Morus nigra
Morus nigra

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Black mulberry (Morus nigra) is a long-lived deciduous fruit tree belonging to the Moraceae family. It is widely valued for its flavourful fruit and medicinal uses and has long been cultivated across western Asia (Kılınçer et al., 2024). The species typically grows to a height of 10–15 metres, with a crown diameter of up to 10 metres, forming a dense canopy that provides both shade and ornamental value in urban and rural landscapes (Švagr et al., 2023). Although it is not cultivated for timber, the heartwood is occasionally used, and the tree is often planted in gardens, parks, and green belts for aesthetic and ecological purposes (Švagr et al., 2023). 

Black mulberry thrives in warm temperate to subtropical regions and adapts well to a range of soil conditions, contributing to its successful spread and naturalization in many parts of southern and central Europe. The species’ centre of origin is believed to be in Türkiye (Anatolia, Asia Minor) and Iran, where numerous ancient, massive trees can still be found, indicating long-standing cultivation (Kılınçer et al., 2024). Black mulberry is naturalized and widely cultivated throughout Europe, North Africa, and the Americas, where it continues to be valued for its nutritious fruit, medicinal properties, and adaptability to different environments (Švagr et al., 2023). 

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Acknowledgements

This distribution map has been developed by the European Commission Joint Research Centre (partly based on the EUFORGEN map) and released under Creative Commons Attribution 4.0 International (CC-BY 4.0)


Caudullo, Giovanni; Welk, Erik; San-Miguel-Ayanz, Jesús (2017). Chorological maps and data for the main European woody species. figshare. Collection. https://doi.org/10.6084/m9.figshare.c.2918528

 

The following experts have contributed to the development of the EUFORGEN distribution maps:

