic vari ability node. The simple diagram of the FBM model is constructed as
ables, and light variables, which are used to simulate the distribution Fig. 2a by learning the network structure with FMC information (see
suitability of A. sinensis. For bio-climatic variables, we collect 19 SEVs Table 2) in the machine Hugin (Hugin Expert 8.4, 2016). The network
(Bio1-Bio19) from the WorldClim database (www.worldclim.org/curr structure of model A is obtained by machine learning of 90 sampling
ent). For soil variables, we collect 5 SEVs from the Harmonized World sites without adding operator experience information.
Soil Database (HWSD, http://www.fao.org/soils-portal/en). For topo In the process of model building (Fig. 2b and Fig. 2c), we constructed
graphic variables, we collect 3 SEVs from the Resource and Environment temperature, precipitation, and soil as three IEVs nodes. Habitat suit
Data Cloud Platform (RESDC, http://www.resdc.cn). In addition, 4 SEVs ability is a node supported by precipitation, temperature and soil nodes.
related to light are acquired from the Chinese Ecosystem Research The common ground of model B and model C: (1) The networks are
Network (CERN, http://www.cern.ac.cn). roughly divided into three layers. (2) The suitability of temperature
Variables data selection is a three-step process, the first step is to set (Bio1 and Bio10), precipitation (Bio12, Bio15, and Bio18), and soil
up membership function between all SEVs and the content of ferulic acid (SOM) directly determine the habitat suitability of A. sinensis. (3) The
with 90 sampling sites of A. sinensis in Matlab software (Matlab software effects of topography (EL) regard as potential impacts on temperature
9.1, 2016), retained SEVs were those with an R-squared greater than 0.8. and precipitation (Russak, 2009; Marini et al., 2011). For the difference
The second step is to do the correlation analysis with environmental between model B and model C, model B has a clear hierarchy (Fig. 2b).
variables that we have chosen, in order to remove the high correlation Considering the importance of annual mean temperature (Bio1) and
variables and prevent overfitting of the model. If two or three variables annual precipitation (Bio12) for the growing of A. sinensis, the model C
are highly correlated, choose the one that has more biological signifi add two connection arrows directed from nodes of Bio1 and Bio12 to the
cance and value for A. sinensis growth. Finally, we obtain eight SEVs, node of habitat suitability (Fig. 2c), based on the model B.
including Bio1 (annual mean temperature), Bio10 (mean temperature of For an FBM network, the probability distribution of each node must
warmest quarter), Bio12 (annual precipitation), Bio15 (precipitation be described by a set of mutually exclusive “states”, such as Table 2.
seasonality), Bio18 (precipitation of the warmest quarter), SOM (soil Before obtaining the probability distribution of the entire network, we
organic matter), Totalrad13 (annual total solar radiation), and EL need to set the states of all nodes.
(elevation above sea level). The eight SEVs included two kinds of tem
perature (Bio1 and Bio10), three kinds of precipitation (Bio12, Bio15, 2.4. The states of SEVs nodes in FBM
and Bio18), one soil variable (SOM), one light variable (Totalrad13),
and one terrain variable (EL). BNs are usually modeled on discrete domains, so continuous vari
ables need to be discretized first (Uusitalo, 2007). Hence, we classify
2.3. Bayesian network for FBM eight SEVs by FMC and EIC discrete method. In order to set the state of
SEVs through FMC, we establish the membership function between the
In the field of Bayesian theory, the Bayesian paradigm is based on a marker compound ferulic acid of A. sinensis and environmental variables
concept that the probability of an event can be corrected by the of 90 sampling sites based on the Matlab software to determine vari
knowledge of additional information (Smith et al., 1985), and the ables’ thresholds (Lu et al., 2012; Guo et al., 2016). The value of each
modification of this probability is called the condition. BN is usually a membership function (degree of membership) indicates that the higher
common method when Bayesian theory applies to practical research the value of the function, the higher the content of the marker com
(Havron et al., 2017). More specifically, BN is causal network that as pound ferulic acid. According to the degree of membership, we classify
signs probabilities to all internal nodes (“child nodes”). The parent the states of the environmental variables. We divide the numerical range
nodes and the child nodes connect by directional arrow. The states of for FMC so that each SEVs node could classify into four states (Table 2).
their parent nodes are used as the condition for probability transfer. “1”, “2”, “3”, and “4” respectively represent node state 1, 2, 3, and 4 of
4
Q. Zhang et al. Ecological Modelling 450 (2021) 109560
Table 3 Table 5
Calculate numerical intervals by the membership function of variable Bio1. A serial number of learning documents for FBM’s machine learning. We extract
Values of F(x) Numerical intervals of x
from 90 sampling sites, and divided them into 9 groups randomly, according to
the increasing sequence number from 1 to 9, with the interval of 10 sampling
0 ~ 0.30 x < 4.61; x >10.36 sites to obtain the learning documents of 9 sets of sampling sites. Finally, we
0.30 ~ 0.50 4.61 ≤ x < 5.31; 9.67 < x ≤ 10.36
could get 9 learning documents.
0.50 ~ 0.70 5.31 ≤ x < 5.92; 9.05 < x ≤ 9.67
0.70 ~ 1.00 5.92 ≤ x ≤ 9.05 The learning documents Number of sampling sites Sequence number
1 10 Seq 1st
2 20 Seq 2nd
3 30 Seq 3rd
Table 4
4 40 Seq 4th
The eight SEVs’ states divided by equal interval classification (EIC) method.
