Nwab 032

National Science Review
REVIEW 8: nwab032, 2021

https://doi.org/10.1093/nsr/nwab032
Advance access publication 23 February 2021
ENVIRONMENT/ECOLOGY
1
State Key Laboratory
of Vegetation and
Environmental Special Topic: Ecological Civilization—Insights into Humans and Nature
Change, Institute of
Botany, Chinese
Academy of Sciences,
The global significance of biodiversity science in China:
Beijing 100093,
China; 2 Ministry of an overview
Education Key
Laboratory of Ecology
Xiangcheng Mi1,† , Gang Feng2,† , Yibo Hu 3 , Jian Zhang 4 , Lei Chen1 ,
Downloaded from https://academic.oup.com/nsr/article/8/7/nwab032/6147049 by guest on 22 May 2023

and Resource Use of
the Mongolian
Plateau and Inner Richard T. Corlett5,6 , Alice C. Hughes5,6 , Stuart Pimm7 , Bernhard Schmid8 ,
Mongolia Key
Laboratory of Suhua Shi9 , Jens-Christian Svenning10 and Keping Ma 1,11,∗
Grassland Ecology,
School of Ecology and
Environment, Inner
Mongolia University, ABSTRACT
Hohhot 010021,
China; 3 Key Biodiversity science in China has seen rapid growth over recent decades, ranging from baseline biodiversity
Laboratory of Animal studies to understanding the processes behind evolution across dynamic regions such as the
Ecology and Qinghai-Tibetan Plateau. We review research, including species catalogues; biodiversity monitoring; the
Conservation Biology,
Institute of Zoology, origins, distributions, maintenance and threats to biodiversity; biodiversity-related ecosystem function and
Chinese Academy of services; and species and ecosystems’ responses to global change. Next, we identify priority topics and offer
Sciences, Beijing suggestions and priorities for future biodiversity research in China. These priorities include (i) the ecology
100101, China;
4
Zhejiang Tiantong and biogeography of the Qinghai-Tibetan Plateau and surrounding mountains, and that of subtropical and
Forest Ecosystem tropical forests across China; (ii) marine and inland aquatic biodiversity; and (iii) effective conservation
National Observation and management to identify and maintain synergies between biodiversity and socio-economic development
and Research Station,
School of Ecological
to fulfil China’s vision for becoming an ecological civilization. In addition, we propose three future
and Environmental strategies: (i) translate advanced biodiversity science into practice for biodiversity conservation;
Sciences, East China (ii) strengthen capacity building and application of advanced technologies, including high-throughput
Normal University, sequencing, genomics and remote sensing; and (iii) strengthen and expand international collaborations.
Shanghai 200241,
China; 5 Center for Based on the recent rapid progress of biodiversity research, China is well positioned to become a global
Integrative leader in biodiversity research in the near future.
Conservation,
Xishuangbanna Keywords: biodiversity inventory, biodiversity maintenance, biodiversity monitoring, biodiversity origins,
Tropical Botanical
Garden, Chinese
biodiversity-ecosystem functioning
Academy of Sciences,
Menglun 666303,
China; INTRODUCTION from primary production, nutrient cycling, purifica-
(Continued on next The most striking feature of life on Earth is its incred- tion of water and air, and mitigation of greenhouse
page) ible diversity. On the one hand, this biodiversity’s gas emissions and climate change, to crop pollina-
mere existence begs many fundamental scientific tion, food and genetic resource provisioning, disease
∗
Corresponding questions: how many species are there on the planet? control, and educational, cultural and spiritual ben-
author. E-mail: How and why does biodiversity vary dramatically efits [3]. Alarmingly, human-caused global changes
[email protected] over the surface of the Earth? Within an ecosystem, in climate and land use are increasingly threaten-
†
Equally contributed
to this work.
how do different species evolve and coexist with each ing biodiversity from the poles to the tropics and
other and their abiotic environments? How does this on land and in the ocean [4]. How is biodiversity
coexistence change over time, and how does this in- responding and adapting to the dramatic changes
Received 24 May fluence species’ ability to evolve? These questions in the Anthropocene? How can scientists accurately
2020; Revised 3
January 2021;
remain challenging despite recent advances in re- evaluate and predict the risks of local diversity loss
Accepted 14 search [1,2]. On the other hand, biodiversity’s func- and global extinctions and so guide the effective
February 2021 tions and services are the basis for sustainable devel- protection of biodiversity? These are the most ur-
opment and human well-being. These services range gent challenges in biodiversity science [5,6]—the

C The Author(s) 2021. Published by Oxford University Press on behalf of China Science Publishing & Media Ltd. This is an Open Access article distributed under the terms of the Creative
Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original
work is properly cited.
Natl Sci Rev, 2021, Vol. 8, nwab032

Figure 1. (A) The number of papers on China’s biodiversity by China’s scholars in each year between 2000 and 2019 published
in science citation index (SCI) journals (searched on Web of Science using ‘biodiversity’ and ‘China’ as the topic and ‘China’
as address); (B) its ratio to the number of papers on biodiversity by global scholars (‘biodiversity’ as the topic); and (C) the
number of papers on China’s biodiversity by China’s scholars published in high-profile journals (Nature, Science, Proc Natl
Acad Sci USA and Nat Commun).
multidisciplinary study of biodiversity in order to 9000 years [15], and human-induced loss of natural
(Continued from conserve it while considering evolutionary and habitat has caused species decline and losses across
previous page) ecological processes, anthropogenic influences and several millennia. This process intensified in the 20th
6
Center of environmental changes affecting it. century. Rapid increases in human population and
Conservation Biology, China harbours over 35 000 higher plant species urbanization during the last 50 years have pushed a
Core Botanical
[7], over 2700 terrestrial vertebrate species [8] large number of species to the brink of extinction or
Gardens, Chinese
Academy of Sciences, and over 28 000 marine species [9], with a high to extinction in the wild, e.g. Yangtze River dolphins
Menglun 666303, proportion of endemic species [10]. Among the or baiji (Lipotes vexillifer), Rhododendron kanehirae,
China; 7 Nicholas 36 global biodiversity hotspots (https://www. and Saiga antelope (Saiga tatarica) [16]. Thus, the
School of the
Environment, Duke conservation.org/priorities/biodiversity-hotspots), combined effects of natural and anthropogenic
University, Durham, the ‘Mountains of Southwest China’ hotspot lies factors have been, and still are, shaping biodiversity
NC 27708, USA; almost entirely (>98%) within China, and the patterns in China.
8
Department of
Geography, Remote ‘Himalaya’ (33%), ‘Mountains of Central Asia’ Over the past two decades, research and pub-
Sensing Laboratories, (29%) and ‘Indo-Burma’ (15%) hotspots lie partly lications on China’s biodiversity have increased
University of Zurich, within China. In general, biodiversity in China is dramatically in quantity and quality. The number of
Zurich 8057,
Switzerland; 9 State
much higher than in any other country covering papers published in international journals has grown
Key Laboratory of similar latitudes. Three factors may explain this from a few dozen in 2000 to over 1700 in 2019
Biocontrol, high biodiversity. First, China is among the few (Fig. 1). The Nature Index 2020 Annual Tables
Guangdong Key countries with the whole spectrum of climate zones, highlighted the Chinese Academy of Sciences and
Laboratory of Plant
Resources, Key from tropical to cold temperate. Around 40% of several other institutions from China as among the
Laboratory of the land surface of China rises above 2000 m in most prolific producers of high-quality research
Biodiversity Dynamics elevation. The high geodiversity in China, in turn, in the natural sciences. Chinese institutions make
and Conservation of
Guangdong Higher creates a wide range of habitats for diverse species. up all top-ten fastest-rising institutions worldwide,
Education Institutes, Its mountainous areas serve as macroclimatic and including those in environmental and life sciences
School of Life microclimatic refugia for many endemics [11]. (http://www.nature.com/collections/chdeajdica).
Sciences, Sun
Yat-Sen University,
Second, the orogeny of the Himalaya, the uplift of Both of these are closely related to biodiversity sci-
Guangzhou 510275, the Qinghai-Tibetan Plateau, and the subsequent ence and conservation. The List of Highly Cited Re-
China; 10 Center for development of the East Asian monsoon had huge searchers 2019 (https://clarivate.com/) also shows
Biodiversity Dynamics
impacts on China’s landforms, topography and a good performance from China. Chinese mainland
in a Changing World
(BIOCHANGE) and climate [12]. The result was an elevated rate of increased its share of Highly Cited Researchers
Section for species diversification and speciation [13]. Third, significantly, from 483 or 7.9% in 2018 (covering
Ecoinformatics and during the ice ages of the Pleistocene, large parts the period 2006–16) to 636 (10.2%) in 2019 (for
Biodiversity,
Department of of China remained relatively warm and humid, the period 2008–18). In 2014, 113 Highly Cited
Biology, Aarhus serving as climatically stable refugia for many relict Researchers were from Chinese mainland, whereas
University, Aarhus C, taxa, such as ginkgo (Ginkgo biloba) and dawn by 2019, there were 347, nearly a 3-fold increase.
DK-8000 Denmark
and 11 University of redwood (Metasequoia glyptostroboides) [14]. Be- There are multiple reasons behind this accel-
Chinese Academy of sides natural diversity-promoting processes, human erating progress, including increasing financial
Sciences, Beijing activities have reshaped China’s biodiversity. Agri- investment in basic research at national and
100049, China
cultural activities in China date back at least 8000– provincial levels and significant improvements in
Page 2 of 25
70°E 80°E 90°E 100°E 110°E 120°E 130°E 140°E
⑥
① · Cataloguing and big biodiversity data
⑥
② · Biodiversity dynamics and monitoring
40°N
40°N
⑦
③
Inventory &
Monitoring
30°N
⑧
30°N
④
⑤
20°N
20°N
⑤
⑤ South China Sea Islands
1:96,000,000
80°E 90°E 100°E 110°E 120°E
Elevation (m)
-136 – 0 2001 – 3000
1 – 500
Processes & Threats &
3001 – 4000
501 – 1000 4001 – 5000
1001 – 1500

