Biological Conservation 285 (2023) 110173
Contents lists available at ScienceDirect 
Biological Conservation 
journal homepage: www.elsevier.com/locate/biocon 
Perspective 
Opportunities and challenges in Asian bee research and conservation 
Natapot Warrit a,*, John Ascher b,*, Parthib Basu c,d, Vasuki Belavadi e, Axel Brockmann f, 
Damayanti Buchori g,h, James B. Dorey i, Alice Hughes j, Smitha Krishnan k, Hien T. Ngo l, 
Paul Williams m, Chao-Dong Zhu n,o,p, Dharam Abrol q, Kamal Bawa r,s, Chet Bhatta t, 
Renee M. Borges u, Silas Bossert v,w, Cleofas Cervancia x, Nontawat Chatthanabun a, 
Douglas Chesters n, Phung Huu Chinh y, Kedar Devkota z, Hanh Pham Duc aa, Rafael Ferrari n,ab, 
Lucas Garibaldi ac,ad, Jin Ge o, Dibyajyoti Ghosh ae, Dunyuan Huang af, Chuleui Jung ag,ah, 
Alexandra-Maria Klein ai, Jonathan Berenguer Uhuad Koch aj, Erin Krichilsky ak,al, 
Krushnamegh Kunte f, Tial C. Ling am, Shanlin Liu an, Xiuwei Liu ao, Arong Luo n, Shiqi Luo an, 
Junpeng Mu ap, Tshering Nidup aq, ZeQing Niu n, A. Mustafa Nur-Zati ar, Shannon B. Olsson f, 
Gard W. Otis as,at, Fang Ouyang o, Yan-Qiong Peng au, Windra Priawandiputra av, 
Maxim Proshchalykin aw, Rika Raffiudin av, Anandhan Rameshkumar ax, Zongxin Ren ay, 
Azhagarraja Suruliraj az, Sanjay Sane f, Xiaoyu Shi n, Palatty Allesh Sinu ba, Deborah R. Smith bb, 
Zestin W.W. Soh bc, Hema Somananthan bd, Tuanjit Sritongchuay ai,be, Alyssa B. Stewart bf, 
Cheng Sun bg, Min Tang bh, Chawatat Thanoosing m,bi, Teja Tscharntke bj,bk, Nico Vereecken bl, 
Su Wang bm, Kanuengnit Wayo bn, Siriwat Wongsiri bo, Xin Zhou an, Zhenghua Xie bp,bq, 
Dan Zhang br, Yi Zou bs, Pengjuan Zu bt, Michael Orr n,bu,** 
a Center of Excellence in Entomology and Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok, Thailand 
b Department of Biological Sciences, National University of Singapore, Singapore 
c Department of Zoology, University of Calcutta, Kolkata, India 
d Centre for Pollination Studies, University of Calcutta, Kolkata, India 
e Agricultural Entomology, University of Agricultural Sciences, Bengaluru, India 
f National Centre for Biological Sciences, Tata Institute of Fundamental Research (NCBS-TIFR), Bengaluru, India 
g Departemen Proteksi Tanaman, Fakultas Pertanian, IPB University, Bogor, Indonesia 
h Center for Transdisciplinary and Sustainability Sciences (CTSS), IPB University, Bogor, Indonesia 
i College of Science and Engineering, Flinders University, Adelaide, Australia 
j School of Biological Sciences, University of Hong Kong, Hong Kong, China 
k Bioversity International, Bengaluru, India 
l Food and Agriculture Organization of the United Nations, Rome, Italy 
m Natural History Museum, London, United Kingdom 
n Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, China 
o State Key Laboratory of Integrated Pest Management, Institute of Zoology, Chinese Academy of Sciences, Beijing, China 
p College of Life Sciences, University of Chinese Academy of Sciences, Beijing, China 
q Division of Entomology, Faculty of Agriculture, Sher- e -Kashmir University of Agricultural Sciences &Technology, Chatha, India 
r Ashoka Trust for Research in Ecology and the Environment, Bengaluru, India 
s Department of Biology, University of Massachusetts, Boston, United States 
t Radford University Carilion, Roanoke, USA 
u Centre for Ecological Sciences, Indian Institute of Science, Bangalore, India 
v Department of Entomology, Washington State University, Pullman, USA 
w Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA 
x Institute of Biological Sciences, College of Arts and Sciences, University of Philippines Los Baños College Laguna, Los Baños, Philippines 
y Mountain Bee Development Center, Hanoi, Viet Nam 
z Agriculture and Forestry University, Rampur, Nepal 
aa Bee Research Centre, Hanoi, Viet Nam 
* Corresponding authors. 
** Correspondence to: Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, China. 
E-mail addresses: Natapot.W@chula.ac.th (N. Warrit), dbsajs@nus.edu.sg (J. Ascher), michael.christopher.orr@gmail.com (M. Orr).  
https://doi.org/10.1016/j.biocon.2023.110173 
Received 22 March 2023; Received in revised form 15 June 2023; Accepted 17 June 2023   
Available online 7 August 2023
0006-3207/© 2023 The Author(s). Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-
nc-nd/4.0/).
