Clonal Diversity, Cultivar Traits, Geographic Dispersal, and the Ethnotaxonomy of Cultivated Qat ( Catha edulis , Celast

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Department of Biology, Colorado State University, Fort Collins, CO, USA United States Department of Agriculture, Agricultural Research Service, National Laboratory for Genetic Resources Preservation, Fort Collins, CO, USA 3 Institute for Integrative Nature Conservation Research, University of Natural Resources and Life Sciences, Vienna, Austria 4 East African Herbarium, National Museums of Kenya, Nairobi, Kenya 5 National Herbarium, Department of Plant Biology and Biodiversity Management, College of Natural Sciences, Addis Ababa University, Addis Ababa, Ethiopia 6 Department of Biology, College of Science and Art, Al Baha University, Baljurashi, Saudi Arabia 7 Agricultural Research and Extension Authority, Dhamar, Yemen 8 Department of Horticulture and Forestry, College of Agriculture, Sana’a University, Sana’a, Yemen 9 United States Department of Agriculture, Agricultural Research Service, Genomics and Bioinformatics Research Unit, Stoneville, MS, USA 10 American Institute for Yemeni Studies, Sana’a, Yemen *Corresponding author; e-mail: [email protected] 2

Farmers and traders have developed a system of names to refer to different qat (Catha edulis) cultivars, using stem color as the primary trait to differentiate them. In this study, we tested if the named cultivars from Ethiopia, Kenya, and Yemen represent genetic clusters. We also quantified clonal reproduction and tracked the geographic dispersal of cultivated–qat genotypes using microsatellite genotypes of specimens collected from across the major qat growing regions (Ethiopia, Kenya, and Yemen). Specimens were mapped to determine whether names, morphology, genetic clusters, or genotypes (in particular clones) were geographically restricted. Stem color was found to be a plastic trait because specimens of the same genotype have different colored stems. A single cultivar name was often applied to numerous clonal groups and genetic clusters, creating widespread homonymy in the ethnotaxonomy of qat cultivars. The East African Rift (EAR) in central Ethiopia was found to be a barrier to the exchange of both clonal groups and certain cultivar names. In Kenya, both cultivar names and clonal groups were broadly dispersed geographically. Nearly all of these clonal groups originated from the Mt. Kenya/Meru area in central Kenya. Key Words:

Catha edulis, Qat, Khat, Clonal reproduction, Ethnotaxonomy, Genetic diversity.

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Received 9 December 2019; accepted 3 August 2020; published online ___________ Electronic supplementary material The online version of this article (https://doi.org/10.1007/s12231-02009501-4) contains supplementary material, which is available to authorized users.

Introduction Qat [Celastraceae: Catha edulis (Vahl) Forssk. ex Endl.], a species of evergreen shrubs and trees, is of major cultural and economic importance in southwest Arabia and East Africa. An estimated 20

Economic Botany, XX(X), 2020, pp. 1–19 © 2020, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

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million people consume qat daily in ea