> 5000
1501 – 2000 Mechanisms Responses
· Origin & Evolution · Drivers of Threats

· Geographic distributions & Drivers · Responses to global change
· Maintenance mechanisms at local scales · Biodiversity & Biosafety
· Biodiversity & Ecosystem functioning
· Biodiversity & Ecosystem services
Figure 2. Research highlights of China’s biodiversity science in three main areas, including 10 key research topics. The map
on the left shows the elevation ranges in China. The lines with different colours in the map divide the country into eight
vegetation regions [19]: cold-temperate deciduous needle-leaved forests; temperate mixed-needle and broad-leaved
forests; warm-temperate deciduous broad-leaved forests; subtropical evergreen broad-leaved forests; tropical
monsoon rain forests and rain forests; temperate grassland; temperate desert; Qinghai-Tibetan Plateau alpine
vegetation. Digital elevational model (DEM) data at 10 arcmin spatial resolution were downloaded from WorldClim [20].
The four pictures present the iconic species Ginkgo biloba (copyright Yunpeng Zhao), Rhinopithecus roxellanae (copyright
Sheng Li), Ailuropoda melanoleuca (copyright Yibo Hu) and Chrysolophus pictus (copyright Sheng Li). The map of China is
from http://bzdt.ch.mnr.gov.cn/.
Chinese ecologists’ and biologists’ research capacity Considering the global significance of China’s
through international collaborations. In addition, biodiversity and the marked improvement in
the recent establishment of multiple excellent biodiversity research by Chinese research institutes
ecological research platforms (e.g. Chinese Forest over the past two decades, this paper provides an
Biodiversity Monitoring Network, the Biodiver- overview of the achievements. Our purpose is to
sity and Ecosystem Functioning Experiment in summarize recent advances in biodiversity science
China (https://bef-china.com/), and the Chi- by scientists from China and to identify potential
nese National Ecosystem Research Network barriers and future directions for further advancing
(http://cnern.cern.ac.cn/en/)) have enhanced biodiversity science, both in China and worldwide.
understanding of biodiversity across spatial and The rich experiences and lessons learned over recent
temporal scales. Building on these successes, there decades also provide useful examples for other coun-
has been an increase in conservation research capac- tries and regions hoping to follow a similar develop-
ity (e.g. the Chinese Union of Botanic Gardens, the ment path. In reviewing publications with Chinese
Germplasm Bank of Wild Species, and the National institutions as the first affiliation in high-profile jour-
Park-centred protected area system). Policies within nals such as Science and Nature (and other journals
various government programmes (e.g. the Grain in their families) or high-impact systematic efforts,
for Green Programme, Returning Grazing Land to we highlight the significant progress of biodiversity
Grassland Project, and Three-North Shelter Forest science in China in three critical research areas:
Programme) [17] and a key national governance (i) inventory and monitoring (what and how many
strategy for ecological civilization promotes national species are present in China, and where are they
conservation efforts. They enable restoration and located? How and why do they change over time?);
conservation to complement biodiversity and (ii) mechanisms and processes (origination, evo-
related research. Notably, the Chinese government lution and adaptation, environmental relations,
has invested US$378.5 billion over the past few and biotic interactions within China’s biota); and
decades in 16 massive sustainable development (iii) threats and responses (how is biodiversity
projects [17] and has thereby substantially slowed threatened, and how does it respond to global
biodiversity decline [18].
Page 3 of 25
Table 1. Number of species, endemic species and threatened species for well-studied revised into the Flora of China with 49 volumes
taxonomic groups in China. in English [24]. Plants of China: A Companion to
Number Number Flora of China followed to answer the question
Number of endemic of threatened of why there are so many species in China and
Taxonomic group of speciesa species species how the Chinese people have traditionally used
them [21]. Recently, both Fauna Sinica, with 162
Bryophytes 3021 524 186 volumes (Table S2), and Florarum Cryptogamarum
Ferns 2129 842 182 Sinicarum, with 96 volumes, have been published
Gymnosperms 237 88 148 (Table S3). The compilations of floras of most
Angiosperms 28 996 14 693 3363 provinces and dozens of provincial-level faunas have
Amphibians 408 272 176
been completed, providing detailed descriptions of
Reptiles 461 142 137
regional biodiversity. Moreover, the Fossil Flora of
Birds 1372 77 146

China (four volumes) summarizes the distributions,
Mammals 673 150 178
research history and characteristics of China’s 2248
a
The number of species and endemic species is from Gao et al. (2018) [10], and the number of threatened fossil plant species [25]. The Catalogue of Exotic
species is from Zang et al. (2016) [29]. and Cultivated Plants in China appeared in 2018 and
documents 13 635 exotic plant species and 13 941
change?). We divide these three areas into 10 key native cultivated plant species [26] (Fig. S1). Based
research topics (Fig. 2). on the Checklist of Marine Biota of China Seas [27]
and other information, The Living Species in China’s
Seas documented more than 28 000 marine species
BIODIVERSITY INVENTORY belonging to 59 phyla in China’s seas [9].
AND MONITORING The first version of Catalogue of Life: China (CoL-
China) was released in 2008, and it has been up-
Species catalogue and big
dated annually. The 2020 version includes 122 280
biodiversity data species and intraspecific taxa covering different tax-
Species catalogue in China onomic groups (http://www.sp2000.org.cn). The
China’s rich biodiversity poses a challenge for ade- CoL-China also provides a real-time update for new
quately collating, analysing and managing the enor- species to fill the existing gaps in the inventory, par-
mous amount of biodiversity information, effectively ticularly for terrestrial arthropods and fungi. For ef-
communicating the information to the public, and fective conservation and management of biodiver-
facilitating the development of biodiversity-related sity, Chinese researchers adopted the International
policies. Several generations of Chinese scientists Union for Conservation of Nature (IUCN) Red List
have made significant progress in cataloguing categories and criteria to evaluate species threat sta-
China’s species and aggregating big biodiversity tus. There were 4408 seed plant species and 3351
data as a fundamental building block for further vertebrate species in the first assessment in 2004
research to meet these challenges [21]. [28], and 34 450 higher plant species and 4357 verte-
Researchers at the Chinese Academy of brate species in the second assessments in 2013 and
Sciences organized 850 universities and insti- 2015, respectively (Table 1) [29]. The threat sta-
tutes to conduct more than 40 national-scale tus of 9302 out of >10 000 species of macrofungi
surveys of natural resources, including biodiversity. was also assessed in 2016 [30]. Trends in overall
They were national-scale efforts in the 1950s–60s threat status for groups of species are indicated by
and regional-scale surveys in the 1970s–80s. These changes of species numbers in threatened status of
surveys are responsible for generating much of successive Red List assessments (Fig. 3) [31]. Based
our knowledge of China’s species. For example, on the two successive species Red List assessments,
the first Qinghai-Tibet Plateau scientific survey indicators showed that the conservation statuses of
during 1973–76 found seven new plant genera, birds, reptiles and amphibians are declining, while
300 new plant species, 20 new insect genera and mammals, gymnosperms and angiosperms are im-
400 new insect species [22]. These large-scale proving (Fig. 3). These improvements suggest that
biodiversity investigations provide a solid basis current conservation activities are relatively effective
for further species descriptions in China. As the for most mammals, gymnosperms and angiosperms.
world’s largest completed flora, Flora Reipublicae In contrast, urgent actions, which are different from
Popularis Sinicae (FRPS) started in 1958 and current conservation approaches, are needed to re-
was completed in 2004. It documented 31 141 verse the decline of birds, reptiles and amphibians.
vascular plant species in 126 books of 80 volumes Despite recent progress, many taxonomic
(Table S1, Fig. S1) [23]. Subsequently, FRPS was groups, such as invertebrates and fungi, have not
Page 4 of 25

Figure 3. Trends in IUCN listings of threatened species in China. (A) Changes in the threatened ranks of gymnosperms,
angiosperms, amphibians, reptiles, birds and mammals in China. The numbers at the end of bars were the numbers of species
assessed. (B) Red List Index for gymnosperms, angiosperms, amphibians, reptiles, birds and mammals in China. The first
assessment was in 2004, the second in 2013 for plant species and 2015 for animal species. Red List Index is an indicator
of overall extinction risk for a set of species. It is used to track meaningful trends in biodiversity status simply by looking at
overall changes in threat status of the same set of species between updates [31].
been thoroughly inventoried. We still lack sufficient in our knowledge may bias further biogeographic
data to assess the threat to them. Second, some analysis and biodiversity conservation planning.
remote areas such as south-eastern Tibet and Further research should explore poorly inventoried
border areas in Yunnan province have not been well taxonomic groups and areas and check questionable
inventoried for most taxonomic groups. Finally, identifications of herbarium specimens using DNA
only 10% of specimens in Chinese collections barcoding. Such research may elucidate mechanisms
have precise geographic localities. Moreover, some of biodiversity formation at biogeographic scales
species are certainly misidentified [32]. These gaps by integrating the big data from species catalogues
Page 5 of 25
with data about the tree of life, geological history Atlas of China (1 : 1 000 000) and a vegetation di-
and other factors influencing species distributions vision map (1 : 10 000 000) were both published in
over time. 2001, with 11 groups of vegetation types and 796 for-
mations [37]. The two maps were subsequently up-
National Specimen Information Infrastructure dated with 960 formations and subformations and
and big biodiversity data 116 vegetation regions [38]. Since the two vegeta-
In recent decades, increasing effort and financial tion maps were based on vegetation surveys in the
investment have enabled the development of 1980s, China’s vegetation map was recently updated
many biodiversity databasing platforms. Dozens with 12 vegetation types and 866 vegetation for-
are now online [33]. The Chinese National mations/subformations. About 3.3 million km2 of
Specimen Information Infrastructure (NSII; China’s vegetated area has changed vegetation type
http://www.nsii.org.cn/) was established in 2003 since the first vegetation map in the 1980s [39]. The