N. Warrit et al.                                                                                                                                                                                                 B  i o  l o g  i c a  l  C  o  n s  e r  v a  t i o  n 285 (2023) 110173
ab Environmental Science Training Center, Federal University of Southern Bahia, Porto Seguro, Brazil 
ac Universidad Nacional de Río Negro, Instituto de Investigaciones en Recursos Naturales, Agroecología y Desarrollo Rural, Río Negro, Argentina 
ad Consejo Nacional de Investigaciones Científicas y Técnicas, Instituto de Investigaciones en Recursos Naturales, Agroecología y Desarrollo Rural, Río Negro, Argentina 
ae Southern Regional Centre, Zoological Survey of India, Chennai, India 
af Key Laboratory of Conservation and Utilization of Pollinator Insect of the Upper Reaches of the Yangtze River, Ministry of Agriculture and Rural Affairs, Chongqing, 
China 
ag Department of Plant Medicals, Andong National University, Andong, Republic of Korea 
ah Agriculture Science and Technology Research, Institute, Andong National University, Andong, Republic of Korea 
ai University of Freiburg, Nature Conservation and Landscape Ecology, Freiburg, Germany 
aj United States Department of Agriculture - Agricultural Research Service, Pollinating Insect Research Unit, Logan, USA 
ak Columbia University Ecology, Evolution, and Environmental Biology (E3B), New York, USA 
al American Museum of Natural History, Invertebrate Zoology (IZ) and Richard Gilder Graduate School (RGGS), New York, USA 
am Bee Protection Laboratory, Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai, Thailand 
an Department of Entomology, College of Plant Protection, China Agricultural University, Beijing, China 
ao Institute of Agro-Products Processing, Yunnan Province Academy of Agricultural Science, Kunming, China 
ap Ecological Security and Protection Key Laboratory of Sichuan Province, Mianyang Normal University, Mianyang, China 
aq Sherubtse College, Royal University of Bhutan, Kanglung, Bhutan 
ar Entomology Branch, Forest Research Institute Malaysia, Kepong, Malaysia 
as School of Environmental Sciences, University of Guelph, Guelph, Ontario, Canada 
at Institute of Bee Health, Vetsuisse Faculty, University of Bern and Agroscope, Schwarzenburgstrasse, Bern, Switzerland 
au CAS Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, China 
av Department of Biology, Faculty of Mathematics and Natural Sciences, IPB University, Bogor, Indonesia 
aw Federal Scientific Center of the East Asia Terrestrial Biodiversity, Far East Branch of the Russian Academy of Sciences, Vladivostok, Russia 
ax Hymenoptera Section, Zoological Survey of India, M-Block, New Alipore, Kolkata, India 
ay Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, China 
az Department of Entomology, Dr. Rajendra Prasad Central Agricultural University, Pusa, India 
ba Central University of Kerala, Kasaragod, India 
bb University of Kansas, Lawrence, KS, USA 
bc National Parks Board, Singapore Botanic Gardens, Singapore 
bd School of Biology, Indian Institute of Science Education and Research, Thiruvananthapuram, India 
be Department of Computational Landscape Ecology, Helmholtz Centre for Environmental Research-UFZ Leipzig, Leipzig, Germany 
bf Department of Plant Science, Faculty of Science, Mahidol University, Bangkok, Thailand 
bg College of Life Sciences, Capital Normal University, Beijing, China 
bh Department of Biological Sciences, Xi’an Jiaotong-Liverpool University, Suzhou, China 
bi Department of Life Sciences, Silwood Park Campus, Imperial College London, UK 
bj Agroecology, University of Goettingen, Goettingen, Germany 
bk Centre for Biodiversity and Sustainable Land Use, University of Goettingen, Goettingen, Germany 
bl Agroecology Lab, Université Libre de Bruxelles (ULB), Brussels, Belgium 
bm Institute of Plant Protection, Beijing Academy of Agriculture and Forestry Sciences, Beijing, China 
bn The Native Honeybee and Pollinator Research Center, King Mongkut’s University of Technology Thonburi, Ratchaburi, Thailand 
bo Agricultural Interdisciplinary Program, Graduate School, Maejo University, Chiang Mai, Thailand 
bp Institute of Highland Forest Science, Chinese Academy of Forestry, Kunming, Yunnan, China 
bq Key Laboratory of Cultivating and Utilization of Resources Insects of State Forestry Administration, Institute of Highland Forest Science, Chinese Academy of Forestry, 
Kunming, China 
br Characteristic Laboratory of Forensic Science in Universities of Shandong Province, Shandong University of Political Science and Law, Jinan, China 
bs Department of Health and Environmental Sciences, Xi’an Jiaotong-Liverpool University, Suzhou, China 
bt Department of Environmental Systems Science, ETH Zürich, Zürich, Switzerland 
bu Entomologie, Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany  
A B S T R A C T   
The challenges of bee research in Asia are unique and severe, reflecting different cultures, landscapes, and faunas. Strategies and frameworks developed in North 
America or Europe may not prove applicable. Virtually none of these species have been assessed by the IUCN and there is a paucity of public data on even the basics 
of bee distribution. If we do not know the species present, their distribution and threats, we cannot protect them, but our knowledge base is vanishingly small in Asia 
compared to the rest of the world. To better understand and meet these challenges, this perspective conveys the ideas accumulated over hundreds of years of cu-
mulative study of Asian bees by the authors, including academic, governmental, and other researchers from 13 Asian countries and beyond. We outline the special 
circumstances of Asian bee research and the current state of affairs, highlight the importance of highly social species as flagships for the lesser-known solitary bees, 
the dire need for further research for food security, and identify target research areas in need of further study. Finally, we outline a framework via which we will 
catalyze future research in the region, especially via governmental and other partnerships necessary to effectively conserve species.   
1. Special challenges for pollinator research in Asia treating the region was published in 1896 (Dalla Torre, 1896), over a 
century ago, demonstrating a severe gap in our knowledge of Asian bee 
Concerns about potential insect declines and the subsequent loss of biodiversity. 
essential ecosystem services such as pollination are increasing world- The knowledge gaps in Asia are alarming, as bee extinctions have 
wide (IPBES, 2016; Harvey et al., 2020; Wagner, 2020). Bees are widely been only recently documented in well-studied areas such as Singapore 
acknowledged as the most important terrestrial pollinators, with many (Ascher et al., 2019). In another worrying example, the largest bee in the 
plants reliant on their actions (Klein et al., 2007; Ollerton et al., 2011; world, Megachile pluto Smith, has been sold online multiple times despite 
Requier et al., 2022); their conservation depends on our knowledge of conservation concerns (Fig. 1K; Vereecken, 2018). The remaining Asian 
all facets of their life history and distribution (Hortal et al., 2015). Asia bee fauna remains virtually unexplored from a conservation perspective, 
holds 15 % of total bee diversity, with many unique lineages (Fig. 1), but because monitoring bees and verifying declines is challenging and re-
these many species comprise only 1 % of public global bee specimen quires baseline data that do not yet exist (Orr et al., 2021). Most research 
data (Orr et al., 2021). In addition, the most recent complete catalog and pollination conservation actions or policies rely heavily on, or may 
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be augmented by, information from specimen databases and, when reduction of pesticide use and pollution of water, light, and air; increase 
available, IUCN Red List status assessments, but we are only beginning in landscape heterogeneity and semi-natural areas; implementation of 
to build these foundational resources. pollinator-friendly practices across landscapes; protection and restora-
No-regret solutions can be enacted now as a first step, including tion of biodiverse areas and threatened species; and control of alien 
Fig. 1. Asian bee biodiversity. Apidae: A. Xylocopa insularis Smith; B. Xylocopa myops Ritsema; C. Ceratina collusor Cockerell; D. Ctenoplectra chalybea Smith; E. Apis 
dorsata Fabricius; F. Geniotrigona thoracica (Smith); G. Bombus supremus Morawitz; H. Nomada adusta Smith; and I. Amegilla andrewsi (Cockerell). Colletidae: J. Hylaeus 
penangensis (Cockerell). Megachilidae: K. Megachile pluto Smith; L. Megachile atrata Smith; and M. Euaspis polynesia Vachal. Halictidae: N. Nomia iridescens Smith and 
O. Lasioglossum adonidiae (Cockerell). Photos (©) were taken by ZWW Soh (A,B,C,D,E,F,H,I,J,L,M,N,O), PH Williams (G), and C Bolt (claybolt.com; K). 
