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Taxonomic Revision of Oxalis Subsect. Oxalis (Oxalidaceae)

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Taxonomic Revision of Oxalis Subsect. Oxalis (Oxalidaceae) ISSN 1346-7565 Acta Phytotax. Geobot. 70 (3): 159–172 (2019) doi: 10.18942/apg.201906 Taxonomic Revision of Oxalis subsect. Oxalis (Oxalidaceae) 1,* 2,† 2 SatoShi aoki , tetSuo ohi-toma and Jin murata 1Graduate School of Science, The University of Tokyo, 3-8-1 Komaba, Meguro-ku, Tokyo 153-8902, Japan. *[email protected] (author for correspondence); 2Botanical Gardens, Graduate School of Science, The University of Tokyo, 3-7-1 Hakusan, Bunkyo-ku, Tokyo 112-0001, Japan. † Present address: Nature Fieldwork Center, Okayama University of Science, 1-1 Ridaicho, Kita-ku, Okayama 700-0005, Japan The classification of Oxalis subsect. Oxalis (Oxalidaceae) is revised based on phylogenetic, cyto- logical and morphological evidence. For the flora of Japan, one new species and one new variety are described, and five new combinations are proposed. The new species, O. nipponica, resembles O. griffithii, but is distinguished by its unique phylogeny, larger genome and distribution. Oxalis nipponica is subdivided into two subspecies, nipponica and kantoensis based on phylogeny and fruit length. Subsp. kantoensis is further divided into two varieties based on ploidy level, length of the fruit and seeds and habitat. While diploids correspond to var. kantoensis, the tetraploids cor- respond to the newly described var. alpigena. Variety alpigena resembles O. acetosella and has been confused with it. Fourteen taxa, including one hybrid, are recognized in subsect. Oxalis. Key words: new species, new variety, Oxalidaceae, Oxalis nipponica subsp. kantoensis var. alpigena The taxonomic history of Oxalis L. sect. Ox- cally, have caused frequent misidentifications in alis subsect. Oxalis based on Hara (1954, 1955), subsect. Oxalis, especially of the species in East- Terao (1979), Huang & Liu (2003), Huang et al. ern Asia (Liu & Watson 2008). (1998), Lourteig (2000), Amano (2001), and Liu To reassess the taxonomy of subsect. Oxalis, & Watson (2008) was summarized previously we compared the morphology, genome size and (see Table 1 in Aoki et al. 2017). In the Flora of molecular phylogeny of samples of Oxalis from a Japan (Amano 2001), this section was treated as wide geographical area (Aoki et al. 2017, 2018). sect. Acetosellae (Reiche 1894) R. Knuth 1914, In the phylogenetic tree (Fig. 1), some taxa were but it should have been sect. Oxalis, because this not monophyletic. section includes O. acetosella L., the type of the In our studies, Oxalis griffithii var. griffithiiin genus Oxalis (Turland et al. 2018, International Japan (including the indistinguishable O. aceto- Code of Nomenclature Art. 22.1). Lourteig (2000) sella var. longicapsula) formed a clade (Clade C) treated all taxa in the subsection and recognized independent of those in China (Clade A). Oxalis O. magellanica G. Forst. in Oceania and South acetosella var. longicapsula was also in Clade C, America, O. griffithiiEdgew. & Hook.f., O. obtri- and O. griffithii var. kantoensis was placed in angulata Maxim. and O. leucolepis Diels in the Clade D. Most Japanese plants of O. acetosella Far East and eastern Asia, O. oregana Nutt. ex var. acetosella, formed a clade with O. acetosella Torr. & A. Gray and O. trilliifolia Hook. in North var. acetosella in China and Europe (Clade B), America and the widespread O. acetosella. Leaf- but a few Japanese plants were in Clade D. Addi- let shape and the spacing of the petioles on the tionally, the American O. acetosella var. aceto- rhizome, which are difficult to express numeri- sella occupied a unique phylogenetic position (as 160 Acta Phytotax. Geobot. Vol. 70 Oxalis montana in Fig. 1). The comparison be- In the present taxonomic treatment, we made tween the previous classification (Lourteig 2000, species correspond to the confident clades. Intra- Amano 2001) and the results from our analysis is specific taxa were assigned based on cytological summarized in Table 1. or minor phylogenetic differences accompanied Our cytological analysis showed differences by their ecological and/or morphological differ- in genome size among Clades A–D, but Clades C ences. By minor phylogenetic differences, we and D had genomes of nearly the same size. mean monophyly with low confidence or mono- Clades C and D also had genomes of two sizes, typic lineages that render a species paraphyletic which corresponded to the diploids and tetra- or polyphyletic if they are considered as species. ploids reported by Terao (1979). We examined specimens in the herbaria of Kyoto Besides examining specimens, we measured University (KYO), Tokyo Metropolitan Univer- the length of the capsule, the length to width ratio sity (MAK), Makino Botanical Garden (MBK), of the capsules and the length of