land

Article
Spatio-Temporal Changes and Habitats of Rare and Endangered
Species in Yunnan Province Based on MaxEnt Model
Yiwei Lian 1,2,3 , Yang Bai 1,2,3, *, Zhongde Huang 1,2,3 , Maroof Ali 1 , Jie Wang 1,2,3 and Haoran Chen 4

1 Center for Integrative Conservation & Yunnan Key Laboratory for Conservation of Tropical Rainforests and
Asian Elephants, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences,
Mengla 666303, China; [email protected] (Y.L.); [email protected] (Z.H.);
[email protected] (M.A.); [email protected] (J.W.)
2 Yunnan International Joint Laboratory of Southeast Asia Biodiversity Conservation, Jinghong 666303, China
3 University of Chinese Academy of Sciences, Beijing 100049, China
4 Institute of International Rivers and Ecological Security, Yunnan University, Kunming 650500, China;
[email protected]
* Correspondence: [email protected]

Abstract: Biodiversity is crucial for ecosystem functioning, but it is rapidly declining due to human
activities and climate change. Protecting biodiversity has become a key priority for global environ-
mental conservation actions. Rare and endangered species have a great impact on the ecosystem, yet
due to their limited survival capacity, they are more prone to extinction, thus exerting a significant im-
pact on biodiversity. However, current research reveals a lack of information concerning the potential
distribution and changes of these species. This study used the maximum entropy model to predict
the present and future potential habitats of rare and endangered species in Yunnan Province. After
superimposing model results, four richness regions are divided by the natural breakpoint method
and analyzed. Existing protected areas are compared with hotspots, and the land-use composition of
hotspots is also analyzed. The results revealed that, in both current and future scenarios, rare and
endangered species in Yunnan Province are primarily found in the western mountainous region,
the Xishuangbanna–Wenshan high temperature area, and the Kunming–Qujing dense vegetation
cover area. These species are also expanding their distribution towards the western mountainous
Citation: Lian, Y.; Bai, Y.; Huang, Z.; area. However, under the low carbon emission scenario (RCP2.6), these species will spread from
Ali, M.; Wang, J.; Chen, H.
the high abundance regions to the low altitude hotspots by 2070. In the high carbon emissions
Spatio-Temporal Changes and Habitats
scenario (RCP8.5), there will be fewer high abundance areas in 2070 than in 2050. The transfer matrix
of Rare and Endangered Species in
analysis reveals regional richness variations over time. Furthermore, the analysis revealed significant
Yunnan Province Based on MaxEnt
conservation gaps and found that existing hotspot areas were heavily affected by human activities.
Model. Land 2024, 13, 240. https://
doi.org/10.3390/land13020240
To improve conservation efficiency, it is necessary to enhance the protection of existing hotspots in
Yunnan Province. Climate change plays a significant role in species migration, with precipitation
Academic Editor: Guillermo J.
levels being a key factor. The necessary actions should be taken to address the insufficient protection,
Martinez-Pastur
resolve conflicts between human activities and land use in critical areas, and formulate effective
Received: 22 December 2023 strategies for adapting to future climate changes. Yunnan Province, with its rich species resources, has
Revised: 28 January 2024 the potential to become a global innovator in biodiversity conservation by implementing improved
Accepted: 10 February 2024 conservation strategies.
Published: 16 February 2024

Keywords: rare and endangered species; MaxEnt model; biodiversity prediction; climate change;
gap analysis

Copyright: © 2024 by the authors.

Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
1. Introduction
conditions of the Creative Commons With global environmental changes and explosive population growth, various factors,
Attribution (CC BY) license (https:// such as habitat loss, habitat fragmentation, resource overuse, invasion of alien species,
creativecommons.org/licenses/by/ environmental pollution, and human-induced climate change, along with barriers to bio-
4.0/). logical reproduction, have led to the degradation of ecosystems and loss of biodiversity [1].

Land 2024, 13, 240. https://doi.org/10.3390/land13020240 https://www.mdpi.com/journal/land

Land 2024, 13, 240 2 of 19

Biodiversity loss is a significant global problem that worsens over time [2]. Currently, bio-
diversity faces five major pressures: climate change, habitat loss and degradation, nutrient
overenrichment and pollution, overexploitation and unsustainable use, and invasive alien
species [3]. Furthermore, a conflict arises between economic level and national consump-
tion level development and biodiversity conservation [4]. The monitoring of endangered
species by the relevant authorities is inadequately implemented [5]. The overall trend of
biodiversity loss has not been effectively controlled, and China continues to cope with
a multitude of issues stemming from its economic development, which in turn puts a
significant strain on biodiversity [6].
Studies have demonstrated the significant impact of losing various life forms on the
structure and function of entire ecosystems. It also affects various ecosystem services [7,8].
This is particularly concerning for rare and endangered species (RESs) which have low
abundances and are more vulnerable to environmental change and extinction [9–11]. These
species often have small and fragmented geographical ranges [12]. These characteristics
correspond to species on the IUCN Red List of species on the six threatened levels of near
threatened, vulnerable, endangered, critically endangered, extinct in the wild, and extinct.
The contribution of RESs to ecosystems is crucial [13]. Furthermore, RESs have a poor ability
to reproduce and spread [14] and will become extinct if they are not protected [15], which
has greater significance for biodiversity. Biodiversity can have catastrophic consequences
for ecosystems, as it is crucial for human health and well-being. Therefore, reducing
biodiversity loss and protecting animal and plant resources are included in the Millennium
Development Goals and Action Goal Three of the Kunming–Montreal Global Biodiversity
Framework (GBF) [16,17].
At present, the Earth’s climate is experiencing rapid dynamic change [18]. These
changes are negatively impacting the habitats of RESs, leading to a decline in biodiversity
indicators [19]. Despite this, biodiversity pressure remains high, creating a correlation
between biodiversity change and climate change. While most organisms have some de-
gree of adaptability to environmental changes, human activity has reduced their ability
to adapt [20]. Therefore, predicting potential hotspots for RESs can help conservation
managers understand how species distributions may change under these circumstances.
Of the many ways to conserve biodiversity, in situ conservation, particularly through
the establishment of protected areas (PAs), is the most effective method for conserving
biodiversity [21,22]. PAs are cost efficient and play a crucial role in addressing biodiversity
loss [23–25], and establishing protection systems is key to all effective biodiversity conser-
vation tools [26]. However, even with the existence of PAs, species populations both inside
and outside PAs continue to decline [27]. Many PAs established before 1992 are facing
increasing human pressure, and changing environmental conditions may compromise
their ability to protect species and ecosystems in the future [28]. Therefore, it is urgent to
optimize the distribution pattern of PAs to effectively protect RESs.
The maximum entropy (MaxEnt) model is a very effective model for predicting a
species’ geographical distribution. It can only use the existing data to run the model, and
the prediction accuracy is high, which avoids data overfitting [29–31]. The MaxEnt model
can achieve the maximum level of randomness in a generic context, without being restricted
by pattern limitations [32]. The Maxent model’s predictions are closer to realistic niches [33],
so after the model passes the validation threshold, the results are highly available. The Max-
Ent model has been widely used to predict the habitat distribution of RESs, and Chinese
scholars have used it to predict the distribution area of Cornus officinalis and Thuja sutchue-
nensis Franch. [34,35]. Additionally, it has been used to study the geographical ranges of
endangered species around the world, like Ctenomys magellanicus, Cryptobranchus alleganien-
sis, and wild Nepeta crispa [36–38]. The forecast results were excellent. Previous studies
used different methods to identify conservation priority areas in Yunnan Province [39–41].
Some of these studies have applied the MaxEnt model to explore the distribution of RESs in
Yunnan Province [42–44]. However, there is still a lack of research that applies the MaxEnt
model to predict priority PAs in Yunnan Province. This study aims to fill the research gap
Land 2024, 13, 240 3 of 19

