J. Jpn. Bot. 92(1): 34–43 (2017)

Harashuteria, a New Genus of Leguminosae (Fabaceae) Subfam. Papilionoideae Tribe Phaseoleae

a a b, Kazuaki Ohashi , Koji Nata and Hiroyoshi Ohashi *

aSchool of Pharmacy, Iwate Medical University, Yahaba, Iwate, 028-3694 JAPAN; bHerbarium TUS, Botanical Garden, Tohoku University, Sendai, 980-0862 JAPAN *Corresponding author: [email protected]

(Accepted on November 25, 2016)

A new genus, Harashuteria K. Ohashi & H. Ohashi, is proposed as a member of the tribe Phaseoleae of Leguminosae (Fabaceae) based on Shuteria hirsuta Baker by comparative morphological observation and molecular phylogenetic analysis of Shuteria and its related genera. Molecular phylogenetic analysis was performed using cpDNA (trnK/matK, trnL–trnF and rpl2 intron) markers. Our molecular phylogeny shows that Shuteria hirsuta is sister to Cologania and is distinct from Shuteria vestita or Amphicarpaea, although the species has been attributed to these genera. A new combination, Harashuteria hirsuta (Baker) K. Ohashi & H. Ohashi is proposed.

Key words: Amphicarpaea, Cologania, Fabaceae, Glycininae, Harashuteria, Hiroshi Hara, new genus, Phaseoleae, Shuteria, Shuteria hirsuta.

The genus Shuteria Wight & Arn. was protologue. Kurz (1877) recognized Shuteria established on the basis of S. vestita Wight & hirsuta as a member of Pueraria, because he Arn. including 4–5 species in Asia (Schrire described Pueraria anabaptista Kurz citing 2005). The genus belongs to the subtribe S. hirsuta as its synonym (hence Pueraria Glycininae in the tribe Phaseoleae and is anabaptista is superfluous). Amphicarpaea closely allied to Amphicarpaea, Cologania, lineata Chun & T. C. Chen is adopted as a and Dumasia especially in the flower structures correct species by Sa and Gilbert (2010) in the (Lackey 1981). Among Shuteria, S. hirsuta recent treatment of Fabaceae for Flora of China, Baker (Fig. 1) is a divergent species from others but is treated by Ohashi and Ohashi (2016) as a of the genus in having a spurred standard, synonym of Shuteria hirsuta. monadelphous stamens and linear pods with Recently, however, Shuteria is consistently septa between the seeds. The species has so separated from Amphicarpaea, Cologania, far been placed in a controversial taxonomic Pueraria and other members of the subtribe position among Amphicarpaea, Pueraria and Glycininae in molecular phylogenetic analyses Shuteria. Baker (1876) described the species (Doyle and Doyle 1993, Lee and Hymowitz based on two syntypes those were referred to 2001, Stefanović et al. 2009, Egan et al. 2016). Amphicarpaea ferruginea Benth. as indicated Shuteria belongs to a Kennediinae-Desmodieae on the original label of the specimens (Fig. clade, while the other genera to a Glycininae 1), although the name was not cited in the clade within the Core-Phaseoleae clade. In these

—34— February 2017 The Journal of Japanese Botany Vol. 92 No. 1 35

Fig. 1. Lectotype of Shuteria hirsuta Baker (©The Board of Trustees of the Royal Botanic Gardens, Kew). 36 植物研究雑誌 第 92 巻 第 1 号 2017 年 2 月

Table 1. Voucher information and GenBank accession numbers on data newly sequenced for this paper Taxon Locality Voucher Shuteria hirsuta (1) Cultivated in Setsunan University, Osaka Hiroko Murata (TUS) (Origin: Myanmar) (2) Nepal. Koshi Zone, Sankhuwasabha Distr. S. Noshiro & al. 9810179 (TUS) (3) Thailand. N. Chiang Mai H. Koyama & al. T-48821. (TUS) Shuteria vestita (2) Nepal. Between Thare and Dhuche H. Tabata & al. 8631 (TUS) Dumasia truncata Japan. Miyagi Pref., Higashi-Matsushima-shi, H. Kasai 3873 (TUS) Miyatojima Toxicopueraria peduncularis China. Yunnan: Li-jiang, Wen-bi-feng Murata & al. 662 (TUS)

