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Iran

Genetic architecture of phenological, morphological, and phytochemical traits in Cannabis landraces

Mehdi Babaei, Davoud Torkamaneh

The Plant Genome· 2026

Genome-wide association study of 145 Iranian Cannabis sativa landrace accessions, male and female, genotyped at roughly 233,000 common SNPs. The analysis resolves the collection into three genetically distinct subpopulations shaped by geography, climate and traditional cultivation, and identifies 91 genomic regions associated with 40 phenological, morphological and phytochemical traits, including 15 pleiotropic loci linked to flowering time, plant architecture, biomass accumulation and cannabinoid biosynthesis. The authors report high heritability for most traits and rapid linkage disequilibrium decay, and present the panel as a resource for high-resolution mapping and marker-assisted selection towards cultivars with tailored cannabinoid profiles. A population-scale genetic-architecture reference for the Iran gene pool, complementing the earlier Mostafaei Dehnavi et al. 2025 population-genomics study of Iranian Cannabis.

Unraveling the Confusion of Flowering Types and Terminology in Cannabis sativa

Madigan Eckels; Royal Heins; Bruce Bugbee

HortScience (American Society for Horticultural Science)· 2026

Perspective/review arguing that cannabis “flowering” terminology is inconsistent across research and industry, which leads to misclassification of photoperiodic behavior and reproductive structures. Proposes clearer categories for female reproductive development (pistillate flowering) and distinguishes multiple male flowering expressions, with descriptive examples and photos to standardize usage across controlled-environment and field contexts.

South Africa

Effect of Propagation Techniques on Growth, Development, Oil Yield, and Quality of Medicinal Cannabis (Cannabis sativa) Found in Lusikisiki, Eastern Cape, South Africa

Azile Dumani, Tembakazi Theodora Silwana, Ifeanyi Moses Egbichi, Adebola Omowunmi Oyedeji, Babalwa Mpambani, Hlabana Alfred Seepe

Horticulturae· 2025

Tests propagation techniques and their effect on growth, oil yield and quality of the Lusikisiki medicinal cannabis material, extending the Dumani Eastern Cape survey into agronomy.

South Africa

Cannabis sativa in South Africa: history, legislation, production, and pharmacological aspects

Michaela Mouton, Minja Gerber, Robert Verpoorte, Frank van der Kooy

Phytochemistry Reviews· 2025

A review of Cannabis sativa in South Africa covering history, legislation, production and pharmacological aspects, synthesising the national picture.

Domesticated cannabinoid synthases amid a wild mosaic cannabis pangenome

Ryan C. Lynch, Lillian K. Padgitt-Cobb, Andrea R. Garfinkel, Brian J. Knaus, Nolan T. Hartwick, Nicholas Allsing, Anthony Aylward, Philip C. Bentz, Sarah B. Carey, Allen Mamerto, Justine K. Kitony, Kelly Colt, Emily R. Murray, Tiffany Duong, Heidi I. Chen, Aaron Trippe, Alex Harkess, Seth Crawford, Kelly Vining, Todd P. Michael

Nature· 2025

The first comprehensive cannabis pangenome, built from 193 genomes (181 new PacBio assemblies plus 12 previously published) representing 144 biological samples spanning use types, history, sex expression and agronomic traits. Includes 78 haplotype-resolved chromosome-scale assemblies and 103 contig-level assemblies covering both male (XY) and female (XX) plants. Documents surprisingly high genetic and structural variation for a single species and proposes a new population structure and hybridisation history. Resolves the heteromorphic X and Y sex chromosomes for the first time, including a variable boundary at the sex-determining and pseudoautosomal regions and male-biased expression of several flowering regulators. Finds that the cannabinoid synthase loci (CBDAS, THCAS) themselves carry very little diversity despite sitting in a region rich in pseudogenised paralogues, structural variation and distinct transposable element arrangements, evidence that the synthases have been recently domesticated against an otherwise wild-mosaic genomic background. Identifies acyl-lipid thioesterase variants associated with fatty acid chain length and with the production of the rarer propyl cannabinoids THCV and CBDV. Confirms that high-CBD hemp lineages such as CBDRx emerged through introgression of the CBDAS locus into a predominantly drug-type genetic background. Concludes that the Cannabis sativa gene pool remains only partly characterised, that wild relatives likely persist in Asia, and that the crop's broader potential is largely unrealised. Provides the new reference framework for placing landrace genotypes in a global population context and underwrites the conservation case that elite modern cultivars have lost diversity that landraces and wild-growing populations still hold.