seeds among our National Taiwan University (TAI), Hokkaido samples. We then compared the measurements. University Museum (SAPS), and the University From the data gathered from samples of O. of Tokyo (TI). We also consulted digital images griffithii subsp. griffithii, we could not distin- of specimens at the herbaria BR, E, G-DC, GH, guish between the samples from China (Clade A) HPSU, ID, K, LINN, M, P, PE, SOC, SRP, TNS, and Japan (Clade C). While diploids and tetra- US and WTU. The herbarium acronyms follow ploids in Clade C were indistinguishable, capsule Thiers (2018). length and seed length were informative for dis- From our results, we recognize two new cryp- tinguishing between diploids and tetraploids in tic taxa. We treated the Japanese plants previous- Clade D. Consequently, we decided to revise sub- ly identified as O. griffithii var. griffithii as a new section Oxalis. In this revision, we excluded O. species, O. nipponica subsp. nipponica. Our magellanica, because it was located phylogeneti- treatment is based on differences in genome size cally outside the outgroup in our phylogenetic and phylogenetic position. analysis (Aoki et al. 2017). We kept the descrip- The second unrecognized cryptic taxon cor- tion of North American taxa simple, because we responds to tetraploids of O. griffithii var. kan- examined few specimens in our previous studies toensis (Terao) T. Shimizu. This cryptic taxon (Aoki et al. 2017, 2018). was previously treated as tetraploids of O. aceto- Oxalis japonica Franch. & Sav. has been con- sella by Terao (1979). In a previous study (Aoki et sidered to be a Japanese endemic distinct from O. al. 2017), we determined its phylogenetic position griffithii (cf. Makino 1908, Hara 1954), probably and ploidy level. Considering its characteristic based on the description and the Japanese name habitat and distribution, we now chose to treat used in the original study. Hara (1955), however, this tetraploid as a new variety distinct from var. determined the holotype, Savatier 2816 (P, kantoensis and from O. acetosella, despite the P00724040) of O. japonica, to be O. martiana difficulty in separating them based on morpholo- Zucc., which had been introduced to Japan from gy. The diploid plant previously identified as O. South America. Hara (1955) identified the Japa- griffithii var. kantoensis is now treated as O. nip- nese plants that had been treated as O. japonica ponica subsp. kantoensis var. kantoensis, and the or O. acetosella subsp. japonica (Franch. & Sav.) cryptic tetraploid is treated as O. nipponica sub- H. Hara as O. griffithii. Huang et al. (1998) ac- sp. kantoensis var. alpigena. We also treat O. cepted O. acetosella subsp. japonica and synony- wulingensis from China, O. acetosella subsp. mized O. obtriangulata under it without consid- griffithiivar. formosana from Taiwan, and O. tai- ering Hara’s (1955) treatment. In addition, they monii from Taiwan as subspecies of O. griffithii. (Huang et al., 1998) did not recognize O. aceto- We treated them as subspecies, since all of them sella subsp. griffithii as occurring in Japan. In were included in the same clade as O. griffithii this study, we follow the treatment of Hara (1955). (Clade A), but did not share identical sequences October 2019 aoki & al. – Taxonomic Revision of subsect. Oxalis 161 table 1. Comparison between previous taxonomy (Lourteig 2000, Amano 2001) and taxonomy proposed here for confusing taxa in subsect. Oxalis. Proposed taxonomy shows scientific names suggested in this article and their corresponding phy- logenetic position as shown in Fig. 1 and ploidy levels for taxa in Clade D. Previously recognized taxa Proposed taxa O. griffithii (Clade A) O. griffithii var. griffithii O. nipponica subsp. nipponica (Clade C) O. griffithii var. kantoensis (diploid) O. nipponica subsp. kantoensis var. kantoensis (Clade D, diploid) O. acetosella (Clade B, diploid) O. acetosella var. acetosella (diploid, tetraploid) O. montana O. nipponica subsp. kantoensis var. alpigena (Clade D, tetraploid) O. acetosella var. longicapsula O. nipponica subsp. nipponica (Clade C) Fig. 1. Abridged phylogeny of Oxalis subsect. Oxalis based on nuclear ribosomal internal transcribed spacer (ITS) and Bayes- ian methods. Numbers near branches indicate posterior probabilities and bootstrap values by maximum likelihood and maximum parsimony methods. Hyphens indicate branches that did not appear in analysis; branches without numbers are monotypic branches based on a single sequence. For details, see Aoki et al. (2018). with Oxalis griffithii subsp. griffithii, and they cluding one hybrid, in subsect. Oxalis: O. aceto- exhibited clear morphological differences from sella, O. obtriangulata, O. nipponica subsp. nip- subsp. griffithii. We also recognize O. montana as ponica, O. nipponica subsp. kantoensis var. kan- distinct from O. acetosella
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