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species (Harvey et al., 2020; Dicks et al., 2021). Where possible, active initiatives. We will discuss case studies from China, India, and Thailand 
restoration of more intact or threatened habitat should also be priori- in depth in this section. 
tized, given dire threats such as land conversion to palm oil and wide- In the last six years, the Chinese Pollinator Forum has regularly 
scale agricultural expansion in the Global South (Grass et al., 2020; brought together regional and overseas experts to advance research and 
Aizen et al., 2022). However, such solutions are generalized and can conservation efforts. Many extensive projects, such as the East China 
conflict with economic aims, with both funds and political will limiting Pollinator Project and the Second Tibetan Expedition Program, have 
their application, making more targeted solutions a priority. To tailor centered on or included strong pollinator components, in an effort to 
these and other efforts most effectively, we must also quickly increase document bee diversity in this world hotspot of bee biodiversity (Fig. 2). 
regional knowledge and tailor conservation planning for the enduring The Chinese Entomological Society’s Special Committee on Pollinator 
challenges faced in Asia (Sodhi et al., 2010; Dicks et al., 2021). Insects has been a guiding force in many of these and other initiatives. 
Local research is needed to generate, consolidate, and verify baseline There is increasing appreciation of the importance of pollinators and the 
biodiversity information; identify and fill information gaps; provide need for their conservation, likely due to international research trends 
appropriate locally-tailored solutions; share information; and establish and the central importance of food security and medicinal plants in 
effective monitoring frameworks (Rabajante et al., 2020; Orr et al., China. Taxonomic and systematics research is also ongoing in China 
2022a). The impediments to pollination research and conservation in (Williams et al., 2017; Niu et al., 2020; Williams et al., 2020; Ferrari 
Asia are unique and severe, and policies or initiatives from North et al., 2021; Zhang et al., 2022), but given the high expected number of 
America or Europe are not necessarily effective elsewhere, as regional undescribed species in China (Orr et al., 2021), much more support is 
habitats, fauna, and cultural and political practices differ. Asia is char- necessary for these ventures, as presently there exist so few data for the 
acterized by immense climatic and topographic heterogeneity, from country that its rank in species richness drops from 6th to 40th in public 
tropics to tundra. Half of the world’s human population exists on this data (Zhu et al., 2022; Fig. 2A–B). This is also the case for many other 
continent—increasingly concentrated in fast-growing cities, with many Asian countries, where metrics such as richness or records per area show 
populations now transitioning to more land-intensive and unsustainable them to be outliers with too few data (Fig. 2E–F). 
diets (e.g., beef) and products (palm oil, rubber, etc.; Grass et al., 2020; Great progress is also being made in India, where there are also 
Marcacci et al., 2022). Public and governmental knowledge and higher numbers of species and endemics (Fig. 2). Conservation efforts on 
appreciation of pollinators are rare, making regulations on practices pollinators saw early support in the All India Coordinated Research 
such as landscape alteration, management, and pesticide use difficult to Project (AICRP) on honeybees, initiated in 1981 under the Indian 
enact or enforce. Trans-boundary research has also been limited by Council of Agricultural Research. In 2007, this initiative became the 
language, political boundary disputes, and cultural practices. Rigid na- broader AICRP on Honeybees and Pollinators. Presently, the project has 
tional restrictions on collecting, sequencing, and sample sharing exac- 26 centers covering 20 states, most of which are affiliated with state 
erbate these issues (Prathapan et al., 2018). agricultural universities. In addition to providing training to farmers in 
In this perspective, we enumerate the unique challenges and urgent sustainable beekeeping, some centers also conduct pollination studies 
needs in Asian bee research, with a focus on East, South, and Southeast involving honeybees, and to a lesser extent, non-honeybees (Krishnan 
Asia (Supplemental Note 1). Following a discussion on the current state et al., 2017; Belavadi et al., 2021). More recently, the Indian Pollinator 
of research, we review the importance of flagship social bee groups of Initiative (InPollin, inpollin.com) was founded as a platform to enhance 
Asia, and highlight the understudied biodiversity of solitary bees. We studies on non-social bees, facilitate interactions between researchers, 
then explore the importance of bees in pollination in agricultural and and to standardize methods that will allow regional studies to be com-
natural ecosystems. We conclude with four sets of recommendations on bined to generate India-wide datasets. The InPollin has organized 
policies and strategies, field-level implementation, community engage- several national meetings, student workshops, and two international 
ment, and priorities for research and monitoring. online seminar series. In-person national meetings have previously 
received international researchers from both nearby and distant coun-
2. The current state of bee research in Asia tries, leading to several collaborative downstream efforts (e.g., this 
paper; Kitnya et al., 2022). One of the initiative’s future goals is to 
In the last decade, many discussions have centered on the need for launch a country-wide effort to document bee diversity in India. 
Asia-wide pollinator research and conservation initiatives. However, Bee research in Thailand has been greatly enhanced by two large 
due to limited funding, organizational struggles, challenges in data projects: (1) the USA-Thai project “Thailand Inventory Group for Ento-
sharing that lead to severe underestimations of true species richness and mological Research” (TIGER, 2022) and (2) the Global Biodiversity In-
endemic species numbers (Fig. 2A–D), and travel restrictions, a unified formation Facility-Biodiversity Information Fund for Asia (GBIF-BIFA). 