to document historical and current distributions of Vegetation of China summarized the floristic compo-
species in China. NSII also includes multiple sub- sition and distribution of the main vegetation types
platforms, such as the Chinese Virtual Herbarium [40]. Chinese vegetation scientists have started to
(http://www.cvh.ac.cn/), National Animal Collec- write monographs about particular vegetation units
tion Resource Center (http://museum.ioz.ac.cn/), defined in their system of vegetation classification.
Specimen Resources Sharing Platform for Edu- They introduced the new term ‘vegegraphy’ for this
cation (http://mnh.scu.edu.cn/), Digital Spec- type of monograph. The first volume of Vegegraphy of
imen Sharing Platform for Nature Reserves China (about spruce forests) was published in 2017
(http://www.papc.cn/), Chinese Field Herbar- [41].
ium (https://www.cfh.ac.cn/) and Plant Photo
Bank of China (http://www.plantphoto.cn/).
NSII has already digitized 15.7 million specimens Biodiversity dynamics and monitoring
from 329 Chinese herbaria and museums and Many regions in China simultaneously display a
13 million colour photographs and other items mix of positive and negative biodiversity trends.
(http://www.nsii.org.cn/). These species cata- Several national-scale biodiversity monitoring net-
logues and digitized specimens provide essential works have been established across major ecosys-
information on the names, taxonomic relationships tems and nationwide environmental gradients to
and distributions of life in China and play critical enable monitoring. These networks include the Chi-
roles in understanding species origins, evolution nese Biodiversity Observation and Research Net-
and biodiversity conservation [13,34,35]. work (Sino-BON), China Biodiversity Observation
In 2018, CAS initiated the Big Earth Data Sci- Network (China-BON), Chinese National Ecosys-
ence Engineering Project, with a budget of US$250 tem Research Network (CNERN), Chinese Ecosys-
million over five years (CASEarth, http://www. tem Research Network (CERN), Chinese Forest
casearth.com). One of the critical elements of Ecosystem Research Network (CFERN) and Na-
CASEarth is to integrate available biodiversity in- tional Forest Inventory. These networks have statis-
formation for users from the academic community, tically rigorous designs with conceptual models of
decision-makers, conservation practitioners and the biodiversity to quantify and understand these trends
general public (http://www.bio-one.org.cn/). Citi- and their drivers. They provide forewarning of bio-
zen science is an increasingly important mechanism diversity loss and enable adaptive management of
for collecting data on species distributions across biodiversity for policymakers [42].
space and time. Citizen science approaches for bird
species have been well developed and have been Sino-BON and China-BON
used to identify almost nationwide potential habi- Sino-BON and China-BON monitor biodiversity
tats of 1111 bird species, including 167 nationally change in multiple taxonomic groups, includ-
protected species and 70 threatened species. Nearly ing plants, animals and microbes. The Chinese
25% of the nationally protected species and 20% of Academy of Sciences initiated Sino-BON in 2013.
the threatened species use farmland as habitat [36]. It is composed of 10 subnetworks, including
Thus, new agricultural systems in China should de- three subnetworks of plant diversity (CForBio,
velop to integrate traditional and scientific knowl- Steppe & Desert Network and Forest Canopy
edge of sustainable intensification towards conserv- Network), six subnetworks of animal diversity
ing focal species in high-priority areas. (Mammal, Bird, Amphibian and Reptile, Insect,
Soil invertebrate, and Freshwater fish), and a
Vegetation maps and ‘vegegraphy’ of China subnetwork of soil microbial diversity. For each
With the efforts of three generations, including over subnetwork, several sites have been established
200 Chinese vegetation scientists, the Vegetation across China [43] (Fig. 4). The Ministry of Ecology
Page 6 of 25
ZYZ
DXAL
FL
Synthesis Center LS
DLS
ML
QLFP QLHG CBS1
Standard & Criteria CBS2
Zoological Botanical Microbial
Sub-Center Sub-Center Sub-Center BTM
Data Management
Mammal & Sharing BDGS
ObNet CFor BiO
WL TT
Remote Sensing
Bird ObNet Steppe & QJY-GTS
Desert ObNet
Soil Microbial YLXS
ObNet MuL
Amphibian Forest Canopy
and Reptile PE
ObNet Desert ObNet
NBH CBL

NG DHS
XSBN
Fish ObNet
Insect ObNet
Soil
Invertibrate
ObNet
Sino-BON CForBio
Qianjingyuan
National Park site
Legend
5 ha plot
+ 1 ha plot
0.04 ha plot
+ 0.09 ha plot
24 ha plot
Survey grid
Functional zones
Traditional utilization zone
Core protection zone
Recreation and exhibition zone
Ecological conservation zone
Figure 4. An illustration of how Sino-BON is organized from sites (Qianjiangyuan National Park, GTS) to subnetwork (CForBio) and national-scale network
(Sino-BON) for monitoring dynamics of species and ecosystems, and multiple trophic interactions through cooperation among the 10 subnetworks.
Top left panel: structure of Sino-BON (Chinese Biodiversity Monitoring and Research Network) including a synthesis centre and zoological, botanical
and microbial sub-centres. The Zoological sub-centre includes six subnetworks (ObNet): Mammal, Bird, Amphibian and Reptile, Fish, Insect and Soil
Invertebrate. The Botanical sub-centre includes three subnetworks: CForBio (Chinese Forest Biodiversity Monitoring Network), Steppe and Desert, and
Forest Canopy. The microbial sub-centre includes only the microbial subnetwork. Top right panel: 20 large forest dynamics plots along latitude in Eastern
China. Bottom right panel: 1 km × 1 km grids in four zones of Qianjiangyuan National Park. Bottom left panel: biodiversity monitoring at the Park from
top to bottom: drone for near-surface remote sensing, forest crane for canopy biodiversity, infrared-triggered camera for mammals and birds, automated
acoustic recording device for songbirds, phylogeny of woody species with molecular data, dendrometer measuring the growth of trees, seed rain trap,
seedling plot, and insect nest trap. The map of China is from http://bzdt.ch.mnr.gov.cn/.
and Environment of China initiated China-BON Sino-BON has established a multidisciplinary

in 2011. It now has four subnetworks (Mammals, research platform to monitor species and ecosys-
Birds, Amphibians and Butterflies). China-BON tems’ dynamics and multiple trophic interactions
now includes more than 440 observation sites and through cooperation among the 10 subnetworks,
9000 line or point transects to cover representative as illustrated in Fig. 4. CForBio has established
ecosystems in China [44]. 23 large forest dynamics plots in one subnetwork,
Page 7 of 25
each with an area around 20 ha, covering forests National Forest Inventory
from boreal to tropical zones. The plots monitor The State Forestry and Grassland Administra-
1893 woody species and more than 2.69 million tion of China has established a national-scale
tree individuals with a total plot area of 658.6 ha forest inventory system to monitor forest re-
[45]. In some of the CForBio plots, the Sino-BON- sources and biodiversity. The national forest
Forest-Canopy subnetwork has equipped eight for- inventory has been conducted at five-year inter-
est cranes to monitor forest canopy microclimate vals since 1973. The ninth inventory was in 2018
and its diversity of plants, arthropods and microbes (http://www.forestry.gov.cn/gjslzyqc.html). A
[46]. The Sino-BON-Mammals subnetwork has set systematic sampling design uses 2 × 2 km to
up 20–150 infrared-triggered cameras in 1 × 1 km 8 × 8 km grids in all provinces to conduct a qual-
grids of 30 representative natural forests to moni- itative assessment. Permanent plots (0.0667 ha
tor terrestrial mammals and ground-dwelling birds. each) have standardized measurements in each plot,

