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Classification and Systematics of the Fagopyrum Species

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Classification and Systematics of the Fagopyrum Species Breeding Science 70: 93–100 (2020) doi: 10.1270/jsbbs.19028 Invited Review Classification and systematics of the Fagopyrum species Takanori Ohsako*1) and Chengyun Li2) 1) Laboratory of Plant Resource Science, Graduate School of Life and Environmental Sciences, Kyoto Prefectural University, 74 Kitainayazuma, Seika, Kyoto 619-0244, Japan 2) State Key Laboratory for Conservation and Utilization of Bio-Resources in Yunnan, Yunnan Agricultural University, Heilongtan, Kunming, Yunnan 650201, China Fagopyrum (Polygonaceae) is a small genus including less than 30 species, mostly endemic to southern China. The genus includes two cultivated species, common buckwheat F. esculentum and Tartary buckwheat F. tataricum. Fagopyrum is clearly distinguished from other genus in Polygonaceae by the central position of embryo in achene. The genus is divided into two major groups, namely the cymosum group and the urophyllum group, based on morphological characters and molecular systematics. In the last three decades the number of species in the genus has doubled by the discovery of new species by Japanese and Chinese groups. Most of them are revealed to be included in the urophyllum group based on morphological and molecular genetic analyses. Molecular systematic surveys have also detected inappropriate treatment of some non-Fagopyrum species as new species or combination in the genus. Key Words: classification, delineation of Fagopyrum, intraspecific variation, molecular systematics, new species. common buckwheat populations distributed in eastern Tibet Overview of the genus Fagopyrum, a group of cul‐ is the most closely related to cultivated forms (Konishi tivated and wild buckwheat species et al. 2005), indicating that cultivated common buckwheat was domesticated in the eastern Tibet to western Sichuan. Fagopyrum (Polygonaceae) is a small genus comprised of Tsuji and Ohnishi (2000, 2001) performed phylogenetic less than 30 species (Jin et al. 2018). Most of the wild spe‐ analyses of cultivated (F. tataricum ssp. tataricum) and cies show narrow distribution areas in Southern Asia, wild (F. tataricum ssp. Potanini) forms of Tartary buck‐ mainly on the southeastern edge of Qinghai-Tibetan Plateau wheat based on RAPD and AFLP data and showed that cul‐ (Ohnishi 1998a). Two species, common buckwheat tivated Tartary buckwheat was domesticated in eastern F. esculentum and Tartary buckwheat F. tataricum, are cul‐ Tibet to northwestern Yunnan, the same area as the birth‐ tivated, and a wild species F. cymosum is utilized as forage place of common buckwheat. and a source of pharmaceutical drugs (Wang et al. 2005). The delineation of Fagopyrum differs among classifica‐ For common buckwheat and Tartary buckwheat, cultivated tion systems of Polygonaceae. Species recognized as the and wild forms are distinguished and classified as sub‐ member of the genus Fagopyrum in the present day are species (Ohnishi 1998b). Wild common buckwheat listed in Table 1. Gross (1913) proposed many new combi‐ F. esculentum ssp. ancestrale was discovered at the first nations for the species in the section Polygoneae. In the time in 1990 (Ohnishi 1990, 1998a) at Yongsheng, Yunnan, study, two genera, Tiniaria and Fagopyrum, were reduced China. Now wild common buckwheat is found in north‐ into a single genus Fagopyrum with respective sections western Yunnan, western Sichuan, and eastern Tibet around Tiniaria and Eufagopyrum in the genus. Samuelsson (1929) the Three Parallel Rivers region where the Jinsha or upper listed 10 species in the section Fagopyrum of the genus Yangtze, Mekong, and Salween rivers flowing down north Polygonum. Steward (1930) mostly followed the system of to south in parallel (Ohnishi and Konishi 2001). Samuelsson (1929) and included 10 species in the section F. esculentum ssp. ancestrale is thought of as wild ancestor Fagopyrum of the genus Polygonum. In the present day, of cultivated buckwheat F. esculentum ssp. esculentum most studies seem to follow the system of Hedberg (1946) (Ohnishi 1998a, 1998b). AFLP analysis revealed that wild that treats Fagopyrum as an independent genus. Fagopyrum has been discriminated from other genera or Communicated by Yasuo Yasui sections by embryo morphology. Dammer (1894) stated a Received March 1, 2019. Accepted July 16, 2019. key to Fagopyrum as large folded cotyledons in achene. First Published Online in J-STAGE on December 17, 2019. *Corresponding author (e-mail: [email protected]) Gross (1913) characterized the section Eufagopyrum by 93 Breeding Science BS Vol. 70 No. 1 Ohsako and Li Table 1. Fagopyrum species Chromosome Mating Group Species Distribution of natural populations Reference number system cymosum F. esculentum