platform has been slow to emerge. The establishment of the Asian sec- They have resulted in an impressive collection of bees. Morphological 
tion of both the IUCN (International Union for Conservation of Nature) study of this collection enables reliable identifications, expediting 
Wild Bee Specialist Group and the Asian Pollinator Initiatives Alliance further research (Chatthanabun et al., 2020; Nalinrachatakan et al., 
(APIA) since 2020 will more effectively bring together the expertise, 2021). A next step for Thailand is to collate molecular diagnostic re-
institutions, stakeholders, and data required for a synergistic Asian sources and to integrate them with morphology using an integrative 
pollinator initiative and bee conservation assessments. taxonomy approach across borders (Orr et al., 2020), helping build a 
Challenges faced include limited resources and a lack in many foundation for Southeast Asian bee research. Broader testing and 
countries of tenure-track university and/or governmental bee re- application of species concepts would ensure that the use of species 
searchers well-positioned to participate in equitable international col- names is standardized across countries. An additional promising avenue 
laborations. This is the case even for many countries with advanced bee for bee research in Thailand and throughout Asia is the Asian Pollinator 
research foundations. Both Japan and Russia have species checklists, for Initiatives Alliance (APIA, 2023), founded in early 2020 by several 
example (Tadauchi and Murao, 2014; Lelej et al., 2017), but there is NGOs and foundations based in Thailand and surrounding countries. It 
little funding available to digitize the many bee specimens housed in has been instrumental in connecting private, public, and educational 
these countries, leading to Eastern Russia being ranked 33rd in Discove sectors (e.g., researchers, political foundations, non-profit organiza-
rLife.org checklist-based species richness yet only 67th based on public tions, farmers and other interested stakeholders). To date, most of 
data resources (Fig. 2). Pollinator research in much of the rest of Asia has APIA’s activities have been based in Thailand, but with increasing 
been relatively recent (Cervancia, 2018; Ren et al., 2018). Thus, region- network connections in other Asian countries the organization is keen to 
wide efforts are also still preliminary, and there is much to build on as achieve broader regional impact. 
evidenced by the success of several recently established national 
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N. Warrit et al.                                                                                                                                                                                                 B  i o  l o g  i c a  l  C  o  n s  e r  v a  t i o  n 285 (2023) 110173
A B
USA #1 USA #1
Mexico #2 Mexico #2
Brazil #3 Australia #3
Turkey #4 Brazil #4
Australia #5 South Africa #5
China #6 Canada #6
South Africa #7 France #7
Argentina #8 Argentina #8
Greece #9 Spain #9
Spain #10
Italy #11 Germany #10
Iran #12 Italy #11
Israel #13 Austria #12
Morocco #14 Turkey #13
France #15 Costa Rica #14
Greece #15
India #21
Region Japan #32
East Russia #33
Asia * China #40
Indonesia #52
Others India #46
Japan #60 Thailand #47
Pakistan #48
Mongolia #67
East. Russia #67
Pakistan #71
Rep. Korea #69
Malaysia #78
Indonesia #84
Thailand #80
Malaysia #86
Philippines #94 Singapore #87
0 1000 2000 3000 0 1000 2000 3000
Species Richness Species Richness
[authorative DiscoverLife checklist]
C [publicly available data] D
USA #1 USA #1
Australia #2 Australia #2
Brazil #3 Brazil #3
Mexico #4 Mexico #4
South Africa #5 South Africa #5
China #6 Argentina #6
Argentina #7 Chile #7
India #8 Japan #8
Chile #9 Turkey #9
Indonesia #10 Madagascar #10
Madagascar #11 Spain #11
Peru #12 China #12
Turkey #13 Peru #13
D.R. Congo #14 Morocco #14
P. N. Guinea #15 Costa Rica #15
Philippines #16
Japan #17 India #17
Thailand #18
Pakistan #19
Malaysia #29
Indonesia #39
Pakistan #32
Rep. Korea #42
East. Russia #35
Mongolia #45
Sri Lanka #42 Philippines #46
Thailand #45 Malaysia #57
0 500 1000 1500 2000 0 500 1000 1500
Endemic Species Endemic Species
[authorative DiscoverLife checklist] E [publicly available data] F
Liechtenstein #1 18.82 . . . Singapore #1
Switzerland #2 10.59 . . . Luxembourg #2
Netherlands #3 Andorra #3
Singapore #4 Malta #4
Luxembourg #5 N. Cyprus #5
Norway #6 Liechtenstein #6
Costa Rica #7 S. Vincent & Gr. #7
Estonia #8 San Marino #8
Sweden #9 Israel #9
Germany #10 Seychelles #10
Finland #11 Trinidad & Tob. #11
Poland #12 Dominica #12
Austria #13 S. Kitts & Nevis #13
Ireland #14 Switzerland #14
Malta #15 Costa Rica #15
Rep. Korea #30 Palau #17
Japan #37 Brunei #45
Palau #42 Rep. Korea #57
Japan
Sri Lanka #68
#46 0.0355 0.0006
Timor-Lest #74 0.0006
Thailand #64 0.0171
Kyrgyzstan #77 0.0005
Kyrgyzstan #79 0.0081
Georgia #80 0.0005
Timor-Leste #86 0.0064
Sri Lanka #89 0.0004
Malaysia #88 0.0061
Georgia #89 0.0061 Bhutan #99 0.0003
0 2.5 5 7.5 10 0 0.025 0.050 0.075 0.100
Relative Sampling Effort Relative Species Richness
[records / km²] † [species / km²] †
Fig. 2. Ranking of countries in terms of biodiversity and data resources. Based on authoritative checklist data (Ascher and Pickering, 2022) and public data (Dorey 
et al., in review), focusing on the areas of East and South Asia listed in Supplemental Note 1. The top 15 global areas are given first, with Asian areas listed thereafter 
with other regions excluded. A. Species richness via checklist. B. Species richness via public data. C. Endemic species via checklist. D. Endemic species via public data. 
E. Relative sampling effort via public data, using specimens per area. F. Relative species richness via public data, using species per area. Underlying data are in 
Supplemental Table 1. Overall, the public data vastly underestimate biodiversity compared to the checklist (for instance, China moving from #6 to #40 in species 
richness, from checklist to public data, and Indonesia going from #10 to #39 for number of endemic species). Based on E and F, only Singapore appears to be 
relatively well-known within Asia. 