The Sino-BON-Birds subnetwork has established including tree diameter at breast height (DBH) and
16 international and 38 domestic sites across the height of trees with ≥5 cm DBH. Since the seventh
Eurasian continent to monitor 2569 migratory in- inventory (2004–8), 415 000 sampling plots were
dividuals of 63 bird species using remote telemetry surveyed for each inventory. From the first to
devices [43]. the ninth inventory, the forest coverage in China
increased from 12.69% to 22.96%, and the total
growing stock volume from 8.66 to 17.56 billion m3
CERN, CFERN and CNERN [49]. This platform monitors forest resources dy-
To observe long-term changes in ecosystem namics in China and accounts ecosystem services,
functions, the Chinese Academy of Sciences ini- such as carbon stocks, and their interactions with
tiated CERN in 1988. It now consists of 44 field biodiversity and human disturbance [50].
stations covering all major ecosystems in China
(http://www.cern.ac.cn). Using a standardized
monitoring and data quality-control system, most
PROCESSES AND MECHANISMS
CERN stations have accumulated data over 30 Origin and evolution of China’s
years. These provide unparalleled insights into biodiversity
ecosystem dynamics and changes. An example is
the long-term observational data on soil organic Uplift of the Qinghai-Tibetan Plateau created
carbon in the top 20-cm layer in old-growth forests a cradle for biodiversity evolution
in southern China from 1973–2003. There was an As an evolutionary cradle of biodiversity, many
accumulation of atmospheric carbon at an unex- lineages in China originated with topographic
pectedly high rate. It highlighted the importance changes and climate shifts, such as the uplift of the
of establishing long-term observational studies on Qinghai-Tibetan Plateau and surrounding regions,
the responses of belowground processes to global the development of the East Asian monsoon, and the
change [47]. The former Ministry of Forestry of aridification of Western China [51]. There is increas-
China initiated CFERN in 1992 and currently ing evidence that a proto-Tibetan Plateau had prob-
has 110 field stations covering nine forest types ably already developed in Cretaceous-Paleogene
(https://cfern.org/). It includes two main forest times (∼60 Ma; ‘Ma’ meaning ‘million years ago’)
transects: the North–South Transect of East- before the collision of India with Asia. A gener-
ern China, along temperature and precipitation alized expansion of the Qinghai-Tibetan Plateau
gradients in Eastern China, and the West-East Tran- started no later than the early Miocene to Pliocene
sect of Southern China, along the Yangtze River (25–5 Ma). It was accompanied by continued
from near sea level to altitudes higher than 6000 m surface uplift of high mountain ranges and the rise of
on the Qinghai-Tibetan Plateau [48]. To integrate, a wide east–west orientated deep central valley until
standardize and coordinate the above-described the Neogene [52,53]. Chinese biologists have pro-
platforms sponsored by different sectors in China, vided striking examples of how these geographical
the Ministry of Science and Technology of China developments continuously shaped lineage diversi-
sponsored CNERN. It seeks to integrate observa- fication for various taxonomic groups. For example,
tion facilities and data resources and standardize a series of geological events such as the growing
research methods, tools and protocols. Currently, Eurasian land from the Tethyan region (about
CNERN has 53 field stations covering China’s 40 Ma), the collision of the Indian and Asian plates
main ecosystems, selected from well-equipped and during the Eocene (37–34 Ma) and the closure of
developed stations in CERN, CFERN and others the Turgai Strait (∼29 Ma) [54] drove the adaptive
(http://cnern.cern.ac.cn/en/). radiations of Holarctic amphipods of the genus
Gammarus.
Page 8 of 25
Similarly, Hynobiidae, an ancient lineage of living of admixture and isolation between heterogeneous
salamanders, diversified with the continuous uplift habitats in sky-islands have driven intraspecific pop-
of the Qinghai-Tibetan Plateau and its neighbour- ulation diversification of the cushion willow (Salix
ing regions [55]. The concept of a ‘proto-Tibetan brachista) and likely many other species in the
Plateau’ with a flat centre is contentious. For exam- Hengduan Mountains [69].
ples, well-preserved fossil palm leaves suggest an ex-
tensive valley system with a floor lower than 2.3 km Refugia acting as an evolutionary museum
above sea level in central Tibet during the late Pa- for relict species
leogene (25 Ma) [56]. Moreover, the south-eastern China is also an evolutionary museum. Qiu et al.
margin of Tibet in the latest Eocene (∼34 Ma) synthesized previous phylogeographic studies to
was likely to have been ∼3000 m above sea level identify refugia of relict species, including those
and attained its present elevation only in the early near the south-eastern edge of the Qinghai-Tibetan

Oligocene [57]. Plateau (Hengduan Mountains, Yungui Plateau),
After India’s collision with Asia, a burst of ter- the Three Gorges Mountains Region, and many
restrial faunal and floral diversification occurred other mountain ranges such as the Qinling and Nan-
in China and neighbouring areas. For example, ling ranges [70]. In addition to these refugia, Zhao
both mountain building and intensification of the et al. also recently identified three refugia for ginkgo
Asian monsoon likely accelerated diversification in south-western, southern and eastern China [71].
and colonization of Asian frogs of the tribe Paini Tang et al. identified long-term climatically sta-
(Anura: Dicroglossidae) in freshwater ecosystems ble refugia for relict plant species in south-western
[58]. Temperate alpine flora emerged from early China and northern Vietnam [14].
Oligocene and its diversification peaked in the At a national scale, China is both a museum and a
middle Miocene in Tibet-Himalaya-Hengduan re- cradle of biodiversity, while it also has multiple bio-
gions [59,60]. Some high-alpine vertebrate species geographic origins from other continents. The analy-
acquired anatomical or physiological adaptations sis of a dated phylogeny of Chinese angiosperms and
to extreme environments [61–64], while modern occurrence data revealed that eastern China (hu-
alpine plant genera started to diversify on the mid to semi-humid areas) has served as a floristic
Qinghai-Tibetan Plateau during the Late Miocene museum, by a signature of older divergence in its
[65]. whole flora, as well as both a museum and a cradle
After the Miocene, along with global cooling in for woody genera [13]. In contrast, western China
the Pliocene and recent glacial cycles, a new period (arid to semi-arid areas) is an evolutionary cradle for
of diversification started on the Qinghai-Tibetan herbaceous genera, shaped by recent diversification
Plateau and its margins. For example, a Pliocene in its flora [13]. Contrastingly, 38% of plant taxa in
mammal assemblage was found from a high-altitude Eastern Asia arrived from other floras in the North-
basin in the western Himalayas, with cold adapta- ern Hemisphere and the Gondwana. Nonetheless,
tions such as the woolly rhinos (i.e. Coelodonta tolo- about 48% of plant taxa in Eastern Asia have an
goijensis and C. thibetana) [66]. These findings show in situ origin, providing additional evidence of
that cold-adapted mammals started to diversify China’s role as a floristic cradle [72].
within this region in the Pliocene [66,67].
Recent studies support species diversification Domestication of plants and animals
through multiple mechanisms such as allopatric spe- The domestication history and rapid evolution of do-
ciation, pollinator-mediated isolation and diploid mestic plants and animals have puzzled evolutionary
hybridization on the Qinghai-Tibetan Plateau [68]. biologists since Darwin. At least 20% of the world’s
Recently, a novel model of speciation may under- cultivated species (136 out of 666 species) origi-
pin the formation of the region’s high biodiver- nated from China, where more than 8000 years of
sity, known as the ‘Mixing-Isolation-Mixing’ (MIM) selection shaped modern varieties of Chinese crops
model. Unlike the classic allopatric model with and livestock [21]. These examples provide an excel-
full geographical isolation between populations, the lent opportunity for the study of domestication [15].
MIM model allows gene flow to shuffle adaptive Modern techniques allow reconstruction of the his-
gene complexes built up during isolation to generate tory of plant and animal domestication. This history
many new combinations and thus diversify exponen- highlights the role China has played in the domesti-
tially. By combining analyses of genomic data and cation of global staple crops and livestock. As exam-
historical geographic information, mangrove specia- ples, dogs were domesticated from wolves in south-
tion appears driven by multiple MIM cycles of open- ern East Asia ∼33 000 years ago [73]. Silkworms
ing or closing of sea straits during historical sea (Bombyx mori) may have been initially domesti-
level fluctuations [68]. Similarly, recurrent cycles cated in China as tri-moulting lines. They spread
Page 9 of 25
independently along the Silk Road, which resulted tively stable glacial-interglacial climate (most no-
in the development of local varieties [74]. tably south-western China) have higher proportions
Studies on domestic plants and animals also of both endemic bird species and endemic plant
unravel the mechanisms of rapid evolution behind species [81,82]. Contemporary temperature is posi-
artificial selection, providing insights into the past. It tively associated with overall tree species richness in
also suggests strategies for future crop and livestock China and North America, perhaps supporting the
breeding, considering conditions to ensure their metabolic theory of ecology [34]. Notably, Chinese-
survival under future climates. For example, under led research has also shown that glacial-interglacial
artificial selection, some of the candidate genes climate change, along with contemporary climate,
highly expressed in the Chinese native dogs’ (Canis topographic heterogeneity, and species traits, deter-
lupus familiaris) brain evolved rapidly. They were mine the range-size of terrestrial vertebrates at the
partially responsible for dogs’ behavioural trans- global scale [83].

formation [75]. Comparative genomics revealed In addition to these natural drivers, to reflect
the role of interspecific gene flow in cattle (Bos China’s long cultural history and dense human
taurus) domestication. For example, Tibetan cattle population, many studies have addressed the con-
obtained the gene for hypoxia adaptation from sequences of anthropogenic activities across past
sympatric yaks (Bos grunniens) [76]. Rice (Oryza decades, centuries and millennia in China. Those ac-
sativa) and wheat (Tricicum aestivum) are major tivities have impacted the range dynamics of mam-
global crops. Through selection and breeding, crops mals and plants and the distribution and population
have undergone significant phenotypic changes in size of threatened species [84–88]. For example, hu-
flowering time, grain size, colour, shattering, seed man disturbance constrains herbivorous waterfowls’
dormancy and tillering. For example, a minor-effect ability to track the ‘green wave’ of food availability
quantitative trait locus, DTH2 (days to heading pro- along their spring migration routes [89]. Extensive
tein on chromosome 2), has been shown to probably artificial shorelines, loss of essential intertidal and
represent a target of human selection for adapta- wetland regions, global warming, and larval trans-
tion to long day length [77]. Likewise, RNA-seq port are the main factors limiting distributions of in-
techniques have revealed the mechanisms for the tertidal invertebrates along the Chinese coast [90].
enhanced environmental adaptability and im- Notably, a large-scale field experiment conducted in
proved grain quality of hexaploid wheat (Tricicum subtropical regions in China and subarctic regions
aestivum, AABBDD) compared with its diploid in Norway found that nutrient enrichment can mod-
progenitor (Aegilops tauschii, DD) and tetraploid ify temperature-biodiversity relationships in aquatic
progenitor (Triticum turgidum, AABB). Expansion microcosm communities [91].
of agronomically relevant gene families in Aegilops
tauschii is associated with disease resistance, abiotic
stress tolerance and grain quality [78]. Phylogenetic and functional diversity
distribution and the underlying drivers
Geographical distribution of biodiversity While taxonomic diversity treats species as ecolog-
As a consequence of differences in evolutionary ically equivalent, phylogenetic diversity and func-
history, topography, contemporary and past cli- tional diversity better reflect species’ evolution-
mate, species richness and diversity patterns vary ary and ecological differences [79,80]. Besides
across China. Many studies have tested the effects higher plant species richness, China also has higher
of these different temporal-spatial factors on Chi- plant phylogenetic diversity than Europe and North
nese biodiversity from different perspectives, includ- America, reflecting their different histories [92,93].
ing taxonomic, phylogenetic and functional diver- Phylogenetic diversity for both gymnosperms and
sity [34,79,80]. Our understanding of these various angiosperms in China also declined with increasing
drivers’ contribution to biodiversity patterns across contemporary climatic stress, consistent with a role
China is growing, providing insights into mecha- for tropical niche conservatism in shaping China’s
nisms and drivers behind diversity patterns else- plant diversity [80]. More frequent phylogenetic
where. clustering than overdispersion in Chinese terrestrial
vertebrates suggests essential roles of both regional
ecological and evolutionary factors, such as envi-
Taxonomic diversity distribution and the ronmental filtering and intra-regional speciation, in
underlying drivers shaping animal assemblages [94].
Species distributions and species richness patterns, As an important facet of biodiversity, func-
and their multiple-scale drivers have been widely tional diversity links to ecosystem function and
studied in China. Specifically, regions with rela- ecosystem services across temporal-spatial scales
Page 10 of 25
[79,95]. Notably, an influential global study found dynamics plots have provided strong evidence for
that root trait diversity was greatest in the tropics. CNDD in temperate, subtropical and tropical forests
It declined sharply in temperate and desert biomes [100]. For instance, density dependence can be a
due to both evolutionary history and soil resource key factor in structuring seedling survival patterns
supply [79]. Another global study showed that wa- at very local scales. This can, in turn, result in a
ter availability influenced the relationship between community showing compensatory trends [101].
maximum plant height and hydraulic traits, e.g. taller In some cases, negative interactions also operate
species from wet regions had greater xylem efficiency among neighbours of closely related species [102],
and lower hydraulic safety [96]. At the national scale, but this effect’s strength is smaller than for con-
precipitation, plant life-form and evolutionary his- specific neighbours [100]. By incorporating multi-
tory together affect the relationship between the ple life stages and numerous influential factors, the
type of leaf margins of Chinese woody plants and strength of CNDD also varies with tree life stages