Integrating target capture with whole genome sequencing of recent and natural history collections to explain the phylogeography of wild-growing and cultivated cannabis

Manica Balant, Daniel Vitales, Zhiqiang Wang, Zoltán Barina, Lin Fu, Tiangang Gao, Teresa Garnatje, Airy Gras, Muhammad Q. Hayat, Marine Oganesian, Jaume Pellicer, Alireza S. Salami, Alexey P. Seregin, Nina Stepanyan-Gandilyan, Nusrat Sultana, Shagdar Tsooj, Magsar Urgamal, Joan Vallès, Robin van Velzen, Lisa Pokorny

Plants, People, Planet· 2025

Global phylogeographic study using a Hyb-Seq approach that combines target capture (with the universal Angiosperms353 enrichment panel) and shotgun whole-genome sequencing on the same data, applied to wild-growing accessions, cultivars and herbarium specimens drawn from natural history collections. Phylogenomic and population genomic workflows on the integrated dataset support treating Cannabis as a monotypic genus and recover three main genetic groups: East Asia, Paleotropis, and Boreal. Paleotropis is further divided into Iranian Plateau, Central and South China–Himalayas, and Indoafrica subgroups. Boreal is divided into Eurosiberia–West Mongolia and Caucasus–Mediterranean subgroups. The strongest signal in the data is geographic rather than use-type, with cultivars clustering by region of origin rather than by hemp/drug distinction. Iranian and Central Asian wild-growing populations emerge as particularly distinctive and morphogenetically diverse, consistent with the idea that this region functions as a genetic melting pot. The study is the first global landrace-inclusive phylogeography of cannabis after Ren et al. 2021 and is methodologically significant for showing that herbarium specimens (including older natural history collections) can be integrated with fresh material to recover phylogenetic signal in a genus where field sampling is legally difficult. Provides the geographic backbone against which any landrace genotype on the wiki can be situated, complements the Lynch et al. 2025 pangenome at the population scale, and supports the conservation case that wild-growing populations from understudied regions hold diversity not represented in modern cultivars.

Unveiling Cannabinoids and Terpenes Diversity in Cannabis sativa L. From Northern India for Future Breeding Strategies

Dipayan Ghosh, Anitika Kundu, Neha Chaudhary, Darshan Gupta, Mridula Dwivedi, Ram Swaroop Verma, Birendra Kumar, Karuna Shanker, Narendra Kumar

Chemistry & Biodiversity· 2025

Cannabinoid and essential-oil profiling of Cannabis sativa collected from five populations across northern India, conducted at CSIR-Central Institute of Medicinal and Aromatic Plants (CIMAP), Lucknow. Histochemical work locates cannabinoids and terpenes in capitate stalked and capitate sessile glandular trichomes. Essential-oil analysis reports (E)-caryophyllene at 10.30%–36.80% as the major terpene, followed by α-humulene at 0.50%–15.29% and α-bisabolol at 0.00%–16.40%. Cannabinoid and terpene content shows significant diversity among and within the five populations. Correlation analysis finds α-pinene, β-pinene and limonene positively correlated with CBD content, while α- and β-selinene positively correlate with tetrahydrocannabinolic acid content. The authors frame the work as identifying candidate cultivars and chemotypes for Indian cannabis breeding programmes.

Population genomics of a natural Cannabis sativa L. collection from Iran identifies novel genetic loci for flowering time, morphology, sex and chemotyping

Mahboubeh Mostafaei Dehnavi, Annabelle Damerum, Sadegh Taheri, Ali Ebadi, Shadab Panahi, George Hodgin, Brian Brandley, Seyed Alireza Salami, Gail Taylor

BMC Plant Biology· 2025

Genotyping-by-Sequencing (GBS) study of 228 male and female individuals drawn from 35 natural Iranian Cannabis sativa populations spanning multiple climatic zones, grown in a randomised complete block design field experiment at the University of Tehran. Recovers approximately 23,266 high-quality SNPs and links them via association analysis to inflorescence characteristics, flowering time, plant morphology, THC and CBD content, and sex. Resolves the Iranian collection into five fineSTRUCTURE groups against a higher-level two-cluster PCA structure, and finds that Iranian populations are globally distinctive when compared to international reference data. Identifies novel candidate genetic loci for flowering time, morphology, sex determination and chemotype, providing markers that can underpin future cannabis breeding from Iranian germplasm. Frames Iranian natural populations as wild relatives and progenitor-adjacent material that retain genetic variation lost from highly bred modern cultivars. Critical regional reference for the Iran/Persian gene pool on the wiki, complementing the broader Balant et al. 2025 phylogeography (which places the Iranian Plateau as a distinct Paleotropis subgroup) with population-scale resolution and a marker-trait association layer.