5
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3. Eusocial bees as flagship groups in Asia change (Thapa et al., 2018). However, even the much more widespread 
A. cerana, adapted to a wide range of climates (Ji et al., 2020), faces 
Eusocial species are abundant in many ecosystems and often threats of competition and pests from the continued use of the intro-
managed for agricultural and horticultural crops, making them familiar duced and managed A. mellifera (Theisen-Jones and Bienefeld, 2016), 
to all. Such flagship species are often essential for effective conservation and in some places they might be largely replaced. Further, highly 
messaging and support. Asian ecosystems harbor the richest diversity of migratory and forest-dependent species, such as A. dorsata (Fig 1E) have 
honeybee species (Apis L.; Apidae), both managed and wild (Smith, been shown to be impacted by urbanization and air pollution (Thim-
2021). The Western Honeybee, Apis mellifera L., is the most prominent megowda et al., 2020) and isolation from natural habitats (Klein et al., 
managed bee worldwide (Garibaldi et al., 2013), even in many parts of 2003). Some of the island-endemic Apis species likely face similar 
Asia where it has been introduced. Contrastingly, recent analyses sug- threats, but this requires further study (Raffiudin et al., 2022). Stingless 
gest there may be 14 or more Asian honeybee species (Smith, 2021; bees may also face threats due to unsustainable harvesting practices, or 
Kitnya et al., 2022; Su et al., in review). Few of these taxa are regularly even through species introductions and subsequent competition (Car-
managed for honey production, and they receive much less research or valho, 2022; Qu et al., 2022). Competition is again a potential factor 
conservation consideration (Abrol, 2020). The full potential of endemic threatening native bumblebees, in light of the continued use of the 
Asian honeybees to pollinate crops remains unrealized, despite their potentially invasive European Bombus terrestris (L.) (Orr et al., 2022b). 
importance (Klein et al., 2003; Krishnan et al., 2017; Garibaldi et al., Co-introduced parasites and pathogens could prove even more 
2013; Marcacci et al., 2022). However, there has been substantial dangerous when introducing non-native species (Cameron and Sadd, 
progress in certain fields, and the genomes of three species (A. florea 2020). The spread of such introduced parasites and pathogens can 
Fabricius, A. dorsata Fabricius (Fig. 1E), and A. cerana Fabricius) were escalate widely through movement of affected native species within the 
recently sequenced and are being actively explored (Karpe et al., 2016; region (Owen, 2017), making disease monitoring imperative. The effects 
Ji et al., 2020; Fouks et al., 2021). Still, many knowledge gaps persist. of climate change are poorly understood for bees in general but are 
Ultimately, the combination of problems facing honeybees and polli- expected to be much more severe in the case of montane species such as 
nation services in Asian ecosystems are specific to the region and require bumblebees (including Bombus supremus Morawitz, Fig. 1G; Williams 
local expertise and research. et al., 2015), and could be compounded by grazing (Xie et al., 2008), 
Another eusocial group, the stingless bees (Meliponini (Apidae)), has warranting urgent investigation and actions. 
gained more attention in the Asian research and management commu-
nities in the last 20 years. More than 57 species from 10 genera are 4. Solid taxonomy of native bees of Asia is imperative 
described from Asia (Ascher and Pickering, 2022; Fig. 1F). They are 
some of the most common bees in subtropical and tropical environ- Non-eusocial species comprise most bee species worldwide (85–90 
ments, including in human-modified landscapes such as agricultural %, Danforth et al., 2019), but they are exceptionally understudied in 
(Klein et al., 2003; Sritongchuay et al., 2019, 2022) and urban (Stewart Asia, as evidenced by limited coverage of regional taxa in the most 
et al., 2018; Stewart and Waitayachart, 2020) habitats, which makes recent global treatise (Michener, 2007; notably, some social taxa such as 
them attractive management candidates. They have long been used for Lasioglossum (Curtis; Halictidae) are also less known; Fig. 1O). Case in 
indigenous meliponiculture in Asia, but have been poorly studied when point, the public data rankings include only one Asian country in the top 
compared to the Neotropical fauna (Engel et al., 2019). Stingless bees 10 for number of endemics (Japan), yet the Discoverlife.org checklist 
are increasingly managed commercially in many parts of Asia, including lists three different Asian countries in that top 10 ranking (China, India, 
in Bhutan, China, India, Indonesia, Malaysia, the Philippines, and Indonesia; Fig. 2C–D). The limited study and knowledge of non-eusocial 
Thailand (Chuttong et al., 2014; Rattanawannee and Duangphakdee, Asian bee species impairs our ability to protect them, given that most 
2019; Salatnaya et al., 2021; Nidup, 2021; Qu et al., 2022). Although IUCN listings of insects rely on distributional data. In China, for instance, 
less studied than honeybees, the medicinal properties claimed of sting- nearly 1300 species have been recorded, but estimates from Yan-Ru Wu 
less bee propolis and honey have increased public perception and and others anticipate the presence of at least 1000 additional species. 
funding for research in Asia in recent years (Al-Hatamleh et al., 2020). Orr et al. (2021) also demonstrated that China was one of the three 
Bumblebees (Bombus Latreille; Apidae) also reach their highest major under-described regions globally (plus Australia and Chile- 
richness in Asia, with half of described species found in China (Williams, Argentina). In addition, many described species are unidentifiable as 
1998, Fig. 1G). There has been a great deal of study, historical and fully verified ranges are typically fragmentary or erroneous. Further-
ongoing, on the systematics of this challenging group (Williams, 1998; more, many data remain unshared, at times a consequence of local 
Williams et al., 2017; Williams et al., 2020; Williams, 2022), resulting in regulations, or due to the difficulty of generating data and a hesitancy of 
tens of thousands of verified specimens in China. These should be taxonomists to share it without proper recognition (Zhu et al., 2022). All 
leveraged for conservation assessments (e.g., IUCN Red Listing). Their this greatly complicates bee identification in the region, especially when 
highly-variable color patterns, often attributable to mimicry (Williams, the undescribed species belong to difficult groups such as the sweat bee 
2007), make identification especially challenging despite publication of genus Lasioglossum. These challenges are exacerbated because many 
identification resources for regions such as the Himalaya (Williams, species-defining type specimens are held outside of their countries of 
2022). The group has seen further study across fields, including the origin, which reduces the identifiability of Asian species and highlights 
genomic and the molecular bases of color patterns (Tian et al., 2019; Sun the moral imperative for such institutions to fully image their type 
et al., 2021). However, behavioral and physiological studies are rela- specimens, including extensive imaging of all useful characters (Orr 
tively scarce in the region (Ge et al., 2021). Although native bumblebees et al., 2020). Further, many institutes focus too much on “high-impact” 
are promising managed pollinators similar to other eusocial bees and are publications in hiring and promotion (Zhu et al., 2022), reducing the 
being reared in some countries (Hannan et al., 1997; Yoon et al., 2002), appeal to students. All of this contributes to a dramatic deficit in bee 
the use of commercialized, non-native species in parts of Asia poses a experts in Asia. Knowledge of the life histories of Asian bee species is 
potential major threat for native pollinators (Matsumura et al., 2004; lagging still further behind, limiting our ability to develop effective and 
Orr et al., 2022b), as seen in parts of South America (Aizen et al., 2022; targeted conservation strategies. 
