Active metabolites of stingless bee nest materials used for meliponitherapy: Bibliometrics, impact of biodiversity in conservation, and emerging microbiome

Patricia Vit, MSc, PhD^1*, Vassya Bankova, MSc, PhD^2, Gina Meccia, MSc^3, David S Nogueira, MSc, PhD^4, Christopher Mduda, MSc, PhD^5, Megan T Halcroft, PhD^6, Zhengwei Wang, MSc, PhD^7, Enrique Moreno^8, Qibi Wang, MSc^9, Amelia Nicolas, MSc, PhD^10, María Araque, MD, PhD^11, and Jason E Stajich, PhD¹²

1. Apitherapy and Bioactivity (APIBA), Food Science Department, Faculty of Pharmacy and Bioanalysis, Universidad de los Andes, Mérida 5101, Venezuela
2. Institute of Organic Chemistry with Centre of Phytochemistry, Bulgarian Academy of Sciences, 1113 Sofia Bulgaria.
3. Apitherapy and Bioactivity (APIBA), Research Institute, Faculty of Pharmacy and Bioanalysis, Universidad de los Andes, Mérida 5101, Venezuela
4. Instituto Federal de Educação, Ciencia e Technologia do Amazonas, Cacaochiera, São Gabriel da Cachoeira, 69750-000, Amazonas, Brazil.
5. Department of Crop Science and Beekeeping Technology, University of Dar es Salaam Dar es Salaam, Tanzania.
6. Beec, Business, 13 Walkerty War Road, Hampton, NSW, 2790, Australia.
7. CAS Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Kunming 650000, China
8. Smithsonian Tropical Research Institute, Balsa, Arnam, Republic of Panama
9. School of Ecology and Environment, Yunnan University, Kunming 650500, China
10. Central Bicol State University of Agriculture, Camarines Sur, Bicol, Philippines
11. Laboratory of Microbiology, Department of Microbiology and Parasitology, Faculty of Pharmacy and Bioanalysis, Universidad de Los Andes, Mérida 5101, Venezuela.
12. Institute for Integrative Genome Biology; Department of Microbiology and Plant Pathology, University of California Riverside, Riverside, CA 92521, United States of America.

[email protected]

 

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PUBLISHED: 30 November 2025

CITATION: Vit, P., Bankova, V., et al., 2025. Active metabolites of stingless bees nest materials used for meliponitherapy: Bibliometrics, impact of biodiversity in conservation, and emerging microbiome. Medical Research Archives, [online] 13(11).
https://doi.org/10.18103/mra.v13i11.7096

COPYRIGHT: © 2025 European Society of Medicine. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

DOI https://doi.org/10.18103/mra.v13i11.7096

ISSN 2375-1924

Abstract

Stingless bees (Hymenoptera: Apidae: Apinae: Meliponini) collect biotic and abiotic resources from nature to be transformed into nest materials with diverse functions such as structural, immune, defense, and nutritional. The 605 species of stingless bees collecting natural resources processed with associated microbiota are a spectacular biodiversity forming pot-honey, pot-pollen, cerumen and propolis valued in meliponitherapy. The bioactive metabolites have botanical, entomological, and microbial origins. Only pot-honey has been regulated since 2014 in Bahia, Brazil and further national standards. Forecasts of climate change affect stingless bee distribution, their productivity, and may influence the diversity of active metabolites in the nest. A sequence of researches serving meliponitherapy illustrated the ancient use of pot-honey eye drops to the latest cerumen components, reducing oxidative stress, and recent synergism with antibiotics to overcome antimicrobial resistance. Besides the chemical composition, the antioxidant and antimicrobial activities are fundamental added values, supporting a medicinal approach for both nutritional and pharmaceutical applications. Increased aliphatic organic acid contents in fermented pot-honey is not a defect, but a microbial biotransformation to preserve their wet honey with active metabolites. Characterizing the microbiome of stingless bees and their nest materials assists in identifying potential active biomolecules of microbial origin. Authenticity and chemical variability were discussed for quality control. Bibliometrics complemented this review for medicinal stingless bees (2004–2023) and stingless bees in climate change (2010–2023). Neotropical biodiversity of stingless bees was evidenced with the 259 stingless bee species richness in Brazil, 95 of them used in meliponiculture. Nest materials of 64 stingless bee taxa in 14 countries were reviewed for their flavonoid and polyphenol contents, and 13 biological activities. Conservation of stingless bees’ biodiversity has been addressed in the face of climate change and the chemical pool represented for meliponitherapy. Active metabolites from the stingless bee nest are not envisaged to be extracted but to be used in their original matrices: pot-honey, pot-pollen, cerumen, or propolis. A synthesis of most active metabolites could be an option for pharmaceutical developments to reproduce a bioactive chemical repertoire of stingless bees in nature, with a role on Sustainable Development Goals SDG2 food security and SDG3 good health and well-being.

Keywords: antimicrobial resistance, biological activity, cerumen, climate change, flavonoids and polyphenols, medicinal, meliponitherapy, microbiome, pot-honey, pot-pollen, propolis, stingless bee.

1. Introduction

As the largest Neotropical country, Brazil has been leading conservation policies to protect biodiversity, and meliponine conservation is one example of that. The first official standard for stingless bee honey was approved in the state of Bahia for Melipona honey. Promoting nutraceutical uses of stingless bee pot-honey, pot-pollen, cerumen, and propolis, considers rational exploitation protecting natural resources for a sustainable meliponine industry. The diversity of active metabolites for meliponitherapy rely on the bee flora; the bees harvesting and transporting resources to the nest; colony and microbial processing; and chemical transformations.

The global decrease in rainforests has adverse impact on biodiversity of stingless bees and plants used as nesting sites, nest materials, and food sources for their colonies. Particularly, Brazilian deforestation in the Neotropical region, and the greatest 2020 deforestation rate of the Brazilian Amazon in a decade need our attention. Rocha et al. (2020) analyzed the loss of functional diversity. Knowing the vital role of honeydew, nectar, pollen, oil, and resin use of plant resources by stingless bee colonies, any altered tree landscape would affect bee density and performance. Having diverse requirements and food preferences, different stingless bee species would adapt better than others to the deforestation stress in a dynamic forest community, causing shifts in abundance and diversity. Bees and forests have a vital synergism. Therefore, distinctive conservational objectives are linked to the utilization of stingless bee products. The Atlantic Forest is the second largest tropical rainforest in the American continent after the Amazon, home of stingless bees, their nest products, and communities exploiting these natural resources managed for meliponiculture.

Stingless bee keepers harvest pot-honey, pot-pollen, cerumen, and propolis from stingless bee nests for their unique sensory attributes and beneficial medicinal values. Stingless bee studies have become a hotspot in the international bee product research, with a progressive expansion to characterize their entomological biodiversity, and to propose therapeutic applications.

Main topics of the Apitherapy section in the Apimondia Congress held in Santiago de Chile, September 2023, were:

1. Scientific-based evidence supporting the nutritional, physiological, and health claims of bee products,
2. Preclinical research – safety, pharmacology, and toxicology of bee products. Guidelines for medical applications,
3. Clinical trials in apitherapy – doses, interactions, side effects (human and veterinary medicine).
4. Update on the use of apitherapy in infectious diseases, and
5. Regulatory issues and clinical ethics related to the integration of apitherapy as TCM in healthcare systems.

The first topic on bee products, particularly stingless bee nest materials, was reviewed because diverse nest materials have different chemical composition and added values on bioactive properties for pharmaceutical design. Their attributes caused by the botanical origin is an investigation initiated with Apis mellifera, and further variations caused by the entomological origin were mandatory for the meliponine biodiversity—Engel et al. (2023) recognize 605 stingless bee species—scientific attention was more recently addressed to the microbial origin of.

The support of traditional knowledge, such as to Scaptotrigona mexicana pot-honey by ethanolic fermentation was demonstrated with Tetragonisca angustula pot-honey in Venezuela, and recent studies exploring the meliponine rich microbiome producing pot-honey metabolites of microbial origin quantified by targeted 1H-NMR.

Another approach to differentiate nest materials is using non-invasive techniques to study volatile organic compounds (VOCs) with diverse chemical structures, ecological roles, and origins. For example, a diversity of 95 VOCs in Tetragonisca angustula cerumen types and propolis was detected, identified by HS-SPME/GC-MS, and grouped in chemical classes: 1. Acids (11), 2. Alcohols (16), 3. Aldehydes (7), 4. Esters (16), 5. Ketones (8), 6. Monoterpenes (17), 7. Oxides (5), 8. Sesquiterpenes (11), and 9. Others (4) by Betta et al. (2024). Their transformations are fascinating, waiting for suggested biochemical and microbial pathways. The acetic acid accumulated in the cerumen of empty honey pots, was esterified into methyl acetate in the entrance tube, and five acetates in the Tetragonisca angustula honey pots.

Microbial metabolites of stingless bee nest materials were reviewed by Vit (2024) for alcohols (ethanol, glycerol, isoamylic), aliphatic organic acids (AOA) (acetic, gluconic, lactic, oxalic, succinic, tartaric), amino acids (phenylalanine, proline, pyroglutamic acid), antibiotics (meliponamycin A, B, recently discovered as microbial metabolites in Melipona scutellaris), diphenylether (asterric acid), polyketide pigments (monascin), phenolic acids (3-phenyllactic acid), polyols (2,3-butanediol), statins (lovastatin), steroids (ergosterol), sugars (dihydroxyacetone, maltose, raffinose, trehalulose, turanose), surfactants (suspected sophorolipids), and vitamins (ascorbic acid). Surprisingly, a honey authenticity test revealed suspected microbial associations with the Scaptotrigona vitorum pot-honey from Ecuador.

These studies are beneficial for multifactorial medicinal stingless bee science and applications in integrative medicine as a good source of bioactive natural compounds with therapeutic and nutritional value. Salomon et al. observed significant reduction of cotton pellet-induced granuloma weights at all doses tested (27.34%, 35.53% and 47.53% granuloma inhibition) in Wistar rats treated daily with injected intravenously Tetragonisca fiebrigi honey (1000 mg/kg b.w.) for a week. In contrast, significant reduction in hind paw edema (44.44%) was achieved with Tetragonisca fiebrigi honey oral administration, causing analgesic responses in the three models used (acetic acid, formalin, tail immersion). Antioxidant activity, melissopalynological, physicochemical, phenolics, and sugars HPLC assessments of the honey were provided.

The tropical uses of pot-honey, pot-pollen, cerumen and propolis in meliponitherapy are traditional knowledge/constitute traditional knowledge gaining scientific interest for their biological activities and the most studied flavonoids and polyphenols active biomolecules. The scientific literature was tabulated for that approach, and additionally complemented with bibliometrics on medicinal stingless bees, and a further evaluation of stingless bees and climate change to understand the impact on biodiversity conservation. Protection of stingless bee and associated microbiome biodiversity is considered vital for chemical diversity of nest materials used in meliponitherapy.

This wide-ranging review integrates the chemical, biological, ecological, and therapeutic dimensions of stingless bee nest materials within the framework of meliponitherapy. It encompasses a detailed exploration of the diversity of active metabolites originating from botanical, entomological, and microbial sources found in pot-honey, pot-pollen, cerumen, and propolis of stingless bees. The review includes a bibliometric analysis of global research on medicinal stingless bees and their relationship with climate change, highlighting publication trends, collaborative networks, and research impact. It further synthesizes current evidence on the antioxidant, antimicrobial, and pharmacological activities of these bioactive compounds and their implications for nutraceutical and pharmaceutical development. It also assesses the importance of biodiversity conservation and related microbiomes, as well as their promising medical applications. The primary objective is to establish a science-based framework to promote the rational and sustainable use of these natural products for pharmaceutical and nutraceutical applications, considering the potential effects of environmental changes on their chemical composition and availability. Our aim is to provide a sound background on the science supporting meliponitherapy, to propel discoveries on microbial biomolecules as potential medicinal natural resources for health, alone or combined with drugs.

2. Bibliometrics

Science mapping from the citation network and a tool to visualize bibliometric networks such as Bibliometrix are needed to describe structures of research. This section has two bibliometric reviews on stingless bees, one on their medicinal uses (107 documents) in the period 2004 to 2023, and another on climate change (25 documents) in the period 2010 to 2023. The annual growth rate of both datasets is compared in Figure 2. There is a tendency to increase the number of documents with time. The annual growth of medicinal stingless bees recorded with the first paper in 2004, escalated in 2012 and 2013, with a sudden drop followed by a second peak in 2018, and a further drop with a steady growth in the last three years up to 15 documents in 2023. The annual growth of stingless bees and climate change shows the first paper in 2010, the second in 2015, a yearly paper from 2017 to 2019, growing interest reaching a peak of 7 papers in 2022. However, production decreased to 3 documents in 2023, that may recover in two weeks before the end of the year.

