Domestication of date palms in Siwa oasis and across the Middle East and North Africa : articulate the scales of ethnography and domestication over the long term Vincent Battesti, Muriel Gros-Balthazard

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Vincent Battesti, Muriel Gros-Balthazard. Domestication of date palms in Siwa oasis and across the Middle East and North Africa : articulate the scales of ethnography and domestication over the long term. Harlan III, : the Third Jack R. Harlan International Symposium Dedicated to the Origins of Agriculture and the Domestication, Evolution, and Utilization of Genetic Resources, Jun 2019, Montpellier, France. ￿hal-02270215￿

HAL Id: hal-02270215 https://hal.archives-ouvertes.fr/hal-02270215 Submitted on 27 Aug 2019

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Distributed under a Creative Commons Attribution - NonCommercial - NoDerivatives| 4.0 International License Domestication of date palms in Siwa Oasis and across the Middle East and North Africa: Articulating the scales of ethnography and of domestication over the long term Data : under publication (Gros-Balthazard & Battesti et al.) Vincent Battesti & Muriel Gros-Balthazard (CNRS/Musée de l’Homme, MNHN & Center for Genomics and Systems Biology, NYU Abu Dhabi)

Studying agrobiodiversity in the oasis of Siwa (Egypt), by combining ethnobotany and genetics, we demonstrated: • The existence of “ethnovarieties,” i.e., voluntary collections of multiple clones sharing mostly phenotypic characteristics under the same local name, as a way to manage agrobiodiversity. • Siwa’s cultivated date palms form a third genetic cluster, with a large and unique diversity, compared to the previously described Middle Eastern and North African date palms. • Siwa region's feral date palms are also distinct, sharing alleles with another species (P. theophrasti), a gene pool for local farmers.

INTRODUCTION Possible main location for Phoenix dactylifera domestication The history of the date palm (Phoenix dactylifera L.) as a cultivated is still poorly known, despite some major advances made using genetic tools1. An unexpectedly high differentiation between African and Middle Eastern populations was discovered and its origins was only recently explained by the North African date palms being introgressed by a wild relative: Phoenix theophrasti Greuter2. Moreover, ways of assessing the current agrobiodiversity of the cultivated date palm do not yet seem well established4.

Siwa oasis (Egypt), at the crossroads of ancient OBJECTIVES: To better seize the long- and short-term domestication process of this cultivated and Trans-Saharan routes and of the oriental and emblematic crop, the date palm, our approach combines anthropology (ethnoecology) with population western genetic cluster genetics (using 17 microsatellite markers).

An original folk An original, large and unique categorization system by biodiversity in Siwa Some named types are true Siwa inhabitants Using an existing SSR dataset from 5, 6, 7.

cultivars as supposed to be ● West ● ● East ● ●●● ● Nilotic Egypt ● Siwa cult ● ● Siwa feral desert West 20 ● 20 in oasis phoeniculture: they ● ● ● ● East ● ●● 15 ● ● ● ● ● ● Nilotic Egypt ● 4 ● ● ●●●● ● ●● ● ● ● ●● ● ● Siwa cult share not only a formal ● ● ●● ●

15 ● úšik #1 ● ● ● ● ● Siwa feral desert ● ● ● 10 úšik #2 ● ●● ● ṣaɛidi [tasutet] (siwi) ● ● ● ● ● P. theophrasti Uncultivated date palm identity, important for ● ● ● from the desert 2 ● ● ● ● ● ● ● ● ● ● ● 10 ● ●●● ●●●●●● ● ●● 5 3361_ushik_ezzuwagh ● ● ●● ●● ● ● ● ● ● ● 10 ● ●● ● ●● ●●●● ● ● 3408_alkak_wen_djemb Isiwan people, but also a ● ●●● ● 3352_ushik_azzugagh ● ●● ●● ● ● ● ● 3365_ushik_ezzuwagh ● ●● ● ● ● ●● ● ● ● ● 3364_ushik_ezzuwagh ● ● ● ●●●● ● ●● ●●● ● ●●● ●●●● ● ●● ● 5 ● ●●●●●●● ● ● ● 3373_tadjubart ●● ● ● ●● ●●●● ●●●●●●●● ●●●●●● ●●●●● ● ● ● ●● ● ●●● ● ● ●●●●●●●●●● ●●●●●●●●●●●●●●●●●●●●●●●●●● ●●● ● ● 0 ● ● ● ●●●● ● 0 ● ● ● ● ●●●●●●●●●●●●●●● ●●●●●●●●●●● ● ●● ●●● ●● ● ● ●● ● ● ● ●●●●●●●●●●●●●●●●●● ●●●●●●●●●●●●●●●●●●●●●●●●● ● 3445_ushik_nekwayes genetic identity. ● ● ● ●● ● ●●●●● ●●●● ●●● ●●●●●● ●●●●●● ● ● 5 ● ● ● ● ● ● 3448_ushik_nekwayes ● ● ● ● ● ● ● ●● ●● ● ●● ● ● ●● ● ● ● ● ●●●●●●●●● ● ● 2406_ushik_nekwayes ● ● ● ● ● ●●●●●●●●●●● ● 3359_kaaibi ● ● ● ● ●●●●●●●●●●●●●●●●●●●● ●