temperature [95]. A recent national study also found and fluctuating environments, with the strongest
that contemporary climate affects the geographi- CNDD at early life stages [103]. In addition, recent
cal variation in sexual systems of Chinese woody observational work with a tropical species has shown
plants indirectly through its effects on mature plant that seedlings near closely related conspecific neigh-
height [97]. bours have reduced growth performance [104]. Fur-
ther experimental work suggests that soil microbes’
within-species specialization helps explain such in-
traspecific variation in CNDD [105].
Biodiversity maintenance in local
Although CNDD has been detected widely in
communities plant communities, empirical support for the pre-
How species coexist in diverse communities where diction that CNDD caused by natural enemies can
many species share similar resource requirements translate into enhanced community diversity re-
has been an enduring puzzle for community ecol- mains limited. Key studies in subtropical forests
ogists. Classical theory shows that stable coexis- combined with field surveys and shade-house exper-
tence occurs when stabilizing niche differences out- iments have revealed that adult trees cause density-
weigh fitness differences by causing species to more dependent mortality in conspecific seedlings by reg-
strongly limit themselves than limit their heterospe- ulating the frequency of pathogenic soil fungi [106].
cific competitors [98]. The processes that determine Such pathogen-mediated density dependence could
the distribution and abundance of species include translate into ‘rare species advantage’ in the com-
dispersal limitation, abiotic filtering, resource community [107]. Moreover, tree species’ adverse ef-
petition and trophic interactions with natural ene- fects on seedling survival extend to their close
mies and mutualists. In the context of rapid degrada- relatives because of phylogenetic conservatism in
tion and destruction of natural ecosystems, there is host-specific interactions between plants and their
an urgent need to understand the underlying mecha- pathogens [108]. Similar patterns were also evi-
nisms of species coexistence that may protect species dent in temperate forests, where natural enemies,
from local extinction and inform ecosystem restora- such as plant-associated fungi and insect herbivores,
tion. Furthermore, the resilience of a community to reduce seedling recruitment and survival at high
environmental changes often stems from a level of adult conspecific density. This pattern is especially
redundancy. Understanding the role of species and pronounced for ectomycorrhizal and shade-tolerant
guilds would facilitate a clearer understanding of sys- species [109]. It suggests that plant–pathogen inter-
tems’ resilience and thereby enable restoration to actions and feedbacks through host-specific changes
ensure continued ecosystem function. in soil communities are important for plant species
coexistence. More importantly, a recent empiri-
cal study provided compelling evidence that both
The prevalence of conspecific density harmful pathogenic fungi and beneficial ectomyc-
dependence in plant communities and the orrhizal (EcM) fungi shape interspecific variation
underlying mechanisms in the strength of CNDD (Fig. 5). Pathogenic
Conspecific negative density dependence (CNDD) fungi may play a key role in driving tree interac-
may be a major mechanism for the maintenance of tions’ strength but can be overruled by EcM fungi
plant diversity. Proximity to conspecific adult plants [110]. Further studies evaluating the possible links
reduces seedling survival through resource compe- between CNDD, pathogens, mycorrhizal symbio-
tition and attacks by host-specific pathogens, her- sis and local plant species diversity may explain
bivores or predators [99]. Covering the complete how numerous plant species can coexist in natural
climatic spectrum in China, the analyses of forest communities.
Page 11 of 25
A B to broaden the applicability of functional traits to

0.6 0.6
large-scale patterns and their generating mecha-
Conspecific seedling effect
Conspecific seedling effect

0.4 0.4
nisms [116]. By sequencing the transcriptome of
0.2 0.2 gene ontologies for light use and harvesting among
0 0
co-occurring species, the functional genomic simi-
larity of these genes has the potential to increase the
-0.2 -0.2
breadth and depth of our understanding of how gene
-0.4 -0.4 function influences community assembly [117].
-0.6 -0.6
-4 -3 -2 -1 0 1 2 3 -4 -3 -2 -1 0 1 2 3 Biodiversity and ecosystem functioning

Pathogen accumulation rate EcM fungus accumulation rate
The tremendous diversity in China is at risk from

C 0.9 D 0.9 rapid economic development, its large human pop-
0.6 0.6 ulation, and climate change, destroying biodiversity
Conspecific adult effect
Conspecific adult effect
0.3 0.3 and harming ecosystem functioning [17]. China’s

high biodiversity and its regional heterogeneity pro-
0 0
vide an exceptional opportunity to conduct con-
-0.3 -0.3 trolled experiments and field surveys across natural
-0.6 -0.6 gradients to understand how the processes maintain-
-0.9 -0.9
ing biodiversity contribute to ecosystem functions.
Chinese ecologists have recently strived to under-
-4 -3 -2 -1 0 1 2 3 -4 -3 -2 -1 0 1 2 3 stand the mechanisms behind the relationship be-
Pathogen accumulation rate EcM fungus accumulation rate tween biodiversity and ecosystem function in both
forest and grassland ecosystems. One element of
Figure 5. (A–D) Relationships between species-specific CNDD effect and pathogenic
this research involves biodiversity–ecosystem func-
and EcM fungus accumulation rates demonstrating that species coexistence in sub-
tropical forests depends on the interplay between mutualistic and pathogenic fungi tioning (BEF) experiments across various ecosys-
[110]. Solid lines and dashed lines indicate significant and non-significant effects at tem types. Generally, they found that increasing bio-
P < 0.05, respectively. Each point represents a tree species with different mycorrhizal diversity leads to increased ecosystem functions—
associations. such as productivity and stability—a relationship
that appears across scales. This relationship is likely
mediated by the biotic interactions across trophic
Functional traits, phylogeny and community
levels and interspecific trait variation.
assembly
To understand the processes underlying community
assembly, Chinese ecologists have studied the in- BEF relationships in forest ecosystems
fluence of biotic and abiotic factors on plant phy- The largest BEF experiment in forest ecosys-
logenetic and functional diversity and composition. tems started in 2009/2010 in subtropical China
Resource partitioning, such as interspecific habi- (referred to as BEF-China). Among 26 BEF plat-
tat differences in topography and soil nutrients, forms around the world, BEF-China is the forest
and dispersal limitation, has emerged as an impor- experiment with the largest number of species
tant explanatory factor for multiple biodiversity pat- (42 tree species and 18 shrub species), the
terns in China’s plant communities. The patterns largest area (around 40 ha) and the highest
included species–area relationships, variation in range of species richness (from monocul-
community composition across space or time (β tures to mixed forests with 24 tree species)
diversity) and functional alpha diversity [111,112]. (http://www.treedivnet.ugent.be/). The design of
At a global scale, evolutionary history and the co- the species mixtures considered differences among
evolution of key functional traits, such as root and taxonomic, phylogenetic, functional diversity and
hydraulic traits, have been explored by Chinese genetic diversity [118]. This experiment extrapo-
scientists to determine functional trade-offs along lated the findings of previous grassland experiments.
large-scale environmental gradients [96,113,114]. It showed that biodiversity roughly doubled plant
Chinese ecologists have proposed new research productivity and carbon storage across the tested
pathways. For example, intraspecific trait variation range of species richness. The positive effects of bio-
may predict demographic rates better than species- diversity increased significantly over eight years of
level traits. The latter cannot capture some dimen- plant growth [119]. BEF-China also includes plots
sions of tree function critical for demographic rates in a neighbouring national park’s natural forests
[115]. ‘Community-level traits’ have been proposed to compare with experimental stands [120]. The
Page 12 of 25
current findings suggest that restoring biodiversity dominant species [129]. The combined effects of
based on multispecies afforestation strategies can above- and belowground biodiversity explained 45%
effectively mitigate climate change and provide of the variation in ecosystem multifunctionality
many other benefits and services. [125]. Yet high livestock diversity (sheep and cattle
In subtropical forests, the biotic interactions grazing vs. sheep or cattle or no grazing) increased
across trophic levels and interspecific trait varia- ecosystem multifunctionality by increasing diversity
tion mediated the mechanisms underpinning the of a range of taxonomic groups, including plants, in-
observed positive BEF relationships, rather than insects, soil microbes and nematodes. This relation-
traspecific genetic diversity [121–123]. For exam- ship was much clearer when the analysis correctly
ple, plant diversity had non-random dilution effects accounted for the different components of diversity
on root pathogenic infection [121], while phyloge- and the different functions [122].
netic diversity of herbivore communities decreased At a national scale, carbon storage increased with