2.1 METHODOLOGY AND RESULTS OF THE BIBLIOMETRIC REVIEW ON MEDICINAL STINGLESS BEES

The Scopus database was used to review the scientific literature on medicinal stingless bees. Other words were used in the query string but the number of retrieved documents was lower; for example, using apitherapy only 16 documents, and 37 documents using pharmaceutical. Therefore, we selected medicinal for the search done on the 1st December 2023.

TITLE-ABS-KEY ( stingless bee AND medicinal )

The main information of bibliometric descriptors is presented in Table 1 including publications from the first retrieved document since 2004 to 2023, almost two decades. The 107 documents of the dataset were published in four languages: English (100), Portuguese (3), Spanish (3), and Chinese (2). Note that the addition gives 108 documents, possibly one was bilingual or considered bilingual by the dataset.

Main information of the dataset bibliometric descriptors	Counts of all documents
Timespan	2004:2023
Sources (Journals, Books, etc)	80
Documents	107
Annual Growth Rate %	15.32
Document Average Age	4.84
Average Citations per Document	18.63
References	5419
Document Contents
Keywords Plus (ID)	869
Author’s Keywords (DE)	327
Authors	489
Authors of Single-Authored Documents	6
Authors Collaboration
Single-Authored Documents	6
Multi-Authored Documents	101
Co-Authors per Document	4.97
International Co-Authorships %	24.3
Document Types
Article	73
Book	1
Book chapter	12
Conference paper	4
Editorial	1
Note	1
Review	15
No. of languages	4
English	100
Portuguese	3
Spanish	3
Chinese	2

The first document of this dataset was Quality standards for medicinal uses of Meliponinae honey in Guatemala, Mexico and Venezuela, which was cited 132 times since 2004. The International Honey Commission was not supporting the proposal of medicinal stingless bee honey, but honey eye drops were sold in pharmacies before we started our research, and more recently online pharmacies offer the familiar eye drops, pot-honey, supplements with pot-honey, pot-pollen, and stingless bee propolis. The Vit et al. article was published the same year of the Apidologie special number for European honey, with the seminal article Main European unifloral honeys: Descriptive sheets by Livia Persano-Oddo and Roberto Piro on more than 35,000 unifloral, multifloral and honeydew honeys. The next document on this topic had to wait until 2008, Composition and antioxidant activity of Tetragonula carbonaria honey from Australia, cited as Trigona carbonaria at that time. Medicinal animals as therapeutic alternative in a semi-arid region of Northeastern Brazil, Properties of honey from Tetragonisca angustula fiebrigi and Plebeia wittmanni of Argentina and Antimicrobial activity of honey from the stingless bee Tetragonula carbonaria determined by agar diffusion, agar dilution, broth microdilution and time-kill methodology. The annual growth of medicinal research of stingless bees is plotted in Figure 2. The topic took flight in 2012 with four documents and a prolific peak in 2013 with the book Pot-honey. A legacy of stingless bees, and a further peak of productivity with the second book Pot-pollen in stingless bee melittology. A drop of productivity was observed, the following year of each book publication, and a possible effect of the COVID-19 pandemic in 2020. A spectacular recovery in 2021 shows the inner motivation of authors, with a steady growth in 2022 and 2023, a merit of the multidisciplinary teams of worldwide experts with scientific interest embracing medicinal research of tropical stingless bees.

2.1.1 Most productive authors

The top ten authors in Table 2 are from Argentina, Australia, Brazil, Malaysia, and Venezuela. Hilgert publishes on ethnomedicinal uses of stingless bees, Brooks is microbiologist of cerumen and propolis, Mustafa and Ahmad are veterinarians with papers in multiple subjects, and Vit is a biologist interested in quality control of pot-honey, pot-pollen, and propolis.

Ranking	NP	Stingless bee medicinal research Author	Affiliation, city	Country
1	4	Hilgert, N.I.	Instituto de Biología Subtropical, CONICET, Facultad de Ciencias Forestales, Universidad Nacional de Misiones, Puerto Iguazú	Argentina
2	3	Brooks, P.R.	Faculty of Science, Health, Education and Engineering, University of the Sunshine Coast, Maroochydore	Australia
3	3	Mustafa, M.Z.	Hospital Universiti Sains Malaysia, Kubang Kerian 16150, Kelantan	Malaysia
4	3	Vit, P.	Food Science Department, Faculty of Pharmacy and Bioanalysis, Universidad de Los Andes, Mérida	Venezuela
5	2	Ahmad, H.	Department of Veterinary Preclinical Sciences, Faculty of Veterinary Medicine, Universiti Putra Malaysia, Serdang	Malaysia
6	2	Al Hatamleh, M.A.I.	Department of Immunology, School of Medical Sciences, Universiti Sains Malaysia, Kubang Kerian	Malaysia
7	2	Balestieri, J.P.B.	Research group on Biotechnology and Bioprospecting Applied to Metabolism (GEBBAM), Federal University of Grande Dourados, Dourados	Brazil
8	2	Beux, M.R.	Department of Food Engineering, Federal University of Parana (UFPR), Curitiba	Brazil
9	2	Campos, J.F.	Research Group on Biotechnology and Bioprospecting Applied to Metabolism (GEBBAM), Federal University of Grande Dourados, Dourados	Brazil
10	2	Carollo, C.A.	Laboratory of Natural Products and Mass Spectrometry, Federal University of Mato Grosso do Sul, Cidade Universitária, Campo Grande	Brazil

2.1.2 Impact factor of top ten authors of stingless bee medicinal documents (2004–2023)

Top ten authors	h-index	g-index	m-index	TC	NP	PY start
Mustafa MZ	4	4	0.667	134	4	2018
Brooks PR	3	3	0.231	85	3	2011
Vit P	3	3	0.150	236	3	2004
Ahmad H	2	2	0.667	31	2	2021
Al-Hatamleh MAI	2	2	0.500	73	2	2020
Balestieri JBP	2	2	0.286	57	2	2017
Campos JF	2	2	0.286	57	2	2017
Carollo CA	2	2	0.286	57	2	2017
Dos Santos CM	2	2	0.286	57	2	2017
Dos Santos EL	2	2	0.286	57	2	2017

The most productive institutions worldwide in Table 4 show the Universiti Putra Malaysia in the top position with 7 publications, Instituto Tecnológico de Mérida with 6, Universidad de Los Andes, School of Medical Science, Universiti Sains Malaysia, and Universidad Nacional de Misiones with 5, as the top five institutions.

Ranking	NP	Stingless bee medicinal research	Institution	Country
1	7	Universiti Putra Malaysia	Malaysia
2	6	Instituto Tecnológico de Mérida	Mexico
3	5	Universidad de Los Andes	Venezuela
4	5	School of Medical Science, Universiti Sains Malaysia	Malaysia
5	5	Universidad Nacional de Misiones	Argentina
6	5	Universiti Sains Malaysia	Malaysia
7	5	Universiti Sultan Zainal Abidin	Malaysia
8	3	Universiti Kebangsaan Malaysia	Malaysia
9	3	Universiti Teknologi Malaysia	Malaysia
10	3	Universidad Nacional Autónoma de México	Mexico

The top ten countries engaging with stingless bee medicinal research have 4 to 29 publications in the period (2004-2023), with Malaysia at the top (29), Brazil (19), Mexico (14), Australia and India (7), as the top five.

Ranking	NP	Stingless bee medicinal research	Country
1	29	Malaysia	Malaysia
2	19	Brazil	Brazil
3	14	Mexico	Mexico
4	7	Australia	Australia
5	7	India	India
6	6	Argentina	Argentina
7	5	Venezuela	Venezuela
8	4	Indonesia	Indonesia
9	4	Kenya	Kenya
10	4	United Kingdom	United Kingdom

The most globally cited countries in the Bibliometrix plot were Brazil (480), Malaysia (403), Venezuela (236), Australia (195), India (113), Ethiopia (59), Mexico (51), Bulgaria (50), Argentina (47), and Indonesia (38).

In Table 6, the ranking of stingless bee medicinal research top ten sources selected by authors, hosted from 2 to 6 documents each. The most productive sources were the book Pot-Honey. A Legacy of Stingless Bees, followed by the Journal of Apicultural Research, the book Stingless Bee’s Honey from Yucatan: Culture, Traditional Uses and Nutraceutical Potential, Journal of Ethnobiology, and Ethnomedicine, and Estudios de Cultura Maya, as the top five sources. Journals h-index varied between 8 and 231, 2/8 journals are Quartile 1. The maximum impact score was 5.57 for the International Journal of Molecular Sciences ranked in the 10th position; note that alphabetical order is applied for the same number of publications. The metrics used for the journals are not available for the books in Resurchify, but the h-index was used as an impact factor in the following Bibliometrix plot.

Ranking	NP	Stingless bee medicinal research	Sources (h-index, Quartile, impact score)	Publisher, country
1	6	Pot-Honey: A Legacy of Stingless Bees	Springer, New York, United States
2	5	Journal of Apicultural Research	(h66, Q2, 2.08) Taylor and Francis Ltd., United Kingdom
3	4	Stingless Bee’s Honey From Yucatan: Culture, Traditional Uses AND Nutraceutical Potential
4	3	Journal of Ethnobiology AND Ethnomedicine	(h 84, Q1, 4.27) BioMed Central Ltd., UK
5	2	Estudios DE Cultura Maya	(h 8, Q2, 0.20) UNAM, Instituto de Investigaciones Filologicas, Mexico
6	2	Ethnobiology AND Conservation	(h 18, Q2, 1.54) Universidade Federal Rural de Pernambuco, Brazil
7	2	Food Research	(h15, Q3, 1.03) Malaysia
8	2	Indian Journal OF Traditional Knowledge	(h40, Q2, 0.92) National Institute of Science Communication and Information Resources (NISCAIR), India
9	2	Interciencia	(h 39, Q3, 0.40) Interciencia Association, Venezuela
10	2	International Journal OF Molecular Sciences	(h 230, Q1, 5.57) Multidisciplinary Digital Publishing Institute (MDPI), Switzerland

In the Bibliometrix plot, the local impact of medicinal stingless bee sources by h-index shows the highest impact (6) for the book Pot-Honey. A Legacy of Stingless Bees detached from the nine journal’s impacts (2) of the dataset. Table 6 corresponds with the most relevant sources Bibliometrix plot, not shown here. Note that the top ten Scopus-ranked sources by number of publications in Table 6 differ from the Bibliometrix-ranked sources by h-index in Figure 4.

Bibliometrix produced the most locally cited sources of top ten authors and number of articles. In Table 7, authors are considered sources. Local citations measure how many times an author (or a document) included in the dataset collection have been also cited by other authors in the dataset collection. Further Scopus search was done for each author to know the topic (stingless bee AND medicinal), total number of publication (NP), and total number of citations (NC). Six of these ten authors have zero topical publications, and therefore zero citation in this topic. The number of articles cited by other authors of the dataset (local collection) were lower than the topical citations in the Scopus database for Vit (89/234), Bankova (25/51), and Alves (19/21). Biluca had more citations from other authors in the dataset, than in the Scopus database (32/12), as well as other authors with zero topical citations.

Sources	Document cited by another document of the dataset	NP	Scopus database (topical/total)	NC	Scopus database (topical/total)
Vit P	89	3/66	234/1 698
Biluca FC	32	1/26	12/741
Bogdanov S	26	0/43	0/5 518
Michener CD	26	0/86	0/1 975
Bankova V	25	1/204	51/12
Roubik DW	24	0/138	0/6 297
Crane E	20	0/34	0/326
Alves RRN	19	1/266	21/7 917
Kek S P	17	0/6	0/365
Cortopassi-Laurino M	16	0/4	0/250

In Table 8, the funding sponsors for the stingless bee medicinal research are from Argentina (1), Australia (1), Austria (1), Brazil (2), Bulgaria (1), El Salvador (1), Malaysia (3). Up to 8 projects received support from two Brazilian funding agencies, from two Malaysian Ministry (7) and University (4), and the National Council of Science and Technology from El Salvador (3), as the top five funding sponsors.

Ranking	NP	Stingless bee medicinal research	Funding sponsor	Country
1	8	Conselho Nacional de Desenvolvimento Científico e Tecnológico	Brazil
2	8	Coordenação de Aperfeiçoamento de Pessoal de Nível Superior	Brazil
3	7	Ministry of Higher Education, Malaysia	Malaysia
4	4	Universiti Kebangsaan Malaysia	Malaysia
5	3	Consejo Nacional de Ciencia y Tecnología	El Salvador
6	3	Consejo Nacional de Investigaciones Científicas y Técnicas	Argentina
7	3	Universiti Sains Malaysia	Malaysia
8	1	Australia and New Zealand Banking Group Limited	Australia
9	1	Austrian Development Agency	Austria
10	1	Bulgarian Academy of Science	Bulgarian

The Scopus database covers 4 broad supergroup areas (health sciences, life sciences, physical sciences, and social sciences) categorized into 27 subject areas that are automatically scrutinized in the left side menu by number of publications, and plotted as percentages of documents of a pie chart in the analyzed results report. In Table 9, the top ten studied subject areas on stingless bee medicinal research (2004–2023) had the following top five: Agricultural and Biological Sciences (30.6%), Biochemistry, Genetics and Molecular Biology (10.9%), Medicine (10.4%), Environmental Science (7.8%), and Social Sciences (5.7%).