by considering species distribution patterns under future environmental changes. The fu-
ture environmental prediction model relies on four representative concentration pathways
(RCPs) that depict various emission trajectories. These RCPs include emission substances,
emission concentrations, and land-use trajectory. As the carbon dioxide concentration in
the atmosphere increases, the Earth’s temperature rises. The RCPs are named after the
radiative forcing target levels for 2100, which are estimated based on emissions and other
influencing agents. Four selected RCPs were considered in this study: RCP2.6 (very low
forcing levels), RCP4.5/RCP6 (moderately stable scenarios), and RCP8.5 (very high baseline
emission scenario).
This study assessed the efficacy of existing PAs by comparing the distribution of
simulated hotspots of RESs with that of existing PAs. The MaxEnt model was used to
predict the changes in the potential distribution area of RESs under future climate-change
scenarios (2050, 2070). The study extracted land-use data corresponding to high species-rich
regions. It analyzed the impact of major human activities on the potential distribution
of RESs. In this study, ArcGIS software was used to generate a habitat-change transfer
matrix, enabling the calculation of future increases or decreases in potential distribution
areas. This approach provides an effective means to quantify the potential habitat changes
of vegetation influenced by climate change.
This study aims to address the following questions: (1) are the existing PAs in Yunnan
Province effectively protecting the potential habitats of RESs? (2) Which land-use types
have the most significant impact on the potential habitats of RESs? What are the main
land-use types in the hotspot area? (3) How will the potential distribution of RESs change
under different emission patterns (RCP2.6, RCP4.5, RCP6.0, and RCP8.5) under future
climate change (2050, 2070)?

2. Materials and Methods

2.1. Study Area
The study area is in Yunnan Province, located between 97◦ 310 ~106◦ 110 east longitude
and 21◦ 80 ~29◦ 150 north latitude (Figure 1). The province is adjacent to Guangxi Zhuang
Autonomous Region and Guizhou Province in the east, separated from Sichuan Province
by the Jinsha River in the north, close to Xizang Autonomous Region in the northwest,
adjacent to Myanmar in the west, and bordering Laos and Vietnam in the south and
southeast, respectively. Yunnan Province, along with Guangxi Zhuang Autonomous
Region and Hainan Island, form the Indo–Burma region in the Southeast Asian biodiversity
hotspot [45], which is of paramount significance for global biodiversity conservation.
Yunnan Province has a total area of 394,100 km2 , accounting for 4.1% of the country’s
total area. It is characterized by five major landforms: mountains, hills, basins, plateaus,
and small plains. The diverse regional terrain is the primary reason for the conservation
of numerous species in Yunnan Province. Additionally, the province has a rich climate
belt and a variety of climates. Yunnan Province has a cold, warm, and hot (including
subtropical) climate. Most of Yunnan belongs to a subtropical plateau monsoon climate,
and Northwest Yunnan belongs to a cold zone climate. Eastern and Central Yunnan have
a temperate climate. The south and southwest of Yunnan Province belong to the tropics,
showing the climate characteristics of dry and hot valleys [46]. Therefore, Yunnan creates a
habitat suitable for the growth of many species. As of 2016, a total of 2253 species of seed
plants belonging to 225 families and 2140 genera have been identified in Yunnan, making it
the province with the highest number of plant species in China [47].
According to the Guiding Opinions on Establishing a System of Protected Natural
Areas with National Parks as the main body [48], China’s protected natural areas, which
are primarily composed of national parks, can be categorized into three categories: national
parks, nature reserves (including national, provincial, municipal and county levels), and
natural parks. This study focuses on five types of natural parks: wetland parks, forest parks,
scenic areas, parks, and stone desert parks [49]. The distribution of these protected-area
types is shown in Figure 1.
 Areas with National Parks as the main body [48], China’s protected natural areas, which
are primarily composed of national parks, can be categorized into three categories: na-
tional parks, nature reserves (including national, provincial, municipal and county levels),
and natural parks. This study focuses on five types of natural parks: wetland parks, forest
Land 2024, 13, 240 parks, scenic areas, parks, and stone desert parks [49]. The distribution of these protected-
4 of 19
area types is shown in Figure 1.