Orr et al., 2022b). A major challenge to taxonomic work is that there are dwindling 
Among these selected flagship social bee groups, some species such career incentives to do taxonomy (Orr et al., 2020; Zhu et al., 2022). The 
as A. laboriosa Smith, the Himalayan Giant Honeybee, are threatened for delimitation and description of new species and the building up of 
reasons that may include habitat specialization, limited distribution and essential identification resources have historically been slow processes, 
nesting opportunities, unsustainable honey harvesting, and climate exemplified by the ongoing centuries of taxonomic research in areas 
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such as Europe, while Asia has seen far less focus. Coupled with quickly- but diverse small-holdings still dominate many Asian landscapes 
increasing career advancement expectations, there is a huge risk that (Krishnan et al., 2017; Zou et al., 2017; Sritongchuay et al., 2022). Less 
hiring committees and funding agencies will fail to recognize the sustainable systems will require active restoration to facilitate recovery 
importance of supporting basic, especially taxonomic, research (Zhu of wild pollinator populations. Land-use pressures in Asia preclude the 
et al., 2022). conversion of large arable areas into flower-rich habitats. Hence, there is 
To counter these challenges, we must better integrate new technol- a need to identify local wildflower species that efficiently attract and 
ogies into taxonomic research programs (Orr et al., 2021; Hartop et al., support pollinators to small-agricultural holdings in various agro- 
2022). Streamlining and expediting training, when coupled with shared ecological regions (Laha et al., 2022). Furthermore, the impacts on 
data on species (Orr et al., 2022b), would stabilize the taxonomic pollinators of increased land fragmentation and urbanization remain 
foundation necessary for regional and trans-boundary research on bees, understudied in rapidly expanding Asian urban and peri-urban spaces 
even enabling monitoring via newer molecular technologies once reli- (Thimmegowda et al., 2020; Wenzel et al., 2020). There is still potential 
able DNA libraries are available (Tang et al., 2015). Additionally, as a to better align modern practices with new research on sustainability and 
community we should actively seek to recognize and cite taxonomic with indigenous practices that have been honed over millennia and may 
hypotheses and works wherever possible. Given that many fields (e.g., prove to be more sustainable (IPBES, 2016). There is an urgent need to 
ecology, conservation biology) are currently hindered by taxonomic identify and understand the impacts of different interacting drivers of 
uncertainty in many Asian bee taxa, increasing research effort, training, pollinator decline across the region. 
and funding in taxonomy could be an investment beneficial to all. Natural ecosystems have seen markedly less research and conse-
quently remain less understood, as food security demands have made 
5. The understudied importance of bees for pollination in agricultural landscapes higher priority targets for study (Requier et al., 
agricultural and natural systems in Asia 2022). Yet bee pollinators are critical across settings, as an estimated 87 
% of flowering plant species depend on animal pollinators (Ollerton 
If practitioners and researchers cannot identify bees beyond a small et al., 2011), with bees being by far the most common (Willmer, 2011). 
subset of eusocial species, we cannot measure, monitor, and maintain Bees thus impact natural systems across scales, from individual plants 
pollinator services. Although methods may be transferable, knowledge (offspring quantity and quality) to populations and species (plant ge-
itself is not, and many endemic crops and growing systems in Asia netic diversity, gene flow, and evolution), to entire communities and 
cannot be assumed to function in the same way as elsewhere, even for ecosystems (structures and function) (Willmer, 2011). Most bee polli-
the same species. Apple, for instance, requires hand pollination in nation research to date conducted in natural and semi-natural envi-
numerous countries in Asia due to pollinator shortages, as do numerous ronments in Asia has focused on either specific plant taxa or specific bee 
less-studied crops in Southeast Asia (Wurz et al., 2021). Key to any re- taxa (Corlett, 2004; Ren et al., 2018). Community-wide studies are 
gion, however, will be sustainable land management and the preserva- scarce, in large part due to the difficulty of collecting data. However, 
tion of semi-natural and natural areas as pollinator habitat (Krishnan recent technological advances such as using environmental DNA (eDNA) 
et al., 2012; Krishnan et al., 2017; Chatterjee et al., 2020; Garibaldi or metagenomics to study plant-pollinator interactions may improve 
et al., 2021), because biodiversity generally begets food security more prospects (Ruppert et al., 2019), but will remain a challenge for regions 
sustainably than conventional agricultural intensification (Murray Las- with insufficient technological bases. Knowledge of bee-plant in-
ley et al., 2013; Requier et al., 2022). Compared to much of North teractions in natural habitats is essential for conservation efforts, not 
America and Europe, considerable natural landscape heterogeneity still only for assessing bee species with specific habitat, foraging, and/or 
exists in Asia, and urgent evidence-based actions are needed to preserve nesting requirements, but also for the conservation of plant species that 
habitats and the vital pollination services they enable. depend on them. Such research is imperative in the face of rapidly dis-
The management of the three aforementioned eusocial groups is a appearing natural habitats and rising species extinctions (Hughes, 
cornerstone of agriculture in Asia. However, their importance differs 2017). 
regionally. Apis are used in various systems throughout, stingless bees 
are restricted to subtropical and tropical regions, and bumblebees are 6. Looking forward – building a foundation and catalyzing bee 
used elsewhere in greenhouses (Orr et al., 2022b). Even for native research in Asia 
honeybees, much more effort is needed to maximize their potential as 
managed pollinators (Gupta, 2012; Schreinemachers et al., 2017). There is no silver bullet or panacea for the myriad of challenges faced 
Relatively little research has been directed to quantifying the biology by stakeholders and researchers trying to achieve bee conservation in 
and relative importance of non-honeybees in pollination of the large Asia. Both multisectoral, cross-border regional initiatives and 
number of crop species grown in the region (Requier et al., 2022), nationally-tailored approaches will be needed to advance bee research 
despite the growing body of evidence of their importance as essential in the region. We provide the following suggestions to improve bee 
crop pollinators (Klein et al., 2003; Klein et al., 2007), irrespective of research efforts in Asia, with cross-border capacity building and 
managed honeybee abundance (Garibaldi et al., 2013). Such research on knowledge sharing as paramount goals. 