Ranking	NP	%	Stingless bee medicinal research	Subject area
1	59	30.6	Agricultural and Biological Sciences
2	21	10.9	Biochemistry, Genetics and Molecular Biology
3	20	10.4	Medicine
4	15	7.8	Environmental Science
5	11	5.7	Social Sciences
6	10	5.2	Chemistry
7	10	5.2	Pharmacology, Toxicology and Pharmaceutics
8	8	4.1	Chemical Engineering
9	5	2.6	Earth and Planetary Sciences
10	5	2.6	Engineering

2.1.2.1 Author’s keywords

Compared with a word cloud, the tree map is structured in fields with visualized descending order of frequent keywords, both representations use bright colors. The frequencies of author’s keywords, and their percentages in the Figure 5 tree map are visible: stingless bee (21, 11%), honey (17, 9%), stingless bees (17, 9%), propolis (14, 7%), meliponini (7, 4%), antioxidant (6, 3%), antioxidant activity (6, 3%), chemical composition (6, 3%), meliponiculture (6, 3%), antibacterial (5, 3%), antimicrobial activity (5, 3%), stingless bee honey (5, 3%), natural products.

2.1.2.2 Country collaborative map

The collaboration between countries sharing publications on medicinal stingless bee research in the period 2004 to 2023 was visualized in Figure 6 using red connectors in a worldwide map. The frequencies of collaboration between two countries are available in the corresponding Excel file. The highest collaborative frequency was between Brazil and Portugal with 3 documents, and for Jordan and Malaysia with two documents.

2.1.2.3 Most globally cited documents

The plot of most globally cited documents in Figure 7, shows the top ten documents cited from 147 to 50 times in publications of medicinal stingless bees (2004–2023), silva iaai 2013 food chem (147 citations), vit p 2004 bee world (132), boorn kl 2010 j appl microbiol (110), oddo lp 2008 j med food (101), avila s 2018 trends food sci technol (89), zulkhairi amin fa 2018 adv pharmacol sci (82), choudhari mk evid-based complement altern med 2013 (74), al-hatamaleh mai 2020 biomolecules (64), popova m 2021 phytomedicine (50), and massaro cf 2011 naturwissenschaften (50). The journal types were on food (2), medicine (2), a combined medicinal food (1), bees (1), biomolecules (1), microbiology (1), natural science (1), and pharmacy (1). All these documents were distributed in two clusters of the conceptual structure by factorial analysis in the next plot.

2.1.2.4 The conceptual structure for most cited articles

Concepts are embedded in a network of associations and contexts, having partial meaning based on links formed between them. Bibliometrix uses factorial analysis of correspondence analysis (CA) as a graphical method to compare variables. Scientific researchers use a conceptual framework to understand a problem and develop the analytical approach, a roadmap to conceptualize an outline that connects different ideas, concepts, and theories within a scientific field.

In this plot of research impact, the most cited medicinal stingless bee documents of the dataset (2004–2023) plotted in Figure 7 generated two clusters like principal component analysis (PCA) in a CA factorial map, clustering bipartite network of terms extracted from closeness of keyword, title or abstract fields. Factorial analysis is a data reduction technique. CA is used to represent the rows and columns of a two-way dimensional space, with Dim 1 explaining 61.21% of the variations, and Dim 2 explaining 15.81%. A red cluster 1 for boorn kl 2010, choudhari mk 2013, zulkhairi amin fa 2018, and ualem b 2013, and vit p 2004 was separated by the second dimension in the upper quadrants except vit p 2004. A blue cluster 2 for popova m 2021, silva iaai 2013, al-hatamleh mai 2020, oddo lp 2008, and massaro cf 2011. This cluster was separated by Dim 1 in the right quadrant except massaro cf 2011 located to the left, all of these documents were separated by Dim 2 in the lower quadrants.

2.1.2.5 Simple and multiple country publications

Publications based on two categories of corresponding author simple country publication (SCP) and multiple country publication (MCP), are represented with bars of two colors for 19 countries in Figure 9 Bibliometrix plot. Argentina, India, China, Ethiopia, Brunei, Bulgaria, and Colombia publications of the dataset were intra-country SCP. Malaysia (24) and Brazil (19) published major number of documents with most SCP. Australia and Mexico also had more SCP than MCP, and Indonesia was balanced SCP-MCP in four documents. Kenya and Venezuela had more MCP than SCP in three documents. Austria, Bolivia, Japan, Jordan, and Nigeria had one document MCP, indicating compulsory multiple country interaction.

2.2 METHODOLOGY AND RESULTS OF THE BIBLIOMETRIC REVIEW ON STINGLESS BEES IN CLIMATE CHANGE

A second bibliometric review focused for a search of stingless bees in climate change. After reviewing the retrieved documents, we found that three of them only used climate change in the abstract, in general sentences such as: “Ecosystem services provided by such communities may be more greatly affected by environmental changes (anthropogenic activities and climate change) than are services provided by communities with greater functional redundancy”; “Used in folk medicine as antiseptic, antioxidant and antimicrobial agent, the composition is due to bee species, climate changes, local flora, and soil type”; and “Basic ecological knowledge is essential to inform agricultural management policies and to foresee preventable food scarcity problems, especially in view of climate change scenarios that predict drastic alterations in plant geographical distributions”. The dataset was retrieved with the Scopus database in the “TITLE-ABS-KEY” field query string the 1st December 2023. The operator AND was used for stingless bee AND climate change, with the operator AND NOT for guild, chayote and geopropolis, as follows.

TITLE-ABS-KEY ( stingless bee AND climate change AND NOT guild AND NOT chayote AND NOT geopropolis )

The first and most cited document of stingless bees and climate change was by Batalha-Filho et al., it was used to explain the Pleistocene estimated divergence time of Melipona quadrifasciata subspecies in Brazil, in a period of climatic and geomorphological changes in the Neotropics, causing subspecies distribution of Melipona quadrifasciata quadrifasciata to the south, and Melipona quadrifasciata anthidioides to the north. The second document was on the same species of stingless bee, a key pollinator of the Atlantic flora in Brazil, and approach to mitigate the effect of climate change in habitat fragmentation by identifying key conservation areas and strategies after a forecast to year 2080.

The main information of bibliometric descriptors is presented in Figure 11.1 including publications from the first retrieved document since 2010 to date in 2023, almost two decades. The 25 documents of the dataset were published in English. Three documents were excluded with the AND NOT operator because although climate change was in the abstracts, the investigations were not about climate change.

A total of 19 sources were used to disseminate the research on stingless bees and climate change. The metrics and publishers of the most prolific journals were Apidologie (5 documents, h-index 96, Quartile 1, impact factor 2.41, Springer Science + Business Media, United States), PLoS ONE (2 documents, h-index 404, Quartile 1, impact factor 3.75, Public Library of Science, United States), and Regional Environmental Change (2 documents, h-index 82, Quartile 2, impact factor 4.30, Springer Verlag, Germany).

The types of retrieved documents were 15 articles, one book chapter, one conference papers, and 4 reviews, all published in English. The first and most cited document was published by Batalha-Filho et al. in the journal Apidologie. Leading countries were Brazil (12), United States (4), Indonesia (3); Australia, Colombia, Kenya, and Thailand with 3 documents each, and one document for Argentina, Botswana, and China. The top seven authors published three documents each, and two the last three authors. Universidade de São Paulo led with 5 publications, five institutions with three, and two publications for the last in the top ten list. The top five Scopus subject areas of research on stingless bees and climate change were Agricultural and Biological Sciences (44.4% of the documents), Environmental Sciences (19.4%), Biochemistry, Genetics and Molecular Biology (13.9%), Multidisciplinary (8.3%), Earth and Planetary Sciences (5.6%), followed by Chemistry (2.8%), Computer Science (2.8%), and Engineering (2.8%).

Research for publications was sponsored by ten top agencies: Coordenação de Aperfeiçoamento de Pessoal de Nível Superior CAPES (Brazil), Conselho Nacional de Desenvolvimento Científico e Tecnológico CNPq (Brazil), Australian Research Council (Australia), Fundacao de Amparo a Pesquisa do Estado de Sao Paulo FAPESP (Brazil), National Science Foundation (United States), University of Kansas (United States), Bayer CropScience (Germany), Biotechnology and Biological Sciences Research Council (United Kingdom), Department of Atomic Energy, Government of India (India), and Direktion fur Entwicklung und Zusammenarbeit (Switzerland).

Correlations and classifications of authors’ keywords were investigated with multivariate graphical tools. Metrics of scientific literature and other Bibliometrix plots, such as topical dendrograms by Hierarchical Cluster Analysis (HCA), word clouds, country collaboration maps, most global cited documents, co-authors networks, and corresponding authors country publications were illustrated in Figure 11.

The metrics for 25 documents in the scientific literature of stingless bees and climate change (2010:2023) cited 1863 references, showed an average citation per document of 13.4%, used 89 author’s keywords, had an average author number of 6.52, and had 32.00% international co-authorship. Two clusters were produced in the topical dendrogram by HCA after factorial analysis of author’s keywords. The red cluster grouped traditional knowledge, traditional and recent beekeeping, propolis, medicinal honey, bumble bees, honey hunters, honeybees, pests, and social bees. The large blue cluster was visualized with two smaller clusters. The left blue cluster was mainly based on thermal biology, tetragonula, critical thermal maxima, austroplebeia, chill coma, from 30 and further branches with keywords from other papers; for example, single nucleotide polymorphism, local adaptation, isolation by resistance; and environmental associations, gene flow, species distribution modelling, hybridization, climate niche, habitat suitability.

The country collaborative map showed red line interactions with at least two papers shared between Colombia and the United States. A word cloud plot visualized higher frequencies of author’s keywords with larger letters in central positions, stingless bees (7) was the most frequent, then, climate change (6), pollination (4), honey bees (3), stingless bee (3), sustainability (3), conservation (2), deforestation (2), ecological niche modeling (2), meliponiculture (2), meliponini (2), pests (2), melipona (2), social bees (2), and one for the remaining 35 keywords of the plot, anthropogenic change, biodiversity loss and biotic interaction among others. The most global cited documents were ten documents cited from 77 times (batalha-filho 2010) to 14 (gonzalez2021), in five different journals Apidologie, Food Chemistry, Evol Appl, Perspect Ecol Conserv, PLoS ONE, and one book chapter Asian Beekeeping in the 21st Century.

3. Scanning medicinal stingless bee resources, research, and efforts for wellness

3.1 SIGNIFICANCE OF THE ENTOMOLOGICAL BIODIVERSITY OF STINGLESS BEES IN APITHERAPY

3.1.1 Richness of stingless bees in Brazil as resources for meliponiculture

The conservation of stingless bee biodiversity is of paramount importance for this natural resource with medicinal traditional uses. Brazil has a great wealth of stingless bees, which makes this country stand out from the rest of the Neotropical region, since the last survey carried out by Nogueira, counted a total of 259 species, distributed in the five regions of this country. The Amazon region with 197 of these species, 128 of which occur in the Amazonas state. Although there are many species in Brazil, the most familiar for management are used for honey production, the vast majority belonging to the genus Melipona Illiger, as M. seminigra, M. interrupta, M. flavolineata, M. subnitida, M. scutellaris and M. quadrifasciata. Species from other genera that also deserve to be highlighted in terms of productivity for Brazil are Tetragonisca angustula, Tetragona clavipes, and species of Scaptotrigona.

In a proportion of 95 managed stingless bees over the total of 259 species of stingless bees in Brazil, around half of these species are used for meliponiculture in the Northeast and Central-West regions, and a little more than half in the Southeast and South regions. On the contrary, the richest North region uses a third of these resources, with the remaining 137 species of stingless bees unexploited or not selected for meliponiculture, or which do not have data about nesting.

3.1.2 Managed stingless bee species in Brazil

Good practices of sustainable stingless bee keeping do not represent a risk for the stingless bee biodiversity conservation, on the contrary, more colonies are kept, and divided to increase productivity. However, feral nest hunting is a common practice in rural areas; moving colonies from their natural substrates to stingless bee hives. On the other hand, pot-honey hunting is more destructive, its intensity may affect the natural populations of stingless bees, but is the traditional method especially for non-domesticated underground species. For example, developing conventional stingless bee farming is an opportunity to increase availability of pot-honey in demand for ethnomedicinal use in Baringo County, Kenya. Some Brazilian stingless bee species used in meliponiculture are illustrated with the images and size of the Brazilian Association of Bee Studies.

