Medicinal food plants of Sabah (Eastern Malaysia): a source of potential natural products and nutraceuticals for the fight against cancer

Carynn Tanbuda; Mazdida Sulaiman; Pauline Yong Pau Lin; Fiffy Hasnidah Saikim; Nor Azizun Ruzdi; Mogana Rajagopal; Nicholas Pang Tze Ping; Nor Hayati Abdullah; Jhonnel Villegas; Sanen Marshall; Mukesh Singh Sikarwar; Veeranoot Nissapatorn; Prapairat Seephonkai; Marcelo Iriti; Mark S Butler; Christophe Wiart

doi:10.1080/13880209.2026.2663269

. 2026 May 7;64(1):668–724. doi: 10.1080/13880209.2026.2663269

Medicinal food plants of Sabah (Eastern Malaysia): a source of potential natural products and nutraceuticals for the fight against cancer

Carynn Tanbuda ^a, Mazdida Sulaiman ^b, Pauline Yong Pau Lin ^c, Fiffy Hasnidah Saikim ^a, Nor Azizun Ruzdi ^a, Mogana Rajagopal ^d, Nicholas Pang Tze Ping ^e, Nor Hayati Abdullah ^f, Jhonnel Villegas ^g,^h, Sanen Marshall ⁱ, Mukesh Singh Sikarwar ^j, Veeranoot Nissapatorn ^k, Prapairat Seephonkai ^l, Marcelo Iriti ^m, Mark S Butler ⁿ, Christophe Wiart ^b,^✉

^{^a}Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, jalan UMS, Kota Kinabalu, Sabah, Malaysia

^{^b}Department of Chemistry, Faculty of Science, Universiti Malaya, Kuala Lumpur, Malaysia

^{^c}Faculty of Social Sciences and Humanities, Universiti Malaysia Sabah, Jalan UMS, Kota Kinabalu, Sabah, Malaysia

^{^d}Faculty of Pharmaceutical Sciences, UCSI University, Cheras, Kuala Lumpur, Malaysia

^{^e}Faculty of Medicine, Universiti Malaysia Sabah, jalan UMS, Kota Kinabalu, Sabah, Malaysia

^{^f}Natural Product Division, Forest Research Institute of Malaysia, Kepong, Kuala Lumpur, Malaysia

^{^g}Faculty of Teacher Education, Davao Oriental State University, Davao Oriental, Philippines

^{^h}Faculty of Tropical AgriSciences, Czech University of Life Sciences Prague, Praha-Suchdol, Czech Republic

^ⁱCenter for the Promotion of Knowledge and Language Learning, Universiti Malaysia Sabah, jalan UMS, Kota Kinabalu, Sabah, Malaysia

^{^j}Amity Institute of Pharmacy, Amity University, Gwalior, Madhya Pradesh, India

^{^k}Futuristic Science Research Center (FSRC) - School of Science, Research Excellence Center for Innovation and Health Products (RECIHP) and World Union for Herbal Drug Discovery (WUHeDD), Walailak University, Nakhon Si Thammarat, Thailand

^{^l}Department of Chemistry, Faculty of Science, Mahasarakham University, Maha Sarakham, Thailand

^{^m}Department of Biomedical, Surgical and Dental Sciences, University of Milan, Milan, Italy

^ⁿMSBChem Consulting, Brisbane, QLD, Australia

^✉

CONTACT Christophe Wiart asianjpharmcog@gmail.com Institute of Tropical Biology and Conservation, University Malaysia Sabah, 88400, Kota Kinabalu, Malaysia.

Roles

Carynn Tanbuda: Methodology, Software

Mazdida Sulaiman: Investigation

Pauline Yong Pau Lin: Investigation

Fiffy Hasnidah Saikim: Methodology

Nor Azizun Ruzdi: Writing – original draft

Mogana Rajagopal: Software

Nicholas Pang Tze Ping: Writing – review & editing

Nor Hayati Abdullah: Conceptualization

Jhonnel Villegas: Writing – review & editing

Sanen Marshall: Investigation

Mukesh Singh Sikarwar: Investigation

Veeranoot Nissapatorn: Writing – original draft

Prapairat Seephonkai: Writing – review & editing

Marcelo Iriti: Writing – review & editing

Mark S Butler: Project administration, Writing – review & editing

Christophe Wiart: Supervision, Writing – review & editing

PMCID: PMC13159609 PMID: 42095890

Abstract

Context

The pharmaceutical and nutraceutical industries are seeking structurally and pharmacologically novel anticancer molecules, as well as onco-protective nutraceuticals. One approach to achieving this goal is to study traditional pharmacopoeias, particularly those from regions where cancers are less common. Certain ethnic groups in Sabah (East Malaysia) appear to have a low incidence of cancer, and the study of their pharmacopeia could lead to the discovery of original anticancer molecules or nutraceuticals.

Objectives

This review presents a selection of 64 plants used for medicinal food in Sabah their potential for clinical uses.

Methods

The data for this focused narrative review were gathered from Google Scholar, PubMed, ScienceDirect, Web of Science, PubMed, the Internet Archive, and Google books. For each plant the search string included the binomial denomination and the words “cytotoxic” or “tumors.” of the binomial denomination of each plant and “cytotoxic” and “tumors” was employed. Each result was examined and articles that did not contain information relevant to the topic or coming from non-peer-reviewed journals were excluded.

Results

Eight plant species, of which Helminthostachys zeylanica (L.) Hook., Pycnarrhena tumefacta Miers, Myrmecodia platytyrea Becc., and Mangifera pajang Kosterm, demonstrate activities in vitro and in vivo, which call for further research. Others constitute a source of cytotoxic natural products that warrant further investigation.

Conclusion

There is currently a need to find oncopreventive nutraceuticals as well as original natural products for developing anticancer drugs. Such products could potentially be found among the medicinal and edible plants of Sabah. Further studies are needed.

Keywords: Borneo, ethnopharmacology, natural products, oncology

Introduction

Cancer is the second leading cause of death in most regions of the world (Bray et al. 2021), and was responsible for approximately 10 million deaths in 2020 (WHO 2022). By 2050, the number of cancers worldwide will reach 35.3 million, with developing countries being the most affected (Bizuayehu et al. 2024). Although some cancers are treatable when diagnosed early, the prognosis for some remains poor, and it is more necessary than ever to find anticancer drugs.

Several anticancer drugs originate from the plant kingdom. These tumoricidal compounds include vinblastine (Hodgkin’s lymphoma), podophyllotoxin (breast cancer), paclitaxel (Taxol) (ovarian and breast cancer), and homoharringtonine (leukemia) (Yang et al. 2025). These substances, derived from the secondary metabolism of plants, have been used in the semi synthesis of anticancer drugs such as vinorelbine (breast and ovarian cancer) from vinblastine, cabazitaxel (prostate cancer) from paclitaxel (Yang et al. 2025), and Sacituzumab Tirumotecan (breast cancer) from camptothecin (Butler et al. 2026). Many semi-synthetic camptothecin derivatives are the subject of clinical trials (Butler et al. 2026) including ABBV-969, for the treatment of prostate cancer (Tolcher et al. 2025).

Another important aspect in the fight against cancer is prevention, or what could be called “onco-prevention”. It is known that some cancers develop after chronic exposure to carcinogenic substances such as 7,12-dimethylbenz[a]anthracene (tobacco, industrial chemicals) (Rodgman and Perfetti 2006), glyphosate (food) (Wogan et al. 2004; Bai and Ogbourne 2016), as well as through chronic inflammation (Multhoff et al. 2011), oxidative stress (Jelic et al. 2021), and in cases of immunodeficiency (Pai et al. 2021). Therefore, consuming medicinal foods or plants with antimutagenic, anti-inflammatory, and/or immunostimulatory properties could be a way to prevent cancer or even extend the life expectancy of patients.

In some communities, the incidence of cancer is low, or even non-existent, even among the elderly. This has been observed in Sudan, Gambia, Sri Lanka, and among American Indians (Levin 1910). This phenomenon also appears to exist in Sabah (Eastern Malaysia) among local populations belonging to Bornean-speaking ethnic groups, including the Dusun, Kadazan and Murut, as well as among non-Bornean-speaking ethnic groups such as the Lundayeh, Brunei and Rungus (King and King 1984, Kroeger and Kroeger 1986; Wiart 2024; Tanbuda et al. 2025). These ethnic groups are distributed across areas of high biodiversity and high rates of plant endemism (Beaman and Beaman 1990). To date, approximately 696 species of medicinal plants have been recorded in Sabah, including a significant number of endemic species.

In this context, this review aims to present 64 species of medicinal food plants from Sabah whose phytochemical study could lead to the development of anticancer drugs or whose consumption in the form of nutraceutical products could prevent the appearance of cancer or allow people to live longer with cancer.

Methodology

In a previous study, we reviewed the 696 species of medicinal plants in Sabah, analyzing their taxonomical distribution and utilization among ethnic groups (Tanbuda et al. 2025). Out of these species we selected 64 medicinal food plant species for which there are few or no phytopharmacological studies and which belong to families known to produce cytotoxic natural products. This study is a focused narrative review, based on structured literature searches conducted for each selected species. The approach aims to provide an integrated overview of ethnobotanical uses, phytochemistry, and pharmacological activities, rather than meeting the criteria of a formal systematic review. For each plant covered, a search was conducted using Google Scholar, PubMed, ScienceDirect, Web of Science, Internet Archive, and Google Books. The search string included the binomial denomination of each plant and the words “cytotoxic” or “tumors.” If entering a binomial denomination and the words “cytotoxic” or “tumors” did not yield any results, the process was repeated with the genus name. The inclusion criteria focused on articles, conference proceedings, and books specifically addressing the selected medicinal food plant species. Exclusion criteria included articles, conference proceedings, and books related to medicinal plants from regions or countries outside Sabah, as well as non-English works or those not peer-reviewed.

Plants used by multiple ethnic groups

General observations

Most of the plants employed for medicinal purposes by multiple ethnic groups are not endemic, except for some of those used by both Dusun and Kadazan (Table 1 and 2).

Table 1.

Medicinal plants of Sabah.

[Subclass]	Genus, species, authority	Symptoms/diseases (ethnic group)	local names	(references)
[Superorder]
Order
Family
[Lycopodiidae Bek. (1862) (Lycophytes)]
Selaginellales Prantl (1854)
Selaginellaceae Willk. (1854)	Selaginella argentea (Wall. ex Hook. & Grev.) Spring^†ψ	Asthma, body aches, fever, headaches (Murut)	sondotnulogo	(Ahmad and Raji 19 1992; Kulip 2003)
		Medicinal (Bonggi-Molbog)	ipa ipa puteh	(Lin 2022)
Ophioglossidae Klinge (1832)
Ophioglossales Link. (1833)
Ophioglossaceae Martinov (1820)	Helminthostachys zeylanica (L.) Hook.	Cancer, wounds (Lundayeh)	pajerok	(K Kulip 2007)
		Postpartum (Bonggi-Mogbol)	onitug	(Lin 2022)
		Medicinal food (Dusun)	aruk aruk
[Polypodiidae Cronquist, Takht. & W. Zimm. (1966)]
Blechnales Pic. Sem. ex Reveal (1993)
Blechnaceae Newmann (1844)	Stenochlaena palustris (Burm. f.) Bedd.	Medicinal food, postpartum, fever, skin diseases (Dusun)	lambiding	(Maid et al. 2017)
		Postpartum, fever, skin diseases (Kadazan)	lambiding
Gleicheniales Link (1825)
Gleicheniaceae C. Presl (1825)	Gleichenia truncata (Willd.) Sprain^†ψ	Sore eyes (Dusun)	laputong	(Kulip 2014)
Polypodiales Link (1833)
Athyriaceae Alston (1956)	Diplazium cordifolium Bl.^†ψ	Cold, fever (Bajau)	giman	(Wiart 2024)
	Diplazium esculentum (Retz.) Sw.^ψ	Medicinal food (Dusun)	pakis	(Maid et al. 2017)
Polypodiaceae Link J. Presl & C. Presl (1822)	Drymoglossum piloselloides (L.) C. Presl^†ψ	Diuretic, gallstones, hypertension (Brunei)	sisik naga	(Mahmud and Razali 2016)
	Drynaria sparsisora (Desv.) T. Moore ^ψ	Asthma, heart diseases (Dusun)	tapako	(Wiart 2024)
		Asthma, heart diseases (Kadazan)	tapako	(Wiart 2024)
Schizaeales Schimp. (1869)
Lygodiaceae M. Roem. (1840)	Lygodium circinnatum Sw.^†ψ	Venereal diseases (Lundayeh)	waratang
		Womb diseases (Dusun)	taribu mianai	(Voeks and Nyawa 2 2006)
Nephrolepidaceae Pic. Serm. (1975)	Lygodium salicifolium C. Presl^†ψ	Chickenpox, smallpox (Lundayeh)	ubat amur
Pteridaceae E.D.M. Kirchn. (1831)	Nephrolepis acutifolia (Desv.) Christ ^ψ	Medicinal food (Dusun)	paku puteh
	Acrostichum aureum L.	Medicinal food (Dusun)	paku besar
[Gnetidae Pax (1894)]
Gnetales Blume (1835)
Gnetaceae Blume (1833)	Gnetum macrostachyum Hook.f.^ψ	Fatigue, postpartum (Brunei)	kokos	(Kulip 1997)
[Magnoliidae Novák ex Takht. (1967)]
(Austrobaileyanae Doweld ex M.W. Chase & Reveal (2009))
Austrobaileyales Takht. ex Reveal (1992)
Schisandraceae Blume (1830)	Kadsura borneensis A.C. Sm^†ψ	Cramps (Lundayeh)	putu urat
	Kadsura lanceolata King^†ψ	Bacterial skin infection, swelling (Dusun)	topis	(Wiart 2024)
[Magnolianae Takht. (1967)]
Laurales Juss. ex Bercht. & Presl (1820)
Lauraceae Juss. (1789)	Eusideroxylon zwageri Teijsm. & Binn.^†ψ	Blow-gun darts poison (Murut)	belian	(Kulip 2003)
	Litsea garciae Vidal. ^ψ	Dislocation, sprains (Murut)	pengolaban	(Wiart 2024)
		Medicinal food (Dusun)	pengolaban	(Maid et al. 2017)
Magnoliales Bromhead (1838)
Annonaceae Juss. (1789)	Artabotrys roseus Boerl.^†ψ	Medicinal (Dusun)	gangon
		Medicinal (Kadazan)	gangon
	Goniothalamus roseus Stapf^†ψ	Fatigue, fever (Dusun)	limpanas	(Foo et al. 2016)
	Goniothalamus velutinus Airy Shaw ^ψ	Magic rituals (Dusun)	kalampanas	(Voeks 2007)
	Goniothalamus woodii Merr ^†ψ	Magic rituals (Murut)	tampaliu	(Wiart 2024)
	Polyalthia tenuipes Merr.^†ψ	Sick children (Dusun)	kabanking	(Wiart 2024)
[Lilianae Takht. (1967)]
Commelinales Mirb. ex Bercht. & J. Presl (1820)
Eriocaulaceae Martinov (1820)	Eriocaulon longifolium Nees ex Kunth^†ψ	Canker sores (Dusun)	kumpau sambangau	(Voeks and Nyawa 2006)
Poaceae Barnhart (1895)	Dendrocalamus asper (Schult. f.) Backer ex K. Heyne	Medicinal food (Dusun)	buluh betong
	Garnotia acutigluma (Steud.) Ohwi^†ψ	Venereal diseases (Lundayeh)	udu bulu
	Panicum palmifolium J. Koenig^†ψ	Malaria (Dusun)	tandaki	(Maid et al. 2017; W Wiart 2024)
		Malaria (Kadazan)	tandaki	(Wiart 2024)
Zingiberaceae Martinov (1820)	Boesenbergia pulchella (Ridl.) Merr.^†ψ	Skin diseases (Dusun)	lipat	(Kulip 2007)
		Skin diseases (Kadazan)	lipat	(Kulip 2007)
	Plagiostachys albiflora Ridl.^†ψ	Medicinal food (Dusun)	wongking	(Kulip 2007)
	Etlingera elatior (Jack) R.M. Sm.	Fever, flatulence, medicinal food (Dusun)	topu	(Kulip 2007a)
		Fever, flatulence (Kadazan)	topu	(Kulip 2007a)
[Ranunculanae Takht. ex Reveal (1992)]
Ranunculales Juss. ex Bercht. & J. Presl (1820)
Menispermaceae Juss. (1789)	Pycnarrhena tumefacta Miers^†ψ	Bacterial skin infection (Lundayeh)	fatagah
		Medicinal food (Murut)	apa	(Wiart 2024)
		Medicinal food (Dusun)	apak
		Medicinal food (Kadazan)	apak
[Rosanae Takht. (1967)]
Vitales Juss. ex Bercht. & J. Presl (1820)
Vitaceae Juss. (1789)	Ampelocissus polita (Miq.) Pelser^†ψ	Medicinal food (Dusun)	mban ambuk	(Voeks and Nyawa 2006)
Fabales Bromhead (1838)
Fabaceae Lindley (1836)	Airyantha borneensis (Oliv.) Brummitt^†ψ	Fatigue (Dusun)	barayung	(Voeks and Nyawa 2006)
		Fever, hypertension, toothaches (Murut)	matamis	(Wiart 2024)
	Koompassia malaccensis Maing^†ψ	Allergy, asthma, bloating, body aches, convulsions, Blood in stools, gastritis, stomachaches, swollen gums, toothaches (Bajau)	raja kayu	(Foo et al. 2016)
	Millettia nieuwenhuisii J.J. Smith^†ψ	Thrush (Murut)	ramus	(Kulip 2003)
Pandaceae Engl. & Gilg (1913)	Galearia fulva (Tul.) Miq.^†ψ	Medicinal food (Dusun)	sanggara	(Voeks and Nyawa 2006)
Phyllanthaceae Martinov (1820)	Bridelia stipularis (L.) Bl.	Fever, diabetes, postpartum, thrush (Dusun)	belingkut	(Kulip et al. 2003)
		Fever, diabetes, postpartum, thrush (Kadazan)		(Kulip et al. 2003)
		Diabetes, thrush (Murut)	bolingkut	(Kulip et al. 2003)
Rosales Bercht. & Presl. (1820)
Moraceae Link (1831)	Ficus retusa L. ^ψ	Shivers (Dusun, Kadazan)	hintotobu	(Kulip 1997)
		Medicinal (Murut)	sialbon rindoh	(Wiart 2024)
Malvales Juss. ex Bercht. & J. Presl (1820)
Dipterocarpaceae Blume (1825)	Shorea parvistipulata F. Heim^†ψ	Fatigue (Murut)	roloi	(Kulip 2003)
Myrtales Juss. ex Bercht. & J. Presl (1820)
Melastomataceae Juss. (1789)	Dissochaeta monticola Bl.^†ψ	Blowgun darts poison (Murut)	bina	(Wiart 2024)
	Melastoma beccarianum Cogn.^†ψ	Blemishes (Dusun) ?	duduk abai	(Voeks and Nyawa 2006)
Sapindales Juss. ex Bercht. & J. Presl (1820)
Anacardiaceae R.Br. (1818)	Mangifera pajang Kosterm. ^ψ	Cancer, high cholesterol, medicinal food (Dusun)	bambangan	(Maid et al. 2017)
Burseraceae Kunth (1824)	Canarium littorale Bl.^†ψ	Medicinal food (Dusun)	adal	(Voeks and Nyawa 2006)
	Dacryodes incurvata (Engl.) H.J. Lam^†ψ	Medicinal food (Dusun)	nguluon	(Voeks and Nyawa 2006)
Sapindaceae Juss. (1789)	Guioa bijuga (Hiern) Radlk.^†ψ	Medicinal food (Rungus)	anggil	(Wiart 2024)
	Nephelium macrophyllum Radlk.^†ψ	Medicinal food, magic rituals (Dusun)	mbokot	(Voeks and Nyawa 2 2006)
	Nephelium uncinatum Radlk. ex Leenh.^†ψ	Medicinal food (Dusun)	kamanggis	(Voeks and Nyawa 2 2006)
[Caryophyllanae Takhtajan (1967)]
Caryophyllales Juss. ex Bercht. & J. Presl (1820)
Nepenthaceae Dumort. (1829)	Nepenthes ampullaria Jack ^ψ	Respiratory diseases (Lundayeh)	telungau becuk
[Asteranae Takht. (1967)]
Ericales Bercht. & J. Presl (1820)
Primulaceae Batsch ex Borkh (1797)	Embelia dasythyrsa Miq.^{σ ψ}	Fever, medicinal food (Dusun)	sowolikan	(Wiart 2024)
Symplocaceae Desf. (1820)	Symplocos odoratissima Choisy ex Zoll. ^ψ	Fever, malaria (Lundayeh)	lobo
Gentianales Juss. ex Bercht. & J. Presl (1820)
Asclepiadaceae Borkh (1797)	Dischidia rafflesiana Wall.^†ψ	Cancer, skin diseases (Bajau)	(Foo et al. 2016)
Rubiaceae Juss. (1789)	Chassalia chartacea Craib ^ψ	Blurred vision (Dusun)	lansi	(Wiart 2024)
	Hydnophytum formicarum Jack ^ψ	Cancer, diabetes, hypertension (Lundayeh)	sarang semut betina	(Wiart 2024)
	Ixora capillaris Bremek^†ψ	Medicinal (Rungus)	tagandap timulu	(Wiart 2024)
	Neonauclea gigantea (Valeton) Merr.^†ψ	Diarrhea, stomach aches, thrush (Dusun)	mahitap	(Wiart 2024)
	Myrmecodia platytyrea Becc.	Drunkenness, hypertension, poison antidote (Dusun)	rajah ubat	(Wiart 2024)
		Hypertension (Lundayeh)		(Wiart 2024)
		Diabetes (Murut)	sarang semut	(Wiart 2024)
		Cancer, diabetes, fever, headaches,	sarang semut	(Wiart 2024)
		hypertension, kidney diseases,
		poison antidote, sinusitis, tuberculosis (Bajau)
	Paederia verticillata Bl.^ψ	Intestinal worms (Dusun)	taud	(Wiart 2024)
		Intestinal worms (Kadazan)	taud	(Wiart 2024).
	Praravinia suberosa (Merr.) Bremek^†ψ	Medicinal (Murut)	kingkimu	(Kulip 2003)
	Psychotria gyrulosa Stapf^†ψ	Headaches (Dusun)	siroromuk	(Wiart 2024)
	Rennellia borneensis Baill.^†ψ	Medicinal	Sabah ginseng	(Wiart 2024)
Lamiales Bromhead (1838)
Oleaceae Hoffmannsegg et Link (1809)	Jasminum aculeatum Blco^†ψ	Flatulence (Murut)	onsom onsom	(Kulip 2003)
	Jasminum bifarium Wall^†ψ	Sore eyes (Lundayeh)	bunga melor	(Wiart 2024)
Solanales Juss. ex Bercht. & J. Presl (1820)
Convolvulaceae Juss. (1789)	Merremia gracilis E.J.F. Campb. & Argent^†ψ	Asthma, diarrhea, fatigue, jaundice, pancreatitis (Dusun)	malagatas	(Kulip 1997)
		Asthma, diarrhea, fatigue, jaundice, pancreatitis (Kadazan)	malagatas	(Kulip 1997)
	Merremia peltata (L.) Merr.	Diarrhea, flatulence, hair loss, babas (Kulip 2003; Kulip)	stomach aches, wounds (Dusun)	2014; Wiart 2024)
		Diarrhea, wounds (Kadazan)		(Kulip 2003)
Asterales Link (1829)
Asteraceae Martinov (1820)	Crassocephalum crepidioides (Benth.) S. Moore	Cancer (Murut)	kinsau	(Awang-Kanak and F Foo 2023)
		Ageing, medicinal food (Dusun)	koyundou	(Awang-Kanak and Foo 2023)