5 50 Seq 5th
SEVs (unit) Four states of SEVs node in FBM network 6 60 Seq 6th
I II III Ⅳ 7 70 Seq 7th
8 80 Seq 8th
Bio1 ( C)
◦
~7.88 7.88~11.65 11.65~15.43 15.43~
9 90 Seq 9th
Bio10 ( ◦ C) ~16.18 16.18~19.35 19.35~22.53 22.53~
Bio12 (mm) ~593.50 593.50~892 892~1190.50 1190.50~
Bio15 ~67.75 67.75~79.50 79.50~91.25 91.25~
Bio18 (mm) ~308.50 308.50~452 452~595.50 595.50~
2.5. The states of IEVs and habitat suitability nodes in FBM
SOM (‰) ~89.18 89.18~104.23 104.23~119.29 119.29~
Totalrad13 ~4150.18 4150.18~4697.98 4697.98~5245.77 5245.77~ For the three IEVs and habitat suitability nodes, we set the states as
(MJ/m2) “unsuitable”, “lowly suitable”, “moderately suitable”, and “highly suit
EL (m) ~1084.75 1084.75~1743.50 1743.50~2402.25 2402.25~
able”. We have three steps to confirm the state of the IEVs and habitat
I, II, III, and Ⅳ represent node state I, II, III, and Ⅳ of SEVs, respectively. suitability node in each sampling sites. Firstly, we calculate and obtain
membership function with all eight SEVs. Secondly, the weight of
SEVs by FMC, and different states represent respective numerical environmental variables is confirmed by the weight of the evaluation
intervals. index, which determined by the maximum entropy principle of infor
The process of dividing numerical intervals has the following three mation (Zhao et al., 2017). Thirdly, the comprehensive suitability
steps. Take Bio1 as an example, firstly, according to Bio1 values and the evaluation index Rk is determined by the fuzzy mathematics compre
marker compound ferulic acid of 90 sampling sites, the relationship hensive evaluation, calculated as Eq. (2). Here, Rw is the information
between the Bio1 and ferulic acid is fitted using Matlab software. Eq. (1) entropy weight vector of the index, Rξ is the fuzzy matter element (Lu
is the membership function. et al., 2012).
( ) [ ]
(1) M1 M2 ⋯ Mm
F(x) = exp − ((10x − 74.86)/26.18)2 Rk = R w R ξ = (2)
Kj K1 K2 ⋯ Km
5
Q. Zhang et al. Ecological Modelling 450 (2021) 109560
Table 6 Table 7
The occurrence rate of “ × ” and “–” was compared under three models (A, B, and The results of 10 testing sites based FME and FBM.
C), two data discrete methods (FMC and EIC), and different learning stages (Seq ID of testing FME FBM by fuzzy mathematics FBM by equal
1st, Seq 2nd-5th, and Seq 6th-9th). sites classification (FMC) interval
Model States of Fuzzy mathematics Equal interval classification (EIC)
habitat classification (FMC) classification (EIC) Model Model Model Model Model
suitability Seq Seq Seq Seq Seq Seq A B C B C
node 1st 2nd- 6th- 1st 2nd- 6th- 1 h h h h m h
Seq 5th Seq 9th Seq 5th Seq 2 m m m m h h
9th 3 h h h h h h
A “×” 82% 60% 31.5% 4 h h h h h h
“–” 14% 33.5% 55.5% 5 h h h h h h
B “×” 60% 11.5% 0 70% 49.5% 0 6 l – l l l l
“–” 0 0 0 0 0 0 7 n – n n l l
C “×” 60% 11.5% 0 70% 49% 0 8 l l l l l l
“–” 6% 12.5% 1% 6% 10.5% 14% 9 l l l l l l
10 l l l l l l
“ × ” represents an impossible distribution in the process of network probabi
listic inference, beyond the predicted range, which is called “conflict”. “–” h, m, l, and n represent highly suitable, moderately suitable, lowly suitable, and
represents the probability value of highly, moderately, lowly suitable, and un unsuitable, respectively. “–” represents the probability value of highly, moder
suitable is equal to 25%, which is called “equal probability”. The “conflict” and ately, lowly suitable, and unsuitable is equal to 25%, which is called “equal
“equal probability” are the performance of the model’s poor extraction of probability”.
effective information and reasoning ability, after learning the data of the cor
responding A. sinensis sampling sites. 2.8. Sensitivity analysis of environmental variables
corresponding likelihood function, and conversely, the fewer explana In this paper, the state of each node and CPT of all nodes are obtained
tory variables, the smaller the likelihood function (Kharroubi and according to the A. sinensis sampling sites, and the sensitivity analysis of
Sweeting, 2010). Here, the log-likelihood of the data given the model is environmental variables affecting the habitat of A. sinensis is carried out.
simply the sum of the log-likelihoods of the individual cases. The size of Starting from the parent node, the state of the node is input Hugin
the formula value takes from the sum of squared residuals (and the software for inference. Each state of a node is preset to 100%, and the
sample size). And, the formula is negative. Hence, the smaller the ab remaining nodes are not set. The remaining nodes are probabilistic
solute value and the smaller the sum of squared residuals, the higher the transmitted according to Bayesian inference. The probability difference
effectiveness of parameter fitting (Lie et al., 2018). However, as the between each child node and its initial state is calculated. The larger the
value of the log-likelihood function becomes smaller, we cannot judge difference, the stronger the sensitivity.
that the model with reduced variables is better than the previous model.