South Africa, Zimbabwe

Comparative Analysis of Cannabis Legalization in South Africa and Zimbabwe: Trajectories, Commonalities, and Divergences

Simon Howell, Clemence Rusenga

Journal of Illicit Economies and Development· 2025

A comparative analysis of cannabis legalisation in South Africa and Zimbabwe, examining the trajectories, commonalities and divergences of reform. Documents how licensing costs and commercial frameworks have excluded smallholders in both countries.

South Africa

Finally Freed—Cannabis in South Africa: A Review Contextualised within Global History, Diversity, and Chemical Profiles

Valencia V. Ndlangamandla, Adeola Salawu-Rotimi, Vuyiswa S. Bushula-Njah, Nompilo L. Hlongwane, Gugu F. Sibandze, Fikisiwe C. Gebashe, Nokuthula P. Mchunu

Plants· 2024

A review contextualising cannabis in South Africa within global history, diversity and chemical profiles. States that no study has characterised southern-African cannabis landraces and that the NCBI genome assemblies include none of African origin, documenting the characterisation gap.

South Africa

Identification of Medicinal Cannabis landraces found in Lusikisiki, Eastern Cape Province, South Africa

Azile Dumani, Tembakazi Theodora Silwana, Babalwa Mpambani, Adebola Omowunmi Oyedeji, Ifeanyi Moses Egbichi, Hlabana Alfred Seepe

Journal of People, Plants, and Environment· 2024

Identifies eight vouchered Cannabis sativa specimens collected around Lusikisiki in the Eastern Cape by morphology only, with no chemotype or genetic work. The first vouchered landrace survey of the Mpondoland heartland.

Unveiling Colombia's medicinal Cannabis sativa treasure trove: Phenotypic and Chemotypic diversity in legal cultivation

Diego J. Enríquez, Julio C. Alonso, Lucas Hille, Stefan Brand, Ulrike Holzgrabe, Daniela Vergara, Guillermo Montoya, Yesid A. Ramírez

Phytochemical Analysis· 2024

Phytochemical and phenotypic characterisation of 156 legally cultivated Cannabis sativa plants grown across diverse ecological regions of Colombia. Ten cannabinoids and 23 terpenes were quantified by liquid and gas chromatography alongside other phenotypic traits. The data resolve four distinct chemotypes based on cannabinoid profile. Type I varieties show significantly higher terpene content (>0.03%) and Type IV varieties show lower content (<0.03%). β-myrcene emerges as the dominant terpene in balanced and CBD-dominant varieties. The authors report that several normally uncommon terpenes appear in significant quantities and suggest that Colombian environmental conditions may favour their expression.

Morocco· Rif Mountains· Rif Mountains

International property rights for Cannabis landraces and terroir products. The case of Moroccan Cannabis and hashish

Pierre-Arnaud Chouvy

International Journal of Drug Policy· 2024

Argues that appellations of origin offer the best available intellectual property protection for cannabis landraces and terroir products, on grounds that what needs protecting is tradition and collective ownership rather than innovation and individual ownership. Finds UPOV plant variety protection unsuited to landraces, which cannot meet the distinctness, uniformity and stability criteria required of cultivars. Develops the case through Moroccan hashish and the kif landrace of the Rif, whose typicity derives from the region's geography and the sociotechnical itinerary of cultivation. Concludes that only hashish made traditionally from kif could qualify for an appellation, since recent imported hybrids have produced a resin of Moroccan provenance but not Moroccan origin, and notes that an appellation cannot legally control third party use of a landrace but extends protection in practice to the agroecosystem on which the terroir depends.

South Africa

The use and potential abuse of psychoactive plants in southern Africa: an overview of evidence and future potential

Norman Zimunda Nyazema, Jonathan Tinotenda Chanyandura, Bronwyn Egan

Frontiers in Pharmacology· 2024

An overview of the use and potential abuse of psychoactive plants in southern Africa, situating cannabis among healer-used species and reinforcing that Leonotis is not true dagga.