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^{^†}

: no cytotoxic study; ψ: no toxicological study.

Table 2.

Geographical distribution, habitat and cultivability.

[Subclass]
(Superorder)
Order
Family	Genus, species, authority	habitat	rarity	cultivability	(references)
[Lycopodiidae Bek. (1862) (Lycophytes)]
Selaginellales Prantl (1854)
Selaginellaceae Willk. (1854)	Selaginella argentea (Wall. ex Hook. & Grev.) Spring^σ	mossy grounds	not rare	cultivable	(Suranga et al., 2018)
Ophioglossales Link. (1833)
Ophioglossaceae Martinov (1820)	Helminthostachys zeylanica (L.) Hook.	Forest	rare	cultivable	(Um, 2010)
[Polypodiidae Cronquist, Takht. & W. Zimm. (1966)]
Blechnales Pic. Sem. ex Reveal (1993)
Blechnaceae Newmann (1844)	Stenochlaena palustris (Burm. f.) Bedd.	near mangroves	not rare	cultivable	(Uda et al., 2020)
Gleicheniales Link (1825)
Gleicheniaceae C. Presl (1825)	Gleichenia truncata (Willd.) Sprain	roadsides	not rare	?
Polypodiales Link (1833)
Athyriaceae Alston (1956)	Diplazium cordifolium Bl.	forest	rare	?
	Diplazium esculentum (Retz.) Sw.	riverbanks	not rare	cultivable	(Singh and Johari, 2018)
Polypodiaceae Link J. Presl & C. Presl (1822)	Drymoglossum piloselloides (L.) C. Presl	roadside	not rare	?
	Drynaria sparsisora (Desv.) T. Moore	roadside	not rare	?
Schizaeales Schimp. (1869)
Lygodiaceae M. Roem. (1840)	Lygodium circinnatum Sw.	edges of forests	not rare	cultivable	(Cunningham and Brinckmann, 2023)
	Lygodium salicifolium C. Presl	roadsides	not rare	?
Nephrolepidaceae Pic. Serm. (1975)	Nephrolepis acutifolia (Desv.) Christ	seashores	not rare	?
Pteridaceae E.D.M. Kirchn. (1831)	Acrostichum aureum L.	mangroves	not rare	?
[Gnetidae Pax (1894])
Gnetales Blume (1835)
Gnetaceae Blume (1833)	Gnetum macrostachyum Hook.f.^σµ	forest	rare	?
[Magnoliidae Novák ex Takht. (1967)]
(Austrobaileyanae Doweld ex M.W. Chase & Reveal (2009))
Austrobaileyales Takht. ex Reveal (1992)
Schisandraceae Blume (1830)	Kadsura borneensis A.C. Sm^β	mountain	rare	?
	Kadsura lanceolata King^σϕ	mountain	rare	?
(Magnolianae Takht. (1967))
Laurales Juss. ex Bercht. & Presl (1820)
Lauraceae Juss. (1789)	Eusideroxylon zwageri Teijsm. & Binn.^σ	forest	rare	cultivable	(Irawan 2012)
	Litsea garciae Vidal.^βπ	forest	rare	cultivable	(Ekamawanti et al., 2023)
Magnoliales Bromhead (1838)
Annonaceae Juss. (1789)	Artabotrys roseus Boerl.^β	forest	rare	?
	Goniothalamus roseus Stapf^β	forest	rare	?
	Goniothalamus velutinus Airy Shaw^β	forest	rare	?
	Goniothalamus woodii Merr ^β	forest	rare	?
	Polyalthia tenuipes Merr.^βπ	forest	rare	?
(Lilianae Takht. (1967))
Commelinales Mirb. ex Bercht. & J. Presl (1820)
Eriocaulaceae Martinov (1820)	Eriocaulon longifolium Nees ex Kunth	rice paddy	not rare	?
Poaceae Barnhart (1895)	Dendrocalamus asper (Schult. f.) Backer ex K. Heyne	village	not rare	cultivable	(Singh et al., 2004)
	Garnotia acutigluma (Steud.) Ohwi	mountain slopes	not rare	?
	Panicum palmifolium J. Koenig	open forest	not rare	?
Zingiberaceae Martinov (1820)	Boesenbergia pulchella (Ridl.) Merr.^β	forest	rare	cultivable	(Saensouk et al., 2025)
	Plagiostachys albiflora Ridl.^σ	forest	rare	?
(Ranunculanae Takht. ex Reveal (1992))
Ranunculales Juss. ex Bercht. & J. Presl (1820)
Menispermaceae Juss. (1789)	Pycnarrhena tumefacta Miers^σπϕµ	forest	rare	cultivable
(Rosanae Takht. (1967])
Vitales Juss. ex Bercht. & J. Presl (1820)
Vitaceae Juss. (1789)	Ampelocissus polita (Miq.) Pelser	forest	rare	?
Fabales Bromhead (1838)
Fabaceae Lindley (1836)	Airyantha borneensis (Oliv.) Brummitt^βπ	forest	rare	?
	Koompassia malaccensis Maing^σ	forest	rare	?
	Millettia nieuwenhuisii J.J. Smith^β	forest	rare	?
Pandaceae Engl. & Gilg (1913)	Galearia fulva (Tul.) Miq.	Forest	rare	?
Phyllanthaceae Martinov (1820)	Bridelia stipularis (L.) Bl.	Forest	rare	?
Rosales Bercht. & Presl. (1820)
Moraceae Link (1831)	Ficus retusa L.	gardens	not rare	cultivable	(Adeoluwa et al., 2014)
Malvales Juss. ex Bercht. & J. Presl (1820)
Dipterocarpaceae Blume (1825)	Shorea parvistipulata F. Heim^β	forests	rare	cultivable	(Susanty, 2019)
Myrtales Juss. ex Bercht. & J. Presl (1820)
Melastomataceae Juss. (1789)	Dissochaeta monticola Bl.^σ	forest	rare	?
	Melastoma beccarianum Cogn.^β	open land	rare	?
Sapindales Juss. ex Bercht. & J. Presl (1820)
Anacardiaceae R.Br. (1818	Mangifera pajang Kosterm.^β	forest	cultivable		(Tinggal and Tee, 1994)
Burseraceae Kunth (1824)	Canarium littorale Bl.^†	swamp forest	rare	?
	Dacryodes incurvata (Engl.) H.J. Lam^σπ	forest	rare	?
Sapindaceae Juss. (1789)	Guioa bijuga (Hiern) Radlk.^σπ	forest	rare	?
	Nephelium macrophyllum Radlk.^β	forest	rare	?
	Nephelium uncinatum Radlk. ex Leenh.^σ	forest	rare	cultivable	(Matius et al., 1998)
(Caryophyllanae Takhtajan (1967])
Caryophyllales Juss. ex Bercht. & J. Presl (1820)
Nepenthaceae Dumort. (1829)	Nepenthes ampullaria Jack^σπϕµ	peat swamp forest	rare	cultivable	(Isnaini, and Novitasari, 2023)
(Asteranae Takht. (1967))
Ericales Bercht. & J. Presl (1820)
Primulaceae Batsch ex Borkh (1797)	Embelia dasythyrsa Miq.^σ	forest	rare	?
Symplocaceae Desf. (1820)	Symplocos odoratissima Choisy ex Zoll.^σπϕ	forest	rare	?
Gentianales Juss. ex Bercht. & J. Presl (1820)
Asclepiadaceae Borkh (1797)	Dischidia rafflesiana Wall.	Swamps	rare	cultivable	(Scott and Sargant, 1893)
Rubiaceae Juss. (1789)	Chassalia chartacea Craib	forest	rare	?
	Hydnophytum formicarum Jack	seashores	rare	cultivable	(Huxley, 1978)
	Ixora capillaris Bremek^β	coastal forest	rare	?
	Neonauclea gigantea (Valeton) Merr.^β	open forest	rare	?
	Myrmecodia platytyrea Becc.	open forest	not rare	cultivable
	Paederia verticillata Bl.^σπϕ	forest	rare	?
	Praravinia suberosa (Merr.) Bremek^β	forest	rare	?
	Psychotria gyrulosa Stapf^β	forest	rare	?
	Rennellia borneensis Baill.^β	forest	rare	?
Lamiales Bromhead (1838)
Oleaceae Hoffmannsegg et Link (1809)	Jasminum aculeatum Blco^βπϕ	coastal forest	rare	?
	Jasminum bifarium Wall	forest	rare	?
Solanales Juss. ex Bercht. & J. Presl (1820)
Convolvulaceae Juss. (1789)	Merremia gracilis E.J.F. Campb. & Argent	forest	rare	?
	Merremia peltata (L.) Merr.	Forest	rare	cultivable	(Kirkham, 2004)
Asteraceae Martinov (1820)	Crassocephalum crepidioides (Benth.) S. Moore	roadsides	not rare	cultivable	(Dossou et al., 2019)

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β: Endemic solely in Borneo; σ: Sundaland; ϕ: Wallacea; π: Philippines; µ: Sahuland.

Selaginella argentea (wall. ex-hook. & grev.) spring (Selaginellaceae)

Plants in the genus Selaginella P. Beauv. (1804) produce cytotoxic phenolic principles (Figure 1). We can cite the biflavone (2S)-2,3-dihydroametoflavone 5,4′-dimethyl ether (1) and flavanone seladoeflavone E (2) (K562; IC₅₀: 8.1 µM) from Selaginella doederleinii Hieron (Zou et al. 2017; Liu et al. 2021). Other examples are (2″S)-2″,3″-dihydroochnaflavone (3) and robustaflavone 7,5′′-dimethyl ether (4) from Selaginella trichoclada Halston active against the breast cancer cell line (MCF-7) with IC₅₀ values of 7.7 and 6.9 μM, respectively (Xie et al. 2022). From the aerial parts of Selaginella delicatula (Desv.) Alston (collected in Taiwan), robustaflavone 7,4′,4′′′-trimethyl ether (5), robustaflavone 4′,4″′-dimethyl ether (6), and 2,3-dihydroamentoflavone 7,4′-dimethyl ether (7) were found to be cytotoxic to murine lymphocytic leukemia cell line (P388) with IC₅₀ values of 4.7, 1.4, and 3.5 µg/mL, repectively (Chen et al. 2005). We can also cite isocryptomerin (8) from the leaves of Selaginella willdenowii (Desv.) Baker (collected in Panama), (ZR-75-1; IC₅₀: 0.5 µg/mL) (Silva et al. 1995), some chalcone-flavanone biflavonoids from Selaginella trichoclada Alston (Xie et al. 2022), and ginkgetin (9) from Selaginella moellendorffii Hieron (OVCAR-3: IC₅₀:1.8 µg/mL) (Sun et al. 1997).

Structural formulas of various flavonoids and related compounds, including seladoeflavone E and robustaflavone derivatives. The figure showcases a collection of chemical structures representing various flavonoids and related compounds arranged in a grid. Key compounds include seladoeflavone E (2), robustaflavones (4-6) with distinct substitutions, isocryptomerin (8), and ginkgetin (9). Other notable structures are 7ß-hydroxycholesterol (12), selaginedorffone B (10), and selaginellin M (11). Each structure features functional groups such as hydroxyl (-OH) and methoxy (-OCH3), illustrating the diversity in their chemical connectivity and configuration. — Cytotoxic natural products of plants of the genus *Selaginella.*

Other cytotoxic principles in this genus are abietane-type diterpenes (Wu et al. 2020) such as selaginedorffone B (10) from S. moellendorffii (MCF-7; IC₅₀: 9 μM) (Ke et al. 2018), phenolic compounds as in selaginellin M (11) (HeLa; IC₅₀: 28.5 μM) (Yang et al. 2012), 7β-hydroxycholesterol (12) (HT-115; IC₅₀: 2.7 μg/mL) (Roh et al. 2010), selagibenzophenone E (13) (a benzophenone) (SMCC-7721; IC₅₀: 15.8 µM) (Long et al. 2023), 8-methyleugenitol (14) (a chromone) from Selaginella siamensis Hieron. (HuCCA-1; IC₅₀: 20.8 µM) (Thamnarak et al. 2022), and cyclopeptides (Yan et al. 2022). To date, no phytochemical, pharmacological, or toxicological studies appear to have been conducted on S. argentea. This plant is cultivable (Table 2).

Helminthostachys zeylanica (L.) hook. (Ophioglossaceae)

Anticancer properties have been demonstrated in vitro and involve a series of cytotoxic and anti-inflammatory prenylated flavonoids, rare monoterpene-flavonoid meroterpenes (Figure 2). An ethyl acetate fraction, at the concentration of 80 µg/mL, induced the apoptosis of gastric adenocarcinoma cell line (AGS) with cleavage of poly (ADP-ribose) polymerase, decreased expression of Bcl-2 and cyclooxygenase 2 (Tsai et al. 2021). The plant produces anti-inflammatory prenylated flavanones such as neougonin A (15), which inhibited lipopolysaccharide-induced nitric oxide production in mouse monocyte macrophages (RAW264.7) cell line, with an IC₅₀ value of 3.3 µM (Cao et al. 2016).