Figure 1. Spatial distribution of PAs in the study area, with terrain and landforms.
Figure 1. Spatial distribution of PAs in the study area, with terrain and landforms.
2.2. Species Occurrence Data
2.2.This
Species Occurrence
study focused Data
on the conservation of vulnerable, near threatened, endangered,
This
critically study focused
endangered, on and
extinct, the conservation of vulnerable,
extinct wild species neararea.
in the study threatened, endangered,
The classification of
critically endangered, extinct, and extinct wild species in the study area.
endangered species is based on Table S1. The selection criteria for these species were basedThe classification
onofthe
endangered
IUCN Redspecies is based on Species
List of Threatened Table S1. TheSpecies
[50]. selection criteria for
distribution these
data werespecies were
obtained
based
from theonIUCNthe IUCN Red
Red List ofList of Threatened
Threatened SpeciesSpecies
and the[50]. Species
Global distribution
Biodiversity data were
Information
obtained
Facility [51].from
Afterthe IUCN Red List
downloading the of
dataThreatened
package fromSpecies
the and the Global
website, Biodiversity
a massive amount ofIn-
formation
data Facility
was screened to[51]. After
select plantdownloading
records withthecleardata package
spatial from theinformation,
distribution website, a massive
while
removing
amount erroneous
of data was and duplicate
screened to records. Therecords
select plant taxonomic withnames
clear were cross-checked
spatial distributionwith
infor-
the latest valid
mation, whilespecies names
removing in the Catalogue
erroneous of Liferecords.
and duplicate [52]. Plant-acceptable
The taxonomicnames nameshave
were
been compared with the latest
cross-checked Worldvalid
Floraspecies
onlinenames
database [53].
in the A total of
Catalogue of 604
Lifespecies from
[52]. Plant-ac-
164 familiesnames
ceptable and species
have beengroups, along with
compared with the
3926 distribution
World points
Flora online for representative
database [53]. A total
RESs,
of 604were collected
species fromin164this research
families and(Table S2).groups, along with 3926 distribution points
species
for representative RESs, were collected in this research (Table S2).
2.3. Environment Variables
2.3.The specific sources
Environment Variablesof the environmental elements used in this study are shown
in Table S3.specific
The The digital elevation
sources model (DEM) elements
of the environmental data wasused
sourced from
in this the are
study Geospatial
shown in
Data Cloud [54], and 19 biological climate variables were sourced from the
Table S3. The digital elevation model (DEM) data was sourced from the Geospatial Data WorldClim
2.1Cloud
version [55],
[54], andrepresenting
19 biologicalaverage values
climate from 1970
variables were to 2000, with
sourced froma spatial resolution2.1
the WorldClim
of 30 seconds/km2 . Climate-prediction data were obtained from WorldClim, based on
BCC-CSM1-1 [56]. (Table S3). Data for four emission pathways (RCP2.6, RCP4.5, RCP6.0,
and RCP8.5) and two time points (2050 and 2070) were downloaded. By 2100, the rising
radiative forcing pathway led to ~3 W/m2 , 4.5 W/m2 , 6.0 W/m2 , and 8.5 W/m2 [57,58]. To
avoid overfitting, a correlation analysis of 19 bioclimatic variables was carried out (Table S4).
Finally, 10 variables were selected, and a total of 14 variables, including elevation, slope,
slope direction, and normalized difference vegetation index (NDVI) [59], were input into
the model. The downloaded data were preprocessed with the relevant tools of ArcGIS10.2.
The 14 biological climate variable layers are extracted from Yunnan Province using the mask
Land 2024, 13, 240 5 of 19

extraction tool of ArcGIS10.2, and, then, random points are created for these extracted layers.
Interpolation (Kriging interpolation) is performed using random points, and, finally, the
interpolated study-area range is converted to an ASCII file for saving.

2.4. Distribution Modeling

Using ArcGIS 10.2 as the platform, environmental variable raster files were filtered
for specific variables. In addition, the coordinate system, boundary, and resolution of all
layers were unified. To avoid low AUC values caused by the large research area or small
distribution points, species data were imported into the MaxEnt model by families.
In order to ensure the model’s normal operation and performance, species with
loci ≥ 6 were imported as a whole. Species with loci less than 6 loci were integrated based
on their distribution characteristics. The integrated species were categorized into three
groups: animals, plants, and fungi.
In this study, 75% of the species distribution information was used to build the model,
while 25% was used for model verification. The background points were set to 10,000 and
heterogeneous threshold parameters were used to transform the model output results. The
other parameters were set to the model default values [42].
This research evaluates the prediction accuracy of the MaxEnt model using the area
under the receiver operating characteristic (ROC) curve (area under curve, AUC). The
evaluation of AUC values generally adheres to the following standards: 0.9 < AUC < 1.0,
the model prediction result is extremely accurate; 0.7 < AUC < 0.8, the prediction accuracy is
high; 0.6 < AUC < 0.7, the prediction accuracy is low; and 0.5 < AUC, the model prediction
fails [60]. For model verification and testing, 25% of the species distribution points were
selected; when AUC ≥ 0.7, the simulated species potential habitat of the model was used
for further analysis for biodiversity conservation planning. An AUC greater than 0.75 was
then considered acceptable. To create a biodiversity hotspot distribution map, the species
distribution probability map was transformed into a floating-type raster using ArcGIS tools.
The distribution probability map was then converted into a hotspot distribution map based
on the minimum training threshold.

2.5. Model Accuracy and Validation

The model used to simulate the distribution of RESs in Yunnan Province is acceptable
for both current and future scenarios (RCP2.6, RCP4.5, RCP6.0, RCP8.5, 2050, and 2070).
The average AUC values for RESs in all scenarios were above 0.75 (Table S5). This indicates
the success of the model in evaluating species distribution within the study area. The
specific AUC values are listed in the Supplementary Materials) in this article.

2.6. Gap Analysis

The protection gap in this study refers to the insufficient space available for RESs
in the existing protected-area system. The identified priority protection hotspots and
the established protected areas were superimposed on the space analysis. Those space
areas without overlap are protection gap areas. The results of this gap analysis method in
previous studies have substantial implications for species conservation [61,62].
In order to evaluate the richness of the potential habitat of creatures identified by the
model, creature hotspots were identified. The results of the model operation are binarized
according to the 10% training threshold generated by the model. The part larger than the
training threshold is selected as the potential distribution area of the family. The layers of
each family/species group after binarization are overlaid by a raster calculator, and the
layers are divided into four levels using the natural breakpoint method [63], in order of
richness from high to low. And then, count the area and proportion of each land use and
land cover in the hotspot areas. This study obtained land-use and land-cover data. The
data set is provided by the Data Center for Resources and Environmental Sciences, Chinese
Academy of Sciences (RESDC) [64]. Then, based on the area and proportion of hotspots,
Land 2024, 13, 240 6 of 19

our research determined the natural community types that RESs rely on. It also identified
the human development that poses the greatest threat to potential habitats.
In order to evaluate the protection effect of existing reserves on representative creature
communities in Yunnan Province, the study examined the simulated hotspots with the
existing reserves. Then, extract the hotspots inside and outside the protected area, calculate
their area and proportion, and evaluate the protection efficiency of the current protected
area. Finally, this study selected regions with a predicted abundance greater than 25% of
RESs from the simulation results as the priority protection area.
In addition, this research also uses data on RESs downloaded from IUCN to build the
fishing net tool to evaluate the number of RESs in each grid memory. Visualize this quantity
of data on a map to get the result. Using this result, we compare the existing protection of
the RESs in IUCN and the MaxEnt model to predict the distribution of RESs.