Akaike information criterion (AIC) commonly used for model se 3. Results
lection (Burnham and Anderson, 2004). AIC is a standard to measure the
goodness of fitness, characterizing how well the model matches the data 3.1. The prediction results of model A, B, and C
(Akaike, 1987). For FBM, the fitting validity of the model represents the
matching degree of the model to the data. Here, AIC is computed as l-k, The predicted results of model A, B, and C with 50 predicted sites of
where l is the log-likelihood and k is the number of free parameters in the nine kinds of sites sequences are show in Appendix A. There are four
model. Using Hugin software, the result of simplified AIC formula is states of habitat suitability of A. sinensis, including highly suitable,
negative. The smaller the absolute value, the better the model perfor moderately suitable, lowly suitable, and unsuitable. There are some
mance (Burnham and Anderson, 2004). special results, like “ × ” and “–”, which can characterize the perfor
mance differences between three models. The “ × ” means the impos
2.7. Modelling process sible distribution in the network probability reasoning process, beyond
the predicted range, which is called “conflict”. The “–” means the
The models are ordered from the standpoint of their adequacy probability of highly, moderately, lowly suitable, and unsuitable is equal
(Halfon and Reggiani, 1978), therefore, we choose three kinds of to 25%, which is called “equal probability”. When calculating the rele
network models (Fig. 2), and two methods of information classification vant indicators in the results, we find that Model A is learned from the
(FMC and EIC) with nine kinds of learning sequence of sampling sites network structure based on the original data, which performed poorly
(Table 5) to compare the suitability and uncertainty of the three models. (Appendix Table A1). In the two discretization methods, whether it is
In the study area, 50 predicted sites (Fig. 1) are randomly selected, model B (Appendix Table A2, A3) or model C (Appendix Table A4, A5),
which are used as the prediction sites for the suitability of A. sinensis, and the performance of FMC is better than EIC. All models learn data from 10
the accuracy and uncertainty of the model are evaluated. sampling sites (Seq 1st), and in the prediction results, “conflict” has a
Even if it is difficult to collect data on the occurrence of A. sinensis higher occurrence rate (Table 6). Obviously, it caused by too little
with complete information including ferulic acid content, we still collect sample information. Consider Seq 1st as the first stage, Seq 2nd to Seq
additional 10 testing sites information (including ferulic acid content 5th as the second stage, and Seq 6th to Seq 9th as the third stage. In these
information), to compare the results of three FBM model (Fig. 2) con three stages, with the prediction results of models A, B, and C, the
structed by the two methods of information classification (Table 2, incidence of “conflict” is gradually decreasing (Table 6). For “equal
Table 4) with the FME. probability”, the occurrence rates are different due to different models.
Model B does not have the phenomenon of “equal probability” (Ap
pendix Table A2, A3), while model A has been increasing in these three
6
Q. Zhang et al. Ecological Modelling 450 (2021) 109560
Fig. 3. Conditional probability distribution table (CPT) in B model after learning 90 sampling sites with SEVs’ states divided by FMC (Seq 9th), IEVs’ states divided
by comprehensive suitability evaluation, in which h, m, l, and n respectively represent node state highly suitable, moderately suitable, lowly suitable, and unsuitable
of IEVs. And 1, 2, 3, and 4 respectively represent node state 1, 2, 3, and 4 of SEVs. They correspond to the states 1–4 in Table 2.
stages (Table 6). When model C is based on EIC, the occurrence rate of Based on the fitting validity, stabilization of the model and predicted
“equal probability” has been increasing (Appendix Table A5), and basis results, the model B with SEVs’ states divided by FMC is the best. Fig. 3
of FMC, it first rises and then decreases (Appendix Table A4, Table 6). shows the conditional probability distribution of model B after learning
the information by FMC from environmental data. Finally, we confirm
that the predicted results of Seq 9th of B model with SEVs’ states divided
3.2. Test results
by FMC and IEVs’ states divided by comprehensive suitability evalua
tion is accurate for A. sinensis, showed in Fig. 5. Among them, if the
We use 10 testing sites to evaluate the performance of FBM (models
predicted sites have the same probability of suitability (Appendix
A, B, and C) and FME. The result shows that models A, B, and C are in a
Table A2), we show the lowest level of suitability (Fig. 5). Generally, it
stable after learning the information of 90 sampling sites (Appendix
can be seen that the highly and moderately suitable habitats of A. sinensis
A1–5). Thus, FBM chose CPT after learning the information of 90 sam
are mainly distributed in Gansu and Shaanxi, which is consistent with
pling sites (Seq 9th) to infer the results of the test sites. The FME also
genuine producing areas of A. sinensis (Shang et al., 2015). Lowly suit
used information from 90 sampling sites to model. Compared with FBM
able habitats are located in Yunnan, Sichuan, Chongqing, and Hubei.
and FME (Table 7), the test results of model B and C are consistent with
the results of FME by FMC. Model A has two “equal probability” oc
currences. Except for ID1 of test sites, the prediction results of models B
3.4. Sensitivity analysis of SEVs and IEVs on habitat suitability of A.
and C test sites are consistent by EIC. In terms of FME prediction results,
sinensis
there is 30% difference. Model B has an unclear boundary in the
discrimination between highly suitable and moderately suitable through
The sensitivity analysis is provided based on CPT (Fig. 3) of B model
EIC. From the results, the operation of model B and model C through
after learning 90 sampling sites with SEVs’ states divided by FMC (Seq
FMC is more stable and good.