with increasing tree phylogenetic diversity [122]. increasing biodiversity, supporting the generality of
Species richness effects were more related to in- the relationship. Based on 6098 forest, shrubland
terspecific trait variation than intraspecific genetic and grassland sites across China, soil organic carbon
diversity [123]. storage was substantially enhanced by increasing
species richness and belowground biomass. Species
richness, aboveground net primary productivity, and
BEF relationships in grassland ecosystems belowground biomass can be as important as envi-
China has established a series of manipulative BEF ronmental drivers’ direct effects such as precipita-
experimental platforms in glasslands. They include a tion and temperature [130].
BEF experiment with plant removal in Inner Mon-
golia [124], a BEF platform on the Qinghai-Tibetan
Plateau [125] and an experiment studying the ef- Ecosystem services and socio-economic
fects of livestock diversity on grassland ecosystem development
functioning in north-eastern China [126]. These ma-
Biodiversity enhances many ecosystem services,
nipulative experiments provide the platforms for a
such as buffering climate extremes, regulating pests
deeper understanding of the ecological mechanisms
and producing food and wood, in addition to cul-
of BEF relationships and the response of plant com-
tural identity and aesthetic inspiration [3]. However,
munities to global change.
recent biodiversity loss due to anthropogenic activ-
The positive biodiversity-stability relationship
ities has weakened ecosystem services by altering
appears in different grasslands, likely caused by
ecosystem functioning and stability at broad scales
asynchronous dynamics among ecosystems’ com-
[131]. In China, scientists have recently made sev-
ponents. Using a 24-year study of Inner Mon-
eral comprehensive assessments of ecosystem ser-
golian grassland, Bai et al. documented plant di-
vices at both regional and national levels, including
versity’s positive influence on ecosystem stability
evaluating the complex relations between ecosystem
[127]. In species-rich communities, they found that
services and socio-economic development. They
ecosystem stability increased from species-level to
propose adding the results of ecosystem services
functional-group to whole-community level due to
assessments into ecological planning to enable
asynchronous dynamics among major components
integrated assessments and planning.
at both species and functional-group levels. Sim-
ilarly, in alpine grassland on the Qinghai-Tibetan
Plateau, a long-term climatic warming experiment The assessment of ecosystem services
over 30 years (1983–2014) showed increased grass To reduce the risks of natural disasters and re-
abundance and decreased sedge abundance. It did store natural capital, China has invested over US$50
not affect the total aboveground net primary produc- billion through the Natural Forest Conservation
tivity [128]. This demonstrated that in species-rich Programme and the Sloping Land Conversion Pro-
ecosystems shifting species composition in response gramme between 2000–2009. These programmes
to climate warming may stabilize primary produc- have enabled the continued provision of various
tion by shifting from aboveground to belowground ecosystem services, including food production, car-
productivity. bon sequestration, soil retention, flood mitigation,
In contrast, the BEF experiment in an alpine site sandstorm prevention and water retention [132].
on the Qinghai-Tibetan Plateau found that warm- Another assessment of China’s six key ecological
ing, rather than precipitation change, reduced tem- restoration projects suggested that the total annual
poral stability over five years. It did so by alter- carbon sink in the regions covered by these projects
ing the asynchronous population dynamics of the between 2001–2010 was estimated to be 132 Tg
Page 13 of 25
carbon per year. Over half of this carbon was from protected area category that includes integrated con-
implementing these projects [133]. Notably, the sideration of biodiversity, ecosystem services and
value of ecosystem services provided by the giant human activities. National level redline programmes
panda (Ailuropoda melanoleuca) and its reserves is have also been developed to maintain ecosystem ser-
about 10–27 times higher than the cost of support- vices and biodiversity. Such approaches provided an
ing these reserves [134]. Ecosystem services pro- additional mechanism to maximize ecological secu-
vided by other taxa are likely understudied. Exam- rity. They also illustrate how environmental policy
ples include pollination services provided by birds, within China has a solid scientific basis.
bats and insects.
The balance between ecosystem services THREATS AND RESPONSES

and socio-economic development Species endangerment is often the result of a combi-

Ecosystem services are the benefits nature pro- nation of extrinsic and intrinsic factors, which lead
vides to people. That said, strategies for improving to declines in populations and long-term viability.
ecosystem service provision may conflict with local Extrinsic factors include habitat loss, degradation
socio-economic development. For example, the and fragmentation, overexploitation, global climate
revegetation programmes in China’s Loess Plateau change, diseases, environmental pollution, and bi-
increased the net primary productivity to an up- ological invasion. Intrinsic factors include genetic
per limit for the water resources. This limit may diversity loss and inbreeding depression. Here, we
threaten the provision of sufficient water for human only focus on three aspects that have shown signif-
demands [135]. The Relocation and Settlement Pro- icant progress in China over recent decades.
gramme of Southern Shaanxi Province would bene-
fit local governments, downstream water consumers
and global beneficiaries over the long term. Short- Species endangerment and adaptation
term costs to residents and government may out- Over the past decade, conservation genetics and ge-
weigh these benefits, however [136]. In south-west nomics have evolved and progressed significantly
China, strict conservation of rain forest would pro- in China. The genetic diversity, endangerment his-
vide greater values from many ecosystem services tory and causes, and survival and adaptation strate-
than ‘cash forest monocultures’. Nonetheless, local gies of many threatened species in China have been
people still prefer to plant monoculture forests, e.g. investigated, especially those of threatened verte-
for rubber and tea [137]. Yet, well-planned initia- brates. This work has enabled the development of
tives can provide mutual environmental and eco- science-based conservation strategies. Two new sub-
nomic benefits. An example is the Paddy Land- disciplines within conservation biology have been
to-Dry Land programme in Beijing. It successfully proposed recently. The first is conservation evolu-
improved water quantity and quality more than it tionary biology [140]. The second is conservation
reduced agricultural output, creating benefits that metagenomics [141], based on the incorporation of
were more than five times greater than the costs evolutionary ideas and metagenomic technology.
[138]. Exploring synergies provides a mechanism to
maximize benefits and enable both the satisfaction of
Genetic diversity and evolutionary potential
human needs and biodiversity maintenance.
High genetic diversity often implies high genetic
evolutionary potential to respond to environmental
Adding ecosystem services into ecological changes. Genome-wide studies provide comprehen-
planning sive insights into the evolutionary potential of threat-
Ecosystem services matter to people, and therefore ened species. Recent studies revealed at least five
programmes for landscape planning should incorpo- cryptic phylogenetic species in Chinese giant sala-
rate them. For example, in a framework for drawing manders (Andrias davidianus) [142] and two phylo-
the ecological redline areas in Shanghai, the inclu- genetic species in red pandas [143], which require
sion of ecosystem services in planning would sub- subsequent lineage-specific conservation strategies.
stantially increase terrestrial habitat protection and Consistent with their small population sizes, the
decrease the trade-offs between development and threatened Baiji river dolphin, Myanmar snub-nosed
environmental quality [139]. A national assessment monkey (Rhinopithecus strykeri), Himalayan red
of protected areas in China found that China’s na- panda (Ailurus fulgens), ironwood tree (Ostrya re-
ture reserves have not played a large enough role in hderiana) and Cercidiphyllum japonicum [144] had
protecting both biodiversity and key ecosystem ser- very low genetic diversities [143,145,146]. Crested
vices [35]. These authors suggest establishing a new ibis (Nipponia nippon) populations have lost almost
Page 14 of 25
half of their ancestral genetic diversity [147]. These more effective management strategies to mitigate
examples highlight the urgency of protecting their the future impacts of climate change on species
genetic diversity while simultaneously reducing the diversity.
impacts of habitat degradation and human activ-
ities. In contrast, the threatened giant panda, Ti- Survival and adaptation strategies
betan antelope (Pantholops hodgsonii) and Chinese Species that survived through historical environ-
red panda (Ailurus styani) still harbour high lev- mental change often evolved adaptive strategies
els of genetic variation [71,143,148,149]. Thus, the to cope with the associated challenges. Recently,
assessment of genetic diversity can guide targeted genome-wide studies have provided insights into
conservation decisions. the mechanisms of adaptive strategies in threat-
ened species. For example, comparative genomics
between the giant panda and the two red panda
Endangerment history and causes