GenBank accession number Taxon trnK/matK trnL intron trnF–trnL spacer rpl2 Shuteria hirsuta (1) LC197941 LC197931 LC197935 LC196161 (2) LC197938 — — — (3) LC197939 — — — Shuteria vestita (2) LC197942 LC197932 LC197936 LC196162 Dumasia truncata LC197940 LC197930 LC197934 LC195833 Toxicopueraria peduncularis LC197943 LC197933 LC197937 LC196160 (1), (2), and (3) correspond to the individuals of Shuteria hirsuta or S. vestita in phylogenetic tree in Figs. 1 and 2. Sequence data of S. vestita (1) are obtained from GenBank. Missing data are indicated by a dash (–). molecular analyses, however, Shuteria hirsuta collected from fresh leaves of a cultivated plant has not been examined, and only S. vestita was in The Botanical Garden, Setsunan University. used as the representative of the genus. Voucher specimen of the plant is deposited in This study aims to solve the controversial TUS. Samples listed in Table 1 including other taxonomic position of Shuteria hirsuta, because S. hirsuta individuals were collected from it has such clear morphological differences herbarium specimen kept in TUS. Already between other species of Shuteria as mentioned published DNA sequences of S. vestita and other above. We present here molecular evidence that species were obtained from GenBank (listed in Shuteria hirsuta is distinctly separated from S. Table 2). vestita hitherto examined and is closely related to Cologania. Shuteria hirsuta and Cologania, DNA extraction, amplification and sequencing however, differ in the flower and legume Genomic DNA was extracted from leaf morphology, the chromosome numbers and their tissue using the modified CTAB method of distribution. In consequence, we here describe Doyle and Doyle (1987). Three chloroplast a new distinct genus Harashuteria K. Ohashi & markers were amplified: trnK intron including H. Ohashi to accommodate the single species, S. matK (trnK/matK), trnL–trnF (trnL intron and hirsuta Baker. trnF–trnL spacer) and rpl2 intron. The PCR primers used for trnK/matK were trnK1L and Materials and Methods trnK2R (Hu et al. 2000). The primers for trnL– Materials trnF (trnL intron and trnF–trnL spacer) were One of the samples of Shuteria hirsuta was primer ‘c’ and primer ‘f’ (Taberlet et al. 1991). February 2017 The Journal of Japanese Botany Vol. 92 No. 1 37

Table 2. Sequence data already published and obtained from GenBank GenBank accession number Taxon trnF–trnL spacer/ trnK/matK rpl2 trnL intron Amphicarpaea bracteata EU717399 EU717317 EU717364 Apios americana EU717426 EU717312 EU717392 Bituminaria bituminosa EU717398 EU717349 EU717362 Bolusafra bituminosa EU717413 EU717309 EU717362 Cajanus cajan EU717414 EU717310 EU717361 Campylotropis macrocarpa EU717418 EU717298 EU717384 Clitoria ternatea EU717286 EU717355 EU717528 Cologania angustifolia GQ246140 – – Cologania lemmonii EU717405 EU717319 EU717371 Cologania pallida JQ619980 – – Cologania tenuis JQ619979 – – Desmodium barbatum EU717420 EU717290 EU717386 Erythrina sousae EU717411 EU717313 EU717377 Galactia striata EU717356 EU717428 EU717394 Glycine max AF142700 EU717321 EU717363 Hardenbergia violacea EU717425 EU717331 EU717391 Kennedia nigricans EU717424 EU717335 EU717390 Kummerowia stipulacea EU717417 EU717299 EU717383 Lablab purpureus EU717408 EU717339 EU717374 Macroptilium atropurpureum EU717409 EU717340 EU717375 Macrotyloma uniflorum EU717410 EU717341 EU717376 Neonotonia wightii U717402 EU717323 EU717368 Ophrestia radicosa EU717430 EU717359 EU717396 Pachyrhizus erosus EU717401 EU717324 EU717367 Pseudovigna argentea EU717403 EU717325 EU717369 Psophocarpus tetragonolobus EU717412 EU717343 EU717378 Pueraria phaseoloides EU717404 EU717327 EU717370 Shuteria vestita (1) EU717423 EU717328 EU717389 Teramnus uncinatus EU717400 EU717330 EU717365 (1) corresponds to the individuals of Shuteria vestita in phylogenetic tree in Figs. 1 and 2. Missing data are indicated by a dash (–).