Six labeled chemical structures: neougonin A, ugonin S, ugonin U, ugonin K, ugonin V, and ugonstilbene A. The figure illustrates six labeled chemical structures in two rows: neougonin A (15) features multiple hydroxyl groups; ugonin S (16) shows a spiro structure with hydroxyls; ugonin U (17) contains a benzene system with hydroxyls; ugonin K (18) includes a methoxy group and hydroxyls; ugonin V (19) is a xanthone with hydroxyls; ugonstilbene A (20) displays a trans-stilbene anatomy with complex side chains. Each structure is numbered for identification. — Cytotoxic natural products of *H. zeylanica.*

Other principles are prenylated flavones such as ugonin S (16) and U (17) (Huang et al. 2009; Su et al. 2016), cytotoxic to acute lymphoblastic leukemia cell line (CEM) and lung cancer cell line (H460) (Lin et al. 2025). Ugonin K (18) inhibited the growth of basal cell carcinoma cell line via reduction of mitochondrial membrane potential, the expression of p53, and subsequent activation of caspases-8, −9, and −3 leading to apoptosis (Chan et al. 2013). Ugonin V (19) given intraperitoneally thrice weekly for 4 weeks at a dose of 15 mg/kg inhibited metastasis of chondrosarcoma tumors in lungs of mice, via at the tumor cellular level and inhibition of cathepsin V expression (Tran et al. 2025). Other cytotoxic principles in this fern are prenylated stilbenes such as ugonstilbene A (20) (Lin et al. 2023). This fern is used as a medicinal food by the Dusun and as a remedy for cancer by the Lundayeh (Table 1). In a clinical study, patients receiving rhizome powder (1 g three times daily for 42 days) experienced no adverse effects. (Su et al. 2022). This fern is cultivable (Table 2). Clinical trials are needed to confirm its efficacy and safety.

Stenochlaena palustris (burm. f.) bedd. (Blechnaceae)

The Dusun use this fern as a medicinal food (Wiart, 2024) (Table 1). The ethanolic extract of its leaves (harvested in Malaysia) has been shown to be toxic to cervical cancer cell line (HeLa) (IC₅₀: 5.8 µg/mL) (Arullappan et al. 2017). This fern produces flavone glycosides, including rutin (21), as well as acylated flavone glycosides (Hendra et al. 2024) (Figure 3). Rutin (21) has been shown to be moderately cytotoxic to malignant melanoma cell line (RPMI-7951) and (SK-MEL-28), with IC₅₀ values of 64.4 and 47.4 µM, respectively (Pinzaru et al. 2021). Kaempferol 3-O-(3″-O-E-p-coumaroyl)-(6″-O-E-feruloyl)-β-D-glucopyranoside (22) and kaempferol 3-O-(3″,6″di-O-E-p-coumaroyl)-β-D-glucopyranoside (or ditiliroside) (23) from this fern inhibited the proliferation of breast cancer cell line (MDA-MB-231) with IC₅₀ values of 70 and 21 µM, respectively (Chear et al. 2019). The toxicity of S. palustris remains unstudied. It is cultivable (Table 2).

Chemical structures of rutin (21) and two kaempferol derivatives (22, 23) with distinct functional groups. The image illustrates three chemical structures: rutin (21) features hydroxyl groups and a glucopyranose unit linked to a coumaroyl moiety. The first kaempferol derivative (22) incorporates methoxy substitutions, while the second derivative (23) has hydroxyl groups without additional substitutions. Each structure highlights specific molecular arrangements and bonding characteristics among the compounds. — Cytotoxic flavonoids of *S. palustris.*

Drynaria sparsisora (desv.) T. Moor

Aqueous extract of Drynaria quercifolia (L.) J.Sm. and Drynaria fortunei (Kunze ex Mett.) J.Sm. were found to be cytotoxic to brine-shrimp s (LC₅₀: 7.6 µg/mL) (Runa et al. 2013) and weakly active against MDA-MB-231 cell line (IC₅₀ ≈ 500 µg/mL) (Telang et al. 2025). Methanol extract of D. quercifolia (L.) J.Sm. inhibited the growth of hepatocellular carcinoma (HepG2) cell line (IC₅₀ ≈ 200 µg/mL) (Prasanna et al. 2019). A dichloromethane extract of Drynaria rigidula (Sw.) Bedd. was found to be cytotoxic to MCF-7 cell line (IC₅₀: 18 µg/mL) (Nugraha et al. 2019). From this fern, a chiratane-type triterpene, chiratone (24), was found to inhibit the growth of prostate cancer cell line (PC3) (IC₅₀: 1 μM) (Liang et al. 2010) (Figure 4). Isolated from this fern’s rhizome, the long-chain alkylated lignan (+)-liglaurate A (25) abrogated the survival of cervical cancer cell line (HeLa) (IC₅₀: 0.1 μM) (Wufuer et al. 2022). In Sulawesi, a plant in the genus Drynaria (Bory) J. Sm. (1841) has been used to treat cancer (Nurrani et al. 2014). To date, no phytochemical, pharmacological, or toxicological studies appear to have been conducted on D. sparsisora.

Two chemical structures: chiratone (24) on the left; racemic liglaurate A (25) on the right, with functional groups labeled. The figure presents two chemical structures side by side. On the left, chiratone (24) features interconnected tetracyclic carbon rings with hydroxyl (OH) groups and indicates stereochemistry. On the right, racemic liglaurate A (25) showcases a complex arrangement, including two phenolic hydroxyls, a carbon backbone, and a racemic notation with ester chains. Labels indicate structure names and numbers. — Cytotoxic natural products of plants of the genus *Drynaria.*

Lygodium circinnatum Sw. (Lygodiaceae)

There are a number of studies that demonstrate the cytotoxic activity of organic extracts from ferns of the genus Lygodium Sw. (1801): Lygodium venustum Sw. (ethanol leaves; NCTC929; IC₅₀: 500 µg/mL) (Morais-Braga et al. 2013), Lygodium microphyllum (Cav.) R. Br. (ethyl acetate; P388; IC₅₀: 50.1 µg/mL) (Kuncoro 2017) and Lygodium flexuosum (L.) Sw. (hexane; Hep3B; IC₅₀: 32 µg/mL). In the latter case, apoptosis was observed with cleavage of the poly(ADP-ribose) polymerase (Wills and Asha 2009).

Plants in this genus produce ecdysteroids and their glycosides (Guo-Gang et al. 2012), and flavonol glycosides, including isoquercetin (26) (Kuncoro et al. 2017) (Figure 5). Isoquercetin (26) (Figure 5), given orally to mice, xenografted with colorectal adenocarcinoma cell line (HT-29), at a dose of 17 µg/g over a week, caused a decrease in tumor volume (da Silva et al. 2022). Other principles in these ferns are naphthoquinones (Chen et al. 2010), triterpenes (Han et al. 2012), phenylpropanoid glycosides (Duan et al. 2012), and diterpenes (Yamauchi et al. 1996). To date, no phytochemical, pharmacological, or toxicological studies appear to have been conducted on L. circinnatum. This fern is cultivable (Table 2).

Chemical structure of isoquercetin, featuring hydroxyl groups, benzene rings, and a glucose moiety. The figure illustrates the chemical structure of isoquercetin, showcasing a complex arrangement of two benzene rings linked through ether connections. It includes multiple hydroxyl groups (–OH) attached to the rings, highlighting its polyphenolic nature. A glucose moiety with a six-membered carbon ring is positioned at the bottom, displaying hydroxyl groups at various sites. The structural notation “isoquercetin (26)” is clearly labeled beneath, aiding in identification of the compound and its functional groups. — Cytotoxic flavonol glycoside of plants of the genus *Lygodium.*

Litsea garciae vidal

A methanolic extract of this plant has been shown to be cytotoxic against various cancer cell lines (Kutoi et al. 2012), but to date, its active principles remain unknown. Plants of the genus Litsea Lam (1792) produce different types of natural cytotoxic substances such as litseaone A (27) and B (28) (chalcones) (Kageji et al. 2018; Kageji et al. 2019) from Litsea rubescens Lecomte (HL-60; IC₅₀: of 6.1 and 10.2 µg/mL, respectively) (Li et al. 2011) and from Litsea cubeba (Lour.) Pers, arctigenin (29) (a dibenzylbutyrolactone lignan), erythro-2,3-bis(4-hydroxy-3-methoxyphenyl)-3-ethoxypropan-1-ol (30) (a stilbene) (Guo et al. 2015; Li et al. 2019), (+)-N-(methoxycarbonyl)-N-norbulbodione (31) (an aporphine alkaloid) (HepG2; IC₅₀: 9.5 µM) (Zhang et al. 2012), and N-methoxycarbonyl-norjuziphine (32) (benzylisoquinoline alkaloid) (MCF-7; IC₅₀:15 μM) (Tang et al. 2017) (Figure 6).

Chemical structures of compounds, including litseone A, litseone B, arctigenin, and others, labeled with identification numbers. The figure displays nine chemical structures of organic compounds arranged in a grid. The top row features litseone A (27), litseone B (28), and arctigenin (29), showcasing various functional groups. The second row includes erythro-2,3-bis(4-hydroxy-3-methoxyphenyl)-3-ethoxypropan-1-ol (30), (+)-N-(methoxycarbonyl)-N-norbulbodione (31), and N-methoxycarbonyl-norjuzipine (32). The bottom row presents bisuegenol A (33), N-trans-sinapoyltyramine (34), and N-trans-feruloylmethoxytyramine (35). Each structure is labeled with corresponding numbers for easy identification. — Cytotoxic natural products of plants of the genus *Litsea.*

Other examples of cytotoxic natural products isolated from in this genus are biseugenol A (33) (a phenylpropanoid dimer) from Litsea costalis (Nees) Kosterm. (collected in Malaysia) (HepG2; IC₅₀: 18 μM) (Hosseinzadeh et al. 2013), N-trans-sinapoyltyramine (34) and N-trans-feruloylmethoxytyramine (35) (amide alkaloids) from Litsea acuminata (Blume) Kurata (collected in China) (HeLa) (Tanaka et al. 2009), butanolides (Cheng et al. 2001), and sesquiterpene glycosides (Wang et al. 2018).

Given the numerous cytotoxic natural products in this genus and considering that the Dusun use the edible fruits of L. garciae (Hassan et al. 2013) as a medicinal food (Maid et al. 2017) (Table 1), further studies are warranted. This fruit tree is only found in Borneo, the Philippines, and Taiwan (Table 2). The observation of the limited geographical distribution of this plant could lead one to wonder if its distribution might be an arboreal vestige of an Austronesian population, which would confirm the hypothesis of a Taiwanese origin of the Austronesian peoples (Solheim 1984).

Artabotrys roseus boerl. (Annonaceae)

Plants in the genus Artabotrys R.Br. (1820) produce cytotoxic aporphine alkaloids. Examples are lysicamine (36), isolated from Artabotrys crassifolius Hook. f. & Thomson (collected in Malaysia), toxic to MCF-7 cell line (IC₅₀: 3.9 µg/mL) (Kwan et al. 2016), and liriodenine (37) (Zhao et al. 2024). Other examples of cytotoxic alkaloids include hexapetalines A (38) and B (39) (benzylisoquinolines) (Zhou et al. 2015) and the protoberberine alkaloid 2,10-dihydroxy-3,9-dimethoxy-8-oxo-protoberberine (40) (Nguemdjo Chimeze et al. 2022) (Figure 7). Plants in this genus also produce cytotoxic sesquiterpenes, such as artaboterpenoid A (41) (Xi et al. 2016). The precise medical use of A. roseus, employed by both Dusun and Kadazan (Table 1), is unknown. Phytochemical, pharmacological, or toxicological reports on A. roseus, endemic to Borneo, are nonexistent.

Chemical structures of six compounds arranged in two rows, labeled with names and numbers. The figure displays the skeletal chemical structures of six compounds organized into two rows. The top row includes lysicamine (36), liriodenine (37), and hexapetaline A (38), while the bottom row features hexapetaline B (39), 2,10-dihydroxy-3,9-dimethoxy-8-oxo-protoberberine (40), and artaboterpenoid A (41). Each structure is labeled and exhibits various functional groups and unique features, emphasizing their molecular diversity. — Cytotoxic natural products of plants of the genus *Litsea.*

Boesenbergia pulchella (ridl.) merr. (Zingiberaceae)

Ethanol extract of rhizomes was found to be cytotoxic to MCF-7 cell line (IC₅₀: 93 µg/mL) (Jing et al. 2010). Active cytotoxic principles are not elucidated yet, but available evidence suggest phenolic principles. Ethanol and ethyl acetate extracts of rhizomes of Boesenbergia rotunda (L.) Mansf. were found to be cytotoxic to HeLa cell line (IC₅₀: 56 μg/mL) (Listyawati et al. 2016) and small duct intrahepatic cholangiocarcinoma cell line (RMCCA-1) (IC₅₀: 22.6 µg/mL), respectively (Sopitthummakhun et al. 2021). In a subsequent study, an ethyl acetate extract of rhizomes of B. rotunda was found to be cytotoxic to A549 cell line (IC₅₀: 22.5 µg/mL) on account of isopanduratin A (42) (a chalcone derivative), pinostrobin (43) and pinocembrin (44) (flavanones), and cardamonin (45) (a chalcone) which inhibited the growth of human lung cancer cell line (A549) with IC₅₀ values of 10.1, 25.3, 36.1, and 5.2 µg/mL, respectively (Han et al. 2023) (Figure 8). Cardamonin (45) induced apoptosis in nasopharyngeal carcinoma cell line (HK1) (IC₅₀: 27 μg/mL) via the activation of caspases-3 and −8, as well as alterations in mitochondrial membrane potential (Break et al. 2021). The prenylated chalcone boesenbergin A (46) induced apoptosis in A549 cell line with mitochondrial membrane potential alteration (Isa et al. 2013), as well as with human T4-lymphoblastoid cell line (CEM-SS) (IC₅₀: 8 μg/mL) (Ng et al. 2013). A hexane extract of the stems of Boesenbergia violacea (K.Larsen & Triboun) Mood & L.M.Prince inhibited the growth of malignant melanoma cell line (A375) (IC₅₀: 31.4 µg/mL) (Choosub and Samosorn 2021). The pharmacological properties and any possible toxicities of B. pulchella, endemic to Borneo, are currently unknown.

Chemical structures of isopanduratin A, pinostrobin, pinocembrin, cardamonin, and boesenbergin A, labeled with numbers. The figure presents five chemical structures. Isopanduratin A (42) shows a complex arrangement with hydroxyl and methoxy groups. Pinostrobin (43) and pinocembrin (44) feature a shared benzopyranone framework, differing only at a substituent. Cardamonin (45) is an open-chain chalcone with two benzene rings connected by an unsaturated ketone. Boesenbergin A (46) is more complex, with multiple benzene rings, functional groups, and a fused ring structure. Each compound is clearly labeled with its corresponding identifier. — Cytotoxic phenolic compounds of plants of the genus *Boesenbergia.*

Etlingera elatior (jack) R.M. Sm

Organic extracts demonstrated cytotoxic, antimutagenic, immunostimulant, and anti-inflammatory activities. Ethyl acetate extract of rhizome was found to be cytotoxic for CEM-SS and MCF-7 cell line with IC₅₀ values of 4 and 6.2 µg/mL, respectively (Habsah et al. 2005). An ethanol extract of flowers inhibited the growth of MCF-7 and MDA-MB-231 cell line with the IC₅₀ of 173.1 and 196.2 μg/mL, respectively (Ghasemzadeh et al. 2015). An essential oil of the leaves was found to inhibit the growth of mouse melanoma cell line (B16F10) (IC₅₀: 214.8 μg/mL) (Sangthong et al. 2022). A dichloromethane extract (200 µg/mL) protected Raji cell line against mutations caused by 12-O-tetradecanoylphorbol-13-acetate (Habsah et al. 2005). Methanol extract induced the proliferation of human peripheral lymphocytes (Safriani et al. 2021). Furthermore, an ethanol extract of flowers administered intraperitoneally to mice at a dose of 100 mg/kg induced a decrease in cyclooxygenase 2 expression after 7 days of treatment (Syaify et al. 2024). The flowers of this ginger are consumed to maintain good health among the Dusun and could represent an interesting material for the development of onco-preventive nutraceuticals. Although the toxicity profile of these flowers in humans has not been established, oral administration of a flower extract at a single dose of 600 mg/kg in rats did not cause any lethality (Sholihah et al. 2025). Further studies are needed on this common garden plant (Table 2).

Pycnarrhena tumefacta miers (Menispermaceae)

P. tumefacta is used as a medicinal food by the Dusun, Kadazan, and Murut (Table 1), and is also consumed locally in Sarawak where it is known as “sengkubak” and used as a condiment to impart a salty flavor to dishes (Yusli et al. 2023). An extract from this vine has been shown to be toxic to cervical cancer cell line (HeLa) (Fernandez 2009). An alkaloid extract of the roots of Pycnarrhena. cauliflora (Miers) Diels induced apoptosis in ductal carcinoma of the breast cell line (T-47D) (Masriani et al. 2014). Dichloromethane extract of stems (collected in Indonesia) was found to be cytotoxic to T-47D cell line (IC₅₀: 59.2 µg/mL) and induced apoptosis (Muharini and Enawaty, 2019). Plants in the genus Pycnarrhena Miers ex Hook. f. & Thomson (1855) produce cytotoxic bisbenzylisoquinoline alkaloids such as (+)-2-northalrugosine (47) (Abouchacra et al. 1987) from Pycnarrhena ozantha Diels (KB; IC₅₀: 10.4 µM) (Angerhoferet al. 1999), obaberine (48), limacine (49), aromoline (50), and isotetrandrine (51) (Van Beek et al. 1982) from Pycnarrhena longifolia (Decne. ex Miq.) Becc. (KB cell line) (Angerhoferet al. 1999), and isotetrandrine (51) and berbamine (52) from Pycnarrhena manillensis S. Vidal (KB cell line) (Bruchhausen et al. 1960; Angerhofer et al. 1999) (Figure 9). More experiments are needed.