PC2 (2.93%) ● ● PC4 (2.11%) 3442_ushik_nekwayes ● ● ● ●●●●●●●●●●●●●●●●●●●●●● ●● 0 ● ● ●●●●●●●●●●●●●● ● 3367_ghrom_saaid ● ● ● ● ● ●●●●●●●●●●●●●●●●●●● ● 2423_ushik_niqbel ● ● ● ●●● ● ● ●●●●●●●●●●●●● 3411_alkak_wen_djemb Some are (what we coined) ● ● ● ● ● ●●●●●●●● ● ●●●●●●●●●●●●●● ● 3412_alkak_wen_djemb ● ● ●● ●●●● ●●●●●●●●●● ● ● ● ● ● ● ● ●●●●●●●●●●●●●●● ●●●●●●●●●●● ● ●●●●● ● ● ●●●●●●●●●●●● ●● ●●●●●●●●●●●●●●● 3362_ushik_ezzuwagh ●● ● ● ● ●●●●●●●●●●● ●●●●●●●●●●●● ●● − 2 ● PC2 (2.68%) PC4 (1.91%) ● ● ●● ●●●● ●●●●● ● ● 0 ● ●●●● ●●●●● ●●●● ●●● ● ●●● ● ●●● ●●● ●●●●●●●●●●●●●●●●●●●● ●● ● ●●●●●●●●●●●●●●●●●●●●● “ethnovarieties,” i.e., ● ● ● ● ●●●●●●●●●●●●●●●● 3417_ghrom_aghzal ● ● ●● ● ●●●●●●●●●●● ●● ● ● ●● ● ●● ●●●●●●●●●● 3351_ushik_azzugagh ● ●● ● ● ●●●●●●●● ● ●●● ● − 10 3354_kaaibi ● ● ●● ● 3409_alkak_wen_djemb ●●● ●● 3407_lekrawmet ● ●● ● 3406_lekrawmet voluntary collections of ● ●●●● ● ●● ● 2412_ushik_ezzuwagh − 4 ● 3360_ushik_ezzuwagh ● 3413_ghrom_aghzal ● ●● ● ● − 10 multiple clones sharing ● ● 3457_tazuwaght 3450_ushik_nekwayes ● −5 0 5 ● 10 −6 −4 −2 0 2 4 3404_lekrawmet 3443_ushik_nekwayes

3444_ushik_nekwayes 3410_alkak_wen_djemb mostly phenotypic 3405_lekrawmet 3389_ushik_niqbel 2422_ushik_niqbel 3392_ushik_niqbel 3458_ushik_niqbel PC1 (4.44%) PC3 (2.48%) 3459_ushik_niqbel 3390_ushik_niqbel 3393_ushik_niqbel characteristics under the 3394_ushik_niqbel −4 0 2 4 6 −10 0 5 10 15 3391_ushik_niqbel 3451_ushik_nekwayes 3358_kaaibi 3296_ushik_niqbel 3449_ushik_nekwayes 3398_amenzu same local name. PCA 3356_kaaibi 2419_ushik_ezzuwagh 3416_ghrom_aghzal 3294_ghrom_aghzal PC1 (4.36%) PC3 (2.16%) 3357_kaaibi 3414_ghrom_aghzal 17 SSRs Feral date palms, collected in 3402_amenzu 3353_ushik_am_aghzuz 3415_ghrom_aghzal abandoned oasis around Siwa share alkak (freḥi) While we expected Siwa diversity 2417_alkak_wen_djemb The existence of 3355_kaaibi alleles with Phoenix theophrasti, the aghzāl to be found within the known 2410_ushik_am_aghzuz 3350_ushik_azzugagh "ethnovarieties" results from 3349_ushik_azzugagh 3401_amenzu species that is known to have 3400_amenzu 3297_amenzu African cluster, we found that it 3366_ghrom_saaid a cultural practice of oasis introgressed African date palms2.