For amplification of rpl2 intron, rpl2 intron f products were run on an Applied Biosystems (5ʹ-GCT CTA GCT AAT TGT CCA CCC TT-3ʹ) 3130xl Genetic Analyzer. and rpl2 intron r (5ʹ-AAA ATG GGA AAT GCC CTA CCT T-3ʹ) were used. In case of failure Phylogenetic analyses of amplification, internal primers were used to Sequence alignment was initially performed amplfy the region in two or more fragments. using MUSCLE (Edgar 2004) in Mega version PCR was performed using KOD Fx neo 7.0 (Kumar et al. 2016) and manually adjusted. (TOYOBO, Osaka, Japan) in 20 μL reactions Phylogenetic analyses were carried out using with the following PCR program: 94°C for 5 Bayesian inference (BI), maximum likelihood min; 40 cycles of 98°C for 10 s 52°C 30 s 72°C (ML), and maximum parsimony (MP) 90 s; 72°C 5 min. PCR products were purified approaches. by polyethylene glycol (PEG) precipitation or An appropriate model of sequence evolution ethanol precipitation with NH4OAc and EDTA. for the combined dataset was estimated using the The sequencing reactions were performed program Kakusan4 (Tanabe 2007). The dataset using the BigDye Terminator, v3.1, Cycle was divided into data partitions (rpl2 intron, trnL Sequencing Kit (Applied Biosystems, Foster intron, trnF–trnL spacer, 5ʹtrnK, 3ʹtrnK, and City, California, U.S.A.), and the reaction matK as protein coding sequence). Models of 38 植物研究雑誌 第 92 巻 第 1 号 2017 年 2 月

Fig. 2. The 50% majority-rule consensus tree (Bayesisn analysis) of Shuteria with Paseoleae species based on the cpDNA dataset (trnK/matK, trnL–trnF and rpl2 intron). Support values on the branches are presented as follows: BI-PP/ML- BS/MP-BS. NS indicates node not supported. sequence evolution for each of the partitions and discarded as burn-in from the final tree set that for all combined datasets were determined. was used to determine the posterior probability Bayesian inference (BI) analyses were distribution. conducted using MrBayes version 3.2 Maximum likelihood (ML; Felsenstein 1981) (Ronquist et al. 2012). The GTR+G model and ML bootstrap (Felsenstein 1985) analyses with Proportional_Codon-proportional model were performed using RAxML v.8 (Stamatakis for cpDNA was selected based on AIC. The 2014), with the partitioned dataset. Partitioned GTR+G model with Codon-proportional model equal mean rate Codon partitioned equal mean was selected for matK analysis. Markov Chain rate model was selected for cpDNA analysis. Monte Carlo (MCMC) analysis was executed Codon partitioned equal mean rate model was for five million generations with four chains, and selected for matK analysis. Analysis with other sampled every 500th generation. The average models (HYK etc.) with no partitioned cpDNA standard deviation of split frequencies was data were performed using MEGA7. checked below 0.01. Tracer v.1.6.0 (Rambaut et The maximum-parsimony (MP) analyses al. 2014) was used to examine the convergence were performed using the Tree-Bisection- of model parameters and check the values of Regrafting (TBR) algorithm with search level effective sample size (ESS) were over 200. 2 in MEGA7. The combined dataset of cpDNA The first 10% of the trees from each run were or matK was analyzed. All positions containing February 2017 The Journal of Japanese Botany Vol. 92 No. 1 39