9th) and IEVs’ states divided by comprehensive suitability evaluation
[eq. (2)]. In order to analyze the degree of influence of SEVs and IEVs on
3.3. Model fitting validation, stabilization and optimal model the habitat suitability of A. sinensis, we set the limit values for each node
in the FBM, which is to set the SEVs and IEVs nodes separately. We set
By FMC, the absolute value of AIC of model A tends to be the highest the probability of a node in a certain state to 100%, the other state
among the three models, and the absolute value of AIC of model B is the probability settings of this node are all 0 (Appendix Table A6). At the
lowest (Fig. 4a). The absolute value of log-likelihood of model A is the same time, the remaining nodes retain the original probability state.
lowest among the three models, followed by model B and model C After the probability transfer, the state probabilities of A. sinensis suit
(Fig. 4b). By EIC, the absolute value of AIC of model B is much smaller able habitat nodes are recorded, and these state probabilities are
than that of model C (Fig. 4c), but the absolute value of log-likelihood of compared to the state probabilities of the habitat suitability nodes of
model C is smaller than the value of model B (Fig. 4d). In the experiment, A. sinensis (Fig. 3). The result indicates that different probability setting
the model with the least AIC absolute value is usually chosen. Therefore, methods lead to changes in the probability values of four suitability
the AIC absolute value of model B is the minimum between the two levels (Appendix Table A6). We extract the maximum and minimum
methods of discretization (FMC and EIC). For model B, the AIC absolute values of the probability change of suitability (Fig. 6).
value of FMC (Fig. 4a) is smaller than that of EIC (Fig. 4c). For the result of SEVs, the main environmental variables that
7
Q. Zhang et al. Ecological Modelling 450 (2021) 109560
Fig. 4. AIC and Log-likelihood indexes of model A, B, and C with 9 kinds of learning sequences of sampling sites. (a) and (b) are under the fuzzy mathematics
classification (FMC). (c) and (d) are based on equal interval classification (EIC).
8
Q. Zhang et al. Ecological Modelling 450 (2021) 109560
Fig. 5. Habitat suitability map of 50 predicted sites of A. sinensis obtained from the Seq 9th of B model with SEVs’ states divided by FMC and IEVs’ states divided by
comprehensive suitability evaluation.
absence of current data, it is still feasible for BN to model biodiversity at the suitability of A. sinensis into “unsuitable”, “lowly suitable”,
fine spatial scales (Grafius et al., 2019). The combination of ecological “moderately suitable”, and “highly suitable”. FMC and EIC are used to
process interaction and spatial explicit data in the BN frame can improve preprocess SEVs. FMC uses a mapping relationship between environ
researchers’ clear understanding of the process in space (Froese et al., mental variables and membership degrees. The EIC is used as a control
2019). Here, RF and Maxent have better spatial distribution perfor group corresponding to FMC to participate in model data preprocessing.
mance to a certain extent (Elith et al., 2010). However, only Maxent can Bayesian networks usually require discrete data (Uusitalo, 2007). Dis
use categorical data as input for environmental variables (Guo et al., cretization means that only the rough features of the original distribu
2017, 2019). FBM retains the input advantage of Maxent in data and tion are captured (Friedman and Goldszmidt, 1996), and statistical
allows input of categorical data. In addition, FBM composed of BN al information is lost (Hamilton et al., 2015; Lucena-Moya et al., 2015;
lows the addition of more empirical and decision-making data, which Meyer et al., 2014). However, discretization is beneficial in this study.
can effectively integrate various ecological data. Other models (such as With the increase of learning data, FBM has successfully provided reli
Maxent, RF) are difficult to do. FBM has unique advantages in flexible able results (Table 6), sensitivity analysis (Appendix Table A6), and the
data fusion (including data types that can be added, visualization of data popular FBM has not provided biased results. Therefore, discretization
relationships, etc.). can mitigate the effects of noise data (incorrectly saved values/typing
errors/incorrect values) (Feki-Sahnoun et al., 2018).
9
Q. Zhang et al. Ecological Modelling 450 (2021) 109560
mathematical denoising algorithm. Models B and C both use FMC growth, or the structure of the network model is not reasonable. The
(Table 2) and EIC (Table 4) discretization methods, respectively. results show that the model requires a large amount of original data, and
Observation shows that two models (Models B and C) better fitting and the quality and quantity of the original data have a profound impact on
reasonable reasoning, when learning FMC data (Fig. 4). In the com the prediction results of the FBM model. This is the same as other SDMs,
parison between FBM and FME, the results also show that FMC is su such as GLM, GARP, and RF (Hirzel et al., 2001; Ray et al., 2018; Zhang
perior to EIC (Table 7). Wiest et al. (2019) believes that a target training et al., 2020b).