2.7. Predicting Potential Habitat under Future Climate Change

In order to evaluate the impact of future climate change on creature communities, this
study used the MaxEnt model to predict the potential habitats of RESs in Yunnan Province
in 2050 and 2070 under four emission modes: RCP2.6, RCP4.5, RCP6.0, and RCP8.5. In
addition to comparing the hotspot distributions over time, as well as the changing area and
spatial distribution, the results were plotted on a binary graph. In order to identify future
protection gaps under predicted climate change, hotspots simulated for 2050 and 2070 were
compared with current hotspots for RESs. Finally, by extracting the area of each richness
and the overlap between them, the transfer of richness levels over time scales is obtained.
Using these areas, we created a transfer matrix to quantify the changes in RES hotspots
from current to future points. In this study, the vector area was calculated by the ArcGIS
field calculator, and the raster area was calculated by grid number × grid size (900 m2 ).
For the purpose of analyzing the changing trend of the high-richness region in the
future, 360 radiation lines are made in the average center of the study area, and the overlap
length of each radiation line with the high-richness region in this direction is taken. The
cross length in each direction was used to qualitatively indicate how much or how less
of a high-richness area exists in that direction. This approach can show the direction and
amount of high-richness area transfer under each emission scenario. The impact factor
contribution ranking is generated when the model is run. The number of times that each
factor appeared in the top five contribution rates was counted to find out the driving factors
with higher contribution rates, and the reasons for the changes in each region under climate
change were analyzed through the driving factors.

3. Results
3.1. Current Habitats of RESs
The rich distribution area for endangered species comprises 80.54% of Yunnan Province
(Figure 2). Species distribution in high richness areas (HRA, richness < 30.84 were con-
centrated in the northwest and east of Yunnan Province and distributed in small areas in
Xishuangbanna. This was with a total area of 25,080.80 km2 . As regards the number of RESs
in the grid, Yunnan contained an ‘unsuitable’ area (UA, richness < 7.90) of 1,171,531.15 km2 ,
‘low richness’ area (LRA, richness < 17.84) of 121,575.17 km2 , and ‘medium richness’ area
(MRA, richness < 30.84) of 66,266.61 km2 . Almost half of the UA showed a concentrated
distribution in the central and the north areas of Yunnan (Figure 2).
 RESs in the grid, Yunnan contained an ‘unsuitable’ area (UA, richness < 7.90) of
1,171,531.15 km2, ‘low richness’ area (LRA, richness < 17.84) of 121,575.17 km2, and ‘me-
dium richness’ area (MRA, richness < 30.84) of 66,266.61 km2. Almost half of the UA
showed a concentrated distribution in the central and the north areas of Yunnan (Figure
Land 2024, 13, 240 7 of 19
2).

Figure 2. Potential distribution map after stacking analysis of all species.

Figure 2. Potential distribution map after stacking analysis of all species.
3.2. Future Habitats of RESs
3.2. Future Habitats of RESs
In the future scenario, the concentrated distribution area of RESs is roughly unchanged
In the future scenario, the concentrated distribution area of RESs is roughly un-
(Figure 3). The spatial
changed change
(Figure flow
3). The in the
spatial figure
change starts
flow in theatfigure
the current distribution
starts at the center.
current distribution
The changes in areasThe
center. with high in
changes abundance
areas with in different
high abundancescenarios andscenarios
in different years areand
shown
years in
are
Figure S1. shown in Figure S1.
The distribution
The distribution area of theareahighof the high richness
species species richness
zone zone was calculated
was calculated underfour
under four
different
different scenarios. scenarios.
The resultsThe resultsthat
showed showed that in
in 2050 2050
the the largest
largest hotspot
hotspot areaarea
waswas observed
observed
under the RCP4.5underemission
the RCP4.5scenario.
emission scenario. In contrast,
In contrast, the smallest
the smallest hotspothotspot
areaarea
waswas observed
observed
under the RCP6.0 emission scenario. In the year 2070, the same remains accurate. Among
under the RCP6.0 emission scenario. In the year 2070, the same remains accurate. Among
the four scenarios, only the RCP8.5 scenario decreases hotspot areas between 2050 and
the four scenarios, only
2070. In the RCP8.5
contrast, scenario
the other decreases
scenarios expand the hotspot
area ofareas between
hotspots 2050 and
in the period from2070.
2050
In contrast, theto other
2070 toscenarios expand
a certain extent the area
(Figure 3). of hotspots in the period from 2050 to 2070
to a certain extent (Figure 3).
To represent the distribution of hotspot abundance in different directions, the intersec-
tion length of hotspots in Yunnan Province was calculated under four scenarios (Figure 4).
The results revealed that, under the RCP2.6 emission scenario, the hotspot distribution
in 2070 increased significantly towards the west and south. Under the RCP4.5 emission
scenario, the hotspots towards the west increased in both 2050 and 2070, while the hotspots
towards the south and east increased substantially in 2050. In the RCP6.0 emission scenario,
there was not much difference in the main distribution directions across the three time
periods. However, distribution areas in the east and west significantly increased in future
scenarios. Within the RCP8.5 emission scenario, the geographical distribution of hotspots
will shift from the southeastern region to the eastern region in the future. The extent of
hotspots in the western region will experience a significant increase by 2050.
The model calculates the contribution of bioclimatic variables to species distribution.
The contribution rate of each type of factor is calculated comprehensively. The contribution
rate of the precipitation factor was 31.96%, the contribution rate of the normalized vegeta-
tion index was 16.52%, and the contribution rate of the topographic factor was 14.57%. The
most important contributing factors are annual precipitation, normalized vegetation index,
and slope.
Land 2024, 13, 240
Land 2024, 13, x FOR PEER REVIEW 8 of 19 8 of 19