No.	Stingless bee species	Brazilian states
1	Trigonisca pediculana	CE, MA, PB, PA, BA, PI, AM, RO, RR, PE, MG
2	Trigonisca duckei	AM, PA, CE, MA, MT, RR
3	Trigona pallens	AC, AM, AP, PA, RO, RR, TO, MA, GO, CE, DF, MT, PI
4	Trigona cilipes	AC, AM, AP, PA, RO, MT, RR, MA, GO
5	Tetragonisca weyrauchi	AC, RO, MT
6	Tetragonisca fiebrigi	MS, RS, SP, PR, SC, MT
7	Tetragonisca angustula	AM, AP, PA, RR, BA, CE, MA, PB, PE, GO, MS, MT, ES, MG, RJ, SP, PR, RS, SC, TO, DF, AC, PI, RO
8	Tetragona quadrangula	GO, MA, MG, MT, PA, TO
9	Tetragona kaieteurensis	AM, PA, RR
10	Tetragona goettei	AC, AM, PA, MT, RO, MA, RR
11	Tetragona essequiboensis	AM, RO
12	Tetragona clavipes (syn. T. elongata)	AC, AM, RO, AP, RR, PA, MA, PI, MT, TO, BA, GO, DF, MG, MS, ES, RJ, SP, PR, SC, RS
13	Schwarziana quadripunctata	BA, GO, ES, MG, RJ, SP, PR, RS, SC, DF
14	Scaura longula	AM, AP, PA, MA, GO, MT, MG, SP, AC, BA, MS, RO, RR
15	Scaura latitarsis (syn. Scaura tenuis)	AC, AM, MT, PA, RO, RR
16	Scaptotrigona xanthotricha	BA, ES, MG, SP, SC, PR, RJ
17	Scaptotrigona tubiba	SP, MG
18	Scaptotrigona tricolorata	RO, MT, AM
19	Scaptotrigona postica	PA
20	Scaptotrigona polysticta	AC, PA, RO, TO, MA, GO, MT, MG, SP, AM, DF, PI
21	Scaptotrigona depilis	MS, MG, SP,

92 — Friesella schrottkyi — ES, MG, SP, PR
93 — Duckeola ghilianii — AM, AP, PA, MT, RO
94 — Cephalotrigona femorata — AM, PA, RO, MA, AC, AP, MT, TO
95 — Cephalotrigona capitata — AP, PA, CE, MT, ES, MG, SP, PR, SC, BA, RJ, MS, AL, GO, RO

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¹North region: Acre (AC), Amazonas (AM), Amapá (AP), Pará (PA), Rondônia (RO), Roraima (RR) and Tocantins (TO). Midwest region: Distrito Federal (DF), Goiás (GO), Mato Grosso (MT) and Mato Grosso do Sul (MS). Southeast region: Espírito Santo (ES), Minas Gerais (MG), Rio de Janeiro (RJ) and São Paulo (SP). Northeast region: Alagoas (AL), Bahia (BA), Ceará (CE), Maranhão (MA), Paraíba (PB), Pernambuco (PE), Piauí (PI), Rio Grande do Norte (RN) and Sergipe (SE). South region: Paraná (PR), Santa Catarina (SC) and Rio Grande do Sul (RS).
Source: BRASIL¹

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Some stingless bee species like Scaptotrigona postica (PA), Plebeia wittmanni (RS), Melipona tumupasae (AC), Melipona favosa (RR), Melipona cramptoni (RR), and Melipona capixaba (ES) were managed only in one state, in contrast with Tetragonisca angustula and Tetragona clavipes widely selected for meliponiculture in 24 and 21 of the 26 Brazilian states respectively.

The map of managed stingless bees was used as a geospatial product and a basis for assessing the risk of extinction of Brazilian fauna species². It was prepared through long discussions and by expert teams, and although there is no consensus on some taxonomic identifications and occurrence records, see the occurrence of the same species in Nogueira³¹; it shows promise in trying to organize the biodiversity to prevent the illegal transport of species outside the political boundaries of their occurrence, as is the case in states (Table 15). The illegal transport of nests to places where they do not naturally occur can cause a series of problems such as the transmission of diseases, increased competition for resources with native species, and genetic modification of wild and managed populations, which can compromise the permanence and maintenance of both native species of this new location, as well as species that came from a different location³⁴. In the long term, these environmental imbalances may harm both bee biodiversity and local stingless beekeepers, as species extinction may occur, especially when there is hybridization between species due to the disturbance of ecological barriers promoted by human action³⁵,³⁶. The dangers of interbreeding are well-known threats to wildlife³⁷, both for bees and extinction of rare plant species³⁶.

Direct and online interviews were focused on bees and beekeepers of 25 indigenous provinces who understand the importance of traditional stingless bee keeping in and cultural practices. Data of 272 beekeepers revealed 19 species of stingless bees are reared, mostly Tetragonula laeviceps, and climate change was one of the obstacles besides pesticides, demanding a strategy for stingless bee keeping and bee conservation to adapt and mitigate environmental changes on climate and land-use³⁸.

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2.3 FLORAL AND EXTRAFLORAL NECTAR, FLORAL POLLEN, RESIN, GUM AND LATEX PLANT NATURAL RESOURCES

Tropical stingless bees nest, feed and interact with tropical plants. Tropical bee flora is represented by biodiverse Fabaceae, Asteraceae, Rubiaceae, Malvaceae, Lamiaceae, Euphorbiaceae, Arecaceae, Poaceae, Apocynaceae, and Melastomataceae as the most visited of 221 plant families³⁹. Stingless bee preferences of available tropical resources, pollen create enormous combinations of pot-honey, pot-pollen, cerumen and propolis variables, explained by natural history⁴⁰ and investigated as a healing matrix. For example, secondary metabolites like flavonoids originate from the foraged plants, and having luteolin derivatives as active phytochemicals in ocular cataract models⁴¹ have a significance for the nature of the stingless bee material, the biomolecular richness, the relationship with the environmental resources, and its biodiverse conservation.

Standard terminology for palynology is used for morphological descriptions of pollen grains⁴² and the major pollen grains in the pollen spectrum⁴³. Pollen identifications at plant family, genus and even species, are assigned after comparisons with pollen atlases and pollen reference collections. The taxonomic status of botanical taxa is systematically updated to avoid synonyms.Garden database available online⁴⁴. Extrafloral nectar causes poor pollen spectra of honey because it contains less pollen than floral nectar.

Plant resins sometimes comprise gums, latex and resin exudates from different parts of plants: bark like Dalbergia ecastaphyllum Fabaceae, and Schinus terebinthifolius Anacardiaceae; buds like Populus spp. Salicaceae; flowers like Clusia major and Clusia minor, Clusiaceae, and Dalechampia spp. Euphorbiaceae; fruits like Corymbia torelliana Myrtaceae, and Coussapoa asperifolia Cecropiaceae; and whole plant like Artocarpus heterophyllus Moraceae, Merremia umbellata Convolvulaceae. Anacardiaceae and Fabaceae are two plant resin source families for stingless bees in Brazil, China, Colombia, and India; Euphorbiaceae is common in Brazil, Colombia, and India; and Clusiaceae is a Neotropical source in Brazil, Colombia, and Venezuela⁴⁵.

Plant resins use has an evolutionary meaning of sociality in stingless bees⁴⁶. Diverse plant resin–based functions such as social immunity, cuticular hydrocarbon chemical profiles, defense, and microbial communities are associated with stingless bees⁴⁰,⁴⁶. Cerumen is a vital material in stingless bee nest architecture, composed by admixtures of beeswax and plant resins. For this reason, stingless bee foragers prioritize resin collection and reduce pollen foraging after hive splitting, as observed for the Australian Tetragonula carbonaria⁴⁷.

In Fig. 14, acetolyzed pollen grains used to identify the Coffea arabica unifloral Tetragonisca angustula honey from Costa Rica are illustrated⁴⁸. Taxa were identified as nectariferous sources, and polleniferous, considered contaminants of honey because they do not secrete nectar, the raw material transformed into honey.

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Fig. 14 Pollen grains of a unifloral pot-honey with nectariferous and polleniferous taxa. Nectariferous 1. Coffea arabica, Rubiaceae produces nectar. A frequency of 54.3% of the pollen spectrum featured this as unifloral coffee honey⁴⁸. Polleniferous 2. Polyads of Inga sp. Fabaceae–Caesalpinioideae, 3. Mimosa sp., Fabaceae–Caesalpinioideae, and single grains or monads of 4. Pinus sp. Pinaceae, 5. Paullinia sp. Sapindaceae, and 6. Miconia sp. Melastomataceae. Pollen from nectarless plants is considered contaminant pollen in melissopalynology. Photos: © E. Moreno After: Moreno et al.⁴⁸.

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A unifloral honey has 45% pollen of one taxa, with exceptions for over-represented and under-represented pollen types⁴⁹. Floral pollen, Apis mellifera bee-bread and stingless bee pot-pollen are obviously pollen grains. A recent controversy has raised for a traditional palynological analysis of propolis, the pollen landed on plant resins, latex or gums, collected and processed into propolis or bee glue. Layek et al.⁵⁰ found that the pollen spectra of Tetragonula iridipennis from India was not accurate to identify the floral and non-floral sources of cerumen and propolis, because this nest material is not seasonal like floral nectar and floral pollen when in bloom.

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3.3 DEMONSTRATED BIOMOLECULES, BIOLOGICAL ACTIVITIES, AND PUTATIVE THERAPEUTIC PROPERTIES OF POT-HONEY, POT-POLLEN, CERUMEN AND PROPOLIS OF THE STINGLESS BEE NEST

Flavonoids and polyphenols — and biological activities — antimicrobial and antioxidant — added values to the medicinal uses of stingless bee products, expanding with the putative therapeutic actions that would deserve more bioassays and clinical trials to support apitherapy. The stingless bee species were carefully retrieved for each study. It is recommended to inform the species in the abstracts. The corresponding entomological authority, institution, and collection where the entomological specimens are deposited, is mandatory in the materials and methods. Continuous updating of names arises with research, and this fact also deserves the attention of melittologists. Valid names should be informed as suggested by M. Engel (P. Vit, personal communication). For example, Axestotrigona ferruginea (cited as Meliponula ferruginea by Popova et al.⁵¹)

It is not our aim to provide ranges of concentrations and IC₅₀ for this table, but to summarize the chronological input for studies on biomolecules and biological activities, projecting therapeutic properties, showing the first and the last publications in each category. Timespan years and (number of publications/number of nest materials) varied as follows for the alphabetical order used in Table 11:
Biomolecules Flavonoids 1993–2023 (16/2); Polyphenols 1993–2023 (16/1);
Biological activities Anti-atherogenic 2019–2022 (3/2); Anticancer 2013–2020 (9/4); Anticataract 1997–2008 (4/1); Antihyperglycemic 2015–2023 (8/2); Antimicrobial 2013–2023 (10/4); Antioxidant 2006–2023 (22/4); Anti-inflammatory 2011–2023 (15/4); Antinociceptive 2014–2022 (3/2); Antiprofilactive 2016–2018 (2/3); Chemopreventive 2016 (2/3), Hypocholesterolemic 2021 (1/3); and Modulator of gut microbiota 2019–2022 (3/1).

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Table 11. Selected stingless bees from some studies on active biomolecules and biological activities, first and last publications

Active Biomolecules	Stingless bee species	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
Flavonoids	Frieseomelitta varia, Melipona compressipes, Melipona favosa, Paratrigona anduzei, Scaptotrigona depilis	Venezuela	1993	–	–	–	Tomás-Barberán et al.⁵²
	Geotrigona sp., Tetragonisca fiebrigi	Ecuador	2023	–	–	Ferreirat et al.⁵³	–
Polyphenols	Frieseomelitta varia, Melipona compressipes, Melipona favosa, Paratrigona anduzei, Scaptotrigona depilis	Venezuela	1993	–	–	–	Tomás-Barberán et al.⁵²
	Geotrigona sp., Tetragonisca fiebrigi	Ecuador	2023	–	–	Ferreirat et al.⁵³	–
Biological activities	Anti-atherogenic
	Heterotrigona itama	Malaysia	2019	–	Othman et al.⁵⁴	–	–
	Heterotrigona itama	Malaysia	2022	–	Zakaria et al.⁵⁵	–	–
Anticancer	Tetragonula spp.	India	2013	–	–	–	Choudhari et al.⁵⁶
	Heterotrigona itama	Malaysia	2020	–	Mahmood et al.⁵⁷	–	–
Anticataract	Melipona favosa	Venezuela	2002	–	Vits⁵⁸	–	–
	Commercial flavonoids present in honey¹	Wales, UK	2008	–	–	–	–
Antihyperglycemic	Leipidotrigona ventralis, Leipidotrigona	Thailand	2015	–	–	–	Vongsak et al.⁵⁹

Active Biomolecules	Stingless bee species	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
(continuation from previous row)	terminata, Tetragonula pagdeni
Anti-inflammatory	Heterotrigona itama	Malaysia	2023	–	Cheng et al.⁶⁰	–	–
	Tetragonula sapiens	Indonesia	2023	–	–	–	Farida et al.⁶¹
	Tetragonula carbonaria	Australia	2011	–	–	Massaro et al.²⁴	–
	Tetragonula biroi	Indonesia	2023	–	–	–	Arung et al.⁶²
	Heterotrigona itama, Tetragonula reepeni, Tetragonula testaceitarsis, Tetragonula fuscobalteata, Tetragonula iridipennis, Tetragonula pagdeni	Indonesia	2023	–	Naibaho et al.⁶³	–	–
	Melipona seminigra	Brazil	2013	–	da Silva et al.⁶⁴	–	–
Antimicrobial	Axestotrigona ferruginea, Axestotrigona togoensis, Melipelbea beccarii, Hypotrigona gribodoi, Dactylurina schmidti, Plebeina armata	Tanzania	2023	–	Mduda et al.⁶⁵	–	–
Antinociceptive	Melipona subnitida	Brazil	2014	–	–	–	Silva et al.⁶⁶
	Tetragonisca fiebrigi	Argentina	2022	–	Salomon et al.⁶⁷	–	–
	Tetragonula carbonaria	Australia	2013	–	–	–	Vit et al.⁶⁷
Antiproliferative	Melipona fasciculata, Melipona rufiventris, Melipona scutellaris, Melipona subnitida, Scaptotrigona polysticta	Brazil		–	–	–	–
	Frieseomelitta nigra, Melipona beecheii, Melipona fasciata, Melipona solani, Scaptotrigona hellwegeri, Scaptotrigona mexicana	Mexico		–	–	–	–
	Melipona favosa	Venezuela		–	–	–	–
Antioxidant	Geniotrigona thoracica, Heterotrigona itama	Malaysia	2018	–	Ismail et al.⁶⁸	Ismail et al.⁶⁸	–
	Melipona subnitida	Brazil	2006	–	–	Silveira et al.⁶⁹	–
	Geotrigona sp., Tetragonisca fiebrigi	Ecuador	2023	–	–	Ferreirat et al.⁵³	–

Table 12. Active biomolecules and biological activities of pot-honey, pot-pollen, cerumen, and propolis.