the biology of wild bees and their contribution to pollination, as has On an international scale, pollinator conservation action is currently 
been conducted enacted in at least China and Japan (Sekita and Yamada, framed largely within the Convention on Biological Diversity’s (CBD) 
1993; Men et al., 2018). International Pollinator Initiative and its current, second Plan of Action 
Many crops are grown in Asia, and recent efforts have begun (for 2018–2030). Below we discuss the opportunities for Asia around the 
examining smallholder landscapes (Motzke et al., 2016; Shi et al., 2021; four elements of this action plan: (1) enabling policies and strategies; (2) 
Sritongchuay et al., 2019, 2022; Li et al., 2023), but most systems implementing field-level change; (3) engaging civil society and the 
require further study (Klein et al., 2007). Each region, if not each crop, private sector; and (4) monitoring, research, and assessment. 
in Asia could be the subject of its own review paper. Similarly, the di-
versity and abundance of crop pollinators change across space, time, and 6.1. Enabling policies and strategies 
focal crop (Senapathi et al., 2021). Despite such complexities, active 
research on and optimization of crop systems is paramount, not just for Scientists will need to demonstrate to policymakers the importance 
food security, but also because there is potential for wild pollination in of pollinators. Thus, engagement is central to enacting policy changes 
many systems, which could be hindered by unbridled agricultural and legislative protections. Current modeling shows that by 2050, five of 
expansion (Aizen et al., 2022). Some economically important insect- eight billion people worldwide may be negatively impacted through 
pollinated crops are grown in unsustainable monocultures elsewhere, insufficient nutrition stemming from losses to pollinator-dependent 
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crops (Chaplin-Kramer et al., 2019). In the same study, the impacts species (Koch and General, 2019; Silva et al., 2020; Wilson et al., 2020), 
across all scenarios were the largest for Africa and South Asia. The while also informing on behavior and bee associations with flowers. A 
importance of policies, legislation, and regulations associated with notable example is Singapore, a rare well-documented tropical country 
pollinators should ideally be reflected within national reports submitted (Fig. 2E–F) where over half of its 120 bee species have been recorded on 
by CBD member countries. In a brief review of the most recent national iNaturalist (Ascher et al., 2022); similar usefulness is documented for 
reports submitted by the 48 countries within broader Asia (including the Cambodia (Ascher et al., 2016). Through carefully planned, large-scale 
Middle East), only 22 (46 %) mentioned pollinators. Additionally, survey-style events like BioBlitzes, invaluable standardized data might 
Indonesia, South Korea, and Cyprus were the only countries that even be generated (Orr et al., 2022a). However, caution must be taken 
mentioned pollinators at least four times, highlighting the lack of with highly-polymorphic or cryptic groups such as bumblebees (Wil-
engagement on the importance of pollinators in Asia. The explicit in- liams et al., 2020) and some difficult solitary groups such as Amegilla 
clusion of pollinators or pollination services within National Reports or (Zonamegilla) Popov (Fig. 1I), lest misidentifications be made and spread 
National Biodiversity Strategies and Action Plans signal that these issues widely in the literature. Lastly, local, national, and global experts should 
are a conservation priority and could promote funding and stimulate be mobilized and better credited for validating identifications. 
research in this direction. Further, most mitigation policies for anthro-
pogenic pressures such as pesticides and pollution are determined by 6.4. Monitoring, research, and assessment 
their effects on humans, not plants and animals (Thimmegowda et al., 
2020), highlighting a clear knowledge and policy gap. The conservation Basic capacity building is needed to enable region-wide monitoring, 
profiles of pollinators in Asia could be raised through science commu- as the requisite expertise varies greatly among Asian countries (Cer-
nication and public engagement using selected bee species as flagship vancia, 2018). The overwhelming majority of countries in Asia lack 
representatives, messaging strongly on the benefits of native bees for comprehensive studies and updated information on their fauna or suf-
crop and home garden yields (Klein et al., 2003; Klein et al., 2007; ficient point data (Fig. 2), rendering the assembly of national bee species 
Garibaldi et al., 2013). checklists nearly impossible, excepting herculean efforts such as those 
involved in the DiscoverLife checklist (Ascher and Pickering, 2022). The 
6.2. Field-level implementation other checklists that exist are regional and non-standardized (Rabajante 
et al., 2020), reducing the comparability of studies between countries 
Land use, land management practices, pollution, and pesticide use (Orr et al., 2022a). 
are important drivers influencing pollinator populations in parts of Asia The Asia group of the newly formed IUCN Wild Bee Specialist Group 
(Dicks et al., 2021). Agriculture in Asia has had and will continue to will prove important for Red List Assessment of regional bees and for 
have a relatively high pollinator-dependency, based on current and promoting and connecting regional networks. Ideally, efforts at Red- 
forecasted rates of agricultural expansion and crop diversification Listing will prioritize taxonomic groups that are best-known both in 
(Aizen et al., 2019). Despite some work in agriculture, the prerequisite terms of specimen records and their taxonomic trustworthiness, (e.g., 
data for syntheses on wild pollinator population trends are still largely Apis, Bombus, Meliponini, and select better-known solitary bee groups 
lacking for Asia (IPBES, 2016; Orr et al., 2021). Efforts to fill these gaps such as Trachusa Panzer (Megachilidae)), thereby picking the “lowest- 
must be standardized as best as possible, and balanced with practicality hanging fruit” first before approaching more problematic groups. 
(Rabajante et al., 2020). Farm- and landscape-level research involving However, this is just a first step, and monitoring should entail collection 
pollinator-friendly interventions and sustainable agricultural practices of all bees, as they may provide baseline data later once species can be 
are lacking. Policy incentives for ecologically-friendlier agricultural reliably identified. Special focus should be given to known hotspots such 
practices, and enforcement of existing regulations are also major chal- as the Hengduan mountains where work has been sparse on most bee 
lenges in most countries. Furthermore, few studies exist on pollinator groups (excepting bumblebees; Williams, 2022), and for areas where 
conservation using co-design and participatory approaches in Asia. historical baselines already exist and can be compared to. These regional 
These approaches can be used to promote pollinator conservation in all initiatives should be supported by Asian countries through govern-
human-modified habitats, across developmental gradients. As policy mental and research institution support, as a means to share information 
action is lacking in some countries, direct engagement with stakeholders and knowledge and strengthen regional networks. 
may prove a more immediate solution to the many challenges. Although various government programs, outlined earlier, have been 
initiated by some countries, there are no national assessments of insect 
6.3. Private sector and civil society engagement pollinators in Asia as seen in Brazil, with its first Brazilian Special Report 
on Pollination, Pollinators and Food Production (Wolowski et al., 2019). 