Six chemical structures of bisbenzylisoquinoline alkaloids labeled with their names and numbers. The figure presents six distinct chemical structures of bisbenzylisoquinoline alkaloids, arranged in two rows. The top row features (+)-2-northalrugosine (47), obaberine (48), and limacine (49), each with varying functional groups, chiral centers, and nitrogen atom configurations. The bottom row includes aromoline (50), isotetrandrine (51), and berbamine (52), showcasing differences in hydroxyl/methoxy arrangements and structural motifs. Each structure is labeled with its name and number for identification. — Cytotoxic natural products of plants of the genus of plants of the genus *Pycnarrhena.*

Bridelia stipularis (L.) Bl. (Phyllanthaceae)

This tree produces friedelane-type triterpenes such as friedelin (53) and the lupane-type triterpene lupeol (54), and a phenolic compound identified as 4-(1,5-dimethyl-3-oxo-4-hexenyl)benzoic acid (Limtragool et al. 2024) (Figure 10). Friedelin (53) was found to be cytotoxic to MCF-7 cell line (IC₅₀: 1.8 µM) and induced an increase in cellular reactive oxygen species, DNA damage, and apoptosis (Subash-Babu et al. 2017). Lupeol (54) inhibited the survival of glioblastoma cell line (U87MG; IC₅₀: 13.6 μM) (Nyaboke et al. 2018). 3-Epi-glutinol (55) (a friedelane-type triterpene) and betulinic acid (56) (a lupane-type triterpene) from Bridelia cambodiana Gagnep were toxic to HL-60 cell line with IC₅₀ values of 5.6 and 6.9 µg/mL, respectively (Khiev et al. 2009). Regarding the biological activity of triterpenes, caution is advised as it depends on their purity level (Jaki et al. 2008).

Twelve chemical structures including friedelin, lupeol, and gallic acid with varied rings and functional groups. This figure displays twelve distinct chemical compounds arranged in rows and columns, labeled from 53 to 62. The top row features friedelin (53), lupeol (54), 3-epi-glutinol (55), and betulinic acid (56), showcasing complex carbon structures and various functional groups. The second row includes 4'-demethyl-4-deoxypodophyllotoxin (57), gallic acid (58), methyl gallate (59), and deoxypodophyllotoxin (60), characterized by aromatic rings and hydroxyl groups. The bottom row presents ß-peltatin (61) and ß-peltatin-5-O-ß-D-glucopyranoside (62), highlighting structural similarities and differences among lignans, each clearly labeled in black. — Cytotoxic natural products of plants of the genus *Bridelia.*

Plants in the genus Bridelia Willd. (1806) have the property of producing aryltetralin lignans of the podophyllotoxin type. This type of lignan constitutes a well-known class of cytotoxic agents. Etoposide (still widely used and forming the basis of several combination chemotherapy protocols) and Teniposide (rarely used today, mainly in the treatment of childhood acute lymphoblastic leukemia) are approved drugs. 4′-Demethyl-4-deoxypodophyllotoxin (57), gallic acid (58), and methyl gallate (59) (benzoic acid derivatives) isolated from the roots of Bridelia balansae Tutcher inhibited the growth of HCT-116 cell line with IC₅₀ values of 0.02, 20, and 16.3 µM, respectively (Zhao et al. 2023). Bridelia ferruginea Benth. produces deoxypodophyllotoxin (60) and β-peltatin (61) (Pettit et al. 2016). Deoxypodophyllotoxin (60) was found to be cytotoxic to A549 and B16F10 with IC₅₀ values of 6 and 6.8 ng/mL, respectively. Given intraperitoneally at a dose of 20 mg/kg/day for 14 days to mice xenografted with Lewis lung carcinoma cell line, deoxypodophyllotoxin (60) caused a decrease in tumor growth (Kim et al. 2002). β-Peltatin (61) induced apoptosis in pancreatic cancer cell line (MIA PaCa-2; IC₅₀: 2.5 nM), with cleavage of poly (ADP-ribose) polymerase, activation of caspase-3, and increased expression of cyclin B1. Given intraperitoneally at a dose of 15 mg/kg/day for 40 days to mice xenografted with prostatic adenocarcinoma cell line (PC-3), β-peltatin (61) reduced the volume of tumors and increased lifespan (Wu et al. 2023). β-Peltatin-5-O-β-D-glucopyranoside (62) was found to be cytotoxic to A2780 cell line (IC₅₀: 4.9 μM) (Liu et al. 2015). Ethanol extract of leaves of B. stipularis given orally to mice at a dose of 500 mg/kg for 14 days to rodents did not cause liver toxicity (Acharyya et al. 2022).

Ficus retusa L. (Moraceae)

Ethyl acetate extract of F. retusa was found to be cytotoxic to HepG2 cell line (IC₅₀: 68.4 µg/mL). The plant produces the oleanane-type triterpene β-amyrin (63) (Caco-2; IC₅₀: 81 µg/mL) (Maiyo et al. 2016), the friedelane-type triterpene friedelinol (64) (THP-1) (Gonçalves Pereira et al. 2020), the flavone luteolin (65) (NCI-H460) (Kyoung-Ah et al. 2006), vitexin (66) (a flavone C-glycoside), the flavanes (+)-afzelechin (67) and (+)-catechin (68) (Sarg et al. 2011) (Figure 11). Vitexin (66) induced the apoptosis of HCT-116 cell line (IC₅₀ ≈ 50 µM) with increased expression of cytochrome c, cleavage of caspases-3 and −9. Given orally at a dose of 50 mg/kg/day (3 days a week) to mice xenografted with drug-resistant HCT-116 cell line, vitexin (66) reduced the growth of tumors (Bhardwaj et al. 2018).

Chemical structures of twelve organic compounds including ß-amyrin, luteolin, and others, labeled with unique identifiers. The figure displays twelve chemical structures arranged in three rows. Top row: ß-amyrin (63), friedelinol (64), luteolin (65), and vitexin (66). Middle row: (+)-afzelechin (67), (+)-catechin (68), and ficuseptine (69). Bottom row: tylocrebrine N-oxide (71), 10S,13aR-isotylolecrbine N-oxide (72), fistulopsine A (73), and fistulopsine B (74). Each structure showcases unique functional groups, rings, and is labeled with a numerical identifier. — Cytotoxic natural products of plants of the genus *Ficus.*

Plants in the genus Ficus L. (1753) produce cytotoxic phenanthroindolizidine alkaloids such as ficuseptine (69) and tylophorine (70), from Ficus septica Burm.f., which at the concentration of 50 µM were toxic to gastric cancer cell line (NUGC) (Wu et al. 2002). 10S,13aR-Tylocrebrine N-oxide (71), 10S,13aR-isotylocrebrine N-oxide (72), and tylophorine (70), from F. septica, at the concentration of 10 µM, inhibited the proliferation of nasopharyngeal carcinoma cell line (HONE-1) by 92, 87, and 80%, respectively (Damu et al. 2009). Other cytotoxic principles in this genus are septicine-type alkaloids such as fistulopsines A (73) and B (74) from Ficus fistulosa Reinw. ex Blume which induced HCT-116 cell line cycle arrest in G1 phase (Yap et al. 2016).

Myrmecodia platytyrea becc. (Rubiaceae)

To date, no cytotoxic principle appears to have been identified in this myrmecophytic epiphyte, although preliminary studies have demonstrated cytotoxic, anti-inflammatory, and immunomodulatory activities. Methanol extracts inhibited the growth of hepatoma cell line (Huh7) (Ju et al. 2018) and HepG2 cell line (IC₅₀: 70 µg/mL), while being less toxic for Vero cell line (Zakaria and Aziz 2022). An extract was found to be cytotoxic to HepG2 cell line (IC₅₀: 5.7 μg/mL) and induced apoptosis with cell cycle arrest in G₀/G₁ phase, increased expression of CDK2 and CDK5, Bax, and caspase-3. In mice, this extract given orally, was able to decrease the growth of hepatocellular carcinoma (Ibrahim 2021). The aqueous extract administered at a dose of 400 mg/kg induced anti-inflammatory effects and increased the immune system of mice (Mohd Zin, 2019). This activity could be due to polysaccharides and potentially β-glucans (Hasan et al. 2019). In line, Triana Hertiani and Sumardi (2010) observed the immunostimulant properties of an ethanol extract of domatia (or ant-tubers) of Myrmecodia tuberosa Jack and Myrmecodia pendens Merr. & L.M.Perry (collected in West Papua) (Ulfah et al. 2013). Plants in the genus Myrmecodia Jack (1823) produce iridoids (Hanh et al. 2016). The use of M. platytyrea by the Bajau for cancer treatment (Table 1) and the non-acute toxicity of its aqueous extract administered orally to rats (400 mg/kg/day) (Hasan et al. 2020) call for further experiments.

Paederia verticillata Bl. (Rubiaceae)

Polar organic extracts of plants of the genus Paederia L. (1767) have demonstrated cytotoxic properties. This was observed with methanol extracts of leaves of Paederia foetida L. (MCF-7; IC₅₀ : 550.1 µg/mL) (Morshed et al. 2012; Priyanto et al. 2022), and an ethanol extract of leaves of Paederia lanuginosa Wall. (HT-29; IC₅₀: 28.8 µg/mL) (Hoang Phu et al. 2025).

The cytotoxic principles isolated from these plants are mainly iridoids (Figure 12). For example, paederosidic acid (75) from P. scandens was found to be cytotoxic to gastric adenocarcinoma cell line (SGC-7901; IC₅₀: 30.5 μM) and induced apoptosis with upregulation of caspases-3 and −9 as well as upregulation of Bax, and downregulation of Bcl-2 (Chen et al. 2015). We can also cite 10-O-trans-p-coumaroyl-(4R,6R)-3,4-dihydro-3α-methylthiopaederoside (76) toxic to five endocrine tumor cell lines (IC₅₀ ˂ 20 µM) (Hu et al. 2024), as well as trans-p-coumaroyl-(4S,6R)-3,4-dihydro-3β-ethoxypaederoside (77) from the aerial parts of Paederia yunnanensis (H. Lév.) Rehder (Hu et al. 2025). The presence of cytotoxic iridoids in P. verticillata is probable and these remain to be identified.

Three chemical structures: paederosidic acid (75), 10-O-trans-p-coumaroyl-3,4-dihydro-3a-methylthiopaederoside (76), and 10-O-trans-p-coumaroyl-3,4-dihydro-3ß-ethoxypaederoside (77), showing various functional groups. The image illustrates three chemical structures labeled (75), (76), and (77). Structure (75) is paederosidic acid with a furan-dihydropyran core, featuring acetylthioester and carboxylic acid groups. Structure (76) has a methylthio group and a lactone, while structure (77) has an ethoxy substituent; both share a coumaroyl group linked to the sugar. Each structure is annotated, demonstrating differences in functional groups and stereochemistry. — Cytotoxic iridoid glycosides of plants of the genus *Paederia.*

Rennellia borneensis baill. (Rubiaceae)

Plants in the genus Rennellia Korth. (1851) produce cytotoxic anthraquinones (Figure 13). Examples are nordamnacanthal (78), rubiadin (79), and rubiadin-1-methyl ether (80) (Osman et al. 2010, 2016a). Nordamnacanthal (79) was found to be cytotoxic to oral squamous cell carcinoma cell line (H400 OSCC; IC₅₀: 6.8 µg/mL) (Shaghayegh et al. 2017). Rubiadin (79) from Morinda umbellata L. inhibited the growth of HepG2 cell line (IC₅₀: 3.6 µM) (Chiou et al. 2014) and from this plant rubiadin-1-methyl ether (80) inhibited the growth of MCF-7 cell line (IC₅₀: 30 µg/mL) (Ali et al. 2000). Likewise, 2-formyl-3-hydroxy-9,10-anthraquinone (81), 2-methyl-3-hydroxy-9,10-anthraquinone (82), and 1,2-dimethoxy-6-methyl-9,10-anthraquinone (83) from the roots of Rennellia elliptica Korth. were active against MCF-7 cell line with IC₅₀ values of 33, 50.1, and 38.7 µg/mL, respectively (Osman et al. 2016b). A preliminary study revealed the presence of anthraquinones in R. borneensis (Rushdan et al. 2023), endemic to Borneo, whose cytotoxic active principles remain to be identified. Since anthraquinones are mutagenic (Tikkanen et al. 1983), this shrub is unsuitable for any medicinal use.

Chemical structures of nordamnacanthal, rubiadin, and 1,2-dimethoxy-6-methyl-9,10-anthraquinone with functional group variations. The figure presents three chemical structures from left to right: nordamnacanthal (78) with a formyl (-CHO) and hydroxyl (-OH) groups; rubiadin (79) with dual hydroxyl groups and variable substituent positions (R¹ and R²); and 1,2-dimethoxy-6-methyl-9,10-anthraquinone (83) with methoxy (-OCH3) and methyl (-CH3) groups. Each structure is labeled clearly and showcases unique functional group variations, contributing to their chemical identities. — Cytotoxic anthraquinones of plants of the genus *Rennellia.*

Crassocephalum crepidioides (benth.) S. Moore

Aqueous extract of this herb (collected in Japan) given orally at a dose of 5 g/kg/day for 29 days to mice xenografted with S-180 cell line caused a decrease in tumor volume by about half. This extract did not cause cytotoxic effects In vitro, but when RAW264.7 cell line were treated with the extract at a concentration of 250 µg/mL, the culture supernatant could inhibit the growth of S-180 cell line because of the presence of nitric oxide derived from activation of NF-κB and increased expression of iNOS. From this extract isochlorogenic acid (a caffeic acid derivative) (84) (Figure 14) activated NF-κB in RAW264.7 cell line (Tomimori et al. 2012). Ethanol extract of the plant (collected in Malaysia) at the concentration of 25 µg/mL protected HepG2 against tert-butyl hydroperoxide (Wijaya et al. 2011).

Chemical structure of isochlorogenic acid with hydroxyl groups and carbon rings. The diagram illustrates the chemical structure of isochlorogenic acid (84), showcasing a hexagonal carbon ring linked to a phenolic group with two hydroxyl (–OH) groups. It features a carbon-carbon double bond extending to a carboxylic acid group, which also contains a hydroxyl group. The structure highlights various functional groups and connections, underlining its organic chemistry characteristics. — Cytotoxic natural products of plants of the genus *Kadsura.*

C. crepidioides is used by the Murut against cancer and as a medicinal food by the Dusun (Table 1). The aqueous leaf extract administered orally gave an LD₅₀ greater than 5 g/kg in rats (Nguemfo et al. 2021). However, the plant produces hepatotoxic pyrrolizidine alkaloids (Wachenheim et al. 1992).

Merremia gracilis E.J.F. Campb. & argent (Convolvulaceae)

Plants of the genus Merremia Dennst. ex Endl. (1841) are known to be poisonous (Brito et al. 2019). They produce glycolipids some of which are able to increase the vulnerability of KB cell line to vinblastine such as merremin A (85) (Figure 15) from the aerial parts Merremia hederacea (Burm. f.) Hallier f. (Wang et al. 2014; Li et al. 2023). Ethanol extract of Merremia emarginata (Burm. f.) Hallier f. inhibited the growth of HT-29 cell line by 58% at the concentration of 200 µg/mL and induced apoptosis via increased expression of caspase-3, decreased expression of Bcl-2 and Bcl-xL (Benedict et al. 2024). Hexane extract of M. emarginata inhibited the growth of A549 cell line (IC₅₀: 18.4 µg/mL) (Baskar et al. 2012).

Structural diagram of merremin A (85), showcasing multiple carbon rings and hydroxyl groups. The diagram illustrates the complex chemical structure of merremin A (85), featuring interconnected carbon rings and five sugar units. It includes multiple hydroxyl (-OH) groups, ester groups, and a macrocyclic lactone ring. Each saccharide unit is linked with specific orientations, shown with stereochemical indicators, highlighting the molecule's intricate connectivity and functional groups. The overall representation emphasizes the organic nature of the compound through its diverse arrangement. — Cytotoxic caffeic acid derivative of *C. crepidioides.*

Merremia peltata (L.) merr. (Convolvulaceae)

The ethyl acetate extract of leaves was found to be cytotoxic to brine-shrimp s (LC₅₀: 22 ppm) (Djamaan, 2016).

Other plants used by multiple ethnic groups

Airyantha borneensis (Oliv.) Brummitt and Koompassia malaccensis Maing, have not been the subject of any phytochemical or pharmacological studies, as have all the plants of their respective genera.

Plants used by the Lundayeh (Kelabit group)

The Lundayeh migrated from other parts of Borneo to Sabah about 100 years ago where they represent a minority. The Lundayeh are found in Sabah, and they are also known as Lunbawang in Sarawak and Kalimantan (King and King 1984). The wealth of their medicinal flora (especially the use of endemic plants) suggest that they should perhaps belong to the Bornean-speaking ethnic group close to the Murut (Wiart et al. 2025).

Kadsura borneensis A.C. (Schisandraceae)

Plants in the genus Kadsura Juss. (1810) produce a series of cytotoxic triterpenes rare in nature (Figure 16). Examples are heteroclitalactone D (86) (HL-60; IC₅₀: 6.7 μM) (Wang et al. 2006), longipedlactone A (87) and F (88) (Xu et al. 2010), and kadheterin A (89) (HL-60; IC₅₀: 14.5 μM) (Luo et al. 2017) from Kadsura heteroclita (Roxb.) Craib. Other examples are longipedlactone M (90) (Yang et al. 2010), ananosic acid B (91) (HeLa; IC₅₀: 0.5 μg/mL), and ananosic acid C (92) (HeLa; IC₅₀: 0.4 μg/mL) from Kadsura ananosma Kerr (Chen et al. 2004) (Figure 16). We can also cite kadlongilactone A (93) from Kadsura longipedunculata Finet & Gagnep. (Pu et al. 2007), xuetonglactone F (94) from K. heteroclita (BGC 823; IC₅₀: 2 μM) (Shehla et al. 2020), and Schisandronic acid (95) from Kadsura coccinea (Lem.) A.C. Sm. Schisandronic acid (95) induced apoptosis in MCF-7 cell line (IC₅₀: 8 μM) with activation of caspase-3 and cleavage of poly (ADP-ribose) polymerase (Tasneem et al. 2021).