Active Biomolecules	Stingless bee species	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
Flavonoids	Frieseomelitta varia, Melipona compressipes, Melipona favosa, Paratrigona anduzei, Scaptotrigona depilis	Venezuela	1993	–	–	–	Tomás-Barberán et al.⁵²
	Melipona subnitida	Brazil	2006	–	–	Silva et al.⁶⁹	–
	Melipona spp.	Venezuela	2011	Truchado et al.¹³⁷	–	–	–
	Tetragonisca angustula	Venezuela	2013	Pérez-Pérez et al.¹³⁸	–	Pérez-Pérez et al.¹³⁸	Pérez-Pérez et al.¹³⁸
	Melipona seminigra	Brazil	2013	da Silva et al.⁶⁴	–	–	–
	Melipona quadrifasciata, Tetragonula clypearis, Scaptotrigona spp.	Brazil	2017	–	–	Pazin et al.¹³⁹	–
	Melipona subnitida	Brazil	2018	Tukistah et al.¹⁴⁰	–	–	de Souza et al.⁶⁷
	Geniotrigona thoracica, Heterotrigona itama	Malaysia	2018	–	–	–

Active Biomolecules	Stingless bee species	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
	Heterotrigona erythrogastra, Tetrigona apicalis, Heterotrigona itama, Geniotrigona thoracica	Malaysia	2019	–	–	–	Asem et al.⁸⁰
	Tetragonula biroi	Philippines	2019	–	Belina-Aldemita et al.¹⁴²	–	–
	Melipona quadrifasciata, Melipona asilvai, Melipona subnitida, Melipona scutellaris	Brazil	2019	–	Oliveira et al.¹⁴³	–	–
	Scaptotrigona bipunctata, Melipona marginata, Tetragonisca angustula, Trigona hypogea, Melipona quadrifasciata, Tetragona clavipes	Brazil	2020	–	Biluca et al.¹⁴⁴	–	–
	Heterotrigona itama	Malaysia	2020	–	Majid et al.¹⁴⁵	–	–
	Melipona seminigra	Brazil	2021	–	–	–	Rebelo et al.⁷³
	Tetrigona apicalis, Tetrigona binghami, Heterotrigona fimbriata	Malaysia	2021	–	–	–	Syed Salleh et al.⁸²
	Geotrigona sp., Tetragonisca fiebrigi	Ecuador	2023	–	–	Ferreirat et al.⁵³	–
	Tetragonisca fiebrigi	Brazil	–	–	–	–	–

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Polyphenols

Active Biomolecules	Stingless bee species	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
Polyphenols	Frieseomelitta varia, Melipona compressipes, Melipona favosa, Paratrigona anduzei, Scaptotrigona depilis	Venezuela	1993	–	–	–	Tomás-Barberán et al.⁵²
	Tetragonula carbonaria	Australia	2011	–	–	Massaro et al.²⁴	–
	Tetragonisca angustula	Venezuela	2013	Pérez-Pérez et al.¹³⁸	–	Pérez-Pérez et al.¹³⁸	Pérez-Pérez et al.¹³⁸
	Melipona seminigra	Brazil	2013	–	da Silva et al.⁶⁴	–	–
	Melipona fasciculata	Brazil	2014	–	–	–	Dutra et al.¹⁰⁶
	Melipona quadrifasciata, Tetragona clavipes, Scaptotrigona spp.	Brazil	2017	–	–	Pazin et al.¹³⁹	–
	Melipona subnitida	Brazil	2018	–	–	de Souza et al.¹⁴⁰	–
	Geniotrigona thoracica, Heterotrigona itama, Heterotrigona erythrogastra	Malaysia	2018	–	Tukishta et al.¹⁴¹	–	–
	Tetrigona apicalis, Heterotrigona itama, Geniotrigona thoracica	Malaysia	2019	–	–	–	Asem et al.⁸⁰
	Melipona quadrifasciata, Melipona asilvai	Brazil	2019	–	Oliveira et al.¹⁴³	–	–

Active Biomolecules	Stingless bee species	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
	Melipona subnitida, Melipona scutellaris	Brazil	2020	–	Biluca et al.¹⁴⁴	–	–
	Scaptotrigona bipunctata, Melipona marginata, Tetragonisca angustula, Trigona hypogea, Melipona quadrifasciata, Tetragona clavipes	Brazil	2020	–	Biluca et al.¹⁴⁴	–	–
	Heterotrigona itama	Malaysia	2020	–	Majid et al.¹⁴⁵	–	–
	Melipona seminigra	Brazil	2021	–	–	–	Rebelo et al.⁷³
	Tetrigona apicalis, Tetrigona binghami, Homotrigona fimbriata	Malaysia	2021	–	–	–	Syed Salleh et al.⁸²
	Geotrigona sp., Tetragonisca fiebrigi	Ecuador	2023	–	–	Ferreira et al.⁵³	–
	Tetragonula laeviceps	Thailand	2023	–	–	Iesa et al.¹⁴⁷	–

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Biological activities

Anti-atherogenic

Stingless bee taxa	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
Heterotrigona itama	Malaysia	2019	–	Othman et al.⁵⁴	–	–
Geniotrigona thoracica	Malaysia	2020	–	Mohd Suib et al.¹⁴⁸	–	–
Heterotrigona itama	Malaysia	2022	–	Zakaria et al.⁵⁵	–	–

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Anticancer

Stingless bee taxa	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
Tetragonula spp.	India	2013	–	–	–	Choudhari et al.⁵⁶
Homotrigona apicalis, Tetragonula fuscibasis, Tetragonula fuscobalteata, Wallacetrigona incisa	Indonesia	2014	–	Kustiawan et al.¹⁴⁹	–	Kustiawan et al.¹⁴⁹
Tetragonula laeviceps	Thailand	2015	–	–	–	Nugitrangson et al.¹⁵⁰
Lepidotrigona terminata	Malaysia	2016	–	Omar et al.⁷¹	–	–
Melipona orbignyi	Brazil	2017	–	–	–	dos Santos et al.¹⁵¹
Heterotrigona itama	Malaysia	2019	–	Ahmad et al.¹⁵²	–	–
Tetragonula biroi	Philippines	2019	–	–	–	Desamero et al.¹⁵³
Heterotrigona itama	Malaysia	2020	–	Mahmood et al.⁵⁷	–	–
Homotrigona fimbriata, Heterotrigona itama, Heterotrigona bakeri, Tetragonula sarawakensis, Tetragonula testaceitarsis, Tetragonula fuscobalteata, Tetragonula laeviceps	Indonesia	2021	–	Arung et al.¹⁵⁴	–	Arung et al.¹⁵⁴

Active Biomolecules	Stingless bee species	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis

Anticataract

Melipona favosa, Tetragonisca angustula, Commercial luteolin derivatives present in honey¹

Venezuela

1997

Vit¹⁵⁵

–

–

–

Commercial luteolin derivatives present in honey¹

Wales, UK

2001

–

–

–

–

Melipona favosa

Venezuela

2002

Vit⁵⁸

–

–

–

Melipona favosa, Scaptotrigona mexicana, Tetragonisca angustula Commercial luteolin derivatives present in honey

Brazil, Mexico, Venezuela

2004

Vit et al.⁹

–

–

–

Commercial flavonoids present in honey²

Wales, UK

2008

–

–

–

–

Lepidotrigona ventralis, Lepidotrigona terminata, Tetragonula pagdeni

Thailand

2015

–

–

–

Vongsak et al.⁵⁹

Geniotrigona thoracica

Malaysia

2017

–

Abdul Aziz et al.¹⁵⁶

–

–

Antihyperglycemic

Heterotrigona itama, Trigona apicalis

Malaysia

2018

–

Na et al.¹⁵⁷

–

–

Tetragonula sapiens

Indonesia

2019

–

Pujiarahayu et al.¹⁵⁸

–

–

Tetragonula biroi, Tetragonula leytebesis

Indonesia

2019

–

Rahmawati et al.¹⁴⁷

–

–

Heterotrigona itama

Malaysia

2020

–

Ali et al.¹⁶⁰

–

–

Heterotrigona itama

Malaysia

2023

–

Cheng et al.⁶⁰

–

–

Tetragonula sapiens

Indonesia

2023

–

–

–

Farida et al.⁶¹

Anti-inflammatory

Tetragonula carbonaria

Australia

2011

–

–

Massaro et al.²⁴

–

Tetragonisca fiebrigi

Brazil

2015

–

–

–

Campos et al.¹⁶¹

Tetragonula carbonaria

Australia

2016

–

–

–

Hamilton et al.¹⁶²

Melipona orbignyi

Brazil

2017

–

–

–

dos Santos et al.¹⁵¹

Tetragonula carbonaria

Australia

2017

–

–

–

Hamilton et al.¹⁶³

Melipona fasciculata

Brazil

2019

–

–

Lopes et al.¹⁶⁴

–

Scaptotrigona bipunctata, Melipona marginata, Tetragonisca angustula, Trigona hypogea, Melipona quadrifasciata, Tetragona clavipes

Brazil

2020

–

Biluca et al.¹⁴⁴

–

–

Tetragonula spp.

Malaysia

2020

–

–

Badrulhisham et al.¹⁶⁵

–

Melipona fasciculata

Brazil

2020

–

–

–

Active Biomolecules	Stingless bee species	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
Antimicrobial	Heterotrigona itama	Malaysia	2020	–	–	–	Zhang et al.¹⁶⁷
	Heterotrigona itama	Malaysia	2021	–	Ooi et al.¹⁶⁸	–	–
	Tetragonula carbonaria	Australia	2022	–	–	Hamilton et al.¹⁶⁹	–
	Tetragonisca fiebrigi	Argentina	2022	–	Salomon et al.¹⁶⁷	–	–
	Heterotrigona itama, Tetragona binghami	Malaysia	2022	–	Wu et al.¹⁷⁰	–	–
	Tetragonula biroi	Indonesia	2023	–	–	–	Arung et al.¹⁵⁴
	Heterotrigona itama, Tetragonula reepeni, Tetragonula testaceitarsis, Tetragonula fuscobalteata, Tetragonula iridipennis, Tetragonula pagdeni	Indonesia	2023	–	Naibaho et al.⁶³	–	–
	Geotrigona sp.	Ecuador	2023	–	–	Ferreira et al.⁵³	–
	Tetragonisca fiebrigi	Brazil	–	–	–	–	–
	Melipona seminigra	Brazil	2013	da Silva et al.⁶⁴	–	–	–
	Tetragonisca fiebrigi	Brazil	2015	–	–	–	Campos et al.¹⁵⁹
	Melipona quadrifasciata, Tetragonisca angustula	Brazil	2017	–	–	–	dos Santos et al.¹⁵¹
	Geniotrigona thoracica, Heterotrigona itama, Heterotrigona erythrogastra	Malaysia	2018	–	Tukishta et al.¹⁴¹	–	–
	Geniotrigona thoracica, Heterotrigona itama, Tetragona binghami	Brunei	2020	–	–	Abdullah et al.¹⁷¹	–
	Axestotrigona ferruginea	Tanzania	2021	–	Popova et al.⁵¹	–	Popova et al.⁵¹
	Tetragonisca fiebrigi	Argentina	2022	–	Dallagnol et al.¹⁷²	–	–
	Heterotrigona itama, Tetragonula binghami	Malaysia	2022	–	Wu et al.¹⁷⁰	–	–
	Axestotrigona ferruginea, Axestotrigona togoensis, Meliplebeia beccarii, Hypotrigona gribodoi, Dactylurina schmidti, Plebeina armata	Tanzania	2023	–	Mduda et al.⁶⁵	–	–
	Heterotrigona itama, Tetragonula reepeni, Tetragonula pagdeni, Tetragonula iridipennis, Tetragonula fuscobalteata, Tetragonula testaceitarsis	Indonesia	2023	–	Naibaho et al.¹⁷³	–	–