3447_ushik_nekwayes constitutes a separate population. 3446_ushik_nekwayes farmers towards date palm ḥalu en ghanem

3377_tattagt 3403_amenzu 3378_tattagt 2405_tattagt trees and from their way to 3379_tattagt 3456_tattagt 3374_tattagt 3375_tattagt 3376_tattagt think of them3, 4. Allelic richness Private allelic richness Siwa date palms show a very 3388_ushik_niqbel c c a b d e d c a b large diversity and a diversity 6 ● ● ● 3363_ushik_ezzuwagh ● ● 2.0 ● 3399_amenzu The same idea applies to the 5 ● ● that is unique to this oasis. ● 4 ● 1.5 notion we call "categories", a ● ● ● 3 way to name other date 1.0 ● The origins of Siwa date palms is ● ● ● ● 2 ● ● ● palms that are not or poorly ● ● a mystery. Genomic data and The most direct consequence is the 0.5 reproduced by off-shoot by 1 our island theory of oases will demonstration that we had until then a clear ● 0 0.0 further help us understanding underestimation of the richness in genetic the local community. this unique diversity. resources of date palm: about fifteen named Siwa feral Siwa feral Siwa Siwa oasis Siwa oasis Siwa Middle East Middle East North AFrica North AFrica

types identified are not about fifteen genetic P. theophrasti P. theophrasti

combinations of alleles. Rarefaction method, haploid sample size = 18 Letters indicate Tukey’s groups derived from Tukey’s test Why such a complex local system in Siwa for categorizing living organisms and in particular date palms? 1 - Gros-Balthazard, Muriel, Khaled M. Hazzouri & Jonathan M. Flowers (2018), Genomic Insights into Date Palm Origins. Genes, 9 (10), 502. https://doi.org/10.3390/genes9100502 2 - Flowers, Jonathan M., et al. (2019), Cross-species hybridization and the origin of North African date palms. PNAS, 116 (5), p. 1651-1658. https://www.pnas.org/content/116/5/1651 - To stock a huge amount of information and to be able to mobilize it easily. 3 - Battesti, Vincent & Gros-Balthazard, Muriel et al. (2018), Date palm agrobiodiversity (Phoenix dactylifera L.) in Siwa oasis, Egypt : combining ethnography, morphometry, and genetics. Human Ecology, 46 (4), p. 529–546. https://hal.archives-ouvertes.fr/hal-01716788 - To allow a flexible management of agrobiodiversity in this local context. 4 - Battesti, Vincent (2013), L’agrobiodiversité du dattier (Phoenix dactylifera L.) dans l’oasis de Siwa (Égypte) : entre ce qui se dit, s’écrit et s’oublie ». Revue d'ethnoécologie, 4. http://dx.doi.org/10.4000/ethnoecologie.1538 5 - Zango, O. et al. (2017), Genetic diversity of Southeastern Nigerien date palms reveals a secondary structure within Western populations. Tree Genet. Genomes 13, 75. https://doi.org/10.1007/s11295-017-1150-z With the 6 - Zehdi-Azouzi, S. et al. (2015), Genetic structure of the date palm (Phoenix dactylifera) in the Old World reveals a strong differentiation between eastern and western decisive populations. Ann. Bot. 116, 101–112. https://doi.org/10.1093/aob/mcv068 collaboration 7 - Moussouni, Souhila et al. (2017), Diversity of Algerian oases date palm (Phoenix dactylifera L., ): Heterozygote excess and cryptic structure suggest farmer 9 of the farmers of the Siwa management had a major impact on diversity. PLOS ONE, 12 (4), e0175232. https://doi.org/10.1371/journal.pone.0175232 201 Oasis (Egypt) who shared

June their practices and Vincent Battesti knowledge. Musée de l’Homme — UMR 7206 Éco-anthropologie (CNRS MNHN P7) 17, place du Trocadéro - 75016 Paris — France http://vbat.org — [email protected] Montpellier, –

III Muriel Gros-Balthazard NYU Abu Dhabi, Saadiyat Campus — Experimental Research (C1), RL1-D

Harlan Contacts P.O. Box 129188 Abu Dhabi — United Arab Emirates [email protected]