On the origin and dispersal of cultivated spinach ( Spinacia oleracea L.)

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RESEARCH ARTICLE

On the origin and dispersal of cultivated spinach (Spinacia oleracea L.) Arnau Ribera . Rob van Treuren

. Chris Kik . Yuling Bai . Anne-Marie A. Wolters

Received: 21 April 2020 / Accepted: 8 October 2020 Ó The Author(s) 2020

Abstract Spinach (Spinacia oleracea L.) is an economically important crop that is cultivated and consumed worldwide. Spinach is interfertile with the wild species S. tetrandra Steven ex M. Bieb. and S. turkestanica Iljin that therefore are presumed to include the most likely crop ancestor. Here we studied variation in 60 Single Nucleotide Polymorphisms (SNP) previously identified in S. oleracea to address the issue of crop ancestry and domestication region. For this purpose we investigated 95 accessions, including 54 spinach landraces from a wide geographic area in Europe and Asia and 16 S. tetrandra and 25 S. turkestanica populations of which the majority had only recently become available. Compared to S. tetrandra substantially higher levels of amplification success and higher levels of variation were detected for S. turkestanica, indicating that S. oleracea is genetically closer to S. turkestanica than to

Electronic supplementary material The online version of this article (https://doi.org/10.1007/s10722-020-01042-y) contains supplementary material, which is available to authorized users. A. Ribera  Y. Bai  A.-M. A. Wolters Plant Breeding, Wageningen University & Research, P.O Box 386, 6700 AJ Wageningen, The Netherlands A. Ribera  R. van Treuren (&)  C. Kik Centre for Genetic Resources, the Netherlands, Wageningen Plant Research, P.O. Box 16, 6700 AA Wageningen, The Netherlands e-mail: [email protected]

S. tetrandra. Our phylogenetic and population structure analysis supported the conclusion that S. turkestanica is the most likely ancestor of cultivated spinach. In addition, these analyses revealed a group of S. oleracea landraces from Eastern and Southern Asia with a strong genetic resemblance to S. turkestanica. This group includes landraces from Afghanistan and Pakistan, which are part of the native distribution range of S. turkestanica. The domestication of spinach may therefore have occurred more eastwards than generally assumed. Furthermore, our study provides support for the hypothesis that after domestication, spinach was introduced into China via Nepal. Additional collecting of spinach landraces is recommended in order to allow the more precise reconstruction of the crop migration routes. Keywords Ancestry  Crop wild relatives  Dispersal  Domestication  Phylogeny  Spinach

Introduction Spinach (Spinacia oleracea L., 2n = 2x = 12) is an economically important leafy vegetable that is consumed worldwide (Morelock and Correll 2008). Although the origin of cultivated spinach is uncertain, it is believed to have been domesticated in the area of present Iran, former Persia, around 2,000 years ago

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Genet Resour Crop Evol

(Rubatzky and Yamaguchi 1997; Morelock and Correll 2008). It is presumed that