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Antinociceptive

Active Biomolecules	Stingless bee species	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
Antinociceptive	Melipona subnitida	Brazil	2014	–	–	–	Silva et al.⁶⁶
	Melipona fasciculata	Brazil	2019	–	–	–	Lopes et al.¹⁶⁴

Active Biomolecules	Stingless bee species	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
Antioxidant	Tetragonisca fiebrigi	Argentina	2022	Salomon et al.¹⁶	–	–	–
	Melipona subnitida	Brazil	2006	–	Silva et al.⁶⁵	–	–
	Tetragonisca angustula	Venezuela	2007	Pérez-Pérez et al.¹¹⁰	–	–	–
	Melipona sp., Tetragonisca sp.	Venezuela	2007	Rodríguez-Malaver et al.¹⁷⁴	–	–	–
	Melipona seminigra	Brazil	2013	da Silva et al.⁶⁴	–	–	–
	Tetragonisca angustula	Venezuela	2013	Pérez-Pérez et al.¹³⁸	–	Pérez-Pérez et al.¹³⁸	–
	Melipona fasciculata	Brazil	2014	–	–	Dutra et al.¹⁰⁶	–
	Tetragonula carbonaria	Australia	2016	–	–	Hamilton et al.¹⁶²	–
	Geniotrigona thoracica, Heterotrigona itama, Tetrigona apicalis	Malaysia	2016	–	Nurdianah et al.¹⁷³	–	–
	Tetragonula carbonaria	Australia	2017	–	–	Hamilton et al.¹⁶³	–
	Tetrigona apicalis, Heterotrigona itama, Geniotrigona thoracica	Malaysia	2019	–	Hanif Fadzilah et al.¹⁷⁵	–	–
	Melipona subnitida, Tetragona clavipes, Scaptotrigona spp.	Brazil	2017	–	–	Pazin et al.¹³⁹	–
	Melipona quadrifasciata, Tetragonisca angustula	Brazil	2019	–	–	dos Santos et al.¹⁵¹	–
	Geniotrigona thoracica, Heterotrigona itama, Heterotrigona erythrogastra	Malaysia	2018	–	Tukisita et al.¹⁴¹	–	–
	Tetrigona apicalis, Heterotrigona itama, Geniotrigona thoracica	Malaysia	2019	–	–	–	Asem et al.⁸⁰
	Tetragonula biroi	Philippines	2019	–	Belina-Aldemita et al.¹⁴²	–	–
	Melipona compressipes	Brazil	2019	–	–	Carneiro et al.¹³⁷	–
	Melipona quadrifasciata, Melipona asilvai, Melipona subnitida, Melipona scutellaris	Brazil	2019	–	Oliveira et al.¹⁴³	–	–
	Geniotrigona thoracica, Heterotrigona itama, Tetragonula binghami	Brunei	2020	–	–	Abdullah et al.¹⁷¹	–
	Scaptotrigona bipunctata, Melipona marginata, Tetragonisca angustula, Trigona hypogea, Melipona quadrifasciata, Tetragona clavipes	Brazil	2020	–	Biluca et al.¹⁴⁴	–	–
	Heterotrigona itama	Malaysia	2020	–	Majid et al.¹⁴⁵	–	–

Table 12 – Antiproliferative, Chemopreventive, Hypocholesterolemic, and Modulator of Gut Microbiota)

Active Biomolecules	Stingless bee species	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
(continued)	Tetrigona apicalis, Tetrigona binghami, Homotrigona fimbriata	Malaysia	2021	–	–	–	Syed Salleh et al.⁸²
	Axestotrigona ferruginea, Axestotrigona togoensis, Meliplebeia beccarii, Meliponula bocandei, Liotrigona spp., Plebeina armata	Kenya	2022	Mokaya et al.¹⁷⁸	–	–	–
	Heterotrigona itama, Tetragonula binghami	Malaysia	2022	Wu et al.¹⁷⁹	–	–	–
	Geotrigona sp., Tetragonisca fiebrigi	Ecuador	2023	–	–	Ferreira et al.⁵³	–
	Tetragonula laeviceps	Thailand	2023	–	–	Iesa et al.¹⁴⁷	–
	Axestotrigona ferruginea, Axestotrigona togoensis, Meliplebeia beccarii, Hypotrigona gribodoi, Dactylurina schmidti, Plebeina armata	Tanzania	2023	Mduda et al.⁷⁰	–	–	–
	Tetragonula carbonaria	Australia	2013	Vit et al.⁶⁷	–	–	–
	Melipona fasciculata, Melipona rufiventris, Melipona scutellaris, Melipona subnitida, Scaptotrigona polysticta, Frieseomelitta nigra, Melipona beecheii, Melipona fasciata, Melipona solani, Scaptotrigona hellwegeri, Scaptotrigona mexicana	Brazil / Mexico	–	–	–	–	–
	Melipona favosa	Venezuela	–	–	–	–	–

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Chemopreventive

Active Biomolecules	Stingless bee species	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
Chemopreventive	Lepidotrigona terminata	Malaysia	2016	–	–	Omar et al.⁷¹	–
	Geniotrigona thoracica, Heterotrigona itama	Malaysia	2018	–	Ismail et al.⁶⁸	–	Ismail et al.⁶⁸
	Lepidotrigona terminata	Malaysia	2016	–	–	Omar et al.⁷¹	–
	Tetragonula spp.	Malaysia	2016	–	–	Yazan et al.⁷²	–

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Hypocholesterolemic

Active Biomolecules	Stingless bee species	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
Hypocholesterolemic	Melipona seminigra	Brazil	2021	–	–	–	Rebelo et al.⁷³

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Modulator of gut microbiota

Active Biomolecules	Stingless bee species	Country	Year	Pot-honey	Pot-pollen	Cerumen	Propolis
Modulator of gut microbiota	Heterotrigona itama	Malaysia	2019	–	–	–	Zulkahiri Amin et al.⁷⁴
	Heterotrigona itama	Malaysia	2020	–	–	–	Mohamad et al.¹⁷⁹
	Tetragonula sarawakensis, Heterotrigona itama, Tetragonula testaceitarsis, Tetragonula minangkabau, Geniotrigona thoracica, Tetragonula binghami	Indonesia	2022	–	–	–	Melia et al.⁷⁵

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Text below the table

The list of stingless bee taxa of this revision was tabulated in Table 13 with their Neotropical, Afrotropical, Indo-Malaysian, and Australian geographical distribution. Pot-honey, pot-pollen, cerumen and propolis from 31 Neotropical (Argentina, Brazil, Ecuador, Mexico, and Venezuela), 9 Afrotropical (Kenya and Tanzania), 22 Indo-Malaysian (Brunei, India, Indonesia, Malaysia, Philippines, and Thailand), and 1 Australian stingless bee taxa were studied for their flavonoid.

Most biological activities were studied for materials of stingless bee species from Indo-Malaysian Heterotrigona itama (9) and Geniotrigona thoracica (7), the Neotropical Tetragonisca angustula (6) and Tetragonisca fiebrigi (6), and the unique medicinal Australian bee Tetragonula carbonaria (5) in our search.

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Table 13. Stingless bee taxa of the geographical region (Neotropical, Afrotropical, Indo-Malaysian, Australian) used in medicinal stingless bee research of Table 12.

No.	Geographical Region / Stingless Bee Taxa	Country	Biomolecules and Biological Activity
Neotropical
1	Frieseomelitta nigra	Mexico	Antiproliferative
2	Frieseomelitta varia	Brazil	Flavonoids, Polyphenols
3	Geotrigona sp.	Ecuador	Flavonoids, Polyphenols, Antioxidant
4	Melipona asilvai	Brazil	Flavonoids, Polyphenols, Antioxidant
5	Melipona beecheii	Mexico	Antiproliferative
6	Melipona compressipes	Brazil	Flavonoids, Polyphenols, Antioxidant
7	Melipona fasciculata	Mexico	Antiproliferative
8	Melipona fasciculata	Brazil	Polyphenols, Anti-inflammatory, Antinociceptive, Antioxidant, Antiproliferative
9	Melipona favosa	Venezuela	Flavonoids, Anticataract, Antiproliferative, Chemopreventive
10	Melipona marginata	Brazil	Flavonoids, Polyphenols, Antioxidant
11	Melipona orbignyi	Brazil	Anticancer, Anti-inflammatory
12	Melipona quadrifasciata	Brazil	Flavonoids, Polyphenols, Anti-inflammatory, Antimicrobial, Antioxidant
13	Melipona rufiventris	Brazil	Antiproliferative
14	Melipona scutellaris	Brazil	Flavonoids, Polyphenols, Antioxidant
15	Melipona seminigra	Brazil	Flavonoids, Polyphenols, Antimicrobial, Antioxidant, Antiproliferative, Hypocholesterolemic
16	Melipona solani	Mexico	Antiproliferative
17	Melipona subnitida	Brazil	Flavonoids, Polyphenols, Antinociceptive, Antioxidant, Antiproliferative
18	Melipona sp.	–	Flavonoids
19	Paratrigona anduzei	Brazil	Flavonoids, Polyphenols
20	Scaptotrigona bipunctata	Brazil	Flavonoids, Polyphenols, Anti-inflammatory, Antioxidant
21	Scaptotrigona depilis	Brazil	Flavonoids, Polyphenols
22	Scaptotrigona hellwegeri	Mexico	Antiproliferative
23	Scaptotrigona mexicana	Mexico	Anticataract, Antiproliferative
24	Scaptotrigona polysticta	Brazil	Antiproliferative
25	Scaptotrigona spp.	Brazil	Flavonoids, Polyphenols, Antioxidant
26	Tetragona clavipes	Brazil	Flavonoids, Polyphenols, Anti-inflammatory, Antioxidant
27	Tetragonisca angustula	Brazil	Flavonoids, Polyphenols, Anticataract, Anti-inflammatory, Antimicrobial, Antioxidant
		Venezuela	Flavonoids, Polyphenols, Anticataract, Antimicrobial, Antioxidant
28	Tetragonisca fiebrigi	Brazil	Flavonoids, Polyphenols, Anti-inflammatory, Antimicrobial, Antinociceptive, Antioxidant

No.	Geographical Region / Stingless Bee Taxa	Country	Biomolecules and Biological Activity
30	Tetragonisca sp.	Venezuela	Antioxidant
31	Trigona hypogea	Brazil	Flavonoids, Polyphenols, Anti-inflammatory, Antioxidant
		5 countries

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Afrotropical

No.	Stingless Bee Taxa	Country	Biomolecules and Biological Activity
1	Axestotrigona ferruginea	Tanzania	Antimicrobial, Antioxidant
		Kenya	Antioxidant
2	Axestotrigona togoensis	Tanzania	Antimicrobial, Antioxidant
		Kenya	Antioxidant
3	Dactylurina schmidti	Tanzania	Antimicrobial, Antioxidant
4	Hypotrigona gribodoi	Tanzania	Antimicrobial, Antioxidant
5	Liotrigona sp.	Kenya	Antioxidant
6	Meliplebeia beccarii	Tanzania	Antimicrobial, Antioxidant
		Kenya	Antioxidant
7	Meliponula bocandei	Kenya	Antioxidant
8	Meliplebeia lendliana	Kenya	Antioxidant
9	Plebeina armata	Tanzania	Antimicrobial, Antioxidant

| | | 2 countries | Antioxidant |

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Indo-Malaysian

No.	Stingless Bee Taxa	Country	Biomolecules and Biological Activity
1	Geniotrigona thoracica	Malaysia	Flavonoids, Polyphenols, Anti-atherogenic, Antihyperglycemic, Antimicrobial, Antioxidant, Antiproliferative
		Brunei	Antimicrobial, Antioxidant
		Indonesia	Modulator of gut microbiota
2	Heterotrigona bakeri	Indonesia	Anticancer
3	Heterotrigona erythrogastra	Malaysia	Polyphenols, Antimicrobial
4	Heterotrigona itama	Malaysia	Flavonoids, Polyphenols, Anti-atherogenic, Antihyperglycemic, Anti-inflammatory, Antimicrobial, Antioxidant, Antiproliferative, Modulator of gut microbiota
		Indonesia	Anticancer, Anti-inflammatory, Antimicrobial, Modulator of gut microbiota
		Brunei	Antimicrobial, Antioxidant
5	Homotrigona apicalis	Indonesia	Anticancer
6	Homotrigona fimbriata	Indonesia	Flavonoids, Polyphenols, Anticancer, Antioxidant
7	Lepidotrigona terminata	Malaysia	Anticancer, Antiproliferative, Chemopreventive
		Thailand	Antihyperglycemic, Antiproliferative, Chemopreventive
8	Lepidotrigona ventralis	Thailand	Antihyperglycemic
9	Tetragonula fuscibasis	Indonesia	Anticancer
10	Tetragonula biroi	Philippines	Flavonoids, Anticancer, Antioxidant