data set can build a model probability structure, which helps a more
robust cross-validation analysis. We developed FBM on a regional scale
4.4. Case and features of FBM
rather than a smaller sub-region or statistical scale. The three FBM
frameworks and the corresponding target training data sets can build the
In China, A. sinensis is widely known as a CHM (Committee of Na
model probability distribution, which is the performance of powerful
tional Pharmacopoeia, 2020), the quality of A. sinensis and its marker
cross-validation analysis. Although there are no specific guidelines to
compound ferulic acid are systematic research (Zhang et al., 2012, 2019;
derive the number of cases for the structure and parameters of the FBM,
Ma et al., 2015). Sufficient and perfect basic research data can effec
as the cases increase, the BN model becomes more robust. In all FBM,
tively support our research. This is why we choose A. sinensis instead of
whether used FMC or EIC, it is shows that after the model structure
other medicinal plants as a typical case.
parameters are fixed, as the amount of data increases, the implicit pa
Each model has its own theoretical framework and is different from
rameters required for inference will increase (Fig. 4), and the absolute
the concept definition with species suitability. Different from most
values of the AIC and log-likelihood functions will also increase. By
previous models, such as GLM, GARP, RF, Maxent (Hirzel et al., 2001;
evaluating the FMC data, Model A, Model B, and Model C have very
Ray et al., 2018; Zhang et al., 2020b; Guo et al., 2017), FBM has a unique
different feedback. Model A only comes from machine structure
understanding of species suitability. As described in the above method,
learning. Model A has fewer nodes, so the model has fewer independent
the membership function fits the marker compound ferulic acid of
variables than the other two models, and the absolute value of the
A. sinensis and environmental variables together, and then further pro
log-likelihood is also smaller (Fig. 4b). But model A has too many hidden
cesses to obtain the level of species suitability. Because FBM currently
variables, and the absolute value of AIC is much higher than that of the
mainly targets medicinal plants, to a large extent, the level of the content
other two models (Fig. 4a). Similarly, model B is superior to model C in
of species marker compound, that is, the level of medicinal value, rep
data fitting, as is the model structure, because the implicit parameters of
resents the level of species suitability. FBM emphasizes that the content
model B are less than model C, it can prevent the model from overfitting
of species marker compound determines the suitability of the species’
during the mapping process.
habitat. The original data required by the FBM model includes the
Although the results contain a number of “conflict” and “equal
content of species marker compound, the location of the species
probability” events, the probability of this happening decreases with the
occurrence, and environmental data. We have compared FBM with FME
increase of the learning data (Table 6). The “conflict” indicates that the
in the manuscript. However, models such as GLM, GARP, RF, and
software cannot obtain results after calculation. The reason is that the
Maxent combine observations of species occurrence or abundance with
network’s inference oversteps the scope of reasoning. This means the
environmental estimates (Elith and Leathwick, 2009). There is no data
network probability distribution selected by researchers cannot exist in
on the content of species marker compound for simulation. Therefore,
the current network. The predicted sites are not suitable for the species’
for medicinal plants, FBM’s prediction mechanism is more effective and
10
Q. Zhang et al. Ecological Modelling 450 (2021) 109560
can help us quickly predict where there will be high-quality medicinal explore relationships between environmental variables, thereby
materials. Because of the different definitions of habitat suitability and improving the model’s ability to identify and fit data and the efficiency
different data sources, the structure of FBM and other models (such as of using species data.
RF and Maxent) are very different, and the evaluation methods are also
different. The reliability from different model systems actually comes 5. Conclusion
from different statistical models (Strandén and Christensen, 2011). The
comparison between FBM and other models (such as RF and Maxent) Combining fuzzy mathematics, expert experience, machine learning,
cannot help FBM to improve model prediction accuracy, reliability and and Bayesian reasoning, taking A. sinensis as a case, a species habitat
robustness. suitability model FBM is established in this study. Model A is generated
Through a series of reliability and robustness analyses, this study by machine learning. Models B and C combine previous research and
adopted the results of Seq 9th of B model with SEVs’ states divided by experience. Compared with model A, the model frameworks B and C are
FMC, and IEVs’ states divided by comprehensive suitability evaluation more accurate and reliable. Model B has simple structure parameters,
(Fig. 5). In this study, 50 predicted sites of A. sinensis were randomly efficient data identification, and data output stability is better than
selected in the study area (Fig. 1). From the prediction results, the dis Model C, and Model A. Comparative experiments of two discretization
tribution of A. sinensis habitat suitability is related to the sampling sites. methods, FMC and EIC, show that the results of FMC is effectively
The prediction result reflects the distribution pattern of ferulic acid recognized by the FBM, and reliable results are output. FBM sensitivity
content in A. sinensis sampling sites. The highly and moderately suitable analysis helps researchers explore in detail the effects of environmental
habitats of A. sinensis are mainly distributed in Gansu and Shaanxi factors on various aspects of species habitat suitability. It is easy for
(Shang et al., 2015). This also further confirms that the model has a researchers to obtain terrestrial plants’ environmental data needed for
reasonable mapping and promotion ability. FBM operation, and many plants have species characteristics that can be
FBM adds sensitivity analysis to help researchers further explore the used for FBM. Hence, the concept and framework of FBM may have a
impact of variables on habitat suitability. From Appendix Table A6, we broad application prospect for research on species habitat suitability in
further extract the degree of influence of each node on the habitat status the future.