species revealed the genetic mechanisms underlying
Understanding the processes and drivers behind
the morphological (pseudo-thumbs) and physiolog-
species endangerment are fundamental elements
ical convergence (specialized bamboo diet) [151].
of effective wildlife conservation that genomic in-
The DUOX2 gene’s pseudogenization might be the
formation can advance. For example, based on
genetic cause of the energy-saving low metabolic
the whole diploid genome and population genome
rate of giant pandas [152]. Genomic analysis found
data, giant panda’s evolutionary history traces back
that in the high-altitude snub-nosed monkeys, some
over eight million years. It included two popula-
genes related to lung function, DNA repair and an-
tion expansions, two bottlenecks and two diver-
giogenesis shared identical amino acid substitutions
gences [148]. Further analysis revealed that Pleis-
[153].
tocene climate changes and recent human activities
Besides the host genome itself, gut microbiota
were the major drivers of population fluctuations
may also play an important role in the survival
and divergences [148]. Interestingly, the golden
and adaptation of threatened species. Metagenomic
snub-nosed monkey (Rhinopithecus roxellana) and
analysis of gut microbiota identified crucial bacte-
Chinese red panda are largely sympatric in the
ria and digestive enzymes for helping digest cel-
Hengduan Mountains. They demonstrate a similar
lulose and hemicellulose in the bamboo diet of
pattern of historical population fluctuation as the gi-
giant pandas [154] and the leaf diet of golden snub-
ant panda [143,150]. This pattern implies that Pleis-
nosed monkeys [150]. They also help other species
tocene climate changes might have affected the de-
to adapt to the extreme environment of the Qinghai-
mographic histories of some sympatric mammals,
Tibet Plateau [155].
especially large-bodied mammals, in similar ways.
They may also reflect similar historical human pres-
sures on these species. Biosafety and associated mechanisms
In contrast, the narrowly distributed grey, black-
and-white and Myanmar snub-nosed monkeys Biological invasions and transgenic crops are two im-
(Rhinopithecus brelichi, R. bieti, R. strykeri) and portant issues in biosafety. Both could greatly impact
Himalayan red panda have experienced contin- China’s society, economy and biodiversity conserva-
uous population declines during the Pleistocene tion and have been major topics of discussion within
[143,150]. They display demographic histories dif- intergovernmental fora and agreements. In recent
ferent from the relatively widely distributed species decades, as a response to technological advances and
above, highlighting the urgency to reduce genetic increasing capacity, the mechanisms, consequences
diversity loss in these species. For threatened plants, and regulation of biological invasions and the bene-
the widely distributed ginkgo’s population ge- fits and adverse effects of transgenic crops have been
nomics reconstructed multiple cycles of expansion well studied in China.
and reduction during its evolutionary history [71].
In contrast, the critically endangered, narrowly Biological invasions
distributed ironwood tree demonstrated contin- The invasion success of alien species largely depends
uous population decline during the Pleistocene on species characteristics, ecosystem properties, bi-
[146]. Although the demographic patterns of widely otic interactions and human activities [156–158].
and narrowly distributed threatened plants were In addition to life-history traits that link with in-
similar to those of threatened mammals, the specific vasive success, evolutionary or genomic charac-
times of population expansion and reduction were teristics are also important drivers. The genomic
different. Understanding the impact of climate analysis of the widespread invasive plant species
change on different species and guilds, and using Mikania micrantha showed that half of its genome
traits in addition to molecular data, may enable consists of long terminal-repeat retrotransposons,
Page 15 of 25
80% of which derive from a significant expansion benefits in weedy relatives following transgene in-
over the past 1 million years [157]. The evolution- trogression [169]. However, little work has focused
ary study of another highly invasive plant, Ageratina on the direct or indirect biodiversity effects of trans-
adenophora, suggests that it may have evolved to al- genic crops in China. Further work should develop
locate more nitrogen to photosynthesis and reduce better safeguards and better understand the impacts
allocation to cell walls, resulting in stronger growth and trade-offs.
ability [159]. Notably, the nematode ascarosides’
pheromones play an important role in spreading
Biodiversity responses to global change
pine-wilt disease [160]. The symbiotic microbes not
only promote insect invasions but also coevolved Populations and species respond in various ways to
with the invasive insects [158]. In addition, a global global change, including shifts in abundances and ge-
study of alien reptile and amphibian species sug- ographic ranges, changes in phenology, behaviour
and physiological plasticity, as well as evolutionary

gests that both human-assisted dispersal and topo-
graphic heterogeneity increase the rate of spread for adaptations [170–173].
these species [156]. Species composition, phyloge-
netic and functional distances could also affect the Species range shifts
invasion success [161]. In the face of a warming climate, species may move
Adverse consequences of these invasive species to higher latitudes or elevations to find a suitable
on biodiversity are widespread. The exotic Spartina habitat [171]. The large spans of latitude and ele-
alterniflora has homogenized the nematode vation in China provide excellent opportunities to
communities in Chinese coastal wetland across understand range shift dynamics. For example, the
different latitudes [162]. A global study of alien alpine treelines on the Qinghai-Tibetan Plateau have
herpetofauna showed that biodiversity hotspots moved upward due to the climatic warming over the
harbour more biological invasions than other past century [174]. Species interactions, however,
regions [163]. Notably, a quantification for invasion e.g. shrubby densification, may slow such move-
risks of alien terrestrial vertebrates along China’s ments [174]. Similarly, in recent decades hundreds
Belt and Road Initiative countries suggests that of Chinese bird species with small body sizes, large
14 hotspots may be at particular risk of biological geographical ranges, high trophic levels and high
invasion [164]. These risks highlight the need for habitat specificity have moved to regions with higher
additional biosafety measures in these regions. elevation and latitude [175,176]. Studies have also
reported range shifts in snakes, lizards and bats
in the past few decades due to climate change in
Impacts of transgenic crops China [177–179]. A study using distribution data
Transgenic crops could benefit people and nature for 9701 plants across China examined the rela-
by increasing food production per unit-area and de- tionship between human activities and the degree
creasing chemical pesticide use. They also carry risks to which species fill their potential climatic ranges
of harmful ecological impacts and present a signif- [87]. Narrow-ranged and widespread species exhib-
icant threat to biodiversity [165,166]. A long-term ited opposing responses to human activities. Human
(1990–2010) assessment of Bt cotton’s widespread activities reduced ranges of narrow-ranged species,
adoption and reduced insecticide use in north- potentially threatening them. In contrast, human
ern China showed increases in abundance of three activities expanded ranges of widespread species,
arthropod predators and decreases in aphid pests causing biotic homogenization [87].
[167]. In contrast, this regional increase in Bt cot-
ton adoption also progressively increased the popu- Vegetation phenology change
lation size of mirid bugs. They became pests in cot- Phenological change is another mechanism for
ton and other crops [168]. One study considered adapting to climatic change. It includes the advance
the fitness effects of transgenic overexpression of a of spring phenology and delayed autumn phenology.
native gene (EPSP) developed to confer glyphosate These changes have been observed for trees, shrubs,
resistance in rice. It found that transgenic F2 crop– herbs, insects and amphibians in China in the past
weed hybrids produced 48%–125% more seeds per half-century [180]. The high sensitivity of vegetation
plant than non-transgenic controls in monoculture- phenology to global change in the Qinghai-Tibetan
and mixed-planting designs without glyphosate ap- Plateau has been studied in detail [181]. Specifically,
plication. The findings suggest that over-expression growing seasons of trees are extending, showing
of a native rice EPSP gene can lead to fitness earlier starts and later ends in response to climate
advantages, even without glyphosate exposure. The warming based on tree-ring data from the Plateau
over-expressed EPSP gene might result in fitness [182].
Page 16 of 25
In contrast, a study of meadow and steppe vege- should also focus on fundamental scientific aspects
tation on the Plateau suggests that, although warm to promote a deeper understanding of the problems.
springs cause an advance in the start of the growing To achieve breakthroughs, we must identify prior-
season, warm winters can also delay spring phases ity topics and develop enabling conditions to meet
due to later fulfilment of chilling requirements. This these challenges. Such an approach should promote
results in delayed spring phenology [183]. Similarly, more China-led international collaborations to ex-
using observations on seven European tree species, it trapolate results to the continental and global scale,
was demonstrated that the response of leaf unfolding and broader generalization and theory develop-
to climate warming has significantly decreased in the ment. Below, we offer some general suggestions af-
past three decades, possibly linked to reduced chill- ter considering practical guidance for scientists and
ing [184]. Notably, climate change effects on global policy-makers.
vegetation phenology along elevational gradients are

not consistent across different regions. This is poten-
tially due to the differences in human disturbance, Advancing priorities in Chinese
vegetation sensitivities to climate change and tem- biodiversity research
perature lapse rates [185]. We selected four specific research fields which play
critical roles in advancing our fundamental under-
Behavioural and physiological plasticity and standing about the origin, evolution and mainte-
evolutionary adaptation nance of biodiversity. First, as the Third Pole of
In addition to range shifts and phenological changes, the Earth, the Qinghai-Tibetan Plateau and the
species may also survive climate change through surrounding mountains serve as a perfect natural
rapid behavioural or physiological plasticity or rapid laboratory for biodiversity studies. There is some
evolutionary adaptation in situ [172]. An experimen- excellent research in this region (see our detailed
tal study showed that even early-stage turtle embryos review in section ‘Origin and evolution of China’s
could move within the egg to adjust to small-scale biodiversity’). Nonetheless, there are still many re-
thermal heterogeneity in their environment [186]. search gaps and areas with relatively little survey
This thermoregulatory behaviour is widespread in data. Examples include the southern slope of the
reptile and bird species [187]. The embryo could Himalayas and the high mountains and valleys of
also adjust its physiology to reduce the fitness penal- the region. Moreover, recent global climate and
ties of adverse thermal conditions [172], as even land-use changes are dramatically changing the en-
small changes in egg temperature can impact off- vironment of the Qinghai-Tibetan Plateau [128].
spring viability. Further, Ciona savignyi, an invasive Therefore, further monitoring of the environment
marine tunicate, showed rapid adaptations to envi- is needed to protect its biodiversity and fragile
ronmental changes through DNA methylation mod- ecosystems more effectively.
ification, causing significant epigenetic signatures Second, subtropical forests in China cover over
[188]. 25% of the total land area (Fig. 2), ranging from
23o N to 33o N in latitude and 93o E to 123o E in
longitude. China harbours nearly 70% of the to-
tal global area of subtropical forest, while other re-
A PERSPECTIVE ON FUTURE gions at these latitudes are deserts or semi-deserts
BIODIVERSITY RESEARCH IN CHINA [189,190]. Most subtropical forests are evergreen
Whilst there has been significant recent progress, broadleaved, with high levels of biodiversity and en-
considerable knowledge gaps remain. They include demism. Over one-third of China’s vascular plant
the geographic distributions and the conservation species and nearly 35% of national nature reserves
status of most species in China (e.g. small inver- are in this region [189]. Various biodiversity and
tebrates, fishes, insects, bats, amphibians, reptiles), ecological studies have been carried out in these
the role of trophic interactions (e.g. food webs regions [13,47,69,119]. Further studies should ad-
and pollination networks) in biodiversity mainte- dress both the serious anthropogenic threats by
nance, the links between biodiversity and ecosystem intensive agriculture and industrial activities and
services, and community-level responses to global the high degrees of urbanization, and the impor-
change. The majority of the studies we have sum- tance of subtropical forests for biodiversity globally
marized focused on specific research questions in and ecosystem functioning and services at local to
China. Many of these studies focused on application regional levels [189].
rather than conceptual or theoretical advances or Third, with 32 000 km of coastline, China’s seas
methodological breakthroughs in basic biodiversity cover an area of 4.73 million km2 [9]. The last
science. In the future, biodiversity research in China 70 years have seen significant advances in marine
Page 17 of 25
biodiversity research [9]. Nonetheless, there is still versity. It includes the development and implemen-
a large disparity in marine biodiversity studies’ ad- tation of the ecological redline policy [195], the con-
vancement compared to terrestrial biodiversity stud- struction of a national-park-centred protected-area
ies. Further studies in marine biodiversity should system, and the recent enforcement of the perma-
strengthen marine specimens’ collections, monitor nent ban on wildlife trade and consumption. The
marine biodiversity in various habitats, and conduct ecological conservation redline policy aims to in-
research on biodiversity changes in Chinese seas clude a quarter of China’s land to protect most
and nearshore ecosystems. There should be stan- species and their habitats in China [195]. China’s
dard monitoring practices to assess change over time updated national park system was proposed in 2013.
[9]. The recent advances in terrestrial biodiversity It builds the system’s basic framework through 10 pi-
studies also provide excellent examples to develop lot national parks of over 220 000 km2 [196]. Re-
marine biodiversity monitoring networks. Coastal cent advances in identifying biodiversity hotspots,