No.	Geographical Region / Stingless Bee Taxa	Country	Biomolecules and Biological Activity
11	Tetragonula fuscobalteata	Indonesia	Antihyperglycemic, Anti-inflammatory
12	Tetragonula iridipennis	Indonesia	Anticancer, Anti-inflammatory, Antimicrobial
13	Tetragonula laeviceps	Thailand	Polyphenols, Anticancer, Antioxidant
		Indonesia	Antihyperglycemic
14	Tetragonula minangkabau	Indonesia	Modulator of gut microbiota
15	Tetragonula pagdeni	Thailand	Antihyperglycemic, Anti-inflammatory, Antimicrobial
16	Tetragonula reepeni	Indonesia	Anti-inflammatory, Antimicrobial
17	Tetragonula sapiens	Indonesia	Antihyperglycemic
18	Tetragonula sarawakensis	Indonesia	Anticancer, Modulator of gut microbiota
19	Tetragonula testaceitarsis	Indonesia	Anticancer, Anti-inflammatory, Antimicrobial, Modulator of gut microbiota
20	Tetragonula spp.	India	Anticancer
		Malaysia	Anti-inflammatory, Chemopreventive
21	Tetrigona apicalis	Malaysia	Flavonoids, Polyphenols, Antihyperglycemic, Antioxidant
22	Tetragona binghami	Malaysia	Flavonoids, Polyphenols, Anti-inflammatory, Antioxidant
		Brunei	Antimicrobial, Antioxidant
		Indonesia	Modulator of gut microbiota
23	Wallacetrigona incisa	Indonesia	Anticancer
	23 taxa	6 countries

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Australian

No.	Stingless Bee Taxa	Country	Biomolecules and Biological Activity
1	Tetragonula carbonaria	Australia	Polyphenols, Anti-inflammatory, Antioxidant, Antiproliferative, Chemopreventive

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3.4 QUALITY CONTROL OF STINGLESS BEE PRODUCTS. KEY BIOMOLECULES, METHODS AND TECHNIQUES

The developing interest towards use of stingless bee products to support human health makes the issue of their standardization increasingly important and urgent. It is well known that stingless bee honey does not comply with the requirements for honeybee honey in the CODEX Alimentarius⁷⁷, which have been created for Apis mellifera honey. The growing amount of data on the characteristics of stingless bee honey led to the need of creation of specific quality standard for pot-honey. In general, stingless bee honeys do not meet the CODEX Standard for honey moisture, free acidity, and total fructose plus glucose levels. In addition, it has been suggested to apply the presence and amount of a rare reducing sugar, trehalulose, as a marker of authenticity of pot-honey⁷⁸.

However, to produce a universal standard and quality parameters, further information from studies of the chemical composition of stingless bee honey — such as organic acids and polyphenol profiles of pot-honey from stingless bee species from different geographical regions is required.

Targeted ¹H-NMR is adequate to compare sugars, amino acids, aliphatic organic acids, HMF, ethanol, and botanical markers of pot-honey produced by diverse entomological origins¹².

Concerning stingless bee propolis, the chemical variability is much greater than that observed with A. mellifera propolis. This has been a serious hindrance in the case of Apis mellifera propolis which now, after decades of intense research, comes to an at least partial solution. The stingless bees’ propolis poses a much more difficult problem, due to its greater chemical diversity. It is interesting to note that several molecules with significant bioactivities have been found in pot-propolis (gallic acid, alpha-mangostine, propolin A), some of them new chemical entities, such as sulawesin A, mammeoin cinnamoyl ester, etc.⁷⁹ (Fig. 15).

Fig. 15 Important bioactive compounds in stingless bee propolis

The antimicrobial and antioxidant activities of stingless bee propolis are usually reported to depend on its total phenolic content, e.g. Asem et al.⁸⁰. However, it is necessary to remember that different stingless bee propolis often contain different phenolic compounds and equal number of observed gallic acid equivalents could correspond to a very different real concentration of phenolics. For this reason, the chemical type of the propolis samples should be observed by LC-MS⁸¹ or GC-MS analyses⁸². As in the case of pot-honey or stingless bee honey, pot-pollen or stingless bee pollen, and cerumen, many more studies have to be conducted on the chemistry and biological activity of stingless bee propolis before any quality control criteria could be formulated.

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3.5 INFLUENCE OF THE MELIPONA GENUS BAHIA STATE HONEY NORM FROM BRAZIL (2014), AND FURTHER POT-HONEY NORMS IN MEDICINAL USES OF POT-HONEY

In their seminal paper, Gonnet et al.⁸³ evidenced the higher moisture and free acidity of Melipona honey compared to Apis mellifera honey. The first proposal for stingless bee honey standards included honey of the Melipona from Guatemala, Mexico, and Venezuela.⁹ The first Brazilian stingless bee honey norm was created for Melipona honey in the State of Bahia.² Forthcoming Brazilian State standards were established in Amazonas⁸⁴, Paraná⁸⁵, Espírito Santo⁸⁶, and Santa Santa Catarina⁸⁷. The Philippine honey norm⁸⁸ included pot-honey in the last revision, but the most important State National standard was created in 2017 for Kelulut — Malaysian name given to all stingless bees— honey in Malaysia⁸⁹, and the second was for the Argentine stingless bee Tetragonisca fiebrigi known with the ethnic name Yate’i in 2019⁹⁰.

Concomitant with the scientific research on medicinal properties of stingless bee products, regulated pot-honey facilitates administrative procedures to launch pharmaceutical products. The diverse state norms in Brazil have promoted presentations of pot-honey, pot-pollen, and propolis of pharmaceutical quality, available online⁹¹.

The facts of standardized pot-materials of the stingless bee nest are promising:

1. More quality products will be produced and will command fair market prices,
2. Regulated products are safe for human and animal health,
3. Support of apitherapy by providing quality raw materials,
4. Best stingless bee-keeping practices should be strictly followed in order to produce quality products, and especially
5. Stingless bee apiaries should backup the volume of marketed stingless bee products, to prevent falsifications.

Potential categorizations of stingless bee materials may include antioxidant activity grading, as suggested for Czech honey using μmoles equivalents Trolox/100 g honey in a scale of:

• pro-oxidant (<1),
• very low (0–50),
• low (51–100),
• moderately low (101–150),
• medium (151–200),
• moderate high (201–250),
• high (251–300),
• very high (>300)
  by Vit et al.⁹².

A more recent idea of categorization arises after the trehalulose discovery in pot-honey by Fletcher et al., 2020³⁹ and chemical marker analysis⁷⁸ could provide useful gradings for entomological origins.
This unique sugar needs extensive research and the following trend reveals useful for trehalulose concentrations (g/100g honey) graded for preliminary stingless bee species:

• (10.0–39.9) Heterotrigona itama, Tetragonula carbonaria, Tetragonula hockingsi;
• (40.0–49.9) Geniotrigona thoracica, Heterotrigona itama, and
• (50.0–59.9) Geniotrigona thoracica.

Our Quality of the beehive for apitherapy in the VII National Congress of Pharmaceutical Science 2009⁹⁴ has focused to Quality of the stingless bee nest for apitherapy in the Apimondia 2023 Central Symposium for Apitherapy, consistent with good practices of stingless bee keeping to achieve great quality of stingless bee products for direct use or in pharmaceutical preparations⁹⁵. Vit and Simova⁹⁶ reviewed the aliphatic organic acids (AOA) in honey and proposed updated reference values for Apis mellifera and stingless bee honeys.

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3.6 INTEGRATIVE APPLICATIONS OF STINGLESS BEE NEST METABOLITES TO OVERCOME ANTIBIOTIC RESISTANCE

Antimicrobial resistance (AMR) has become a global health and socioeconomic issue requiring urgent attention⁹⁷. In 2019, an estimated 1.27 million deaths worldwide were attributable to antibiotic-resistant bacterial infections⁹⁸. The World Health Organization (WHO) estimated that by 2050, diseases caused by multidrug-resistant bacteria could lead to 10 million deaths annually, surpassing cancer as the leading cause of death⁹⁹.

In 2024, the WHO updated its list of bacterial pathogens considered a priority due to limited treatment options and their significant global health impact. At the top of this list are highly virulent bacteria, including carbapenem-resistant Acinetobacter baumannii, carbapenem-resistant Enterobacter, and third-generation cephalosporin-resistant Enterobacterales¹⁰⁰. Infections caused by these pathogens are difficult to prevent, highly transmissible, and associated with a high mortality rate, making them a major public health threat⁹⁹–¹⁰¹.

The global rise of multidrug-resistant pathogens represents one of the greatest challenges to modern medicine, prompting the search for new therapeutic strategies beyond conventional antibiotics¹⁰². In this context, stingless bee nest derivatives, such as pot-honey, pot-pollen, cerumen, and propolis, are emerging as promising sources of bioactive compounds for developing effective therapies against resistant microorganisms¹⁰³.

Stingless bees produce a wide variety of bioactive substances, including antimicrobial peptides, polyphenols, flavonoids, and enzymes that synergistically inhibit bacterial growth and biofilm formation, modulate immune responses, promote tissue repair, and reduce inflammation¹⁰⁴. Propolis, in particular, has been extensively studied for its high content of phenolic compounds and flavonoids with potent antioxidant and antimicrobial effects, capable of inhibiting both Gram-positive and Gram-negative bacteria, including multidrug-resistant strains¹⁰⁵. These compounds act through multiple mechanisms: altering bacterial cell membranes, inhibiting protein synthesis, preventing biofilm formation, and enhancing antibiotic efficacy through proven synergies¹⁰⁶.

Similarly, pot-honey exhibits antimicrobial activity due to the presence of hydrogen peroxide and other compounds that act synergistically with flavonoids and phenolic acids to destabilize bacterial membranes and prevent biofilm formation as key factors in the persistence of chronic resistant infections¹⁰⁴.

Beyond the intrinsic antimicrobial properties of stingless bee nest metabolites, numerous studies emphasize the synergistic interactions between bee products and conventional antibiotics¹⁰³,¹⁰⁷,¹⁰⁸. This synergy not only enhances antimicrobial efficacy and reduces the required antibiotic dosage but also helps prevent the emergence of new resistances and mitigates potential side or toxic effects¹⁰².

Araque and Vit¹⁰⁹ evaluated the synergistic potential of ethanolic extracts of Tetragonisca angustula pot-pollen combined with amikacin and meropenem—two frontline antibiotics used against resistant infections. Their results demonstrated strong synergistic interactions in 75% of the bacterial isolates tested, including clinically significant multidrug-resistant strains such as Klebsiella pneumoniae, Pseudomonas aeruginosa, and Acinetobacter baumannii. This synergy was evidenced by up to a threefold reduction in the minimal inhibitory concentrations (MICs) of the antibiotics, indicating a potential adjuvant role for pot-pollen in restoring antibiotic susceptibility and reducing therapeutic dosages.

Complementing these findings, Vit et al.¹¹⁰ examined the viable compounds in pot-pollenand explored their bioactivity and synergism with antibiotics. In addition to antimicrobial and anti-inflammatory properties, they reported innovative applications of these volatiles in enhancing food flavors, potentially improving the acceptability and clinical application of pot-pollen-derived products. Their bibliometric analysis revealed increasing scientific interest and strong evidence supporting the anti-AMR potential of pot-pollen as a multifunctional natural resource.

From a critical perspective, despite the favorable consensus, the variability in chemical composition due to botanical origin, geographical location, and technical processing presents a challenge in translating findings into standardized and regulated therapeutic applications¹¹¹. Moreover, most studies remain preclinical or in vitro, highlighting the need for robust clinical trials to confirm efficacy and safety in human populations. Nonetheless, advances in the chemical characterization and functional analysis of these natural compounds are paving the way for their inclusion in innovative formulations such as nanoemulsions, gels, and other controlled-release systems¹⁰⁷–¹¹².

The therapeutic promise of stingless bee nest materials extends beyond their capacity to combat bacterial resistance. Their metabolic and cellular protective effects may enhance overall host health and offer potential for managing metabolic and inflammatory diseases that are increasingly prevalent. This integrative perspective underscores the value of stingless bee nest materials as complementary agents in therapeutic management, where responsible and evidence-based utilization could play a pivotal role in the post-antibiotic era.