of A. sinensis (Fig. 6). Through the maximum and minimum values in the
suitability probability changes (Appendix Table A6), we can find out CRediT authorship contribution statement
that SOM, Bio1, Bio10, and EL affect the increase and decrease of four
suitability probabilities in different degrees (Fig. 6). In terms of the Quanzhong Zhang: Conceptualization, Methodology, Software,
range of suitability probability changes (min to max), almost all four Formal analysis, Data curtion, Writing – original draft, Writing – review
suitability changes are dominated by these environmental variables & editing, Visualization. Haiyan Wei: Writing – review & editing, Su
(SOM, Bio1 and Bio10). For IEVs, the temperature suitability node still pervision. Jing Liu: Investigation, Validation, Writing – review & edit
has a great influence on suitability of A. sinensis (Fig. 6). ing. Zefang Zhao: Data curtion, Writing – review & editing. Qiao Ran:
FBM performs Bayesian inference based on FMC. The role of fuzzy Data curtion, Writing – review & editing. Wei Gu: Conceptualization,
mathematics in this study is for the discretization of nodes. According to Validation, Writing – review & editing, Supervision, Project adminis
the degree of membership, we classify the states of the environmental tration, Funding acquisition.
variables (Table 2). This means that each state of the eight SEVs itself
contains some information that supports or not support the suitability of Declaration of Competing Interest
A. sinensis. For example, Bio1 node (Appendix Table A6), when the
probability of “4′′ is 100%, the probability of highly suitable is the The authors declare that they have no known competing financial
largest 39.82%, and the probability of unsuitable is the smallest 11.58%. interests or personal relationships that could have appeared to influence
In contrast, when the “1′′ probability is 100%, the highly suitable the work reported in this paper.
probability is the smallest 17.29%, and the unsuitable probability is the
largest 21.80%. In summary, the four states of FMC (Table 2), from state Acknowledgments
1 to state 4, imply the gradual support of environmental variables for the
growth of A. sinensis. Obviously, FMC can extract the relationship be We thank everyone in our team, because we summarized our pre
tween the value of environmental variables and the growth of A. sinensis, vious studies to make progress and innovation. We also pay tribute to
while EIC cannot (Fig. 4, Table 6). Hence, FMC significantly improves researchers who engage in the study of species distribution. Also, we
the operating efficiency of FBM. Through FMC and the sensitivity appreciate Miss Yunfei Gu, a doctoral student at the University of
analysis (Appendix Table A6), Bio1 in 5.92 ◦ C-9.05 ◦ C, Bio10 in 14.80 Southampton for assisting us in revising the paper. This work is sup
◦
C-18.60 ◦ C, Bio12 in 568.79 mm-791.33 mm, Bio15 in 69.04–87.48, ported by the Basic Research Programs of Natural Science Foundation of
and Bio18 in 284.28 mm-428 mm can promote the highly suitable Shaanxi Province (No. 2020JM-277), the National Natural Science
habitats of A. sinensis (Table 2), which is consistent with the conclusion Foundation of China (No. 31070293), and the Research and Develop
of previous studies (Shang et al., 2015; Zhang et al., 2017). Consistent ment Program of Science and Technology of Shaanxi Province (No.
with the research results of genuine producing areas, SOM has a pro 2014K14–01–02).
found effect on the growth quality of A. sinensis (Zhao et al., 2002). This
is an interesting finding. We can continue to analyze the range where Appendix A
SEVs have the greatest influence on the moderately suitable, lowly
suitable, and unsuitable habitats of A. sinensis (Table 2, Appendix Table A1, A2, A3, A4, A4, A5, and A6
Table A6). Unlike FME, FBM pre-processes the data while preserving the
adaptability of the species to the environment. It adds a machine
learning process that can perform detailed probabilistic reasoning and
11
Q. Zhang et al. Ecological Modelling 450 (2021) 109560
Table A1
The results of 50 predicted sites based on model A with 9 kinds of learning sequences of sampling sites with SEVs’ states divided by fuzzy mathematics classification
(FMC) and IEVs’ states divided by comprehensive suitability evaluation.
ID of predicted sites Seq 1st Seq 2nd Seq 3rd Seq 4th Seq 5th Seq 6th Seq 7th Seq 8th Seq 9th
1 × – – – – – – – –
2 × × × – – – – – –
3 × × – – – – – – –
4 × × × – – – – – –
5 × × × – – – – – –
6 × × × × – – – – –
7 × × × × × × × × ×
8 × × – – – – – – –
9 × × × × – – – – –
10 × × × × l l l l l
11 × × × × × × × × ×
12 n n n N n n n n n
13 – × × × × n n n n
14 × – – – – – – – –
15 × × × × × × × × ×
16 × × × × × × × × ×
17 × × × × × × × × ×
18 – – – – – – – – –
19 × × – – – – – l l
20 × × – – – – – – –
21 × × × × × × × × ×
22 × × × × × × – × ×
23 × × × × × × × × ×
24 × × × – – – – – –
25 × – – – – – – – –
26 × × × – – – × × ×
27 × × – – – – l l l
28 × × × × – – – – –
29 – × × × – – – – –
30 – × × × × × × × ×
31 × × × × × × × × ×
32 × × × × – – – – –
33 × × × × – – – – –
34 × × × × × × × × ×
35 × × × × × × × × ×
36 × × × × – × – – –
37 × × × × × × × × ×
38 × × × × × – – – –
39 × × × × – – – – –
40 × × × × × × × × ×
41 × × × × – – – – –
42 – – – – – – – h –
43 × h h H h h h h h
44 × × – – – – – – –
45 h h/m h/m h/m m m m h h
46 – – – – – – – – –
47 – × × × – – – – –
48 × × × × × × × × ×
49 × × × × – – – – –
50 × – – – – – – – –
h, m, l, and n represent highly suitable, moderately suitable, lowly suitable, and unsuitable. The “h/m” represents the probability value of highly and moderately
suitable is equal, and is greater than that of lowly suitable and unsuitable probability. “–” represents the probability value of highly, moderately, lowly suitable, and
unsuitable is equal to 25%, which is called “equal probability”. “ × ” represents an impossible distribution in the process of network probabilistic inference, beyond the
predicted range, which is called “conflict”, and the final prediction result was regarded as unsuitable.