Zimbabwe

Business as usual? Cannabis legalisation and agrarian change in Zimbabwe

Clemence Rusenga, Gernot Klantschnig, Neil Carrier, Simon Howell

The Journal of Peasant Studies· 2024

Examines Zimbabwe's legal cannabis sector since its 2018 opening to medicinal and industrial production, with unlicensed uses remaining criminalised, and the implications for agrarian change. The authors argue the formal sector is structured to favour those with substantial resources, marginalising small-scale farmers and illicit cultivators and raising the risk of corporate capture, though several factors undermine agribusiness production. They find that the prohibition of recreational cannabis and the sector's export focus together preserve illicit markets and sustain illicit livelihoods. An agrarian political-economy reference for cannabis commercialisation in Zimbabwe and the broader Global South pattern of export-oriented legal sectors marginalising traditional cultivators.

South Africa

A Focus on Cannabis as an Indigenous Entrepreneurial Activity in South Africa by the Indigenous Khoisan and Bantu

Pfano Mashau, Tshililo Farisani, Adenike Bello

Indigenous Entrepreneurship in Sub-Saharan Africa (Palgrave Macmillan)· 2024

Frames cannabis as an indigenous entrepreneurial activity of the Khoisan and Bantu, arguing for its place in South African indigenous enterprise. A book chapter; cite for the argument rather than for figures.

India

Cannabis in Traditional Indian Alchemy

Patricia Sauthoff

In Maas & Cerulli (eds.), Suhṛdayasaṃhitā: A Compendium of Studies on South Asian Culture, Philosophy and Religion. Dedicated to Dominik Wujastyk (Studia Indologica Universitatis Halensis 28). Heidelberg Asian Studies Publishing, pp. 165–179.· 2024

A philological and historical study of cannabis in the 12th–13th-century Sanskrit rasaśāstra text Ānandakanda, whose chapter 15 (186 verses) is the most extensive premodern Indian monograph on the plant. Updates Wujastyk (2002) with attention to the seven cannabis-mantras of the Ānandakanda, the fifteen Sanskrit synonyms (including the earliest secure attestation of gañjā), the male–female plant distinction, alchemical recipes, signs of intoxication, and the rarity of cannabis as an ingredient elsewhere in the rasaśāstra corpus. Closes with the etymological history of "marijuana" and other modern-language terms.

South Africa

Settling 'Dagga'? Shifting Frontiers of Cannabis Knowledge and Governance in South Africa

Thembisa Waetjen, Perside Ndandu

Journal of Southern African Studies· 2023

Examines the shifting frontiers of cannabis knowledge and governance in South Africa, extending Waetjen's account of how dagga was governed from the colonial period onward.

Morocco· Rif Mountains· Rif Mountains

Moroccan hashish as an example of a cannabis terroir product

Pierre-Arnaud Chouvy

GeoJournal· 2022

Develops operational definitions of terroir and landrace and applies them to cannabis cultivation in the Rif region of Morocco. Argues that hashish produced from the kif landrace meets the typicity, originality and reputation criteria of a terroir product, with the Rif as a delimited area where Berber communities have built up a collective production knowledge over a system of physical, biological and cultural interactions. Treats terroir as historically modern rather than fixist, and traditions as partly invented and reconstructed. Distinguishes site from situation as localisation factors: large-scale cannabis production requires both a biophysical site and a political-territorial situation marked by relative isolation from central authority, conditions the Rif has long met. Traces the etymology of balad, bled, beldi and beldiya to show that territory, terroir and landrace are expressed through variations of a single Arabic root in Moroccan Darija. Examines the contested historical-zone narrative and the five douars said to have been authorised under Moulay El Hassan I, drawing on Bellakhdar's 2021 proposal of Aït Aaksi, Griha, Ighmad, Azila and Talarouak. Concludes that the kif landrace cultivated before the 1960s no longer exists in unmodified form, that the introduction of hybrids in the 2000s and the 2021 therapeutic-cannabis legalisation are accelerating introgression, and that an appellation-style protection coupled with legalisation is the most plausible route to conserving the landrace, the agroecosystem and the regional economy.

South Africa

South Africa's Century of Cannabis Politics, 1922–2022

Thembisa Waetjen

South African Historical Journal· 2022

Surveys a century of South African cannabis politics from the 1922 criminalisation to 2022, framing the arc from prohibition to the 2018 Prince privacy ruling.

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