Grid of 21 labeled chemical structures of various organic compounds. The figure showcases 21 distinct organic chemical structures arranged in a grid. Each structure is labeled with a unique compound name and a corresponding numerical identifier (86-106). The compounds feature various complex structural elements, including polycyclic frameworks, lactone, and ether rings. Prominent functional groups such as hydroxyl (-OH) and acetyl (-OAc) are present, with bond orientations clearly illustrated. Noteworthy examples include heterocitalactone D (86) and taiwankadsurin B (106). The layout facilitates the comparison of chemical features and functional group variation among the compounds. — Cytotoxic glycolipid of plants in the genus *Merremia.*

Other cytotoxic principles in this genus are dibenzocyclooctadiene lignans, also rare in nature, such as heilaohusuin B (96) (Yang et al. 2019) and kadusurain A (97) from Kadsura coccinea (Lem.) A.C. Sm. (Zhao et al. 2014) as well as kadcoccine acid B (98) and H (99) (Hu et al. 2016). Heilaohulignan C (100) from K. coccinea induced apoptosis in gastric cancer cell line (BGC-823) and given to mice xenografted with gastric carcinoma cell line, caused a reduction of tumors (Daniyal et al. 2021). Additional instances of cytotoxic lignans are kadsufolin A (101), kadsufolin D (102), angeloylbinankadsurin A (103), and heteroclitin B (104) from Kadsura oblongifolia Merr. (Huang et al. 2011). In addition, a diarylbutane lignan meso-dihydroguaiaretic acid (105) was isolated from K. heteroclita (HT-29; IC₅₀: 16.2 μM) (Minh et al. 2014) and taiwankadsurin B (106) (a homolignan) from Kadsura philippinensis Elmer (Shen et al. 2005). To date, nothing is known about the toxicity of K. borneensis, a vine endemic to Borneo.

Nepenthes ampullaria jack (Nepenthaceae)

An ethanol extract of the roots of this cultivable pitcher plant was found to be cytotoxic to HT-29 and Caco-2 cell line with IC₅₀ values of 62.3 and 49.3 µg/mL, respectively, while being nontoxic to normal intestinal cell line (CCD841CoN) (IC₅₀ > 512 µg/mL) (Dřímalová, 2024). Cytotoxic principles have not been isolated from this plant. What we know about plants of the genus Nepenthes L. (1753) is that their polar organic extracts are often cytotoxic owing to the presence of naphthoquinones. A methanol extract of stems of a plant in the genus (purchased in Taiwan) was found to be cytotoxic to gingival carcinoma cell line (Ca9-22) (IC₅₀ ≈ 15 µg/mL) and at the concentration of 80 µg/mL induced apoptosis with cell cycle arrest in G₂ phase (Lin et al. 2023). Plumbagin (107) (Figure 17), a naphthoquinone isolated from the roots of Nepenthes alata Blanco (purchased in Korea), induced cell cycle arrest in the G₂/M phase of MCF-7 cell line, as well as apoptosis via increased production of reactive oxygen species, elevation in the ratio of Bax/Bcl-2, and release of cytochrome c. In mice xenografted with MCF-7 cell line, plumbagin (107) caused a decrease in the growth of tumors (De et al. 2019). N. ampullaria probably produces cytotoxic naphthoquinones. These remain to be identified. Naphthoquinones, like plumbagin (107), are poisonous (Teixeira et al. 2024), which renders N. ampullaria unfit for any nutraceutical development.

Chemical structure of plumbagin (107) with two fused rings, hydroxyl and carbonyl groups. This illustration depicts the chemical structure of plumbagin (107), featuring two fused aromatic rings. A hydroxyl group (–OH) is attached to one ring, while a carbonyl group (C=O) is positioned on the adjacent ring. The diagram clearly indicates bond connectivity and the arrangement of functional groups, highlighting the key components of the molecule. — Cytotoxic naphthoquinone of plants of the genus *Nepenthes.*

Symplocos odoratissima choisy ex zoll. (Symplocaceae)

Cytotoxic triterpene saponins have been identified from Symplocos chinensis (Lour.) Druce (Fu et al. 2006), including symplocososides A (108), C (109), and F (110) (Tang et al. 2004) and as well as 2β,3β,19-α,24-tetrahydroxy-23-norurs-12-en-28-oic acid (111) (an ursane-type triterpene), the latter active against BGC-823 cell line (Li et al. 2003) (Figure 18). From the leaves of Symplocos cochinchinensis (Lour.) S.Moore (collected in Japan), symplocosin K (112) was found to be cytotoxic to A549 cell line (IC₅₀: 73.8 μM) (Ohyama et al. 2020).

Chemical structures of symplocososides A, C, F, K, matairesinoside, and matairesinol, labeled with their respective identifiers. This image depicts the chemical structures of several natural products. The top left includes symplocososides A (108) with a methyl group and C (109) with a butyl group, both featuring multiple hydroxyl groups. Adjacent is symplocososide F (110), showcasing a complex glycoside framework with various saccharides. Below, symplocosin K (112) includes hydroxy and carbonyl groups, while matairesinoside (113) and matairesinol (114) display variations in their aromatic ring structures. Each compound is labeled with its name and structural identifiers, highlighting differences in functional groups. — Cytotoxic natural products of plants of the genus *Symplocos.*

Phenolic glycosides have been identified from Symplocos racemosa Wight ex C.B.Clarke (collected in Pakistan) (Choudhary et al. 2004) and may account for the fact that a butanol extract of the bark of this plant inhibited the growth of HL-60 and HeLa cell line with IC₅₀ values of 27.1 and 22.8 μg/mL, respectively (Raval et al. 2009). Other constituents in this genus are seco-iridoids from S. cochinchinensis (Lee et al. 2019), dibenzylbutyrolactone lignan glycosides such as matairesinoside (113) from the roots of Symplocos caudata Wall. ex G.Don (collected in China) (Huo et al. 2008) or dibenzylbutyrolactone lignans as in matairesinol (114) (CaCo-2; IC₅₀: 220 μM) (Shoeb et al. 2007) from Symplocos setchuensis Brand as well as, from this plant, pyridoindole alkaloids (Ishida et al. 2001). S. racemosa produces anthraquinones (Farooq et al. 2017). Matairesinol (114) was found to be cytotoxic to HepG2 cell line (IC_50: 30.8 µM) and induced apoptosis with mitochondrial dysfunction, an increase in reactive oxygen species, and activation of caspase-3 (Arzuk et al. 2024). S. odoratissima has not been studied phytochemically or pharmacologically and could be toxic since plants of the genus Symplocos Jack (1760) are aluminum accumulators (Schmitt et al. 2016).

Hydnophytum formicarum jack (Rubiaceae)

The data available so far indicates the presence of cytotoxic phenolics in this myrmecophyte epiphyte, but these have mild effects in vitro, which suggests that use as a treatment for cancer could be due, at least in part, to the presence of immunostimulant principles. Polar organic extract of domatia inhibited the growth fibrosarcoma cell line (HT-1080; IC₅₀: 9.9 µg/mL) (Ueda et al. 2002) and the activity of histone deacetylase in vitro (4 µg/mL; 40%). The plant produces sinapinic acid (115) (a phenylpropanoid) (Figure 19), which inhibited histone deacetylase activity with an IC₅₀ value of 2.2 mM and inhibited the survival of HeLa cell line (Senawong et al. 2013). Another cytotoxic phenolic identified from H. formicarium is 7,3′,5′-trihydroxyflavanone (116), which caused DNA fragmentation and apoptosis in MCF-7 cell line via increased expression of Bax, decreased expression of Bcl-2 at the concentration of 15 μg/mL (Abdullah et al. 2010).

Two chemical structures: sinapinic acid (115) on the left and 7,3',5'-trihydroxyflavanone (116) on the right. The image features sinapinic acid (115) and 7,3',5'-trihydroxyflavanone (116) side by side. Sinapinic acid shows a benzene ring with a methoxy group (–OCH3), hydroxyl group (–OH), and a carboxylic acid group (–COOH) on its structure. The right side presents 7,3',5'-trihydroxyflavanone with a flavanone core, showcasing two hydroxyl groups and an ether linkage. Each structure includes its name and identifying number beneath. — Cytotoxic phenolic compounds of *H. formicarium.*

Of note, an ethanol extract of domatia of H. formicarum (collected in West Papua) inhibited the growth of T-47D cell line by 53.4% (100 µg/mL) and induced lymphocyte proliferation (Darwis et al. 2014). An aqueous extract of domatia given orally to mice caused some immunomodulatory properties (Putra et al. 2025). Do they produce β-glucans?

H. formicarum is cultivable (Table 2). The immunostimulatory properties of this epiphytic plant could be due to the presence of β-glucans or other polysaccharides. More experiments are needed.

Jasminum bifarium wall (Oleaceae)

Plants in the genus Jasminum L. (1753) produce cytotoxic seco-iridoids. We can cite multifloroside (117) (Figure 20) from Jasminum multiflorum (Burm. f.) Andrews decreased the viability of epidermoid carcinoma cell line (A431) (≈ 80%) at the concentration of 200 µM, cell cycle arrest in S phase, an increase of reactive oxygen species, and induction of apoptosis (Zhang et al. 2019). From Jasminum humile L., jasmoside (118) and isojasminin (119) were found to be cytotoxic to THP-1 cell line with IC₅₀ values of 27.5 and 51 µg/mL, respectively (Mansour et al. 2022). 10-Hydroxyoleoside dimethyl ester (120) from Jasminum lanceolarium Roxb. (Shen et al. 1997) was found to be marginally cytotoxic to A431 cell line (200 µM) (Zhang et al. 2019).

Chemical structures of multifloroside (117), jasmoside (118), isojasminin (119), and 10-hydroxyoleoside dimethyl ester (120). The image shows four chemical structures arranged horizontally. From left to right: multifloroside (117) with multiple hydroxyl groups and a sugar moiety; jasmoside (118) featuring distinctive hydroxyl substitutions; isojasminin (119) showing variations in hydroxyl arrangement; and 10-hydroxyoleoside dimethyl ester (120), characterized by ester groups and hydroxyl placements. Each compound is labeled with an identification number. — Cytotoxic iridoid glycosides of *J. bifarium.*

Another interesting aspect of these plants is that their polar and med-polar organic extracts are capable of preventing the development of tumors in rodents resulting from exposure to 7,12-dimethylbenz[a]anthracene. This was observed with an ethanolic extract of flowers of Jasminum grandiflorum L. (300 mg/kg for 14 weeks) (Kolanjiappan and Manoharan 2005), an ethyl acetate extract of leaves of Jasminum subtriplinerve Blume (14.4 mg/kg for four weeks) (Minh et al. 2024), and the essential oil of Jasminum sambac (L.) Aiton (10 mL/kg; orally for 45 days) (Lakshmi et al. 2024).

Other plants used by the Lundayeh

Lygodium salicifolium C. Presl. (Lygodiaceae) and Garnotia acutigluma (Steud.) Ohwi have not been the subject of any phytochemical or pharmacological studies.

Plants used by the Brunei (Malayic group)

Drymoglossum piloselloides (L.) C. Presl (Polypodiaceae)

An ethanol extract inhibited the growth of MCF-7 cell line (IC₅₀: 83.6 µg/ml) (Endrini 2009), HeLa cell line (IC₅₀: 16.2 µg/ml) (Su’lain et al. 2019), and P388 cell line (IC₅₀: 19.3 µg/ml) (Sahid et al. 2013). The plant produces a series of prenylated phloroglucinols (Socolsky et al. 2010, 2011), which could be examined for their possible cytotoxic effects since the acylphloroglucinols paleacenins A (121) and B (122) from Elaphoglossum paleaceum (Hook. & Grev.) Sledge (Polypodiaceae) were cytotoxic to PC-3 cell line with IC₅₀ values of 1.7 and 2.9 µM, respectively (Arvizu-Espinosa et al. 2019) (Figure 21).

Chemical structures of paleacenin A (121) and paleacenin B (122), highlighting differences in carbon chain length and hydroxyl group positioning. The figure illustrates two chemical structures side by side: paleacenin A (121) on the left and paleacenin B (122) on the right. Both compounds feature a phenolic core with hydroxyl (-OH) groups and distinct carbon chains. Paleacenin A has a seven-carbon isoprenyl side chain, while paleacenin B includes an additional carbon, forming an eight-carbon chain. Variations in double bond locations and hydroxyl group positions further distinguish the two molecules, influencing their chemical properties. — Cytotoxic prenylated prenylated phloroglucinols from plants of the family polypodiaceae.

Gnetum macrostachyum hook.f. (Gnetaceae)

This stout primary forest climber is employed for fatigue (Table 1) which is one of the symptoms of cancer. It produces the stilbenes resveratrol (123) and isorhapontigenin (124), and the stilbenoid gnetin C (125) (Kloypan et al. 2012), 5,7,4′-trihydroxy-3′-methoxyflavanone (126) (Saisin et al. 2009) (Figure 22). Resveratrol (123) and isorhapontigenin (124) inhibited the growth of MCF-7 and T-47D cell line with the IC₅₀ values of 51.1 μM (Alkharashi, 2023) and 40 μM (Subedi et al. 2019), respectively. Gnetin C (125) inhibited the growth of prostate cancer (DU-145) cell line (IC₅₀: 6.6 μM). Given intraperitoneally at a dose of 50 mg/kg/day for 30 days to mice xenografted with prostate cancer cell line (C3M-Luc), gnetin C (125) caused a decrease in tumor growth (Gadkari et al. 2020). A very interesting point is that gnetin C (125), when mixed into the diet of rats with prostate cancer (70 mg/kg) for 17 weeks, induced some levels of protection (Parupathi et al. 2022). If this activity is confirmed by further studies and if this stilbenoid proves nontoxic, there would be grounds for clinical development. The stilbenoid macrostachyol D (127) from the roots was found to be cytotoxic to cervical cancer cell line (HeLa) (IC₅₀: 4.1 μM) (Sri-in et al. 2011).

Five chemical structures: resveratrol (123), isorhapontigenin (124), gnetin C (125), macrostachyol D (127), and 5,7,4'-trihydroxy-3'-methoxyflavanone (126) labeled with identifiers. The image illustrates five chemical structures. From left to right: 1) Resveratrol (123) and isorhapontigenin (124) as stilbene derivatives, showing structural differences in their R-groups; 2) Gnetin C (125) and macrostachyol D (127) as complex stilbene dimers, both with hydroxyl and methoxy groups; 3) 5,7,4'-trihydroxy-3'-methoxyflavanone (126), featuring a chromanone core with various functional groups. Each structure has its corresponding identifier for clarity. — Cytotoxic phenolic compounds from *G. macrostachyum.*

Plants used by the Bajau (Bajau group)

The Bajau are categorized as west coast Bajau (Bajau Samah) or the east coast Bajau and in available data pertaining to the medicinal plant they use the separation does not exist. They tend to use plants growing in mangroves and sea side. These plants are in general well-known and non-endemic (Wiart et al. 2025).

Diplazium cordifolium Bl. (Athyriaceae)

This fern does not appear to have been phytochemically or pharmacologically studied. Toxicological studies on this plant have not yet to be conducted.

Dischidia rafflesiana wall. (Asclepiadaceae)

This cultivable climber affords a remedy for cancer (Table 1 and 2). It is known to produce β-amyrin (63) (Van Hoang et al. 2022). Methanol extract of a plant in the genus Dischidia R. Br. (1810) was found to be cytotoxic to P388 cell line (IC₅₀: 24.8 µg/mL) (Manggribeth et al. 2019). Hexane and dichloromethane extracts of Dischidia nummularia R.Br. were found to be cytotoxic to MDA-MB-231 cell line (Khalil-ur-Rehman et al. 2019). From this plant, β-sitosterol (128) (Figure 23) was found to be cytotoxic to P-388 cell line (IC₅₀: 0.5 µM) (Benu et al. 2023). From Dischidia alboflava Costantin (collected in Vietnam) cytotoxic triterpenes have been identified such as β-amyrin acetate (129), friedelin (53), and lupeol (54) (Linh et al. 2023). β-Amyrin acetate (129) (Figure 23) inhibited the growth of A2780 cell line (Chaturvedula et al. 2002).

Molecular structures of ß-sitosterol (128) and ß-amyrin acetate (129) with labeled components. The image shows two molecular diagrams side by side. On the left is ß-sitosterol (128), characterized by a hydroxyl (–OH) group, a tetracyclic steroid structure with three six-membered and one five-membered ring, and a branched alkyl chain. The right side features ß-amyrin acetate (129), a pentacyclic triterpene, modified by an acetate (–AcO) group. Both diagrams highlight their distinct functionalities and carbon rings, with explicit labeling of the molecular identifiers. — Cytotoxic steroid and triterpene from plants of the genus *Dischidia.*

Plants used by the Dusun (Bornean group, Dusunic family)

General observations

Regarding the medicinal plant species used by the Dusun with chemotherapeutic potential, the following observations can be made: (i) half of them are used as edible plants, (ii) they use four species of ferns, (iii) these plants are often from primary forest, and (iv) six species are endemic.

Gleichenia truncata (Willd.) Spreng. (Gleicheniaceae)

Labdane glycosides are produced by Gleichenia quadripartita (Poir.) T. Moore (Socolsky et al. 2007) and Gleichenia japonica Spreng.(Munesada et al. 1992) and clerodane glycosides by Gleichenia microphylla R.Br. (Wada et al. 1998). Gleichinia alpinia R. Br. (collected in Australia) produces flavonol glycosides such as rutin (21) (Gyeltshen et al. 2022) In the family Gleicheniaceae, Dicranopteris linearis (Burm. f.) Underw. produces quercitrin (130) (Figure 24) toxic to HL-60 cell line (IC₅₀: 4.5 µg/mL) (Chen et al. 2014).

Chemical structure of quercitrin (130) featuring multiple hydroxyl groups and a rhamnose sugar connected through glycosidic bonds. The image illustrates the chemical structure of quercitrin, labeled (130). It comprises a flavonoid core with three fused aromatic rings containing multiple hydroxyl (OH) groups. The left ring has two hydroxyls, the middle ring includes a carbonyl group and an ether-linked rhamnose sugar. The rhamnose, a six-membered pyranose ring, has five hydroxyl groups. The structure emphasizes the complex arrangement of functional groups, foundational to quercitrin's chemical properties. — Cytotoxic flavonol glycoside from plants of the family Gleicheniaceae.

Diplazium esculentum (retz.) Sw

Organic polar extracts of this fern demonstrated cytotoxic effect against MDA-MB-231 cell line (ethanol, 100 µg/mL) (Rahmat et al. 2003), brine-shrimp (methanol, LC₅₀: 1.6 µg/mL) (Akter et al. 2014), and chronic myelogenous leukemia cell line (K562) (methanol, 500 µg/mL) (Salleh and Ab Latif 2022). The plant produces simple phenolics (Gyeltshen et al. 2022), cinnamic acid (131) (phenyl propanoid), protocatechuic acid (132) (a benzoic acid derivative), and rutin (21), as well as ecdysteroids (Watanabe et al. 2021) (Figure 25). Cinnamic acid (131) is an inhibitor of histone deacetylase (IC₅₀: 9.1 µg/mL) (Koyu et al. 2024) and abrogated the survival of A549 cell line. Given orally three times a week for a total of six doses at a dose of 1.5 mmol/kg to mice xenografted with HT-29 cell line, cinnamic acid (131) evoked a decrease in the volume of tumors by about half (Zhu et al. 2016). Protocatechuic acid (132) prevented the development of mutations induced by H₂O₂ in common fruit flies (Anter et al. 2011).