12
Q. Zhang et al. Ecological Modelling 450 (2021) 109560
Table A2
The results of 50 predicted sites based on model B with 9 kinds of learning sequences of sampling sites with SEVs’ states divided by fuzzy mathematics classification
(FMC) and IEVs’ states divided by comprehensive suitability evaluation.
ID of predicted sites Seq 1st Seq 2nd Seq 3rd Seq 4th Seq 5th Seq 6th Seq 7th Seq 8th Seq 9th
1 × l × l l l l l l
2 × l n l l l l l l
3 × l × l l l l l h
4 × l × h h h h h l
5 × m n l l l l l l
6 × l h h h h h h h
7 l l l l l l l l l
8 m m m m m m m m m
9 × × × h h h h/l h l
10 × l × l l l l l l
11 × × × l l h m h/m/l h/l
12 × l l/n l l l l l l
13 l l n n n n n n n
14 l m n n n n n n n
15 × l × l l l l l l
16 × l l/n l l l l l l
17 × l l n n n n n n
18 l l h/l l l l l l l
19 l l h/l l l l l l l
20 l m n l l l l l l
21 × m l l l l l l l
22 × × × l l l l l m/l
23 × m × l l l l l l
24 l l l l l l l l l
25 × l × l l l l l l
26 × l n l l l l l l
27 l l l/n l l l l l l
28 × × × l l l l l l
29 l l l/n l l l l l l
30 l l l/n l l l l l l
31 × m n l l l l l l
32 × l l l l l l l l
33 × l l l l l l l l
34 × × × n n n n n n
35 × l l n n n n n n
36 l l l l l l l l l
37 × l × n n n n n n
38 l l l/n l l l l l l
39 l l n l l l l l l
40 × l × l l l l l n
41 l l n l l l l l l
42 × l h h h h h h h
43 h h h h h h h h h
44 m m m m m m m m m
45 m m h h h h h h h
46 l l l/n l l l l l l
47 h/m h h h h h h h m
48 × l h/l l l l l l l
49 × × × h h h h/l h l
50 × × × l l l l l l
The “h/m/l” represents the probability value of highly, moderately, and lowly suitable is equal, and is greater than that of unsuitable probability. The “h/l” represents
the probability value of highly and lowly suitable is equal, and is greater than that of moderately suitable and unsuitable probability. The “m/l” represents the
probability value of moderately and lowly suitable is equal, and is greater than that of highly suitable and unsuitable probability. The “l/n” represents the probability
value of lowly suitable and unsuitable is equal, and is greater than that of highly and moderately suitable probability. “–” and “ × ” are shown Table A1.
13
Q. Zhang et al. Ecological Modelling 450 (2021) 109560
Table A3
The results of 50 predicted sites based on model B with 9 kinds of learning sequences of sampling sites with SEVs’ states divided by equal interval classification (EIC)
and IEVs’ states divided by comprehensive suitability evaluation.
ID of predicted sites Seq 1st Seq 2nd Seq 3rd Seq 4th Seq 5th Seq 6th Seq 7th Seq 8th Seq 9th
1 l l l l l l l m/l l
2 × × l l l l l m/l l
3 m h h h h h m m m
4 l l l l l l h h h
5 l m m m m m m m m
6 × × × × × l l l h
7 × × × × × l l l l
8 m h m m m m m m m
9 × × × × × l l l l
10 × × × × × l l l l
11 × × × × × l l l l
12 × × × × × l l l l
13 × × × × l l l l l
14 × × × × × l l l l
15 × × × × l l l l l
16 × × × × l l l m/l l
17 × × l l l l l l l
18 l h/l h/m/l l l h h h h
19 l l l l l l l l l
20 l l l l l l l l l
21 × × l l l l l l l
22 m l l l l l l l l
23 × × l l l l l m/l l
24 l l l l l l l l l
25 × × l l l l l l l
26 × × l l l l l l l
27 l l l l l l l l l
28 × × l l l l l l l
29 × × l l l l l l l
30 × × l l l l l m/l l
31 × × × × × l l l l
32 × × × × h h h h h
33 × × × × l l l l h
34 × × × × l l l l l
35 × × × × l l l l l
36 × × × × l l l l l
37 × × × × l l l l l
38 × × × × l l l m/l l
39 × × × × × n l l l
40 × × × × l l l m/l l
41 × × × × l l l l l
42 × × × × × h h h h
43 m h h × h h h h h
44 × × h h h h h h h
45 m h h h h h h h h
46 m h m m m m m m m
47 m h/m h h h h h h h
48 × × × × × l l l h
49 × × × × × l l l l
50 × × × × × l l l l
The letters and symbols in the table are explained in Table A1 and Table A2.
