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Recircumscription of Sections of Hemerocallis (Asphodelaceae) from Japan and Adjacent Regions Based on MIG-Seq Data

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Recircumscription of Sections of Hemerocallis (Asphodelaceae) from Japan and Adjacent Regions Based on MIG-Seq Data ISSN 1346-7565 Acta Phytotax. Geobot. 71 (1): 1–11 (2020) doi: 10.18942/apg.201913 Recircumscription of Sections of Hemerocallis (Asphodelaceae) from Japan and Adjacent Regions Based on MIG-seq Data 1,† 1,† 1 2 3 Sho MurakaMi , koji TakayaMa , Shizuka FuSe , Shun k. hiroTa , SaToShi koi , 4 5,‡ 1,* TakahiTo ideno , Takenori yaMaMoTo and Minoru n. TaMura 1Department of Botany, Graduate School of Science, Kyoto University, Kitashirakawa-oiwake-cho, Sakyo-ku, Kyoto 606-8502, Japan. *[email protected] (author for correspondence); 2Field Science Center, Graduate School of Agricultural Science, Tohoku University, 232-3 Aza-yomogida, Naruko Onsen, Osaki, Miyagi 989-6711, Japan; 3Botanical Gardens, Faculty of Science, Osaka City University, 2000 Kisaichi, Katano, Osaka 576-0004, Japan; 4Botanical Gardens, Graduate School of Science, the University of Tokyo, 3-7-1 Hakusan, Bunkyo-ku, Tokyo 112-0001, Japan; 5Tsushima City Hall, 1441 Kokubu, Izuhara-cho, Tsushima, Nagasaki 817-8510, Japan. ‡Present address: Iriomote Station, Tropical Biosphere Research Center, University of the Ryukyus, 870 Uehara, Taketomi- cho, Yaeyama-gun, Okinawa 907-1541, Japan. †These authors contributed equally to this work Circumscription of three sections of Hemerocallis that were recognized by Matsuoka & Hotta (Acta Phy- totax. Geobot. 22: 25–43, 1966) and Hotta (Fl. Jap. IVb: 130–134, 2016) was evaluated through MIG-seq analysis of 45 samples of seven species and nine varieties of Hemerocallis, covering all taxa native to Japan except H. fulva vars. aurantiaca and pauciflora. The Neighbor-Net network of the MIG-seq data yielded three major clusters corresponding to the three sections of Hemerocallis recognized by Matsuo- ka & Hotta (1966) and Hotta (2016), except that H. hakuunensis and H. major, which were included in H. sect. Fulvae by Matsuoka & Hotta (1966) and Hotta (2016), were embedded in the cluster of H. sect. Hemerocallis. Hemerocallis hakuunensis and H. major usually have fragrant flowers and lack horizon- tally spreading rhizomes in the same manner as H. sect. Hemerocallis, which are not features of H. sect. Fulvae. We propose a transfer of H. hakuunensis and H. major from H. sect. Fulvae to H. sect. Hemero- callis. Hemerocallis minor appears to be conspecific withH. lilioasphodelus and is better regarded as H. lilioasphodelus var. minor. Hemerocallis lilioasphodelus var. thunbergii is placed in synonym under H. lilioasphodelus var. lilioasphodelus. Key words: Asphodelaceae, Hemerocallis, H. hakuunensis, H. lilioasphodelus var. minor, H. major, H. sect. Capitatae, H. sect. Fulvae, MIG-seq, Neighbor-Net network, taxonomy Hemerocallis L. (Asphodelaceae) comprises vae, Fulvae, Aurantiacae, Capitatae and Anthe- about 13 species and 12 varieties (Hu 1969, Chen latae, based on characteristics of the inflores- & Noguchi 2000, Hotta 2016) of perennial herbs cence, perianth, and color of anthers. Matsuoka & ranging from grasslands near seashores to moun- Hotta (1966) and Hotta (2016) grouped Nakai’s tainous regions in eastern Asia (Webb 1980), ex- sections into three, Hemerocallis, Fulvae, and cept for H. lilioasphodelus L., whose distribution Capitatae, based on color and odor of the flowers extends to Europe. and flowering time and additional morphological Infrageneric classifications of Hemerocallis characteristics. Thus, the infrageneric classifica- were proposed by Bailey (1930), Nakai (1932), tion of the genus Hemerocallis has been quite Matsuoka & Hotta (1966) and Hotta (2016) (Fig. controversial (Fig. 1), but it has never been reex- 1). Bailey (1930) divided the genus into two sec- amined in the light of molecular methods. tions, Euhemera and Dihemera, based on charac- Based on the study of Hotta (2016), six spe- teristics of the inflorescence. Subsequently, Na- cies and 11 varieties are attributed to Japan, al- kai (1932) proposed six sections, Citrinae, Fla- though Kitamura (1964) reported only four spe- 2 Acta Phytotax. Geobot. Vol. 71 Bailey Nakai Matsuoka & This study Species assigned to each section (1930) (1932) Hotta (1966) (2019) recircumscribed in this study and Hotta (2016) sect. sect. sect. H. citrina, H. hakuunensis Citrinae Hemerocallis Hemerocallis H. lilioasphodelus (=H. minor) sect. H. major sect. Flavae ** Euhemera sect. * sect. sect. Fulvae Fulvae Fulvae H. fulva sect. Aurantiacae sect. sect. sect. sect. Capitatae Capitatae Capitatae H. middendorffii Dihemera sect. Anthelatae Fig. 1. History of selected infrageneric classifications of Hemerocallis. * Matsuoka & Hotta (1966) considered including Hemerocallis sulphurea in H. aurantiaca. If this were accepted, part of H. sect. Flavae sensu Nakai (1932) would be transferred to H. sect. Fulvae by Matsuoka & Hotta (1966), because Matsuoka & Hotta (1966) included H. aurantiaca (= H. fulva var. aurantiaca) in H. sect. Fulvae. ** Hemerocallis hakuunensis and H. major are transferred from H. sect. Fulvae to H. sect. Hemerocallis in this study. cies and four varieties from Japan. Despite this which has been placed in three sections, were great difference in the treatments between Hotta used in this study (Table 1). Eleven samples were (2016) and Kitamura (1964), the species and their collected in the wild, 24 samples were from the infraspecific taxa have never been evaluated us- Botanical Gardens of Osaka City University and ing molecular methods. 10 samples were from the Koishikawa Botanical The purpose of this study was to use multi- Garden of the University of Tokyo. The leaves plexed inter-simple sequence repeat (ISSR) geno- were dried and stored in silica gel prior to DNA typing by sequencing (MIG-seq) to examine extraction. Identification of the plants followed Hemerocallis from Japan and adjacent regions to Hotta (2016). Voucher specimens have been de- determine if it would be useful in constructing an posited in the herbarium of Kyoto University infrageneric classification ofHemerocallis and to (KYO). reevaluate the species and varieties. This method Total DNA was extracted from the samples was recently developed by Suyama & Matsuki using the CTAB method (Doyle & Doyle 1987). (2015). Our intent was to use this genome-wide Single-nucleotide polymorphisms (SNPs) were method to elucidate the taxonomic issues based detected by using MIG-seq following the