Two chemical structures: cinnamic acid (131) on the left, protocatechuic acid (132) on the right, each labeled with functional groups. The figure displays two molecular structures side by side. On the left is cinnamic acid (131), featuring a benzene ring attached to an unsaturated carbon chain ending in a carboxylic acid group (–CO2H). On the right is protocatechuic acid (132), with a benzene ring and two hydroxyl (–OH) groups attached, also terminating in a carboxylic acid group (–CO2H). Each structure is clearly labeled and highlights the arrangement of functional groups. — Cytotoxic phenolic compounds of *D. esculentum.*

Although this fern is a medicinal food (Table 1) (Maid et al. 2017), caution should be exercised regarding the nutraceutical development of this plant. Preliminary data suggest toxic effects greater than 2.5 g/kg (chloroform extract) in mice (Salleh and Ab Latif 2022), but oral administration of an aqueous extract given to mice for 180 days caused erythrocyte destruction and immunosuppression (Roy et al. 2013).

Nephrolepis acutifolia (Desv.) Christ (Nephrolepidaceae)

Aqueous extract of this epiphyte fern was found to be cytotoxic to K562 cell line (IC₅₀: 190.8 µg/mL) (Chai et al. 2015), and the cytotoxic principles involved have not been identified. Essential oil from the roots of Nephrolepis exaltata (L.) Schott and Nephrolepis cordifolia (L.) C. Presl inhibited the growth of A549 cell line with IC₅₀ values of 24.3 and 23.6 µg/mL, respectively (El-Tantawy et al. 2015), while an acetone extract was found to inhibit the growth of PC-3 cell line (Bobach et al. 2014). N. acutifolia is used as medicinal food (Table 1). More experiments are needed.

Acrostichum aureum L. (Pteridaceae)

An ethyl acetate extract was found to be cytotoxic to HeLa cell line (Dai et al. 2005). This fern produces rutin (21), kaempferol (133), and an N-benzoylphenylalanine derivative patriscabratine (134) (Figure 26). Patriscabratine (134) was found to be cytotoxic to MDA-MB-231 and MCF-7 cell line, with IC₅₀ values of 69.8 and 197.3 μM, respectively (Uddin et al. 2012). Kaempferol (133) inhibited the growth of ovarian cancer cell line (A2780; IC₅₀: 19 μM) (Pham et al. 2018).

Four chemical structures: kaempferol, patriscabratine, (+)-pinoresinol-4-O-sulfate, and (2S,3S)-sulfated pterosin C. This figure presents four chemical structures: Kaempferol (label 133) at top left, a flavonoid with multiple hydroxyl groups; patriscabratine (label 134) at top right, featuring an amide and two aromatic rings; (+)-pinoresinol-4-O-sulfate (label 135) at bottom left, a lignan with sulfonate and methoxy groups; and (2S,3S)-sulfated pterosin C (label 136) at bottom right, showcasing a bicyclic structure with hydroxyl and sulfate functional groups. Each compound is labeled. — Cytotoxic compounds of *A. aureum.*

Other cytotoxic principles include the lignan (+)-pinoresinol-4-O-sulfate (135) (MCF-7; IC₅₀: 7 µM) (Minh et al. 2022) and (2S,3S)-sulfated pterosin C (136) (a norsesquiterpene) (AGS; IC₅₀: 23.9 μM) (Figure 26) (Uddin et al. 2011). The plant has anti-inflammatory properties and this is interesting because chronic inflammation is one of the etiological factors of cancer. An ethanol extract given to mice orally at a dose of 400 mg/kg to assuage the pain caused by injection of acetic acid as effectively as diclofenac (25 mg/kg) (Khan et al. 2013). An aqueous extract of the aerial part given orally to rats at a dose of 400 mg/kg/day for 7 days, prior to oral administration of absolute ethanol, prevented the formation of gastric ulcers (Wu et al. 2018).

A. aureum is taken as medicinal food (Table 1). Preliminary toxicological studies indicate a lack of toxicity in acute (LD₅₀ > 5 g/kg) and sub-acute (750 mg/kg/day for 28 days) studies (Akinwumi et al. 2024). It should be noted, however, that an ethanolic extract administered to rats on days 1 to 7 after coitus prevented all pregnancies (Dhar et al. 1992).

Goniothalamus roseus Stapf

This endemic rainforest treelet produces nephrotoxic aristolactam alkaloids (Xue et al. 2025) and has therefore no potential for the development of a herbal remedy. Rather, it is a source of cytotoxic principles such as the styrylpyrones dehydrogoniothalamin (137) (PC-3; IC₅₀: 90.4 µM) (de Souza et al. 2021), 5-acetoxygoniothalamin (138) (HCT-116; IC₅₀: 8.6 μM) (Meesakul et al. 2020), goniothalamin (139) (HL-60; IC₅₀: 5.6 µM) (Petsophonsakul et al. 2013), and goniothalactam (140) (an aristolactam alkaloid) (P-388; IC₅₀ < 4 μg/mL) (Tsai et al. 2005) (Figure 27).

** Displays 17 chemical structures of organic compounds, including lactones, flavonoids, and alkaloids, arranged in rows with labels indicating compound names and functional groups. ** Chemical structures of organic compounds with distinct names and functional groups. — Cytotoxic compounds of plants in the genus *Goniothalamus.*

Cytotoxic styrylpyrones are rare in nature and mostly found in the genus Goniothalamus (Blume) Hook. f. & Thomson (1855). We can cite (6 R,7R,8R)-8-chlorogoniodiol (141) from Goniothalamus amuyon (Blanco) Merr. (Lan et al. 2003), (–)-goniodiol-7-monoacetate (142) from Goniothalamus repevensis Pierre ex Finet & Gagnep. (ASK; IC₅₀: 10.2 µM) (Chanakul et al. 2022), and goniolanceolatin B (143) and D (144) (styrylpyrone dimers) from Goniothalamus lanceolatus Miq. (Bihud et al. 2019).

Other cytotoxic principles are furanopyrones such as isoaltholactone (145) and aporphine alkaloid such as liriodenine (37) from Goniothalamus gitingensis Elmer (collected in the Philippines), toxic to K562 cell line with IC₅₀ values of 4.3 and 6.1 µg/mL, respectively (Macabeo et al. 2014). From Goniothalamus laoticus (Finet & Gagnep.) Bân (collected in Thailand) acetylaltholactone (146), (+)-altholactone (147), and nordicentrine (148) (an aporphine alkaloid) were toxic to KB cell line with IC₅₀ values of 2.9, 3.5, and 0.4 µg/mL, respectively (Lekphrom, et al. 2009). From this plant, the azaanthraquinone alkaloid laoticuzanone A (149) inhibited the growth of KB and HeLa cell lines with IC₅₀ values of 0.6 and 0.5 μg/mL, respectively (Tip-Pyang et al. 2010). From Goniothalamus marcanii Craib, another azaanthraquinone, marcanine G (150), was found to be cytotoxic to A549 and MCF-7 cell line, with IC₅₀ values of 14.8 and 15.1 μM, respectively (Thanuphol et al. 2018). Other cytotoxic principles in the genus are acetogenins such as annomontacin (151) and gigantrionenin (152) from Goniothalamus giganteus Hook. f. & Thomson (Fang et al. 1992), as well as flavonols such as 5,4′-dihydroxy-8-(2-hydroxybenzyl)-3,7-dimethoxyflavone (153) (HepG2; IC₅₀: 16.7 μM) (Trieu et al. 2021).

Goniothalamus velutinus Airy Shaw

This endemic primary rainforest produces nephrotoxic aristolactam alkaloids (aristolactam I, aristolactam AII, and aristolactam BII) (Iqbal et al. 2018). A methanol extract of bark (collected in Brunei) at a concentration of 50 μg/mL abrogated the survival of brine-shrimp s and inhibited the survival of A549 cell line with an IC₅₀ value of 26.3 μg/mL, and induced apoptosis (Erum et al. 2016). From the stems (collected in Brunei), velutinam (154) (Figure 28) inhibited the proliferation of cervical carcinoma cell line (CaSki) with an IC₅₀ value of 10.9 μg/mL (Iqbal et al. 2018), but is nephrotoxic (Xue et al. 2025). Other compounds found in the bark (collected in Sarawak) were the flavanones, pinocembrine (155) and naringenin (156), as well as goniothalamin (139) (Ahmad et al. 2006). Pinocembrine (155) was found to be cytotoxic to drug-resistant acute lymphoblastic leukemia cell line (CEM/ADR5000) (IC₅₀: 53.5 μM) (Joray et al. 2015). In mice xenografted with mouse fibrosarcoma cell line (S-180), the daily intraperitoneal administration of naringenin (156) at a dose of 300 mg/kg for five days caused a decrease in tumor weight by about 50% (Kanno et al. 2005).

Chemical structures of velutinam (154), pinocembrin (155, R=H), and naringenin (156, R=OH) with labels. The figure presents three chemical structures: velutinam (154) on the left, characterized by a three-ring complex with methoxy (OCH3), hydroxyl (OH), and amine (NH) groups. On the right, pinocembrin (155) and naringenin (156) are shown, both featuring a flavanone backbone. Pinocembrin (155) has R = H, while naringenin (156) has R = OH, with their structures clearly labeled and differentiated. — Cytotoxic compounds of *G. velutinus.*

Polyalthia tenuipes Merr. (Annonaceae)

Plants of the genus Polyalthia Blume (1930) are a prolific source of diterpenes and cytotoxic isoquinoline alkaloids. As for the diterpenes we can cite 16,16-dimethoxy-cleroda-3,13-Z-dien-15-oic acid (157) (Figure 29) from the leaves of Polyalthia simiarum (Buch.-Ham. ex Hook. f. & Thomson) Benth. ex Hook. f. & Thomson (collected in China) (SMMC-7721; IC₅₀: 22.4 μM) (Duan et al. 2020), 16α-hydroxycleroda-3,13(14)Z-dien-15,16-olide (158) from Polyalthia peteloti Merr. (Yang et al. 2016) and Polyalthia barnesii Merr. (Ma et al. 1994), and polylauiamide C (159) (a clerodane diterpene dimer derivative) isolated from the roots of Polyalthia laui Merr. (HeLa: IC₅₀: 25.1 μM) (Yu et al. 2016).

Eight chemical structures of compounds labeled 157 to 164, each with distinct functional groups and features. The image shows eight labeled chemical structures arranged in two rows: 1) "16,16-dimethoxy-cleroda-3,13Z-dien-15-oic acid" (157) with a carboxylic acid; 2) "16a-hydroxycleroda-3,13(14)Z-dien-15,16-olide" (158) featuring a cyclic structure; 3) "polylauiamide C" (159) with an amide linkage; 4) "longitriol" (162) characterized by hydroxyl groups; 5) "(-)-5-hydroxygoniothalamin" (163) showing an aromatic ring; 6) "(-)-anonaine" (160) containing nitrogen; 7) "consanguine B" (161) featuring multiple hydroxyl groups; 8) "(+)-rumphiin" (164) with methoxy groups. Each compound is distinctly labeled for identification. — Cytotoxic natural products from plants of the genus *Polyalthia.*

An aporphine alkaloid, (–)-anonaine (160), from the leaves of Polyalthia longifolia var. pendula Benthall (collected in Taiwan), abrogated the survival of AGS cell line (IC₅₀: 8.6 µM) (Chen et al. 2000). From the stems and leaves of Polyalthia obliqua Hook.f. & Thomson (collected in China), consanguine B (161) (an oxoprotoberberine alkaloid) was found to be cytotoxic to HeLa and MCF-7 cell lines with IC₅₀ values of 24.1 and 33.5 μM, respectively (Wu et al. 2016).

Longitriol (162), a cycloartane-type triterpene from the leaves of P. longifolia var. pendula, inhibited the growth of uterine cancer cell line (C33A) and A549 cell line with IC₅₀ values of 10 and 13.1 µg/mL, respectively (Sashidhara et al. 2010). Other compounds are (−)-5-hydroxygoniothalamin (163) (a styrylpyrone) from the leaves of Polyalthia parviflora (collected in Vietnam) (A549, IC₅₀: 7.9 μM) (Liou et al. 2014) and (+)-rumphiin (164) (a phenylpropanoid dimer) isolated from the stems of Polyalthia rumphii (Blume ex Hensch.) Merr. (collected in China) (K562, IC₅₀: 63.2 µg/mL) (Wang et al. 2013).

Organic polar and mid-polar extracts of plants in this genus have been reported to be cytotoxic from Polyalthia cerasoides (Roxb.) Bedd. (ethanol; L929; IC₅₀ ≈ 40 µg/mL) (Ravikumar et al. 2008), Polyalthia debilis (Pierre) Finet & Gagnep. (chloroform extract; HepG2; 23 µg/mL) (Prachayasittikul et al. 2009), Polyalthia evecta (Pierre) Finet & Gagnep. (ethanol; HepG2; IC₅₀: 62.8 µg/mL) (Macana et al. 2012). Essential oil of stems of Polyalthia suberosa (Roxb.) Thwaites was found to be cytotoxic to MCF-7 cell line (IC₅₀: 66.7 μg/mL) (The et al. 2021).

Eriocaulon longifolium Nees ex Kunth (Eriocaulaceae)

Ethyl acetate extract of the whole plant (purchased in China) inhibited the growth of K562 cell line (IC₅₀ ≈ 40 µg/L), and induced apoptosis with cell cycle arrest in G₀/G₁, inhibition of aurora kinases A and B kinase, upregulation of p53 and Bax, downregulation of Bcl-2, activation of caspases-3 and −9, release of cytochrome c, and cleavage of poly (ADP-ribose) polymerase (Fan et al. 2016). From this plant, hispidulin (165) (a flavone), quercetin-3-O-(6″-O-galloyl)-β-D-galactopyranoside (166) (a flavone glycoside), and corilagin (167) (an ellagitannin) inhibited the proliferation of HepG2 cell line with IC₅₀ values of 17.8, 11.2, and 16.6 μM, respectively and induced apoptosis (Fan et al. 2015) (Figure 30).

Four chemical structures of flavonoids: hispidulin, jaceosidin, quercetin-3-O-(6''-O-galloyl)-ß-D-galactopyranoside, and corilagin, labeled with their respective identifiers. The figure illustrates the molecular structures of four flavonoids arranged horizontally. From left to right: 1) Hispidulin (165) features various hydroxyl and methoxy groups. 2) Jaceosidin (168) is similar but has a methoxy group instead of hydrogen. 3) Quercetin-3-O-(6''-O-galloyl)-ß-D-galactopyranoside (166) includes multiple hydroxyl groups and a unique sugar moiety. 4) Corilagin (167) features several hydroxyls and a distinct arrangement, all labeled for identification. — Cytotoxic phenolic compounds from plants of the genus *Polyalthia.*

A dichloromethane extract of Eriocaulon cinereum R.Br. (collected in Indonesia) was found to be moderately cytotoxic to MCF-7 cell line (IC₅₀: 101.1 µg/mL) (Nugraha et al. 2021). From the capitula of Eriocaulon australe R.Br. (collected in China), hispidulin (165) and jaceosidin (168) (a flavone) were found to be cytotoxic to MCF-7 cell line with IC₅₀ values of 7.6 and 15.6 µg/mL, respectively (Xu et al. 2013).

Dendrocalamus asper (Schult. f.) Backer ex K. Heyne (Poaceae)

An ethanolic extract of edible young shoots (harvested in Sulawesi) inhibited the growth of MCF-7 cell line (IC₅₀: 1.4 µg/mL) (Ontaha et al. 2021). The active compounds responsible for this activity are not yet known. However, it is known that plants of the genus Dendrocalamus Nees (1835) produce rutin (21) (Luo et al. 2022). The plant, which is cultivable (Table 2), produces cyanogenic compounds (Pattarathitiwat et al. 2021); the Dusun mitigate its toxicity by boiling it prior to consumption.

Panicum palmifolium J. Koenig (Poaceae)

An extract of the plant (collected in the Philippines) showed some activity against KB cell line (Spjut, 2005). The molecules responsible for this activity are unknown, but phenolic compounds, saponins, and cyclotides can be expected (Figure 31). A tetralin derivative (168) isolated from Panicum turgidum Forssk. was found to be cytotoxic against ovarian carcinoma (SK-OV-3) and breast carcinoma cell line (BT-549) with IC₅₀ values of 5.6 and 10.3 µg/mL, respectively (Zaki et al. 2023). P. turgidum also produces cytotoxic steroidal saponins such as pennogenin 3β-O-α-L-rhamnopyranosyl-(1→2)-O-[α-L-rhamnopyranosyl-(1→4)-O-α-L-rhamnopyranosyl-(1→4)]-O-β-D-glucopyranoside (169) (Zaki et al. 2017). From the aerial parts of the South American Panicum laxum Sw., some cyclotides such as panitide L1 (170) were found to be cytotoxic to HeLa cell line (Nguyen et al. 2013). From the seeds of Panicum miliaceum L. (purchased in India), vanillin (171) (a simple phenolic) was found to be weakly cytotoxic to colon cancer cell line at the concentration of 250 µg/mL with DNA fragmentation, cell cycle arrest in the G0/G1 phase, and apoptosis (Ramadoss and Sivalingam 2020). It is not known for sure whether P. palmifolium is toxic, but Merrill (1943) notes that the seeds are used as a famine food in the Philippines. More experiments are needed.

Chemical structures of a tetralin derivative, penogenin, a peptide, and vanillin, all labeled. The figure presents four chemical structures on a white background. On the left is a tetralin derivative (168) with a hydroxyl group. Next is penogenin (169), a complex steroidal saponin with multiple hydroxyl groups and a branched oligosaccharide chain. The third panel features a peptide sequence (170), identified as panitide L1, and noted for containing three disulfide bonds. Lastly, on the right, vanillin (171) is shown, characterized by its methoxy and aldehyde groups attached to a benzene ring. Each compound is clearly labeled. — Cytotoxic compounds from plants of the genus *Panicum.*

Plagiostachys albiflora ridl. (Zingiberaceae)

This cultivable ginger is employed as a medicinal food (Table 1 and 2), and as a member of the Zingiberaceae family, it may represent a promising candidate for nutraceutical development. Its chemotherapeutic properties need to be studied, as well as its toxicity. Note that plants from the genus Plagiostachys Ridl. (1899) do not appear to have been the subject of any phytochemical or pharmacological studies.