Sichuan, Yunnan, Tibet 16 SI Moench (1794), Ohnishi (1998a) F. homotropicum Sichuan, Yunnan, Tibet 16, 32 SC Ohnishi (1998a) F. tataricum Sichuan, Yunnan, Tibet, Nepal, Pakistan 16 SC Gaertner (1791) F. cymosum Southern China, Nepal, India, Bhutan, 16, 32 SI Meisner (1857) Myanmar, Vietnam, Thailand urophyllum F. urophyllum Sichuan, Yunnan 16 SI Gross (1913) F. lineare Yunnan 16 SI Haraldson (1978) F. leptopodum Sichuan, Yunnan 16 SI Hedberg (1946) F. statice Yunnan 16 SI Gross (1913) F. gracilipes Sichuan, Yunnan, Bhutan 32 SC Diels (1901) F. capillatum Yunnan 16 SI Ohnishi (1998a) F. rubifolium Sichuan 32 SC Ohsako and Ohnishi (1998) F. gracilipedoides Yunnan 16 SI Ohsako et al. (2002) F. gilesii Yunnan 16 SI Hedberg (1946) F. jinshaense Yunnan 16 SI Ohsako et al. (2002) F. pleioramosum Sichuan 16 SC Ohnishi (1998a) F. callianthum Sichuan 16 SC Ohnishi (1998a) F. macrocarpum Sichuan 16 SC Ohsako and Ohnishi (1998) F. tibeticum Tibet 48 unknown Tian et al. (2011) F. pugense Sichuan 16 SC Tang et al. (2010) F. qiangcai Sichuan 16 SC Shao et al. (2011) F. luojishanense Sichuan 16 SC Hou et al. (2015) F. crispatifolium Sichuan 32 SC Liu et al. (2008a) F. longzhoushanense Sichuan 16 SC Wang et al. (2017) F. densovillosum Sichuan 16 SC Liu et al. (2008b) Ambiguous F. wenchuanense Sichuan 16 SC Shao et al. (2011) conduplicate cotyledons or embryo in the central region in defined two characters discriminating the two groups: later‐ achene. Nakai (1926) also distinguished Fagopyrum by the ally long cotyledons and achene partially covered with peri‐ centered position of embryo with conduplicate cotyledons. anths as the key character of the cymosum group and round Two subgroups have often been recognized within to longitudinally long cotyledons and achene completely Fagopyrum. Gross (1913) discriminated two subgroups in covered with perianths as the key character of the the section Eufagopyrum based on morphology of inflores‐ urophyllum group (Fig. 1). cence and perianth on achene. The subgroups composed of F. esculentum, F. tataricum, F. cymosum, F. suffruticosum and F. odontopterum (F. gracilipes var. odontopterum) was characterized by cymose inflorescence with many branch‐ ing and dense flowers and the other subgroups including F. maireii (F. urophyllum), F. bonatii (F. gracilipes), F. Grossii (F. leptopodum var. Grossii), F. statice and F. tristachyum (F. cymosum) by racemose inflorescence with sparse flowers. Roberty and Vautier (1964) amalga‐ mated Fagopyrum species into two species based on the achene size. They united F. esculentum, F. cymosum, and F. tataricum into F. esculentum sensu lato and F. gilesii, F. Grossii, F. gracilipes, F. statice and F. urophyllum into F. gilesii sensu lato. These groupings are mostly concordant with the cymosum group and the urophyllum group of Yasui and Ohnishi (1998a). Differentiation between the cymosum and the urophyllum groups is strongly supported Fig. 1. Cotyledons of Fagopyrum species. (a) F. esculentum ssp. by both morphological characters and molecular data esculentum, (b) F. tataricum ssp. tataricum, (c) F. cymosum, (d) (described in detail below). Ohnishi and Matsuoka (1996) F. urophyllum, (e) F. callianthum, and (f) F. leptopodum. 94 Breeding Science Classification of Fagopyrum Vol. 70 No. 1 BS group (Ohnishi 1998a, Ohsako and Ohnishi 1998, Ohsako Diversity of mating system in the genus et al. 2002). F. capillatum, F. rubifolium and F. gracilipedoides are morphologically similar to About half of the Fagopyrum species exhibit heterostyly F. gracilipes. These four species have heavily pubescent with short style-high anther flowers (thrum) and long style- stipule and leaf blade (Ohsako and Ohnishi 1998, Ohsako low anther flowers (pin). Most species showing heterostyly et al. 2002). F. capillatum and F. gracilipedoides are also have self-incompatibility (SI), but some species such diploid with chromosome number 2n = 16 and possess as F. pleioramosum, F. callianthum, and F. macrocarpum heterostylous SI. F. rubifolium is a self-compatible are self-compatible. Other species have homostylous self- tetraploid (Ohsako and Ohnishi 1998). F. pleioramosum, compatibility (SC). Mating system of the two cultivated F. callianthum, and F. macrocarpum have relatively large species is contrasting; common buckwheat exhibits hetero‐ flower and achene in the urophyllum group. Especially, stylous SI whereas Tartary buckwheat homostylous SC. In flowers and achenes of F. callianthum and F. macrocarpum contrast to SI with homomorphic flowers in other families are the largest within the group (Ohsako and Ohnishi such as Brassicaceae and Solanaceae, the degree of SI of 1998). In addition, they share heterostylous SC. This char‐ common buckwheat is extremely high such that self- acteristic is unique to these species in the genus (Ohsako fertilization rarely occurs. The possibility of
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