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3.7 GENOMIC AND METAGENOMIC INSIGHTS INTO THE MEDICINAL POTENTIAL OF STINGLESS BEE NEST MATERIALS

Antimicrobial agents biosynthesized for the survival of microbial communities in honey contribute to its well-known antimicrobial properties¹¹³. Similarly, interactions among microbial populations inhabiting stingless bee nest materials may explain the frequently reported antimicrobial activities of pot-honey¹¹⁴, pot-pollen¹¹⁵, cerumen¹¹⁶, and propolis¹¹⁷. The wide variety of bioactive molecules of microbial origin has been comprehensively reviewed by Vit (2024)¹¹⁸ and Alves et al. (2024)¹¹⁹.

Metabolites generated by the stingless bee nest microbiome represent a vast reservoir of largely unexplored natural compounds that play critical ecological roles and hold strong potential for biotechnological and medicinal applications, including drug discovery. Developing such applications requires understanding the complex microbiota associated with stingless bees to identify and isolate specific bacterial and fungal strains and their metabolites as potential therapeutic agents or dietary supplements. As shown in bumble bees, microbiome assembly and maintenance are dynamic across the lifespan of social bees¹²⁰, and likely also in stingless bees and their nest materials, a factor that must be considered when harvesting high-quality materials.

Current research on stingless bee gut microbiomes remains at an exploratory stage. The core microbiome of Apis mellifera differs markedly from that of the few stingless bee species studied in Australia¹²¹, Brazil¹²²,¹²³, Mexico¹²⁴, and Thailand¹²⁵. Emerging evidence suggests that host-specific and geographical factors influence microbiome composition in stingless bee nest materials, representing an untapped source of bioactive compounds that may contribute both to colony health and to human health applications.

Biotransformations of phytochemicals such as flavonoids and polyphenols by gut microbes involve two-way interactions:

• microbial degradation reduces molecule size, while
• phenolics modulate microbial populations¹²⁶.

Investigating how microbes transform compounds, communicate with each other within communities, interact with the host, and analyzing their microbial products provides deeper insight into host–microbe interactions and their effects on health and disease. Knowing the microbiome helps finding potential active biomolecules by using advanced metabolomics to identify and quantify the chemicals microbes produce. Exploring their bioactive properties reveals molecules that can act as therapeutics, drug targets, or function as biomarkers. By deciphering microbiome-mediated transformations of host-derived molecules and xenobiotics, researchers can uncover novel enzymes and biochemical pathways. Understanding metabolic fingerprints of the microbiome connects microbial communities or a unique microbe from food storage, architectural materials of the nest, or the stingless bee itself with a particular function or a set of roles.depilis, a Brazilian stingless bee, three microbial populations coordinate during pupal metamorphosis: one produces the steroid ergosterol, two modulate its availability¹²⁷, and the symbiotic yeast Zygosaccharomyces sp. promotes metamorphic development¹²⁸. Such examples illustrate the intricate interplay between stingless bees and their symbionts. Our particular interest lies in exploring how these associated microbes may contribute to human health by:

1. Modulating host physiology,
2. Influencing disease dynamics,
3. Interacting with immune responses, and
4. Enabling discovery of novel drugs and therapeutic agents.

Genomics and metagenomics are two technologies with complementary approaches to this pursuit. Metagenomics characterizes microbial communities and their functional genes within complex nest environments, while genomics focuses on individual organisms and their genetic composition. Genomics supports applications in drug discovery, whereas metagenomics advances environmental microbiome research and natural product exploration. To fully realize the therapeutic potential of stingless bee nest materials, metagenomics and genomics are needed in tandem.

Instead of morphological identifications of pollen grains by melissopalynology⁴⁹, DNA metabarcoding was proposed to characterize the botanical origin of honey¹³⁰ but a multi-kingdom resolution is now represented by a major high throughput technology of whole genome shotgun sequencing (WGS) known as shotgun metagenomics¹³¹ covering all DNA instead of primers, unlocking potential applications. A comprehensive view of the entire community of organisms and their genes is achieved by sequencing all DNA from a sample using this powerful tool for studying complex materials, fragmenting all the DNA, sequencing it, and using bioinformatics to identify the organisms present, their functional genes, and metabolic pathways, regardless of whether they are animals, bacteria, fungi, plants or viruses¹³²,¹³³. Amplicon sequencing and shotgun metagenomics are two platforms used in epidemiological research. Larger microbiome datasets characterize pooled amplicon/shotgun data compared to pure shotgun metagenomic. Usyk et al.¹³⁴ harmonized this pooling approach to leverage the exponential amplicon sequencing data produced over two decades.

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3.8 FRAUD CONTROL OF STINGLESS BEE PRODUCTS

Stingless bee products have lower yields than Apis mellifera because their colonies and nests are smaller, with variable size according to the stingless bee species and environmental factors. Productivity varies with years and some colonies are more propolizers than others. In Table 14, colony size and nest type of some stingless bee species from Brazil³³.

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Table 14. Colony size of selected stingless bee species

Stingless bee species (Maximum flight distance)	Bee size (mm)	Colony size Average No. workers (min–max)	Type of nest
Cephalotrigona capitata
[1.7 km]	9.5	1250 (1000–1500)	• Living tree cavities
• Horizontal brood combs with involucrum
• Rather large storage pots, with pot-honey and pot-pollen in different sections
• Permanent deposit of detritus
• Entrance decorated with plant resins and a hardened cerumen landing platform
Frieseomelitta varia
[1.4 km]	5.5	1200 (800–1600)	• Tree cavities
• Horizontal brood combs without involucrum
• Small spheroidal honey pots, larger pollen-pots slightly elongated
• Entrance size of worker head
Geotrigona mombuca
[unknown]	5.0	2500 (2000–3000)	• Underground
• Cylindrical storage pots, draining gallery at nest bottom, permanent deposits of detritus
• Underground channel connects the nest with the entrance in the surface

Due to the lower yields of pot-honey, pot-pollen and propolis, stingless bee products are known to have higher costs than Apis mellifera, and thus pot-honey has been imitated, mixed with honeybee products, sugars and syrups used in the bee fraud industry (P. Vit, personal observation). Pot-honey yields of Melipona scutellaris are 2–15 kg/year (R.M.O. Alves personal observation), Scaptotrigona spp 3 kg/year, and Tetragonisca angustula 1 kg/year¹⁴⁰ at about 100–200 USD/kg. Pot-pollen yields of Melipona scutellaris are 4–5 kg/year, Scaptotrigona spp. 5–6 kg/year, and Tetragonisca angustula 3–4 kg/year with a value from 32 to 257 USD/kg¹⁴¹.

Untargeted ¹H-NMR studies of sugar profiles of stingless bee honey and chemometrics are particularly useful to detect adulterations¹⁶². For certain entomological origins, the recently discovered trehalulose⁹³ is a potential sugar for genuine pot-honey authenticity⁷⁸. Targeted ¹H NMR revealed distinctive composition of pot-honey at genus level¹². A simple palynological screening would detect absence of pollen spectra in sugar syrups manufactured to fake honey¹². However, sophistication requires diverse bioanalytical techniques for pot-honey fraud control. For the expert, a simple sensory evaluation uncovers the adulteration, but consumers are not always acquainted with stingless bee nest materials as meliponicultors or a community with stingless bee apiaries or meliponaries. A honey authenticity test based on interface emulsion produced after shaking a honey dilution with diethyl ether, differentiated genuine from fake honey¹⁸⁴, and recently provided a further interpretation on suspected microbial associations with the Ecuadorian Scaptotrigona vitorum producing biosurfactants in pot-honey¹².

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Conclusions

The biotic materials processed to form pot-honey and pot-pollen have botanical, entomological, and microbial origins, along with plant resins that are essential for producing cerumen, a vital component of stingless bee nests. These nest materials contain diverse compounds and active metabolites that play crucial roles in their biological activities. Their antimicrobial and antioxidant properties contribute significantly to the added value of stingless bee nest materials, supporting their medicinal potential for both nutritional and pharmaceutical applications.

Research on the metabolites present in stingless bee nest materials and their ecological roles provides valuable insights into the complex relationships between stingless bees and their environment. Understanding the chemical composition of these materials may aid conservation efforts targeting both the pollinators and the ecosystems they sustain. The loss of stingless bee biodiversity would also mean the loss of chemical diversity in the bioactive metabolites within their nests, and, consequently, the loss of valuable natural healing molecules.

Our bibliometric analyses evaluated global scientific research on medicinal stingless bees (2004–2023) and stingless bees in climate change (2010–2023), using Bibliometrix to visualize datasets retrieved from the Scopus database. The medicinal dataset included 107 documents, showing greater research interest compared with the climate change dataset, which contained 25 documents. Top researchers, institutions, sponsors were identified. Hilgert N.I. (Argentina) was the most prolific author in medicinal stingless bee research for her contributions in ethnomedicine, while Martins C.F. (Brazil) led climate change–related studies on habitat loss and pollinator protection, each of them contributing four publications in their respective areas.

Malaysia, Brazil, and Mexico were the top three countries contributing with 62 of 107 documents for medicinal stingless bee research; and for stingless bees in climate change, whereas Brazil, the United States, and Indonesia accounted for 22 of 25 documents on stingless bees and climate change. The most prominent publication sources used to disseminate stingless bee research were the book Pot-Honey: A Legacy of Stingless Bees for the medicinal dataset, and the journal Apidologie for climate change research. Initiatives promoting pollinator protection within agricultural frameworks benefit both stingless bee conservation and the use of active metabolites of their nests in melipotherapy.

The role of microbiomes in metabolite transformation, the authenticity and chemical variability of nest materials, and the relevance of stingless bee biodiversity conservation areemphasized as interconnected drivers of medicinal potential and ecosystem sustainability. In bridging ethnomedicine, microbiology, chemistry, and environmental science, this paper provides a multidisciplinary framework that supports meliponitherapy as a promising field contributing to food security, health, and the Sustainable Development Goals (SDG 2 and SDG 3).

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Proposals for Future

Bibliometric reviews on the medicinal uses of stingless bee products for the human body systems need to be periodically updated, and medicinal research funded. Studies on medicinal properties should involve interactive collaboration among entomologists, chemists specializing in stingless bee products, melissopalynologists, and sensory scientists to establish standardized characterizations of raw materials and extracts used by experts in health sciences, experimental biology, and molecular biology. This multidisciplinary approach, which should also include statisticians, can provide a strong foundation for elucidating mechanisms of action over time.

Medicinal and climate change approaches interact over the long term. Brazil, which harbors the greatest stingless bee biodiversity for a country (259 of 605 recognized species worldwide), manages 95 species across its five regions, 12 of which are featured in the A.B.E.L.H.A. catalog, with sizes ranging from 3.5 to 10.5 mm. It is important to note that research on stingless bee nest materials remains ongoing. Understanding the specific active metabolites and their potential applications for human health continues to evolve framed in the Sustainable Development Goals SDG2 food security and SDG3 good health and well-being.

Stingless bees yield increasing amounts of medicinal pot-honey, pot-pollen, cerumen, and propolis, and their derivative products are being progressively developed. The conservation of natural ecosystems and the establishment of stingless bee–friendly managed environments are vital strategies for biodiversity preservation and sustainable meliponiculture, particularly in anticipation of shifting geographical distributions caused by climate change.

Reviews focusing on chemical classes are valuable for developing databases, updating bioanalytical methods, and revising reported units in the scientific literature. For instance, After reviewing aliphatic organic acids in honey and pot-honey, Vit and Simova⁹⁶ provided updated reference concentration values to replace the underestimated 0.5% AOA content commonly cited in scientific literature. Access to public databases that collect, verify, publish, and maintain DNA sequences globally¹³⁵ will be essential for advancing the study of stingless bee nest materials through expert curation, facilitating data exchange, and understanding. Moreover, targeting outer membrane components to enhance antibiotic permeability in Gram-negative bacteria could represent an innovative strategy to curb antimicrobial resistance, similar to recent insights into fungal capsule structures for approaches on novel disease control¹³⁶.

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Conflict of Interest Statement:

The authors have no conflicts of interest to declare.

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Funding Statement:

None.

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Acknowledgements:

To stingless bee keepers, conservationists, and meliponitherapists of the world. To our families for their unconditional support. To Dr. Cristina Mateescu, President of the Apitherapy Commission, for her invitation to the 48th Apimondia Congress September 4–8, 2023 held in Santiago de Chile, and the Chilean hospitality. To Prof. Breno Freitas from Universidade Federal de Ceará, Fortaleza, Brazil for the A.B.E.L.H.A. link. To our academic institutions for their constant support on stingless bee medicinal research. To stingless bee keepers of the world for their commitment with science, and our families for their steadfast encouragement. To native tropical cultures using meliponitherapy following ancestral knowledge. To modern laboratories interested in stingless bee nest materials to advance in the knowledge of their medicinal applications. To the United Nations University–Biotechnology for Latin America and the Caribbean UNU-BIOLAC for the Research Fellowship at the University of California Riverside to P. Vit, and benefactor Edward RH McDowell Jr.

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