Ampelocissus polita (miq.) Pelser

This vine is used as a medicinal vegetable (Table 1). It appears to have not been the subject of any phytochemical, pharmacological, or toxicological studies. It should be noted that Ampelocissus martini Planch is used as a vegetable in Thailand, which could, prima facie, indicate that these plants are harmless. An aqueous extract of the aerial parts of Ampelocissus latifolia (Roxb.) Planch. (collected in India) was found to be cytotoxic to Dalton lymphoma cell line (IC₅₀: 16 µg/mL), induced DNA fragmentation, and apoptosis (Chaudhuri and Ray 2020). Plant in the genus Ampelocissus Planch. (1884) produce cytotoxic stilbenoids such as (–)-α-viniferin (172) (Huang et al. 2021; Thanasansurapong et al. 2022) (Figure 32).

Chemical structure of (-)-a-viniferin (172) with multiple hydroxyl (OH) groups and interconnected aromatic rings. The figure illustrates the chemical structure of (-)-a-viniferin (172), a complex polycyclic polyphenol. It features three fused dihydrobenzofuran units, each substituted with hydroxyl (OH) groups, totaling six hydroxyls. The structure has a pseudo-triangular arrangement and includes branching points with clear representations of carbon rings, hydroxyls, and hydrogen atoms. The molecular formula is displayed, emphasizing the compound’s symmetrical design and potential chemical reactivity in biological contexts. — Cytotoxic stilbenoid from plants of the genus *Ampelocissus.*

Galearia fulva (Tul.) Miq. (Pandaceae)

This small shrub of the primary forest is used as medicinal food (Table 1). Nothing is known about the toxicity, phytochemistry, and pharmacology of this plant. A methanol extract of the leaves (collected in Malaysia) showed no toxicity to Vero cell line (Rizwana et al. 2010).

Melastoma beccarianum Cogn. (Melastomataceae)

This shrubby endemic plant does not appear to have been the subject of toxicological, phytochemical, or cytotoxic studies. The cytotoxic principles of the genus Melastoma L. (1753) are phenolic in nature. Examples include naringenin (156) and kaempferol-3-O-(2ʺ,6ʺ-di-O-p-trans-coumaroyl)-β-glucopyranoside (173) from the flowers of Melastoma malabathricum L. (collected in Malaysia), which inhibited the growth of MCF-7 cell line with IC₅₀ values of 1.3 and 0.2 µM, respectively (Susanti et al. 2007) (Figure 33).

Chemical structure of kaempferol-3-O-(2'',6''-di-O-p-trans-coumaroyl)-ß-glucopyranoside, featuring multiple aromatic rings and hydroxyl groups. The image presents the chemical structure of kaempferol-3-O-(2'',6''-di-O-p-trans-coumaroyl)-ß-glucopyranoside, illustrating a complex arrangement of multiple aromatic rings linked to a central ß-glucopyranoside unit. It highlights several hydroxyl (-OH) groups positioned on the rings and includes two p-coumaroyl groups esterified to the sugar moiety. The structure features distinct bonds and functional groups that characterize the molecular composition and connectivity, providing insight into the compound's chemical nature. — Cytotoxic flavone glycoside from plants of the genus *Melastoma.*

Mangifera pajang Kosterm (Anacardiaceae)

The fruits of this wild mango tree, endemic to Borneo and cultivable (Table 1 and 2), are edible (John et al. 2025) and used as medicinal food (Maid et al. 2017) (Table 1). An ethanolic seed extract inhibited the proliferation of MCF-7 and MDA-MB-231 cell lines with the IC₅₀ values of 23 and 30.5 μg/mL, respectively. For MCF-7 cell line, the extract induced apoptosis via activation of caspases-2, −3, −6, and −8 (Bakar et al. 2010). Likewise, a seed extract (collected in Malaysia) was found to be cytotoxic to colon cancer cell line (IC₅₀: 63 μg/mL) (Fadzelly Abu Bakar et al. 2010), which contained methyl gallate (59) that inhibited the growth of MCF-7 cell line, (IC₅₀: 81 μM), induced cell cycle arrest in G0/G1 and prompted oxidative damages (Yazan et al. 2022).

Consumption of fruit juice for nine weeks improved the plasma antioxidant capacity of subjects (Ibrahim et al. 2013). These results suggest that these fruits may have potential as nutraceuticals to prevent or improve the health of cancer patients; however, further clinical trials are necessary.

Canarium littorale Bl. (Burseraceae)

The fruits of this tree from the swamp forests are consumed for medicinal purposes (Table 1), but their toxicology, phytochemistry, and pharmacological properties remain unknown. Polar organic extracts of plants in the genus Canarium L. (1754) demonstrated cytotoxic activities as in the acetone extracts of the stem bark of Canarium odontophyllum Miq. (collected in Sarawak) (HCT-116; 50 µg/mL) (Basri et al. 2015; 2016; Ishak et al. 2023) and an ethanol extract of bark of Canarium ovatum Engl. (Balbuena et al. 2019).

The biflavone amentoflavone (174) and protocatechuic acid (132) were isolated from the fruits of a plant of the genus Canarium L. (1759) (collected in Taiwan) (Kuo et al. 2019). Amentoflavone (174) was found to be cytotoxic to HCT-116 cell line (IC₅₀ ≈ 100 µg/mL) (Fang et al. 2025) (Figure 34) and administered to mice xenografted with Ehrlich ascites tumor cell line, enhancing natural killer cell line (NK) activity (Guruvayoorappan and Kuttan 2007). Furthermore, administration of amentoflavone (174) to mice (100 mg/kg/week) xenografted with colon cancer cell line (CT26) prompted the reduction of tumor volume (Fang et al. 2025). Orally administered protocatechuic acid (132) prevented tumor formation in mice exposed to various types of carcinogens (Tanaka et al. 2011).

Chemical structure of amentoflavone (174) with interconnected aromatic rings and hydroxyl groups. The image illustrates the chemical structure of amentoflavone (174), featuring two identical flavone units linked by a central hydroxylated benzene ring. It exhibits multiple interconnected aromatic rings with six hydroxyl (-OH) groups attached in various positions. The structure's symmetry and complexity are emphasized, with the molecular name clearly labeled at the bottom. — Cytotoxic biflavone from plants of the genus *Canarium.*

Dacryodes incurvata (Engl.) H.J. Lam (Burseraceae)

The fruits of this primary forest tree are consumed for medicinal purposes (Table 1). These fruits do not appear to have been the subject of toxicological, phytochemical or pharmacological studies. An aqueous extract of Dacryodes edulis (G.Don) H.J.Lam leaves (collected in Cameroon) administered orally (100 mg/kg/day) for two weeks inhibited the growth of breast tumors (by 42%) induced by 7,12-dimethylbenz[a]anthracene in rats (Mvondo et al. 2021). An ethanol extract of fruit peels (collected in Kalimantan) was found to be cytotoxic to T-47D epithelial cell line (IC₅₀: 143 ppm) (Widyanto et al. 2021).

Nephelium uncinatum radlk. ex leenh (Sapindaceae)

The fruits of this cultivable tree are consumed for medicinal purposes by the Dusun (Table 1 and 2) but also by the Dayaks of East Kalimantan (Matius et al. 2018). These fruits do not appear to have been the subject of toxicological, phytochemical or pharmacological studies. An aqueous extract of Nephelium ramboutan-ake (Labill.) Leenh. was found to be cytotoxic to HT-29 cell line (IC₅₀: 16.6 µg/mL) and induced apoptosis with DNA fragmentation, mitochondrial dysfunction, increased reactive oxygen species, increased Bax protein expression, and induced activation of caspase-3, −7 and −9 (Chan et al. 2012). Aqueous extract of peels of Nephelium lappaceum L was found to be cytotoxic to MCF-7 cell line (IC₅₀: 94.1 µg/mL) (Jantapaso and Mittraparp-Arthorn 2022). N. lappaceum is also known to produce geraniin (175) (an ellagitannin) (Figure 35) (Abdul Ahmad et al. 2017), which at concentrations of 80 µM, inhibited the viability of glioblastoma U87MG and LN229 cell line by about 50% and induced apoptosis with increased expression of caspase-3. In mice xenografted with U87MG, geraniin (175) given at a dose of 60 mg/kg/day for 20 days caused a decrease in tumor weight by 44% (Ren et al. 2017). Saponins such as nephelioside I (176) with weak cytotoxic activities (Lu1 ED₅₀: 19.5 µg/mL; LNCaP, MCF-7 and HUVEC >20 µg/mL) were isolated from the bark of Nephelium maingayi Hiern collected in Indonesia (Ito et al. 2004).

Chemical structures of geraniin (175) in two forms and nephelioside I (176), with a reversible reaction arrow. The figure presents three chemical structures: two forms of geraniin (175) on the left, highlighting a large polyphenolic compound with hydroxyl groups and a cyclized isomer. The center shows a reversible reaction arrow connecting them. On the right, nephelioside I (176) is depicted, a triterpene glycoside featuring multiple hydroxyl groups and a distinct carbon structure. The arrangement emphasizes the relationship and transformations among these compounds. — Cytotoxic ellagitannin and triterpene saponin from plants of the genus *Nephelium.*

Embelia dasythyrsa miq. (Primulaceae)

This vine is used as a medicinal food and for fever and the Dusun even eat the leaves raw (personal observation) (Table 1). It does not appear to have been the subject of toxicological, phytochemical, or pharmacological studies. It probably contains long-chain cytotoxic alkyl benzoquinones and alkyl resorcinols, which are often produced by plants of the genus Embelia Burm.f. (1768) and the genus Ardisia Sw. (1788). Examples include embelia-alkylresorcinols C (177) from Embelia ribes Burm.f. (Chen et al. 2018), as well as embelin (178) (MCF-7; IC₅₀: 80 µg/mL) (Kaur et al. 2015), which induced apoptosis in PC-3 cell line through downregulation of the Akt/mTOR/S6K1 pathway (Kim et al. 2013) (Figure 35). Ardisianone (179), cornudentanone (180), and ardisianol (181) from Ardisia virens Kurz were found to be cytotoxic to MCF-7 cell line (Chang et al. 2009) (Figure 36).

Six chemical structures: embeliaalkylresorcinol C, embelin, ardisianone, cornudentanone, ardisianol, and ardisuloside. The image features six chemical structures arranged in two rows. The top row includes embeliaalkylresorcinol C (177), showing a benzene ring attached to a long aliphatic chain; embelin (178), with two carbonyl groups and a hydroxyl; and ardisianone (179), characterized by a methoxy group and a long chain. The bottom row presents cornudentanone (180), similar to ardisianone; ardisianol (181), a polysubstituted benzene; and ardisuloside (182), a complex triterpenoid glycoside with multiple hydroxyls. Each structure is labeled with its name and number. — Cytotoxic compounds from plants of the genus *Embelia.*

Other cytotoxic constituents found in plants of the genus Ardisia are oleanane-type triterpenoid saponins. Such compounds have been identified in Ardisia insularis Mez, including ardinsuloside (182), which inhibits the growth of A-549, HT-29, and ovarian carcinoma cell line (OVCAR) with IC₅₀ values of 8.5, 16.4, and 13.6 μM, respectively (Van et al. 2015). More experiments are needed.

Chassalia chartacea craib (Rubiaceae)

What is known about the phytochemistry and anticancer properties of this shrub remains very limited. The presence of cytotoxic chassatide-type cyclotides, such as cyclodite C8 (183) (Nguyen et al. 2012) (Figure 37), has been reported. A methanolic extract has also been found to be cytotoxic to brine-shrimp (LC₅₀: 27.8 µg/mL) (Runa 2019). An alkaloid extract from the roots induced apoptosis of A549 cell line (IC₅₀: 8.2 µg/mL) with cell cycle arrest in the sub-G0 phase (Gopal et al. 2022). The plant produces indole alkaloids (Schinnerl et al. 2012), whose cytotoxic effects could be evaluated.

Amino acid sequence of cyclodite C8 (183) showing branching connections and labeled disulfide bonds. The image displays the amino acid sequence of cyclodite C8 (183), formatted in a linear arrangement as "AIPCGESCVWIPC-ISTVIGCSCSNKV-CYR." It features branching connections that visually organize the sequence, emphasizing structural hierarchies. Additionally, labels indicate the presence of three disulfide bonds between specific amino acids: C4-C27, C13-C22, and C8-C20. — Cytotoxic cyclotide from plants of *C. chartacea.*

Neonauclea gigantea (valeton) merr. (Rubiaceae)

This endemic tree does not appear to have been the subject of toxicological, phytochemical, or pharmacological studies. Plants of the genus Neonauclea Merr. (1915) are producers of cytotoxic phenolic principles. Examples are p-coumaric acid (184) (a phenylpropanoid), ficusal (185) (a 2-aryldihydrobenzofuran neolignan), and balanophonin (186) (a 2-aryldihydrobenzofuran neolignan) (Figure 38) from the stems of Neonauclea reticulata (Havil.) Merr. (collected in Taiwan), which were found to be cytotoxic to hepatocellular carcinoma cell line (Hep3B) with IC₅₀ values of 85.3, 92.6, and 29.1 µg/mL, respectively (Chang et al. 2018). 6-Dimethoxy-1,4-benzoquinone (187) from the bark of Neonauclea purpurea (Roxb.) Merr. (collected in Vietnam) was toxic to Vero cell line (IC₅₀: 1.1 µM) (Karaket et al. 2012).

Chemical structures of p-coumaric acid, ficusal, balanophonin, 2,6-dimethoxy-1,4-benzoquinone, and neonaucleoside A. The image features the chemical structures of five compounds: p-coumaric acid (184) with a phenolic structure and carboxylic acid; ficusal (185) with a benzofuran core and multiple hydroxyl groups; balanophonin (186), similar to ficusal but with different substituents; 2,6-dimethoxy-1,4-benzoquinone (187) identifiable by its quinone ring; and neonaucleoside A (188) with a complex polycyclic structure that includes an indole and multiple functional groups. Each structure is labeled for reference. — Cytotoxic natural products from plants of the genus *Neonauclea.*

Plants of this genus produce monoterpene indole alkaloid glycosides such as neonaucleoside A (188) from Neonauclea sessilifolia (Roxb.) Merr. (Itoh et al. 2003).

Psychotria gyrulosa stapf (Rubiaceae)

Plants of the genus Psychotria L. (1759) produce clinically useful alkaloids such as vincosamide (189) (Figure 39) from Psychotria leiocarpa Cham. & Schltdl. which inhibited hepatoma cell line (HLE) growth by about 60% at the concentration of 80 µg/mL and induced apoptosis with mitochondrial dysfunction, increased expression of Bax, decreased expression of Bcl-2, increased expression of caspase-3, and decreased expression of phosphorylated Akt. This alkaloid administered intraperitoneally at a dose of 10 mg/kg/day for 21 days to mice xenografted with hepatocellular carcinoma cell line (Bel 7402) cell line, caused a decrease in tumor volume from about 1200 to 400 mm³ (Zhu et al. 2022). We can also mention emetine (190) (a quinoline alkaloid), from Psychotria ipecacuanha (Brot.) Stokes, which inhibited the growth of T cell leukemia (Jurkat) (IC₅₀: 0.1 µM) and induced apoptosis with DNA fragmentation, mitochondrial dysfunction, and activation of caspase-3 (Möller and Wink 2007). Emetine was able to increase the effectiveness of cisplatin on Jurkat cell line (Möller et al. 2007). It is a molecule used in therapy as an antiprotozoal agent and may have clinical application for the treatment of cancer.

Chemical structures of seven compounds including vincosamide, emetine, and quercetin derivatives, labeled by numbers. The figure illustrates seven organic chemical structures arranged in two rows, featuring compounds like vincosamide (189) with complex rings, emetine (190) with aromatic structures, and two quercetin derivatives (191, 192) displaying glycoside features. Other compounds include asparuloside (193), psychorubrin (194), and helenalin (195), each identified by corresponding numbers. Each structure includes various functional groups and is characterized by specific arrangements of rings and atoms. — Cytotoxic natural products from plants of the genus *Psychotria.*

Other cytotoxic principles are quercetin 3-O-rutinoside (191), quercetin-3-O-glucopyranoside (192) (flavonol glycosides), and asperuloside (193) (an iridoid glycoside) (HT-29; IC₅₀ ˂ 50 µM) from the leaves of Psychotria luzoniensis (Cham. & Schltdl.) Fern.-Vill., (Ramil et al. 2021), psychorubrin (194) (a naphthoquinone) (KB; IC₅₀: 3 µg/mL), helenalin (195) (a sesquiterpene lactone) (T-47D; IC₅₀: 4.6 µM) (Barkhordari et al. 2023), and cyclotides from Psychotria leptothyrsa Miq. (collected in Hawaii) (Gerlach et al. 2010). Although plants in this genus are generally not aromatic, the essential oil of the leaves of Psychotria asiatica L. (collected in Vietnam) was found to be cytotoxic to lung adenocarcinoma cell line (SK-LU-1; IC₅₀: 39.7 µg/mL (Tran et al. 2025). Methanol and ethyl acetate extracts of Psychotria serpens L. (collected in Taiwan) inhibited the growth of KB cell line (IC₅₀: 20 µg/mL) (Lee et al. 1988) and HepG2 cell line (Wang et al. 2020). The toxicity, phytochemistry, and pharmacological properties of P. gyrulosa, a shrub endemic to the rainforest of Borneo, have not been studied.

Other plants used by the Dusun

Kadsura lanceolata King has not been the subject of any phytochemical, pharmacological, or toxicological studies

Plants used by the Rungus (Bornean group, Dusunic family)

Guioa bijuga (hiern) radlk (Sapindaceae)

This coastal tree serves as a medicinal food source for the Rungus (Table 1). It does not seem to have been the subject of any toxicological, phytochemical, or pharmacological studies. Extracts from some plants in the genus Guioa Cav. (1798) have shown cytotoxic activities (Balunas et al. 2006).

Ixora capillaris bremek

This endemic shrub does not appear to have been the subject of any toxicological, phytochemical, or pharmacological studies. Extracts of flowers of Ixora javanica (Blume) DC were found to be cytotoxic to DLA and Ehrlich ascites tumor cell line at a concentration of 12 and 65 μg/mL, respectively, while being less toxic to normal lymphocytes (Nair and Panikkar 1990) and given orally at a dose of 100 mg/kg inhibited the growth of soft tissue fibrosarcoma induced by 20-methylcholanthrene (Nair et al. 1991). Other examples of extracts with cytotoxic activities are a chloroform extract of flowers of I. coccinea (collected in Brazil) (HL-60; IC₅₀: 36.9 μg/mL) (da Silva et al. 2016) and an extract of stems of Ixora brevifolia Benth was found to be cytotoxic to glioma (U251) and K562 cell line with IC₅₀ values of 28.6 and 28 μg/mL, respectively (Medina et al. 2018).