proce- on ample data from single nucleotide polymor- dure by Suyama & Matsuki (2015). Approximate- phisms (SNPs). ly 10 pM of the 2nd PCR product (library) was used for sequencing on an Illumina MiSeq Se- quencer (Illumina), using a MiSeq Reagent Kit Materials and Methods v.3, 150 cycle (Illumina). For data processing, low-quality reads were Leaves of Hemerocallis were collected from removed by the quality_filter option in FASTX- 45 samples from seven species and nine varieties, Toolkit [http://hannonlab.cshl.edu/fastx_toolkit/ February 2020 Murakami & al.—Sections of Hemerocallis Recircumscribed by MIG-seq 3 Table 1. Sources of materials. Sample Species / Variety Locality Source Voucher no. specimen Hemerocallis sect. Hemerocallis 1 Hemerocallis citrina Baroni var. Cult. in Beijing, China Bot. Gard., Osaka City Univ. Murakami 28 citrina 2 H. citrina var. vespertina (H. Hara) Japan: Pref. Yamanashi: Takane-cho Koishiwa Bot. Gard., Univ. Tokyo Murakami 50 M. Hotta 3 H. citrina var. vespertina Japan: Pref. Yamanashi: Takane-cho Koishiwa Bot. Gard., Univ. Tokyo Murakami 51 4 H. citrina var. vespertina Japan: Pref. Kyoto: Kyogamisaki Cape, Murakami 52 Kyotango-shi 5 H. citrina var. vespertina Japan: Pref. Wakayama: Rinkai, Shirahama- Bot. Gard., Osaka City Univ. Murakami 36 cho 6 H. citrina var. vespertina Japan: Pref. Nagasaki: Hatakezaki, Tsushima Yamamoto 1007 Isls. 7 H. lilioasphodelus L. var. lilioasphode- Cult. in Cluj-Napoca, Romania Koishiwa Bot. Gard., Univ. Tokyo Murakami 53 lus 8 H. lilioasphodelus var. thunbergii Japan: Hokkaido: Nosappu Cape, Nemuro-shi Bot. Gard., Osaka City Univ. Murakami 25 (Baker) M. Hotta 9 H. lilioasphodelus var. thunbergii Japan: Hokkaido: Shiraoi-cho Koishiwa Bot. Gard., Univ. Tokyo Murakami 54 10 H. minor Mill. China: Mudanjiang Osaka Gakuin Univ. Murakami 45 11 H. minor Cult. in Univ. of Copenhagen Bot. Gard., Bot. Gard., Osaka City Univ. Murakami 55 Denmark 12 H. minor Cult. in Nishinomiya-shi, Pref Hyogo, Japan Bot. Gard., Osaka City Univ. Murakami 10 H. sect. Capitatae Nakai 13 H. middendorffii Trautv. & C. A. Mey. Japan: Hokkaido: Wakasakanai Seashore, Koishiwa Bot. Gard., Univ. Tokyo Murakami 59 var. middendorffii Toyotomi-cho 14 H. middendorffii var. esculenta Japan: Pref. Aomori: Utozaka, Sannohe-machi Koishiwa Bot. Gard., Univ. Tokyo Murakami 56 (Koidz.) Ohwi 15 H. middendorffii var. esculenta Japan: Pref. Toyama: Mt. Yakushi, Toyama-shi Bot. Gard., Kyoto Univ. Tamura et al. 44001 16 H. middendorffii var. esculenta Japan: Pref. Nagano: Shigakogen, Murakami 46 Yamanouchi-machi 17 H. middendorffii var. esculenta Japan: Tokyo: Kyodo-no-Mori, Fuchu-shi Koishiwa Bot. Gard., Univ. Tokyo Murakami 57 18 H. middendorffii var. exaltata (Stout) Japan: Pref. Akita: Otakizawa, Oga-shi Murakami 23 M. Hotta 19 H. middendorffii var. exaltata Japan: Pref. Yamagata: Hoki, Tobishima Isl. Bot. Gard., Osaka City Univ. Murakami 58 20 H. middendorffii var. exaltata Japan: Pref. Yamagata: Arasaki, Tobishima Isl. Bot. Gard., Osaka City Univ. Murakami 16 21 H. middendorffii var. exaltata Japan: Pref. Yamagata: Sai-no-kawara, Bot. Gard., Osaka City Univ. Murakami 17 Tobishima Isl. 22 H. middendorffii var. exaltata Japan: Pref. Yamagata: Sai-no-kawara, Bot. Gard., Osaka City Univ. Murakami 14 Tobishima
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