Land 2024, 13, x FOR PEER REVIEW 9 of 19

FigureFigure 3. The
3. The contribution
predicted
predicted rate of the
distribution
distribution precipitation
ofofRESs
RESsin the factor
future
in the was 31.96%,
environment.
future the contribution
(A)–(H)
environment. represents rate of the
8 differ-
(A–H) represents nor-
8 different
ent emissionmalized
models, vegetation
respectively.index
(A): RCP2.6, 2050; and
was 16.52%, (B): the
RCP4.5, 2050; (C):rate
contribution RCP6.0,
of the2050; (D):
topographic factor
emission models,
RCP8.5, respectively.
2050;was
(E): RCP2.6, The (A):
2070;most RCP2.6,
(F): RCP4.5, 2050;
2070; (G): (B): RCP4.5,
RCP6.0, 2050;
2070; (H): (C): RCP6.0, 2050; (D): RCP8.5,
14.57%. important contributing factors areRCP8.5,
annual2070).
precipitation, normalized
2050; (E): RCP2.6, 2070; (F): RCP4.5, 2070;
vegetation index, and slope. (G): RCP6.0, 2070; (H): RCP8.5, 2070).
To represent the distribution of hotspot abundance in different directions, the inter-
section length of hotspots in Yunnan Province was calculated under four scenarios (Figure
4). The results revealed that, under the RCP2.6 emission scenario, the hotspot distribution
in 2070 increased significantly towards the west and south. Under the RCP4.5 emission
scenario, the hotspots towards the west increased in both 2050 and 2070, while the
hotspots towards the south and east increased substantially in 2050. In the RCP6.0 emis-
sion scenario, there was not much difference in the main distribution directions across the
three time periods. However, distribution areas in the east and west significantly in-
creased in future scenarios. Within the RCP8.5 emission scenario, the geographical distri-
bution of hotspots will shift from the southeastern region to the eastern region in the fu-
ture. The extent of hotspots in the western region will experience a significant increase by
2050.
The model calculates the contribution of bioclimatic variables to species distribution.
The contribution rate of each type of factor is calculated comprehensively. The

Figure 4. Comparison of the distribution of HRAs from different angles in four scenarios (RCP2.6,
RCP4.5, RCP6.0, and RCP8.5).
Figure 4. Comparison of the distribution of HRAs from different angles in four scenarios (RCP2.6,
RCP4.5, RCP6.0, and RCP8.5).

3.3. Regional Transfer Matrix of Different Richness Levels

Future climate change will have a significant impact on the potential habitats of eco-
logically significant species. Under the emission mode of RCP2.6, by 2050, only 45.81% of
potential habitats in LRAs will be retained, approximately 23.20% will become UAs,
Land 2024, 13, 240 9 of 19

3.3. Regional Transfer Matrix of Different Richness Levels

Future climate change will have a significant impact on the potential habitats of
ecologically significant species. Under the emission mode of RCP2.6, by 2050, only 45.81%
of potential habitats in LRAs will be retained, approximately 23.20% will become UAs,
40.67% of potential habitats in MRAs will be retained, 11.67% will become UAs, 53.22% of
potential habitats in HRAs will be conserved, and 1.41% of high adaptation is transformed
into UA (Table 1). The abundance area transfer matrix under the RCP4.5, RCP6.0, and
RCP8.5 scenarios is in Table S6.

Table 1. The abundance area transfer matrix under RCP2.6 scenarios.

Rcp2.6 2050
UA LRA MRA HRA Total (km2 )
UA 59,808.11 94,941.85 16,464.49 316.7 171,531.15
LRA 28,206.38 55,687.57 34,874.07 2807.15 121,575.17
2022
MRA 7730.65 23,720.17 26,950.82 7864.97 66,266.61
HRA 354.54 3856.64 7520.64 13,348.98 25,080.8
Total (km2 ) 96,099.68 178,206.23 85,810.03 24337.8 384,453.74
Rcp2.6 2070
UA LRA MRA HRA Total (km2 )
UA 26,126.3 67,924.49 2043.84 5.00 96,099.63
Rcp2.6 LRA 336.95 63,359.52 111,492.86 3016.89 178,206.23
2050 MRA 85,810.03 501.88 44,774.67 40,533.48 171,620.06
HRA 24,337.8 24,337.8 545 23,760.05 72,980.65
Total (km2 ) 136,611.07 156,123.69 158,856.38 67,315.43 518,906.57

Under future climate change, these areas will undergo significant changes that will
affect the habitats and ecology of the protected areas. Under the emission mode of RCP2.6,
from 2050 to 2070, only 35.55% of potential habitats in LRA will be retained, approximately
0.19% will become UAs, 26.09% of potential habitats in MRA will be retained, 50.00% will
become UAs, 32.56% of potential habitats in HRAs will be protected, and 33.35% of HRA is
transformed into UAs.
The proportion of the shift in richness for every emission mode is shown in Table 1.
Period 1 refers to 2022~2050, and period 2 refers to 2050~2070. It can be found that what
happens in the near time scale is mostly the transition from the LRA to the UA, while
what happens in the distant time scale is mostly the transition from the HRA to the UA
(Table 2 and Table S6).

Table 2. On time scale, the proportion of each richness area retained and turned into UAs.

The
The The Proportion
Emission Retention Retention Retention Proportion
Period Proportion of of the MRA
Mode Ratio of LRA Ratio of MRA Ratio of HRA of the HRA
LRA to UA to UA
to the UA
1 45.81% 23.20% 40.67% 11.67% 53.22% 1.41%
RCP2.6
2 35.55% 0.19% 26.09% 50.00% 32.56% 33.35%
1 30.47% 18.88% 37.20% 9.27% 74.14% 1.11%
RCP4.5
2 68.16% 7.14% 64.40% 0.97% 39.68% 50.01%
1 43.06% 26.71% 39.90% 14.24% 56.32% 2.70%
RCP6.0
2 59.53% 0.92% 52.03% 0.04% 41.41% 50.00%
1 44.57% 25.96% 36.31% 12.90% 57.55% 2.85%
RCP8.5
2 33.86% 9.44% 51.75% 1.92% 26.03% 0.00%

3.4. Gap Analysis and Priority Conservation Identification

Overlapping the existing PAs with the current HRA, it was found that the overlapping
area of the two is 1369.36 km2 , accounting for 5.46% of the total HRA of 23,017.98 km2 , and
Land 2024, 13, 240 10 of 19

thus for 80.66% (Table 3). If the measured area is replaced by the current hotspot area, then
the overlap area of PAs and hotspot area is 18,075.44 km2 , accounting for 19.34% of the
total hotspot area of 93,471.13 km2 , representing 27.57% of the PAs area, and accounting for
4.59% of the area of Yunnan Province (Figure 5).

Table 3. The protection ratio of existing PAs to HRA and the protection efficiency of PAs under
different scenarios.