Development is a constant in much of Asia and the explicit incor- Brazil’s ability to complete a national assessment of pollinators was due 
poration of pollinating invertebrates into mandated environmental in part to financial support from many sources, alongside the monu-
impact assessments for building infrastructure or other activities could mental efforts of contributors. In Asia, to undertake regional or national 
generate substantial baseline data while providing pollinator habitat. If assessments of pollinators and pollination, faunal baselines and moni-
such data were mandated as public, the transparency of such processes toring, and research, infrastructural and grant support is needed, with 
would be enhanced alongside bolstered research prospects. Special ef- clear expectations of results within reasonable timelines, designed to 
forts could be made in appealing to the Traditional Chinese Medicine strengthen enduring partnerships between researchers, stakeholders, 
industry and others that directly depend on pollinators (medicinal and policymakers. 
plants; Ren et al., 2014). Further incentives for sustainable develop-
ment, coupled with better enforced penalties for regulatory violations, 7. Summary 
could together pave the way for biodiversity conservation. 
Community scientists can also play an important role in biodiversity We know that almost all regions lack information and data regarding 
monitoring, providing potential long-term effort without the need for pollinators, and the situation is critical in Asia (Fig. 2). How do we close 
substantial funding and research infrastructure. iNaturalist, a highly- these knowledge gaps? This paper highlights the importance of 
accessible public database for documenting flora and fauna, has more improving basic bee knowledge in Asia, as recording and assessing 
than 64,000 records of nearly 700 bee species in broader Asia (as of biodiversity is fundamental to measuring progress toward and ulti-
February 2023; Dorey et al., in review). This platform provides the first- mately achieving conservation goals, including those of the CBD’s 
ever accessible images for many major regional pollinators and better Kunming-Montreal Global Biodiversity Framework. Efforts at gener-
documented distribution, especially for range extensions and rare ating and sharing data must be cross-border and transdisciplinary to 
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surmount the manifold challenges faced in Asian bee research. This is Chuttong, B., Chanbang, Y., Burgett, M., 2014. Meliponiculture: stingless bee beekeeping 
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by bees to enable further sustainable development across the region. Dalla Torre, C.G., 1896. Catalogus Hymenopterorum: Apidae (Anthophila). 1896, vol. 
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Danforth, B.N., Minckley, R.L., Neff, J.L., 2019. The Solitary Bees: Biology, Evolution, 
org/10.1016/j.biocon.2023.110173. Conservation. Princeton University Press. 
Dicks, L.V., Breeze, T.D., Ngo, H.T., Senapathi, D., An, J., Aizen, M.A., Basu, P., 
Declaration of competing interest Buchori, D., Galetto, L., Garibaldi, L.A., Gemmil-Herren, B., Howlett, B.G., 
Imperatriz-Fonseca, V.L., Johnson, S.D., Kovács-Hostyánszki, A., Kwon, Y.J., 
Lattorff, H.M.G., Lungharwo, T., Seymour, C.L., Vanbergen, A.J., Potts, S.G., 2021. 
The authors declare that they have no known competing financial A global-scale expert assessment of drivers and risks associated with pollinator 
interests or personal relationships that could have appeared to influence decline. Nat. Ecol. Evol. 5, 1453–1461. 
Dorey, J.B., Cheshire, P.R., Bolanos, A.N., O’Reilly, R.L., Bossert, S., Collins, S.M., 
the work reported in this paper. Lichtenberg, E.M., Tucker, E.M., Smith-Pardo, A., Falcon-Brindis, A., Guevara, D.A., 
Ribeiro, B., de Pedro, D., Fischer, E.E., Hung, K.L.J., Parys, K.A., McCabe, L.M., 
Data availability Rogan, M.S., Minckley, R.L., Velzco, S.J.E., Griswold, T., Zarrillo, T.A., Jetz, W., 
Sica, Y.V., Orr, M.C., Guzman, L.M., Ascher, J.A., Hughes, A.C., Cobb, N.S., 2023. 
A globally synthesised and flagged bee occurrence dataset and cleaning workflow. 
Code used to generate area rankings are public at https://github. Sci. Data. https://www.biorxiv.org/content/10.1101/2023.06.30.547152v1 (in 
com/jbdorey/Asian_Bee_Perspective, based on totals from the Discov- review).  
Engel, M.S., Kahono, S., Peggie, D., 2019. A key to the genera and subgenera of stingless 
erLife checklist (Ascher and Pickering, 2022; Dorey et al. (in review)). bees in Indonesia (Hymenoptera: Apidae). Treubia 45, 65–84. 
Ferrari, R.R., Niu, Z.Q., Kuhlmann, M., Zhang, D., Zhu, C.D., 2021. The cellophane bees 
Acknowledgements of Colletes Latreille (Hymenoptera: Colletidae) from Xizang (Tibet), China. Zootaxa 
5022, 1–72. 
Fouks, B., Brand, P., Nguyen, H.N., Herman, J., Camara, F., Ence, D., Hagen, D.E., 
This effort is a product of the IUCN Wild Bee Specialist Group’s Asia Hoff, K.J., Nachweide, S., Romoth, L., Walden, K.K.O., Guigo, R., Stanke, M., 
Section. We thank Michael Kuhlmann for helpful suggestions. We thank Narzisi, G., Yandell, M., Robertson, H.M., Koeniger, N., Chantawannakul, P., 
Clay Bolt and Eli Wyman for providing a picture of Megachile pluto. Schatz, M.C., Worley, K.C., Robinson, G.E., Elsik, C.G., Rueppell, O., 2021. The 
genomic basis of evolutionary differentiation among honey bees. Genome Res. 31, 
Smitha Krishnan/ SK would like to thank all funders who supported this 1203–1215. 
research through their contributions to the CGIAR Trust Fund: Garibaldi, L.A., Steffan-Dewenter, I., Winfree, R., Aizen, M.A., Bommarco, R., 
https://www.cgiar.org/funders/ and the Nature-Positive Initiative Cunningham, S.A., Kremen, C., Carvalheiro, L.G., Harder, L.D., Afik, O., 
Bartomeus, I., Benjamin, F., Boreux, V., Cariveau, D., Chacoff, N.P., Dudenhöffer, J. 
H., Freitas, B.M., Ghazoul, J., Greenleaf, S., Hipólito, J., Holzschuh, A., Howlett, B., 
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