Fig. 3. The 50% majority-rule consensus tree (Bayesisn analysis) of Shuteria with Paseoleae species based on the matK dataset. Support values on the branches are presented as follows: BI-PP/ML-BS/MP-BS. NS indicates node not supported. gaps and missing data were eliminated for BI, Shuteria hirsuta was not involved in the clade ML and MP analysis. One thousand replicates with S. vestita. Our results indicate S. hirsuta is for bootstrap test were performed for ML and sister to Cologania lemmonii (Fig. 1). BI and MP. ML analysis using several other models (e.g., HYK or F81 instead of GTR+G, and other Results partition models) gave trees with almost identical To explore the position of Shuteria species topology supporting polyphyly of Shuteria and in Phaseoleae, molecular phylogenetic analysis the clade comprising S. hirsuta and C. lemmonii was performed using cpDNA (trnK/matK, trnL– (data not shown). trnF and rpl2 intron) markers. The aligned Since matK data were available in GenBank cpDNA dataset was 4859 positions and matK on four species of Cologania (C. angustifolia dataset was 1569 positions. The Bayesian 50% Kunth, C. lemmonii A. Gray, C. pallida Rose majority rule consensus trees of the combined and C. tenuis Rose), phylogenetic analysis cpDNA (Fig. 1) and of the matK (Fig. 2) were on matK data were also performed (Fig. 3). shown with posterior probability and bootstrap The matK phylogeny shows four species of support values from ML and MP analysis. Cologania form a clade sister to S. hirsuta. Three Shuteria was polyphyletic both in the combined independent specimens of S. hirsuta (Shuteria cpDNA tree (Fig. 2) and matK tree (Fig. 3). hirsuta (1)–(3) in Fig. 3) form a clade that is 40 植物研究雑誌 第 92 巻 第 1 号 2017 年 2 月 sister to Cologania species. keels similar to the wings in length and shape and sessile pistil. The genus is also similar to Discussion Amphicarpaea in having auriculate standard Our results of the phylogenetic analysis and the keels similar to the wings in length reveal the polyphyly of Shuteria and that and shape, but differs in having monadelphous S. hirsuta is sister to Cologania. Thus it is stamens and the linear many-seeded pods with appropriate to exclude S. hirsuta from Shuteria. septa between seeds. Although it might be possible to include S. Description: Climbing herb, leaves pinnately hirsuta in Cologania based on the analysis, 3-foliolate, stipulate, stipellate. Inflorescences the matK tree shows that the distance between raceme, axillary, flowers 2 or 3 per node (3 at the S. hirsuta and Cologania is not close in proximal node), subtended by a primary bract. comparison to the distance among Cologania Pedicels short, each subtended by a secondary species. Cologania, although once merged with bract, with 2 bracteoles at tip. Calyx tubular, Amphicarpaea by Taubert (1894), is a sister 4-lobed, lobes 1/4 as long as the calyx, adaxial to Calopogonium included in the Glycininae 2 connate almost to tip, abaxial one slightly clade (Lee and Hymowitz 2001). Cologania is longer. Standard obovate, lamina erect, auricled tetraploid 2n = 44 and occurs mainly in Mexico, at base, apex rounded or retuse; wings long- distributed in central and south America (Lackey clawed, adherent to the keel, lamina oblong, 1981). The genus differs from Shuteria hirsuta with a retrorse auricle towards the base at in having the standard without auricles at the adaxial base of the lamina, apex obtuse; keel base of lamina, keels shorter than the wings as long as the wings, long-clawed, auricled at with smaller lamina than those of the wings, adaxial base of the lamina, apex obtuse or acute. diadelphous stamens, and stipitate ovary. Thus, Stamens 10, monadelphous. Disk present. Pistils we consider that Shuteria hirsuta should be linear, glabrous, almost sessile, ovary linear, ca. separated from Cologania. 10-ovuled, style filiform, stigma small. Pods Phylogenetic analysis in a thesis by Cagle linear, 9–10-seeded, 2-valved, septate between (2013) for the analysis of Pueraria polyphyly the seeds; seeds globose-oblate, hilum broadly showed that Shuteria hirsuta is sister to elliptic. Toxicopueraria by using the data of about 0.7 Distribution: India, Nepal, Sikkim, Bhutan, kb matK fragment, though no data on S. hirsuta Myanmar, Thailand, Vietnam, Laos, and SW was described in Egan et al. (2016). However, China. our data show that T. peduncularis is sister to Etymology: The new generic name is Dumasia (Figs. 2, 3). composed of Professor Dr. Hiroshi Hara (1911–1986) and Shuteria. He had made Taxonomic treatment great contribution to Systematic Botany and Harashuteria K. Ohashi & H. Ohashi, gen. established comparative floristic studies between nov. Japan and the Himalayas in order to study Type: Harashuteria hirsuta (Baker) K. the origin of the Japanese flora. He guided H. Ohashi & H. Ohashi Ohashi for taxonomic studies on the Himalayan Diagnosis: Similar to Shuteria and Cologania legumes in 1963. in general appearance, but differs from Shuteria Harashuteria hirsuta (Baker) K. Ohashi & and Cologania in having auriculate standard and H. Ohashi, comb. nov. monadelphous stamens. Harashuteria differs Shuteria hirsuta Baker in Hook. f., Fl. Brit. from Shuteria by the linear pods with septa India 2: 182 (1876) [Lectotype (van Thuan between the seeds and from Cologania by the 1972): INDIA. Khasia. Regio temp., alt. February 2017 The Journal of Japanese Botany Vol. 92 No. 1 41