Emission Overlapping Protection Unprotected Proportion of

HRA (km2 ) UA (km2 )
Mode Area (km2 ) Ratio Proportions Effective PAs
Land 2024, 13, x FOR PEER REVIEW
Current 25,080.80 1369.36 23,017.98 5.46% 94.54% 6.23%11 of 19
RCP2.6, 2050 24,337.80 919.63 22,093.26 3.78% 96.22% 4.18%
RCP2.6, 2070 67,315.43 3828.49 65,058.48 5.69% 94.31% 17.42%
RCP4.5, 2050 61,505.38 2826.70 59,111.00 4.60% 95.40% 12.86%
RCP6.0,
RCP4.5, 2070
2070 49,940.75
69,112.58 1281.91
2891.65 47,524.70
66,756.81 2.57%
4.18% 97.43%
95.82% 5.83%
13.15%
RCP8.5,
RCP6.0, 2050
2050 32,591.40
29,774.14 1092.00
914.91 30,149.49
27,245.38 3.35%
3.07% 96.65%
96.93% 4.97%
4.16%
RCP6.0, 2070
RCP8.5, 2070 49,940.75
8697.00 1281.91
145.30 47,524.70
5717.27 2.57%
1.67% 97.43%
98.33% 5.83%
0.66%
RCP8.5, 2050 32,591.40 1092.00 30,149.49 3.35% 96.65% 4.97%
RCP8.5, 2070 8697.00 145.30 5717.27 1.67% 98.33% 0.66%

Assessingthe
Figure5.5.Assessing
Figure thedistribution
distributionpattern
patternof
ofthe
theRESs
RESshotspots
hotspotsand
andPAs.
PAs.

InTable
In Table3,3,the
theprotection
protection ratio
ratio of of
thethe existing
existing PAsPAs to the
to the selected
selected species’
species’ HRAHRA in
in the
future scenario is listed. The protection efficiency of the PAs under this scenario is repre-is
the future scenario is listed. The protection efficiency of the PAs under this scenario
represented
sented by theby the proportion
proportion of theof the existing
existing PAs containing
PAs containing hotspots.
hotspots.
The IUCN data prediction results are divided into
The IUCN data prediction results are divided into hotspots and hotspots and nonhotspot
nonhotspot areas
areas
according to the high richness threshold 31 of the MaxEnt model. There is
according to the high richness threshold 31 of the MaxEnt model. There is a big difference a big difference
betweenthe
between thehotspots
hotspotsof ofthe
theMaxEnt
MaxEntmodel
model(Figure
(FigureS2),
S2),the
thehotspots
hotspotspredicted
predictedby byIUCN
IUCN
dataare
data aredistributed
distributed inin patches,
patches, andandthethe hotspots
hotspots are concentrated
are concentrated in theinsouthwest
the southwest
of Yun-of
nan Province (Figure S3). The MaxEnt prediction results and IUCN prediction results
were superimposed on the Pas, respectively, and it was found that their overlapping areas
were also very different (Figure S4). If the MaxEnt prediction results are superposed with
the hotspots predicted by IUCN data, the area of the protection hotspots of the existing
Land 2024, 13, 240 11 of 19

Yunnan Province (Figure S3). The MaxEnt prediction results and IUCN prediction results
were superimposed on the Pas, respectively, and it was found that their overlapping areas
were also very different (Figure S4). If the MaxEnt prediction results are superposed with
the hotspots predicted by IUCN data, the area of the protection hotspots of the existing PAs
is 40,377.51 square kilometers, and the protection ratio is 61.50% (Figure S5, Table S7).

4. Discussion
4.1. The Existing Protection Efficiency Needs to Be Improved
The potential hotspot areas of species in this study are consistent with the hotspot
results analyzed by other scholars using other methods. The differences are the results
of this study found more hotspots in the Kunming–Qujing region than those predicted
by Zhang et al. [39]. The formation of this hotspot is due to the strong influence of NDVI
in the driving factors. Yang et al. (2016) identified more priority protected areas by
combining animals and plants compared to the priority protected areas in Chuxiong [40].
This may be because Yang’s team determines conservation priorities by county. This means
that a small number of high priority areas within a district result in a county becoming
a conservation priority. Our research results also have hotspots in Chuxiong, which is
partly due to the differences in the methods adopted in the data analysis and visualization
process. Yang et al. used the invest model, NPP (Net Primary Production) index, and
topographic index to identify key areas for biodiversity conservation in 2021 [41]. The
results of Yang et al.’s study shows that the key protected area in northwest Yunnan is
wider. This may be due to the greater weight given to the topographic indicators, which
affected the results. Therefore, although different research methods are used in this paper,
the results are not much different from those of previous studies in the same research area.
The reason for the current low conservation efficiency may be because China’s early
PAs were specifically designed to conserve critically endangered species from extinction,
so they lacked top-level design and systematic planning [65]. In addition, environmental
disturbance and human activities threatening wildlife survival were also factors [66]. PAs
in Yunnan Province play an important role in protecting forests and RESs. However, the
specific needs of RESs in the region have not been fully considered. As can be seen from
the distribution map of PAs in Yunnan Province, the distribution of all types of PAs in
Yunnan Province is relatively even (Figure 1). However, the distribution of the most tightly
managed and effectively protected nature reserves is not consistent with the distribution of
RESs and protection hotspots. Our analysis shows that there are currently 156 various-level
PAs in Yunnan Province, covering 28,118 km2 . The overlap area between provincial level
natural reserves and HRAs is 1369.36 km2 . However, the protection rate of provincial level
PAs for HRAs is only 5.46%, and the remaining 94.54% of hotspots are located outside of
PAs without any protective measures.
The model shows us the potential range of RESs in future scenarios. The comparison
shows that existing PAs are slightly better protected when low carbon emissions are
maintained over long time scales. But when carbon emissions rise, current PAs will
struggle to protect RESs. The prediction results of IUCN data and the MaxEnt model differ
greatly in spatial distribution. The reason for this phenomenon is that the IUCN database
does not match the actual situation in China [67–69]. After superimposing the predicted
results of IUCN and MaxEnt models, the area is 136,382.34 km2 (Figure S4), accounting for
34.61% of the total area of Yunnan Province, which is in line with the protection target of
GBF. Therefore, combining the prediction results of IUCN data can protect more RESs and
expand the coverage of the protection system more accurately.
By analyzing the coverage of species hotspots in each protected area, several PAs with
high protection efficiency were found, such as Baima Snow Mountain Nature Reserve,
Gaoligongshan Nature Reserve, Xishuangbanna Nature Reserve, etc. The majority of these
areas are situated in Western Yunnan, consistent with previous research findings [70]. The
conservation efficiency of the national nature reserve is higher compared to other PAs,
which aligns with the findings of Wang et al. [71]. Located outside the three concentrated
Land 2024, 13, 240 12 of 19

distribution areas, the protection efficiency of the PAs in central Yunnan Province is low.
Overall, national protected areas have higher conservation efficiency than other levels,
which aligns with previous research findings [72].