A hexane extract of Ixora coccinea L. (collected in India) inhibited the growth of Dalton’s lymphoma cell line (DLA) (IC₅₀: 25 μg/mL), with inhibition of DNA synthesis. Given intraperitoneally at a dose of 200 mg/kg to mice xenografted with DLA cell line, this extract caused an increase in life span by 59.2% (Latha and Panikkar 1998). The flowers of I. coccinea contain ixorene isovalerate (196) (a dammarane-type triterpene) (Figure 40) which abrogated the survival of HeLa cell line (Ikram et al. 2016).

Chemical structure of ixorene isovalerrate (196) with multiple rings and side chains. The image illustrates the 2D chemical structure of ixorene isovalerrate (196), featuring a central hydrophobic core with three fused six-membered rings and one five-membered ring. A branched isovalerate side chain is attached, showcasing various functional groups. Bonds between carbon atoms are represented by lines and wedges to indicate 3D orientation, with double bonds and stereochemistry marked accordingly, reflecting the molecule's complex architecture. — Cytotoxic triterpene from plants of the genus *Ixora.*

Plants used by the murut (bornean group, murutic family)

General observations

Regarding the medicinal plant species used by the Murut with possible chemotherapeutic potential, the following observations can be made: (i) about half of them are endemic, (ii) these plants are often from primary rainforest.

Eusideroxylon zwageri teijsm. & binn. (Lauraceae)

This timber tree does not seem to have been phytochemically or pharmacologically studied. It is used for the preparation of blow-gun darts, poisons (Kulip, 2003), suggesting the occurrence of bisbenzylisoquinoline alkaloids. An extract of bark was found to be cytotoxic to T-47D cell line (IC₅₀: 237.5 µg/mL) (Kurniawan et al. 2023). This plant is phylogenetically close to plants in the genus Cryptocarya R.Br. (1810) where various classes of cytotoxic natural products have been identified. From the bark of Cryptocarya laevigata Blume was characterized (–)-neocaryachine (197) (a pavine alkaloid), toxic to multidrug-resistant cervical cancer cell line (KB-VIN) (IC₅₀: 0.2 µM), and caused DNA damages and apoptosis (Suzuki et al. 2017) (Figure 41). (–)-Antofine (198) (a phenanthroindolozidine alkaloid) isolated from the wood of Cryptocarya chinensis (Hance) Hemsl. was found to inhibit the growth to ileocecal adenocarcinoma cell line (HCT-8) (IC₅₀: 0.001 µg/mL) (Wu et al. 2012). Other cytotoxic principles in this genus include flavanones such as cryptometcone I (199) from Cryptocarya metcalfiana C.K. Allen (He et al. 2022), chalcones such as 2,4′-dihydroxy-5′,6′-dimethoxychalcone (200) (P388; IC₅₀: 5.7 µM) from Cryptocarya costata Blume (Usman et al. 2006), (–)-grandisin (201) (a lignan) from Cryptocarya crassinervia Miq. (Saad et al. 1991), and α-pyrones such as obolactone (202) from Cryptocarya obovata R.Br. (Dumontet et al. 2004), and arylalkenyl α,β-unsaturated δ-lactones such as cryptobrachytone A (203) from Cryptocarya brachythyrsa H.W.Li (Fan et al. 2019). E zwageri is cultivable (Table 2). Phytochemical and pharmacological studies on this tree are necessary.

Chemical structures of seven compounds labeled 197 to 203, featuring diverse ring systems and functional groups. The image displays the chemical structures of seven compounds arranged in two rows. From top left to bottom right: (-)-neocaryachine (197) with multiple benzene rings and hydroxyl groups; (-)-antofine (198) featuring a nitrogen atom and several methoxy groups; cryptometcone I (199) showcasing complex rings and functional groups; 2',4'-dihydroxy-5',6'-dimethoxychalcone (200) with chalcone structure; (-)-grandisin (201) having aromatic and methoxy groups; obolactone (202) with a lactone ring; and cryptobrachytone A (203) with a long carbon chain and hydroxyl groups. Each structure is labeled with a corresponding name and number. — Cytotoxic natural products from plants of the genus *Cryptocarya.*

Millettia nieuwenhuisii J.J. Smith (fabaceae)

This endemic climber of the rainforest has not been studied. It is used to treat thrush, which may indicate the presence of a cytotoxic compound, as anticandidal agents are often active against cancer cell line (Routh et al. 2011). Plants of the genus Millettia Wight & Arn. (1834) are a source of prenylated isoflavones with cytotoxic properties. We can cite millexatin N (204), scandenone (205), and auriculatin (206) from the young twigs of Milletia extensa Benth. ex Baker f. (collected in Thailand), which were found to be cytotoxic to MDA-MB-231 cell line with the IC₅₀ values of 15.4, 13.9, and 15.3 µM, respectively (Cheenpracha et al. 2022) (Figure 42). From Milletia pachycarpa Benth. (collected in Thailand), euchrenone b₁₀ (207) inhibited the growth of K562 cell line (IC₅₀: 15.1 μM) (Suthiphasilp et al. 2022). From the seeds of M. pachycarpa, barbigerone (208) and millepachine (209) (a prenylated chalcone) were cytotoxic to HepG2 cell line with IC₅₀ values of 0.6 and 1.2 µM, respectively (Ye et al. 2012). From the stems of Millettia pachyloba Drake, (collected from China) was isolated durmillone (210), toxic to HeLa and MCF-7 cell line with IC₅₀ values of 6 and 11 µM, respectively. At the concentration of 20 µM, durmillone (210) induced apoptosis with cleavage of poly (ADP-ribose) polymerase in HeLa and MCF-7 cell line and induced cellular autophagy (Yan et al. 2019). Durmillone (210) isolated from Millettia dura Dunn, was found to be cytotoxic to A549 cell line (IC₅₀: 6.6 μM) while nontoxic to lung fibroblasts (CCD19Lu) cell line (IC₅₀ > 100 μM) (Buyinza et al. 2021). From the seeds of M. pachyloba, 6-methoxybarbigerone (211) and pachylobin (212) inhibited the growth of KB cell line with IC₅₀ values of 2 and 17.6 µM, respectively (Mai et al. 2010).

Chemical structures of ten compounds, identified by labels from 204 to 213. The image presents the chemical structures of ten organic compounds arranged in three rows, each labeled with a name and a corresponding number from 204 to 213. The compounds include millexatin N (204), scandenone (205), auriculatin (206), euchrenone b10 (207), barbigerone (208), millepachine (209), durmillone (210), 5-methoxybarbigerone (211), pachylobin (212), and millettinol (213). Each structure features distinct functional groups and ring systems, showcasing their chemical diversity with various hydroxyl and methoxy groups. The layout aids in the comparison of these unique compounds. — Cytotoxic phenolic compounds of plants in the genus *Milletia.*

Other types of cytotoxic principles in the genus Millettia are phenolic compounds such millettinol (213) (BCA-1; IC₅₀: 3.4 μg/mL) from Millettia leucantha Kurz (Kurz) Z.Q.Song (Rayanil et al. 2011), as well as oleanane triterpene saponins (Pertuit et al. 2022). Phytochemical and pharmacological studies on M. nieuwenhuisii are necessary.

Shorea parvistipulata F. Heim (Dipterocarpaceae)

Cytotoxic compounds have not been identified from this endemic timber tree. Plants in the genus Shorea Roxb. ex C.F. Gaertn. (1805) are known to produce cytotoxic stilbenoids. The bark of Shorea gibbosa Brandis (collected in Indonesia) contain (–)-hopeaphenol-12b-C-β-glucopyranoside (diptoindonesin F) (214), (–)-ampelopsin A (215), ampelopsin E (216), and (–)-hemsleyanol D (217), (–)-α-viniferin (172), and (–)-vaticanol B (218) (Figure 43), which were cytotoxic to P388 cell line with IC₅₀ values of 34.6, 17, 15.3, 94.7, 25.7, and 46.4 µM, respectively (Saroyobudiono et al. 2008) (Figure 43). From the stem bark of Shorea maxwelliana King, maximol A (219), vaticanol A (220), suffruticosol A (221), and vaticanol G (222) inhibited the growth of HL-60 cell line with IC₅₀ values ranging from 2.7 to 78 µg/mL (Zawai et al. 2013). (–)-Ampelopsin A (215) and (–)-hopeaphenol (223) from the stem bark of Shorea hopeifolia (F.Heim) Symington (collected in Malaysia) were active against HepG2 cell line with IC₅₀ values of 22.5 and 4.5 µg/mL, respectively (Rohaiza et al. 2011). From the bark of Shorea roxburghii G.Don, (−)-hopeaphenol (223), (–)-vaticanol B (218), (–)-hemsleyanol D (217), (+)-α-viniferin (224) (enantiomer of 172), and resveratrol (123) inhibited the growth of SK-MEL-28 cell line with IC₅₀ values of 3.6, 16.6, 15.5, 7.1, and 21 μg/mL, respectively (Moriyama et al. 2016). Other examples are hopeafuran (225) (P388; IC₅₀: 112.6 µM) (Sahidin et al. 2005), and isohopeaphenol (226) from Shorea roxburghii G.Don (Patcharamun et al. 2011; Ninomiya et al. 2017).

Structural formulas of 14 complex organic compounds, including hopeaphenol, ampelopsin, and vaticanol with distinct functional groups. The figure presents 14 chemical structures labeled from 214 to 227, featuring various natural products. Key compounds include (-)-hopeaphenol-12b-C-ß-glucopyranoside (214), (-)-ampelopsin A (215), (-)-hemsleyanol D (217), and a secodammarane triterpenoid (227). The structures display multiple hydroxyl (-OH) groups, benzene rings, and intricate connectivity, showcasing the complexity and diversity of organic molecules, with unique features such as fused cyclic systems and various stereochemistries. — Cytotoxic stibenoids and triterpenes of plants in the genus *Shorea.*

Other cytotoxic principles are dammarane-type triterpenes found in the resin of Shorea javanica Koord. & Valeton, which included (20S)-20-hydroxy-3,4-secodammara-4(28),24-dien-3-al (227) that protected mice against skin tumor formation (Ukiya et al. 2010). An ethyl acetate extract of Shorea roxburghii G.Don flowers (collected in Thailand) was found to be cytotoxic to AGS cell line (IC₅₀: 35.5 μg/mL) (Janthamala et al. 2024).

Dissochaeta monticola Bl. (Melastomataceae)

Despite Dissochaeta monticola being used to make blowgun dart poison, implying the potential presence of neurotoxic alkaloids, this climber, and indeed the entire genus Dissochaeta Blume (1831), has not been phytochemically or pharmacologically studied. This genus belongs to the tribe Dissochaeteae, which is phylogenetically close to the tribe Melastomateae (Kartonegoro et al. 2021).

Praravinia suberosa (Merr.) bremek (rubiaceae)

This endemic rainforest tree has not been studied, and the same appears to apply to the whole genus Praravinia Korth. (1842). Nevertheless, this genus is phylogenetically close to the genus Urophyllum Jack ex Wall. (1824) (Koizumi and Nagamasu, 2016) where alkaloids are present (Teo et al. 1990). An ethanol extract of leaves of Urophyllum arboreum (Reinw. ex-Blume) Korth. was found to be cytotoxic to MCF-7 cell line (IC₅₀: 136.3 µg/mL) (Jumaryatno et al. 2022).

Other plants used by the Murut

Goniothalamus woodii Merr. (Annonaceae) and Jasminum aculeatum Blco remain unstudied.

Ecological and ethnological considerations, and possible cultivation

The majority of these plants—with development potential either as a source of anticancer products or with the potential to be developed as nutraceutical or medicinal products—are rare species, originate from primary forests and form part of the traditional pharmacopeia of the Dusun, and to a lesser extent, that of the Murut (Table 2). The Dusun are the largest community in Sabah, followed by the Kadazan, Bajau, and Murut (Reid 1997). Native to Sabah, they are colloquially referred to as “people of the land” or “people of the orchard.” They are skilled farmers with a deep and respectful connection to the land. The Murut, or “people of the hills,” live primarily in southwest Sabah near primary rainforests and rivers (Prentice 1969), as well as in the uplands of southern Sabah (Kulip 2003). They have extensive knowledge of primary rainforests (Ahmad and Holdsworth 1994). Although cultivation information is lacking for most of these plants, some species with nutraceutical potentials can be cultivated, such as H. zeylanica (Un, 2010), P. tumefacta, M. platytyrea, and M. pajang (Tinggal and Tee, 1994) (Table 2).

Conclusions and future direction

Despite the progress made in oncology over the last few decades, the mortality rate from cancer, and in particular from pediatric cancers, due to tumors not responding to chemotherapy, remains unacceptably high. Several anticancer drugs come from the plant kingdom and it can reasonably be anticipated that the complete study of the approximately 374,000 species identified so far (Christenhusz and Byng 2016) will lead to the discovery of molecules that that could significantly reduce this disease burden. In parallel, there is a need to develop nutraceuticals whose consumption could prevent the development of tumors by, at least in part, reducing inflammation, stimulating the immune system, and/or protecting DNA against mutagens and reactive oxygen species.

In this context, we have selected, from among the 696 species of medicinal plants of Sabah recorded to date, 64 medicinal food plant species for which there are few or no phytopharmacological studies and which belong to families known to produce cytotoxic natural products. What emerges from this study can be condensed into the following major points. (i) Most of these plants are used by Bornean ethnic groups, and primarily by the Dusun and the Murut who live in areas with high plant endemism and who for hundreds of years have learned to use plants from their immediate environment to prevent and combat diseases. Although documentation efforts have already been made to document these plants, the available data are preliminary and fragmentary. Sabah’s medicinal flora is in a very precarious situation, and endemic or primary forest species are at risk of disappearing mainly due to incessant deforestation. Furthermore, modernization and Islamization leads to the loss of oral and ancestral knowledge about these plants. (ii) Most of the plants selected deserve to be studied in depth as possible sources of original natural products for the fight against cancer, and in particular the endemic species. (iii) The oral pharmacopeia of Sabah Bornean ethnic groups includes a significant number of plants consumed as food to maintain good health. Among these, H. zeylanica, P. tumefacta, M. platytyrea, and M. pajang are interesting material.

Although some of these plants have interesting activities in vitro and in vivo, there is the need of further experiments. For the plants whose active principles have been identified or even food plants more experiments are needed to confirm their possible oncopreventive or anticancer properties. It can be also mentioned here that in vitro cytotoxicity is not an absolute hallmark of possible anticancer application but a simple demonstration that a natural product kills a cancer cell. It is an early indication that must be used for further in vivo studies and should the compound be well tolerated and effective, clinical studies become necessary. In addition, toxicological data are largely absent, making translational claims premature. Furthermore, there are no available data of the variability of the plant extracts mentioned (seasonal, geographical, preparation methods) and without phytochemical standardization, reproducibility and clinical translation remain questionable.

To date, there is no complete inventory of medicinal plants in Sabah, and the total number of these plants is estimated to be well over 696 species. However, it should be noted that deforestation to make way for palm oil plantations and the gradual loss of ancestral knowledge threaten to cause the disappearance of a large number of plants. In sum, Sabah’s medicinal food plants still represents, but for how long?, a source of potential natural products and nutraceuticals for the fight against cancer. Will we let this opportunity pass us by?

Funding Statement

The author(s) reported there is no funding associated with the work featured in this article.

Disclosure statement

No potential conflict of interest was reported by the author(s).

Data availability statement

Data sharing is not applicable to this article as no data were created or analyzed in this research.

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PERMALINK

Medicinal food plants of Sabah (Eastern Malaysia): a source of potential natural products and nutraceuticals for the fight against cancer

Carynn Tanbuda

Mazdida Sulaiman

Pauline Yong Pau Lin

Fiffy Hasnidah Saikim

Nor Azizun Ruzdi

Mogana Rajagopal

Nicholas Pang Tze Ping

Nor Hayati Abdullah

Jhonnel Villegas

Sanen Marshall

Mukesh Singh Sikarwar

Veeranoot Nissapatorn

Prapairat Seephonkai

Marcelo Iriti

Mark S Butler

Christophe Wiart

Roles

Abstract

Context

Objectives

Methods

Results

Conclusion

Introduction

Methodology

Plants used by multiple ethnic groups

General observations

Table 1.

Table 2.

Selaginella argentea (wall. ex-hook. & grev.) spring (Selaginellaceae)

Figure 1.

Helminthostachys zeylanica (L.) hook. (Ophioglossaceae)

Figure 2.

Stenochlaena palustris (burm. f.) bedd. (Blechnaceae)

Figure 3.

Drynaria sparsisora (desv.) T. Moor

Figure 4.

Lygodium circinnatum Sw. (Lygodiaceae)

Figure 5.

Litsea garciae vidal

Figure 6.

Artabotrys roseus boerl. (Annonaceae)

Figure 7.

Boesenbergia pulchella (ridl.) merr. (Zingiberaceae)

Figure 8.

Etlingera elatior (jack) R.M. Sm

Pycnarrhena tumefacta miers (Menispermaceae)

Figure 9.

Bridelia stipularis (L.) Bl. (Phyllanthaceae)

Figure 10.

Ficus retusa L. (Moraceae)

Figure 11.

Myrmecodia platytyrea becc. (Rubiaceae)

Paederia verticillata Bl. (Rubiaceae)

Figure 12.

Rennellia borneensis baill. (Rubiaceae)

Figure 13.

Crassocephalum crepidioides (benth.) S. Moore

Figure 14.

Merremia gracilis E.J.F. Campb. & argent (Convolvulaceae)

Figure 15.

Merremia peltata (L.) merr. (Convolvulaceae)

Other plants used by multiple ethnic groups

Plants used by the Lundayeh (Kelabit group)

Kadsura borneensis A.C. (Schisandraceae)

Figure 16.

Nepenthes ampullaria jack (Nepenthaceae)

Figure 17.

Symplocos odoratissima choisy ex zoll. (Symplocaceae)

Figure 18.

Hydnophytum formicarum jack (Rubiaceae)

Figure 19.

Jasminum bifarium wall (Oleaceae)

Figure 20.

Other plants used by the Lundayeh

Plants used by the Brunei (Malayic group)

Drymoglossum piloselloides (L.) C. Presl (Polypodiaceae)

Figure 21.