Fazia Krouchi (Algeria), Hasmik Ghalachyan (Armenia), Thomas Geburek (Austria), Berthold Heinze (Austria), Rudi Litschauer (Austria), Rudolf Litschauer (Austria), Michael Mengl (Austria), Ferdinand Müller (Austria), Franz Starlinger (Austria), Valida Ali-zade (Azerbaijan), Vahid Djalal Hajiyev (Azerbaijan), Karen Cox (Belgium), Bart De Cuyper (Belgium), Olivier Desteucq (Belgium), Patrick Mertens (Belgium), Jos Van Slycken (Belgium), An Vanden Broeck (Belgium), Kristine Vander Mijnsbrugge (Belgium), Dalibor Ballian (Bosnia and Herzegovina), Alexander H. Alexandrov (Bulgaria), Alexander Delkov (Bulgaria), Ivanova Denitsa Pandeva (Bulgaria), Peter Zhelev Stoyanov (Bulgaria), Joso Gracan (Croatia), Marilena Idzojtic (Croatia), Mladen Ivankovic (Croatia), Željka Ivanović (Croatia), Davorin Kajba (Croatia), Hrvoje Marjanovic (Croatia), Sanja Peric (Croatia), Andreas Christou (Cyprus), Xenophon Hadjikyriacou (Cyprus), Václav Buriánek (Czech Republic), Jan Chládek (Czech Republic), Josef Frýdl (Czech Republic), Petr Novotný (Czech Republic), Martin Slovacek (Czech Republic), Zdenek Špišek (Czech Republic), Karel Vancura (Czech Republic), Ulrik Bräuner (Denmark), Bjerne Ditlevsen (Denmark), Jon Kehlet Hansen (Denmark), Jan Svejgaard Jensen (Denmark), Kalev Jðgiste (Estonia), Tiit Maaten (Estonia), Raul Pihu (Estonia), Ülo Tamm (Estonia), Arvo Tullus (Estonia), Aivo Vares (Estonia), Teijo Nikkanen (Finland), Sanna Paanukoski (Finland), Mari Rusanen (Finland), Pekka Vakkari (Finland), Leena Yrjänä (Finland), Daniel Cambon (France), Eric Collin (France), Alexis Ducousso (France), Bruno Fady (France), François Lefèvre (France), Brigitte Musch (France), Sylvie Oddou-Muratorio (France), Luc E. Pâques (France), Julien Saudubray (France), Marc Villar (France), Vlatko Andonovski (FYR Macedonia), Dragi Pop-Stojanov (FYR Macedonia), Merab Machavariani (Georgia), Irina Tvauri (Georgia), Alexander Urushadze (Georgia), Bernd Degen (Germany), Jochen Kleinschmit (Germany), Armin König (Germany), Armin König (Germany), Volker Schneck (Germany), Richard Stephan (Germany), H. H. Kausch-Blecken Von Schmeling (Germany), Georg von Wühlisch (Germany), Iris Wagner (Germany), Heino Wolf (Germany), Paraskevi Alizoti (Greece), Filippos Aravanopoulos (Greece), Andreas Drouzas (Greece), Despina Paitaridou (Greece), Aristotelis C. Papageorgiou (Greece), Kostas Thanos (Greece), Sándor Bordács (Hungary), Csaba Mátyás (Hungary), László Nagy (Hungary), Thröstur Eysteinsson (Iceland), Adalsteinn Sigurgeirsson (Iceland), Halldór Sverrisson (Iceland), John Fennessy (Ireland), Ellen O'Connor (Ireland), Fulvio Ducci (Italy), Silvia Fineschi (Italy), Bartolomeo Schirone (Italy), Marco Cosimo Simeone (Italy), Giovanni Giuseppe Vendramin (Italy), Lorenzo Vietto (Italy), Janis Birgelis (Latvia), Virgilijus Baliuckas (Lithuania), Kestutis Cesnavicius (Lithuania), Darius Danusevicius (Lithuania), Valmantas Kundrotas (Lithuania), Alfas Pliûra (Lithuania), Darius Raudonius (Lithuania), Robert du Fays (Luxembourg), Myriam Heuertz (Luxembourg), Claude Parini (Luxembourg), Fred Trossen (Luxembourg), Frank Wolter (Luxembourg), Joseph Buhagiar (Malta), Eman Calleja (Malta), Ion Palancean (Moldova), Dragos Postolache (Moldova), Gheorghe Postolache (Moldova), Hassan Sbay (Morocco), Tor Myking (Norway), Tore Skrøppa (Norway), Anna Gugala (Poland), Jan Kowalczyk (Poland), Czeslaw Koziol (Poland), Jan Matras (Poland), Zbigniew Sobierajski (Poland), Maria Helena Almeida (Portugal), Filipe Costa e Silva (Portugal), Luís Reis (Portugal), Maria Carolina Varela (Portugal), Ioan Blada (Romania), Alexandru-Lucian Curtu (Romania), Lucian Dinca (Romania), Georgeta Mihai (Romania), Mihai Olaru (Romania), Gheorghe Parnuta (Romania), Natalia Demidova (Russian Federation), Mikhail V. Pridnya (Russian Federation), Andrey Prokazin (Russian Federation), Srdjan Bojovic (Serbia) , Vasilije Isajev (Serbia), Saša Orlovic (Serbia), Rudolf Bruchánik (Slovakia), Roman Longauer (Slovakia), Ladislav Paule (Slovakia), Gregor Bozič (Slovenia), Robert Brus (Slovenia), Katarina Celič (Slovenia), Hojka Kraigher (Slovenia), Andrej Verlič (Slovenia), Marjana Westergren (Slovenia), Ricardo Alía (Spain), Josefa Fernández-López (Spain), Luis Gil Sanchez (Spain), Pablo Gonzalez Goicoechea (Spain), Santiago C. González-Martínez (Spain), Sonia Martin Albertos (Spain), Eduardo Notivol Paino (Spain), María Arantxa Prada (Spain), Alvaro Soto de Viana (Spain), Lennart Ackzell (Sweden), Jonas Bergquist (Sweden), Sanna Black-Samuelsson (Sweden), Jonas Cedergren (Sweden), Gösta Eriksson (Sweden), Markus Bolliger (Switzerland), Felix Gugerli (Switzerland), Rolf Holderegger (Switzerland), Peter Rotach (Switzerland), Marcus Ulber (Switzerland), Sven M.G. de Vries (The Netherlands), Khouja Mohamed Larbi (Tunisia), Murat Alan (Turkey), Gaye Kandemir (Turkey), Gursel Karagöz (Turkey), Zeki Kaya (Turkey), Hasan Özer (Turkey), Hacer Semerci (Turkey), Ferit Toplu (Turkey), Mykola M. Vedmid (Ukraine), Roman T. Volosyanchuk (Ukraine), Stuart A'Hara (United Kingdom), Joan Cottrell (United Kingdom), Colin Edwards (United Kingdom), Michael Frankis (United Kingdom), Jason Hubert (United Kingdom), Karen Russell (United Kingdom), C.J.A. Samuel (United Kingdom).
 

Genetic diversity and variation 

Despite its cultural and economic importance, research on the genetics and conservation of black mulberry remains limited (Kılınçer et al., 2024). Across its range, black mulberry has remarkably low genetic diversity and minimal population structure (Kılınçer et al., 2024; Ismail et al., 2022). Studies consistently report that both wild and cultivated accessions, including those from Europe and surrounding regions, show little genetic variation within the species, although wild black mulberry is genetically distinct from other mulberry species (Ismail et al., 2022). 

This low genetic diversity is thought to result from historical genetic bottlenecks, the species’ predominant vegetative propagation, and widespread clonal propagation in cultivation, leading to the formation of clonal populations (Kılınçer et al., 2024). The nearly identical genetic structure observed is highly unusual among domesticated perennial fruit crops and suggests that black mulberry globally lacks genetic variability (Kılınçer et al., 2024). 

In Syria, black mulberry cultivars also displayed low overall genetic diversity, reinforcing this pattern of uniformity across its distribution (Baroudi, Choumane, and Makhoul, 2023). The species’ very high ploidy level, specifically decosaploidy (22x), is a unique feature within the genus Morus and among flowering plants, contributing to its distinctive genetic stability and evolutionary isolation (Baroudi, Choumane, and Makhoul, 2023; Kılınçer et al., 2024). 