4.2. Climate Change Promotes Habitat Migration

Most RESs are more sensitive to environmental changes than common species be-
cause of their small populations [73]. From the potential habitat transfer matrix, it can be
seen that under the carbon emission level roughly consistent with the current emission
pattern (RCP6.0), the area of large adaptation hotspots will continue to increase from the
present to 2050 and 2070. The areas with a large abundance of RESs in Yunnan Province
are concentrated in three regions, namely the western mountainous area of Yunnan, the
Xishuangbanna–Wenshan high temperature area of Southern Yunnan, and the abundant
vegetation area of Central and Eastern Yunnan. The distribution area of Western Yunnan
is in agreement with the findings of Qi et al.’s study on Cinnamomum mairei, Yang et al.’s
study on Alsophila spinulosa, and Liu et al.’s study on Bhutanitis thaidina [74–76]. The high
temperature distribution area aligns with the research conducted by He et al.’s study on
Asian elephants and Daniele et al.’s study on the fern, who studied RESs in Xishuang-
banna’s hotspots [77,78]. Wei et al.’s study about Excentrodendron tonkinense conducted
a focused investigation on RESs in Wenshan, located in the southeastern part of Yunnan
Province [79]. The high precipitation areas in Yunnan Province are distributed in Western
Yunnan and at the low latitudes of the tropics [80]. The most dense vegetation cover areas
in Yunnan are located near Gaoligong Mountain, Xishuangbanna, and Qujing in Western
Yunnan. At the same time, the western mountainous region of Yunnan exhibits significant
and abrupt variations in its topography. The distribution pattern of HRAs was most likely
related to precipitation, vegetation cover, and topographic changes.
As shown in Figure 3, the increase of hotspots in the due west and due east directions is
related to the expansion of hotspots in the Dehong, Baoshan, Kunming, and Qujing regions.
This demonstrates that, in accordance with Zhang et al.’s findings, RESs tend to migrate
towards mountainous regions. Furthermore, our analysis of topographic parameters
supports this conclusion [81]. Simultaneously, the extents of the hotspots in Xishuangbanna
have shrunk. But in the future, even with the relocation of locations, the results of the study
still show that the high abundance of species will remain concentrated in three regions.
This fits Penman’s point according to the statistics of bioclimatic factors that affect species
distribution [82]. The three factors that have great influence are the precipitation factor, the
topographic factor, and the normalized vegetation index. This result is also consistent with
many previous studies [44,83,84]. In particular, in Singh’s study of India, which is adjacent
to Yunnan, the most important factor affecting vegetation distribution is precipitation,
which is consistent with our results [85]. This suggests that, as carbon emissions increase
and the climate warms, species move to more suitable areas rather than becoming extinct.
The results of an evaluation of the land-use composition in hotspot regions are dis-
played in Table S8. The results indicate that in high richness hotspot areas, arable land and
human activity on the land account for 29.01% (27,119.83 km2 ) of potential hotspot areas.
The primary threats to the potential habitat of RESs in Yunnan Province are the conversion
of arable land and the encroachment of private forests. Yunnan has developed agriculture.
Cash crops such as tea, rubber, and coffee are widely planted in hotspot regions. The
expansion of these artificial forests and agricultural land has led to the fragmentation of
RES habitats, leading to the degradation or complete loss of rich habitats for RESs [86–88].
Changes in the transfer matrix suggest that hotspots for RESs are not fixed but change
with time and climate. In addition, the change in the richness degree of a region in a short
period will not be too large. However, the change in the richness degree for RESs on a
longer time scale will be large [89]. Except for the carbon emission model of RCP8.5, there
is no transition from an HRA to an LRA in the period from 2050 to 2070 because the area
of HRA is greatly reduced towards the two centers of east and west during this period,
Land 2024, 13, 240 13 of 19

and the centripetal shrinkage of MRA and LRA is also changed. Therefore, most UAs are
transformed into MRAs and LRAs.
In particular, because plant populations cannot migrate, the spread is extremely
limited [90]. This requires the creators and managers of PAs to pay attention to the protec-
tion of the potential habitats of RESs in the future during the planning phase.
We extracted the distribution area of HRA in each prefecture-level city in Yunnan
and listed the top three cities with hotspot areas under each carbon emission scenario.
Therefore, if the protection priority area is divided according to the administrative region,
Baoshan, Honghe, and Dehong should be selected (Table S9). Both Kunming and Dali
have large concentrations of hotspots, primarily due to their higher vegetation coverage.
Moreover, Dianchi Lake and Erhai Lake serve as significant ecosystems for several avian
species and aquatic organisms. Therefore, it is imperative to allocate additional focus to
these two significant lakes.

4.3. Suggestions for Protection

The GBF, adopted at the fifteenth session of the Conference of the Parties to the United
Nations Convention on Biological Diversity (COP 15), stipulates that at least 30% of the
world’s land and marine areas should be protected by 2030. Therefore, because of the poor
effect of the current conservation system on the protection of RESs, this study puts forward
the following suggestions.
(1) Fill in the existing protection gap. According to the conservation targets set by
COP15, the current system of PAs is far from adequate for the protection of the RESs
selected. At the same time, it is insufficient to protect key areas of ecosystem services [91].
This can be achieved by canceling the PAs of unprotected RESs where ecosystem services
cannot be secured [68]. To solve this, managers need to flexibly set new PAs in the hotspot
area of RESs distribution [92]. At the same time, the adjustment of PAs should also take
into account the actual existence of species data. The governing body should strengthen
the surveillance and investigation of RESs in their natural habitats. Relevant departments
should pay attention to the existence and disappearance of rare and endangered wild
species in reality, especially the migration patterns of rare and endangered animals. They
should promptly modify the location of PAs [93].
(2) Protection and restoration of vegetation and control of the expansion of the scope
of human activities. From our data, we can see that woodlands and grasslands occupy a
very high proportion, regardless of the mode and year of emission. At the same time, the
NDVI driving factor of the distribution of RESs also occupies a high position. We believe
that forests are the most important habitat patches for RESs in Yunnan Province, so it is
necessary to strengthen forest management to protect more RESs.
According to the result analysis of the land-use data, human activities also have a
significant impact on hotspots. There are potential conflicts in the relationship between
man and land in these areas. These conflicts will reduce the protection efficiency of
existing PAs [94]. In order to achieve effective conservation, these areas should be allocated
more management resources. Therefore, the government needs to combine development
with protection. Therefore, this study recommends governments develop alternative
conservation measures. Proactive strategies should be developed to prevent the outbreak
of human–land conflicts, implement standardized management, and achieve a win–win
situation for conservation and development [95]. Among them are effective district-based
measures, participatory governance, and providing alternative livelihoods [96,97].
(3) Develop more effective conservation strategies for future climate change. The
research model clearly indicates that hotspots with a high concentration of RESs will be
greatly affected by future global climate change. And over time, the species listed can also
change [66].
Often adjusting the boundaries of the protected-area system in response to these
changes is difficult and impractical to manage. The government should adopt a flexible
approach to adapt to future changes in the protected list and climate change without
Land 2024, 13, 240 14 of 19