ecosystems are also under-researched. They include species distributions, drivers of threat and habitat
some of the most threatened ecosystems in terms of fragmentation provide a robust scientific basis for se-
loss, degradation and reclamation [191]. lecting the areas for ecological redlines and design-
Fourth, as the world’s most populous country ing and managing national parks [13,197]. Research
with a long history of human activity, China faces a on mechanisms behind biodiversity and the rela-
severe challenge to balance biodiversity and socio- tionship between biodiversity and ecosystem func-
economic development, especially in eastern regions tioning facilitates the conservation and restoration
with intensive human activities [87]. China also pro- of endangered species and degraded ecosystems
vides a key area for understanding how to conserve [110,121]. At present, the connections between sci-
and restore biodiversity in the Anthropocene. That entists, policy-makers, stakeholders and the public
understanding must include the effects of rapid ur- are frequently weak. Communication between sci-
banization on ecological community assembly and entists with the public should be improved. For
plant–animal interactions [192], effects of climate example, using citizen scientists to collect biodi-
change on the phenology and demography of species versity data is pivotal in addressing gaps in bio-
[181], and the basis for rewilding or reintroducing diversity targets. To establish better collaboration
extirpated species back into natural areas [193]. The with policy-makers to enable science-based policy
Yangtze River is the third-longest river in the world. and implementation, the China Council for Interna-
The surrounding regions still face severe threats to tional Cooperation on Environment and Develop-
freshwater and terrestrial biodiversity due to rapid ment (CCICED) has initiated special policy stud-
urbanization in the delta and the construction of ies which work alongside China’s five-year plans
dams along the river. Dams causing waterway frag- (http://www.cciced.net/cciceden/).
mentation within China have reduced biodiversity
both above and below the dams and reduced genetic
connectivity by impairing dispersal and migration of
freshwater organisms [194]. Although the Chinese
Advanced technology application
national government and scientists are making con- in biodiversity research
siderable efforts to monitor and protect this region’s With further innovations in genomics, remote sens-
biodiversity, there is still a long way to go. Calls ing and other technologies, we can expect to see
for more systematic planning of river-basin manage- additional complementary and synergistic research
ment have been slow to be enacted. Effective mon- that uses China’s continually expanding research
itoring of biodiversity changes is urgently needed, capacity to further push the research frontiers in
in addition to exploring the underlying mechanisms biodiversity science.
of species coexistence, quantifying threats to endan- The rapid development of high-throughput
gered species and evaluating the synergism between sequencing technology and genomics has promoted
biodiversity and socio-economic development. insights into species’ genetic diversity, from several
molecular markers to the whole-genome level. The
decrease of sequencing costs enables large-scale
Translating scientific advances genomic diversity surveys at the species level.
Chinese scientists and institutions have played an
into biodiversity conservation indispensable role in this genomics wave. For
and ecosystem restoration practices example, in 2010, international scientists launched
Ecological civilization is an explicit national gover- a G10K genome project to sequence the genomes
nance strategy of the Chinese government to priori- of 10 000 vertebrate species. The Beijing Genomics
tize the protection and restoration of China’s biodi- Institute-Shenzhen played an important role,
Page 18 of 25
including leading the One Thousand Plants and teractions with global networks have enabled global
Animals Genome Project (2010), the One Thou- standards for field census data, but such standards
sand Insects Transcriptome Project (1KITE, 2011) are less frequent for other biodiversity data. Thus,
and the Ten Thousand Birds Genome Project further collaborative actions, shared sites and coor-
(B10K, 2014). Recently, research teams from the dination must ensure that the huge effort going into
Chinese Academy of Sciences also led the Ten biodiversity research within China can be entirely
Thousand Fishes Genome Project (2019) and the complementary.
Ten Thousand Protists Genome Project (2019).
These massive genome projects have provided
unprecedented insights into genomic diversity and Strengthening and expanding
enabled a more in-depth understanding of biodi- international collaborations
versity evolution. In addition, the rapid application Solving many questions in biodiversity requires

of high-throughput sequencing technology has also global collaboration. Chinese scientists have greatly
facilitated the development of the International benefited from close cooperation with foreign scien-
Barcode of Life (iBOL) project. Chinese scientists tists and international organizations in the last few
have substantially contributed to the iBOL project, decades. However, further partnerships are needed
especially for insects and vascular plants [198]. With to maintain and strengthen the existing joint re-
further advancement of these approaches, we can search projects, support the ongoing establishment
expect to transform our understanding of various of regional collaborations (e.g. Mapping Asia Plants
evolutionary processes of biodiversity on regional Project, Sino-Africa Joint Research Center, and
and global scales. Southeast Asia Biodiversity Research Institute), and
Advances in monitoring techniques have allowed develop new cooperative biodiversity research plat-
an understanding of the factors driving generaliza- forms. The recent development of the Belt and Road
tion and specialization from the cellular and molecu- Initiative (BRI), initialized by the Chinese govern-
lar level to the factors governing community compo- ment in 2013, provides the opportunity to expand
sition, ecosystem functioning and ecosystem service the collaborative research in biodiversity between
provision. Advanced remote sensing technologies China and over 60 countries across mainland Eura-
allow almost real-time spatially continuous large- sia, Africa and the Middle East [201] alongside the
scale monitoring of ecosystems using different sen- development of linear infrastructure across these
sors to explore leaf chemical composition, biodi- countries. The Belt and Road science plan aims to
versity and functional traits [199,200]. China has overcome some of the challenges associated with
the technology to implement these approaches, but developing sustainably at an unprecedented scale
frameworks to promote more interdisciplinary re- through the use of big data and remote sensing, cou-
search are still needed. Long-term biodiversity mon- pled with capacity-building initiatives. Attention is
itoring using traditional and novel methods is the needed to ensure that the route itself avoids the key
key to addressing fundamental and practical ecology biodiversity areas. Moreover, supportive infrastruc-
questions. Scientists across China have established ture and raw materials also need to be accounted for
several long-term biodiversity monitoring networks. to prevent a much wider footprint than the new lin-
By taking advantage of the extensive coverage and ear infrastructure alone will cause. New cooperative
uniqueness of China’s ecosystems and biodiversity, mechanisms need to reduce the potential risks of the
we believe that coordinated, distributed experiment BRI on biodiversity, such as habitat fragmentation
and observation networks with long-term monitor- and biological invasions [201].
ing will substantially improve research infrastruc- Meanwhile, China harbours several international
ture and quality. Interdisciplinary facilitation mech- biodiversity hotspots which cross national bound-
anisms to enhance cooperation between groups and aries, such as the Himalaya, Mountains of Central
institutions are also required to enable cross-taxa Asia, and Indo-Burma. Transboundary cooperation
research. The development of standards and pro- through coordinated inventory, monitoring and re-
tocols must ensure the collection and dissemina- search needs to be strengthened substantially to pro-
tion of biodiversity data in a coherent and stan- tect these key areas [202]. They include popula-
dardized way to enable better comparisons between tions of endangered species, such as the critically
and within sites over time. China’s extensive biodi- endangered Amur leopards across China and Rus-
versity science research is starting to provide long- sia [203]. Chinese biodiversity researchers also need
term data on numerous ecosystems across space to become more active in global intergovernmen-
and time. However, data collection standards and tal initiatives with relevance to biodiversity, such as
approaches and data-sharing mechanisms will need IPBES (the Intergovernmental Science-Policy Plat-
further attention to improve analyses. Increasing inform on Biodiversity and Ecosystem Services), CBD
Page 19 of 25
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70°E 80°E 90°E 100°E 110°E 120°E 130°E 140°E

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· Origin & Evolution · Drivers of Threats

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and Environment of China initiated China-BON Sino-BON has established a multidisciplinary

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A B to broaden the applicability of functional traits to

Conspecific seedling effect

-4 -3 -2 -1 0 1 2 3 -4 -3 -2 -1 0 1 2 3 Biodiversity and ecosystem functioning

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Conspecific adult effect

0.3 0.3 and harming ecosystem functioning [17]. China’s

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The balance between ecosystem services THREATS AND RESPONSES

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