Genetic distribution and clustering 

Studies on the genetic structure of black mulberry indicate limited but distinct genetic clustering within certain regions. In Syria, populations have been shown to group into two main genetic clusters (Baroudi, Choumane, and Makhoul, 2023). However, across its broader range, there is a lack of clear correlation between geographical origin and genetic similarity. This pattern results from centuries of human-mediated introduction, cultivation, and later naturalization of mulberry genotypes outside their original centres of diversity (Baroudi, Choumane, and Makhoul, 2023). 

Gene flow 

Black mulberry is wind pollinated, with seeds being dispersed by birds and other animals, facilitating long-distance gene flow. However, in cultivation the species is often clonally propagated. 


The bibliographic review was conducted by James Chaplin of the EUFORGEN Secretariat in August 2025.

Interspecific taxa dynamics 

Black-fruited cultivars of white mulberry (Morus alba) are often mistaken for black mulberry, as both can produce dark-coloured fruits and share similar leaf morphologies (Kılınçer et al., 2024). This misidentification complicates both conservation and breeding efforts, as it obscures true patterns of genetic diversity and species distribution. 

Molecular and cluster analyses have confirmed that black mulberry forms a distinct genetic group separate from other mulberry species, although it has very high genetic similarity between its own accessions, indicating low intraspecific variation (Ismail et al., 2022). Comparative studies show that white mulberry possesses greater genetic diversity than black mulberry, reflecting its broader distribution, extensive cultivation, and hybridization potential (Baroudi, Choumane, and Makhoul, 2023). Despite their genetic similarity, the two species can be clearly distinguished at the molecular level, supporting their classification as separate taxa (Baroudi, Choumane, and Makhoul, 2023). 

 

The bibliographic review was conducted by James Chaplin of the EUFORGEN Secretariat in August 2025.

Threats 

Black mulberry has extremely low genetic diversity due to its clonal propagation, making it highly vulnerable to pests, diseases, and environmental change (Kılınçer et al., 2024). This lack of variability limits its adaptability and long-term survival potential. In Europe, where black mulberry is mainly cultivated rather than naturally regenerating, opportunities for gene flow and recombination are minimal. Climate change creates risks and opportunities; warmer and drier conditions may stress existing populations but could also expand suitable habitats northward into Central Europe (Švagr et al., 2023). Additional threats include fungal pathogens and insect pests, although their current impact in Europe remains limited (Švagr et al., 2023). The species’ restricted genetic base and dependence on vegetative propagation represent major concerns for its conservation and resilience. 

Management 

Conservation of black mulberry should prioritize the preservation of ancient and heritage trees, which may contain rare genetic variants (Kılınçer et al., 2024). Ex situ germplasm conservation and molecular characterization are essential to safeguard remaining diversity and clarify species identification, especially where confusion with white mulberry exists (Baroudi, Choumane, and Makhoul, 2023). Although genetic improvement through cross-breeding or new rootstocks could enhance commercial use, such interventions risk compromising genetic integrity (Kılınçer et al., 2024). In situ protection, propagation from diverse source trees, and genetic monitoring are key to maintaining the species’ limited diversity while supporting its continued cultural and ecological value. 
 

The bibliographic review was conducted by James Chaplin of the EUFORGEN Secretariat in August 2025.

Further reading

Jhansdakshmi, K. and Rao, A.A. 2021. Conservation of mulberry genetic resources to sustain sericulture under climate change. In: M.K. Razdan and D. Thomas, eds. Mulberry, pp. 225–245. Boca Raton, FL, USA, CRC Press. 

References

Baroudi, H.M., Choumane, W.M., and Makhoul, G.M. 2023. Evaluation of genetic diversity in two Morus species in Syria, using molecular markers ISSR and SSR. Journal of Central European Agriculture, 24(3): 736–748. https://doi.org/10.5513/JCEA01/24.3.3933 

Ismail, I.A., El Dessoky, S.D., Attia, A.O., and Saleh, O.M. 2022. Molecular assessment of established clonal propagated mulberry (Morus nigra L.). Academic Journal of Chemistry, 8(3): 190–196. 

Kılınçer, İ., Khanyile, L., Gürcan, K., Şimşek, Ö., Uzun, A., and Nikbakht-Dehkordi, A. 2024. Decosaploid sour black mulberry (Morus nigra L.) in western Asia: Features, domestication history, and unique population genetics. Genetic Resources and Crop Evolution, 71: 2229–2246. https://doi.org/10.1007/s10722-023-01771-w 

Švagr, P., Gallo, J., Vítámvás, J., Podrázský, V., and Baláš, M. 2023. Potential of Morus nigra in Central Europe focused on micropropagation: A short review. Journal of Forest Science, 69(11): 463–469. https://doi.org/10.17221/73/2023-JFS 

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