frequently adjusting the boundaries of PAs, such as creating a protection model that
combines tourism and management. Alternatively, the government might be able to
promote the establishment of national parks [92]. National parks, as a distinct category
of natural protected areas, effectively balance ecological preservation and recreational
activities without compromising the level of protection provided by protected areas [98].

4.4. Research Limitations and Future Prospects

After research by many scholars, it has been confirmed that the MaxEnt model simu-
lates species distribution [99,100]. Upon comparing our findings with the observed species
distribution, we discovered that certain high altitude regions lack any instances of species
branching. However, according to the model simulation results, these areas still hold
potential for species distribution. This phenomenon is where the MaxEnt model deviates
from reality. In further work, some cold regions that are indeed unsuitable for species can
be excluded from the simulation results, improving the accuracy of predicting potential
distribution areas.
One characteristic of the results of this study is that, in future scenario simulations, the
range value of richness is significantly higher than the current predicted species richness.
The emergence of this result is due to the characteristics of the bioclimate variable model
selected for future scenarios. The BCC-CSM1-1 selected for prediction in this study is a
climate-system model developed by the Beijing National Climate Center, which has played
a good role in many studies [101,102]. However, compared to other system models, this
climate-system model has multiple indicators with larger prediction results [103–105]. The
error of a certain bioclimatic variable may have a decisive impact on the prediction results
of this family or species group in the MaxEnt model. Due to the large number of families
(species groups), the selected variable types were not adjusted based on each species, which
is also the reason for this problem. In future research, our team will strive to improve
this issue.
In addition, our analysis and research only conducted model simulations based on
existing species presence data but did not consider the possibility of animal–plant interac-
tions or animal interactions. Future research should also focus on the distribution of other
species closely related to the research object. This prediction will provide a more scientific
analysis of the existence areas and temporal changes of RESs and can also provide more
accurate protection of biodiversity.

5. Conclusions
In situ conservation is a crucial part of biodiversity conservation, so it is particularly
important to establish a precise conservation system. The MaxEnt model predicts the
current and future distribution of RESs in Yunnan Province. The study revealed that the
areas with the highest concentration of endangered species were primarily located in the
northwest and east regions of Yunnan Province. In addition, they were located in the
Xishuangbanna region. The places of “low richness” in Yunnan Province are primarily
found in the central and northern regions. In the future climate model, the distribution
location of hotspots does not change much, but the distribution area expands in the west
direction. The predicted species distribution hotspots have a large area affected by humans
and are also widely distributed in forest areas. Compared with the species distribution
results simulated by IUCN data, the conservation efficiency of the current conservation
system for RESs is low. This may be because the protection area has been established for a
long time, and the protection objects and protection targets in the past are different from
those in the present.
In view of these phenomena, this study suggests the following proposals. Relevant
departments should combine the forecast results and pay attention to the existence and
disappearance of rare and endangered wild species in reality. It is necessary to modify
the protection strategy and address the current protection deficit. Previous studies have
highlighted the potential for human–land conflicts due to the significant impact of human
Land 2024, 13, 240 15 of 19

activities on protected areas. Therefore, governments should develop alternative conserva-

tion measures in these areas, such as other effective district-based measures, participatory
governance, and providing alternative livelihoods. Last but not least, governments need
to develop effective conservation strategies for future climate change, adapting to future
conservation list adjustments and climate changes.

Supplementary Materials: The following supporting information can be downloaded at:

https://www.mdpi.com/article/10.3390/land13020240/s1, Figure S1: Changes in high-richness
areas over time under different emission modes; Figure S2: Hotspot distribution predicted by IUCN
data; Figure S3: Overlap hotspot distribution predicted by MaxEnt model and IUCN data; Figure S4:
Distribution of protected hotspots in MaxEnt forecast and IUCN data forecast results; Figure S5: The
IUCN hotspot area and MaxEnt hotspot area merged region; Table S1: The classification and classifi-
cation criteria of rare and endangered species selected in this study; Table S2: The list of species used
in the article is classified by the family; Table S3: Environmental variables used for modeling Yunnan
province; Table S4: Correlation analysis of 19 biological environmental variables; Table S5: AUC
value in the current and various situational model running processes; Table S6: Richness area transfer
matrix under RCP4.5,RCP6.0 and RCP8.5 scenarios; Table S7: Data comparison of two prediction
models; Table S8: The current proportion of land use types in hotspots; Table S9: Ranking of hot spot
areas in prefecture-level cities.
Author Contributions: Conceptualization, Y.L., Y.B. and Z.H.; Methodology, Y.L., Y.B., Z.H. and
J.W.; Software, Y.L. and Z.H.; Validation, Y.L., Y.B. and Z.H.; Formal Analysis, Y.L.; Investigation,
Y.L. and Y.B.; Resources, Y.L., Y.B., J.W. and H.C.; Data Curation, Y.L.; Writing—Original Draft
Preparation, Y.L.; Writing—Review & Editing, Y.L., Y.B., Z.H. and M.A.; Visualization, Y.L., Y.B. and
Z.H.; Supervision, Y.B.; Project Administration, Y.B.; Funding Acquisition, Y.B. All authors have read
and agreed to the published version of the manuscript.
Funding: This research was funded by the 14th Five-Year Plan of the Xishuangbanna Tropical Botani-
cal Garden, Chinese Academy of Sciences (E3ZKFF7B), Yunnan Province Science and Technology
Department (202203AP140007), Key Research Program of Frontier Sciences of Chinese Academy of
Sciences (ZDBS-LY-7011), Trans-boundary cooperation on biodiversity research and conservation in
Gaoligong Mountains (No. E1ZK251).
Data Availability Statement: The data presented in this study are available in Table S3.
Acknowledgments: The authors would like to thank the anonymous reviewers for their comments
and suggestions.
Conflicts of Interest: The authors declare no conflicts of interest.

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