14
Q. Zhang et al. Ecological Modelling 450 (2021) 109560
Table A4
The results of 50 predicted sites based on model C with 9 kinds of learning sequences of sampling sites with SEVs’ states divided by fuzzy mathematics classification
(FMC) and IEVs’ states divided by comprehensive suitability evaluation.
ID of predicted sites Seq 1st Seq 2nd Seq 3rd Seq 4th Seq 5th Seq 6th Seq 7th Seq 8th Seq 9th
1 × l × n n n n n n
2 × l l/n l/n l/n l/n l/n l/n l
3 × – × h h h/m h/m h/m h
4 × – × h h h/m h/m h/m h/m
5 × – – – – – l l l
6 × h – – – h h h h
7 l – – l l l l l l
8 l l l/n l/n l/n l/n l/n l/n l
9 × × × l l l l l l
10 × l × l l l l l l
11 × × × n n n n n n
12 × l /n l l l l l l l
13 l l n n n n n n n
14 – – h h h h/m h h h
15 × l × n n n n n n
16 × l n n n n n n n
17 × l l l l l l l l
18 l l l l l l l l l
19 l l l l l l l l l
20 l l l l l l l l l
21 × l l l l l l l l
22 × × × l l l l l l
23 × l × l l l l l l
24 l – – l l l l l l
25 × l × l l l l l l
26 × l l l l l l l l
27 l l n l l l l l l
28 × × × l l l l l l
29 l l n l l l l l l
30 l l n n n n n n n
31 × l l l l l l l l
32 × – – l l l l l l
33 × – – l l l l l l
34 × × × n n n n n n
35 × l n n n n n n n
36 l – – l l l l l l
37 × l × n n n n n n
38 l l n n n n n n n
39 – – h h h h/m h h h
40 × l × n n n n n n
41 l l n l l l l l l
42 × h /m h h h h h h h
43 h h h h h h h h h
44 m m h h h h h h h
45 m m h h h h h h h
46 l l l/n l/n l/n l/n l/n l/n l
47 – h h h h h h h h
48 × – – – – – l l l
49 × × × l l l l l l
50 × × × l l l l l l
The letters and symbols in the table are explained in Table A1 and Table A2.
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Q. Zhang et al. Ecological Modelling 450 (2021) 109560
Table A5
The results of 50 predicted sites based on model C with 9 kinds of learning sequences of sampling sites with SEVs’ states divided by equal interval classification (EIC)
and IEVs’ states divided by comprehensive suitability evaluation.
ID of predicted sites Seq 1st Seq 2nd Seq 3rd Seq 4th Seq 5th Seq 6th Seq 7th Seq 8th Seq 9th
1 m m m m m m m l l
2 × × m m m m m l l
3 m m m m m m m m m
4 l l l l l l l l l
5 l m m m m m m m m
6 × × × × × – h h h
7 × × × × × l l l l
8 – – – – – – – – l
9 × × × × × l l l l
10 × × × × × l l l l
11 × × × × × l l l l
12 × × × × × l l l l
13 × × × × n n n n n
14 × × × × × – – – –
15 × × × × l l l l l
16 × × × × l l l l l
17 × × m m m m m m m
18 – l l l l l l l l
19 l l l l l l l l l
20 l l l l l l l l l
21 × × – – l l l l l
22 l l l l l l l l l
23 × × l l l l l l l
24 – – – – – – – – –
25 × × l l l l l l l
26 × × l – l l l l l
27 l l l l l l l l l
28 × × l l l l l l l
29 × × – – l l l l l
30 × × m m m m m l l
31 × × × × × – – – –
32 × × × × l l l l l
33 × × × × l – – – –
34 × × × × l l l l l
35 × × × × l l l l l
36 × × × × l l l l l
37 × × × × l l l l l
38 × × × × l l l l l
39 × × × × × – – – –
40 × × × × l l l l l
41 × × × × l l l l l
42 × × × × × l h h h
43 h/m/l – – – – – h h h
44 × × h h h h h h h
45 h h h h h h h h h
46 l m m m m m m m m
47 l – – – – – m m h/m
48 × × × × × l l l l
49 × × × × × l l l l
50 × × × × × l l – –
The letters and symbols in the table are explained in Table A1 and Table A2.
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Q. Zhang et al. Ecological Modelling 450 (2021) 109560
Table A6
Sensitivity analysis on habitat suitability node in B model after learning 90 sampling sites (Seq 9th) with SEVs’ states divided by FMC and IEVs’ states divided by
comprehensive suitability evaluation.
Nodes Probability setting of node state (%) The state of the node of the habitat suitability
h (%) h (%) change m (%) m (%) change l (%) l (%) change n (%) n (%) change
"1′′ , "2′′ , "3′′ , and "4′′ respectively represent node state 1, 2, 3, and 4 of SEVs. "h", "m", "l", and "n" respectively represent the node state of IEVs as highly suitable,
moderately suitable, lowly suitable and unsuitable habitat.
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Q. Zhang et al. Ecological Modelling 450 (2021) 109560
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