3–5000 ped. J. D. Hooker & T. Thomson s.n. 3–16 cm long; peduncle 1.5–8 cm long; primary (K000556857)]; Prain in J. Asiat. Soc. Beng. 66, bracts ovate to narrowly ovate, 2–4 mm long, pt. 2, 402 (1897); Craib in Kew Bull. Misc. Inf. acuminate, ciliate. Pedicels short, 1–2 mm 1911: 39 (1911); Gagnep. in Lecomte, Fl. Indo- long, glabrous. Bracteoles linear, 2–3 mm long. Chine 2: 405 (1920); van Thuan in Adansonia, Calyx tubular, glabrous or sparsely puberulent, ser. 2, 12: 303 (1972), p.p. excl. type of Glycine 5–6 mm, 4-lobed; lobes unequal, adaxial lobes ferruginea Grah. nom. nud., Wallich 5514 (K); broadly deltate, 1–1.5 mm long, connate to apex, van Thuan in Fl. Cambodge Laos Viet-Nam 17: lateral ones deltate, ca. 1 mm long, abaxial one 67 (1979), p.p.; H. Ohashi, Enum. Fl. Pl. Nepal deltate, ca. 1.5 mm long, slightly longer than 2: 130 (1979); D. A. Huang, Fl. Reip. Pop. Sin. others. Corolla blue; standard obovate, 10–14 41: 244 (1995); H. Ohashi & K. Ohashi in J. mm long, auricled at the lamina-base, lamina Jpn. Bot. 91 Suppl.: 236 pollen & 247 (2016). 7–9 mm long, 6–8 mm wide; wings 10–12 mm [Pueraria anabaptista Kurz in J. Asiat. Soc. long, with a long slender claw, 5–7 mm long, Bengal, Pt. 2, Nat. Hist. 45(4): 253 (1877, not lamina oblong, ca. 5 mm long, 2–3 mm wide, 1876), nom. superfl.] with an auricle at adaxial base, apex obtuse; [Shuteria anabaptis C. Y. Wu in J. W. China keels 9.5–12 mm long, with a long slender Border Res. Soc. 16: 173 (1946), in adnot. as claw, 5.5–7 mm long, lamina, ca. 5 mm long, “(Kurz) C. Y. Wu, comb. nov.”, nom. nud.] 2–3 mm wide, with an auricle at adaxial base, Amphicarpaea lineata Chun & T. C. Chen apex obtuse or acute. Stamens 10–14 mm long, in Acta Phytotax. Sin. 7: 23, pl. 8 (1958) [Type: monadelphous, vexillary one coherent with the CHINA. Hainan. Paoting Hsien. 6 Jan. 1934. H. others except distal free part. Anthers small, Y. Liang 64056 (PE–holotype, ISBC–isotype)]; almost orbicular. Disk small, ca. 0.2 mm long. Chun & Chang, Fl. Hainan. 2: 303 (1965); Sa Pistil linear, 10.5–13 mm long, glabrous; ovary and Gilbert, Fl. China 10: 249 (2010). linear, 9–12-ovuled; style filiform, 2–6 mm long, Shuteria lancangensis Y. Y. Qian in Guihaia stigma minutely capitate. Legumes black, linear, 23: 307 (2003); Sa & Gilbert, Fl. China 10: 206 ca. 5 cm long, 4 mm wide, hirsute, 9–10-seeded, (2010) [Type: CHINA. Yunnan. Lancang, alt. with septa between the seeds. Seeds black, 1900 m, in thickets. 28 Sep. 1992. Y. Y. Qian nearly orbicular, 3 × 2.7 × 2 mm in size; hilum 2683 (HITBC–holotype; IBK–isotype)]; Sa & ca. 1 × 0.8 mm. Pollen grains see Ohashi and Gilbert, Fl. China 10: 206 (2010). Ohashi (2016). [Pueraria ferruginea Kurz ex Sa & Gilbert, Representative specimens examined: INDIA. Khasia. Fl. China 10: 205 (2010), nom. illeg.] Regio temp., alt. 3–5000 ped. J. D. Hooker & T. Thomson [Shuteria ferruginea (Kurz) Baker ex Sa & s.n. (K000556857). SIKKIM. Regio subtrop. J. D. Hooker Gilbert, Fl. China 10: 205 (2010), nom. illeg.] s.n. (K000894837). NEPAL. Dumham - Taplejung. 1 Nov. 1963. Kanai & al. 6301695 (TI, TUS); Papung - Sangrati Herbs, perennial. Stems slender, densely Pati. 26 Aug. 1977. Ohashi & al. 771068 (TI, TUS); Koshi hirsute. Stipules linear to narrowly ovate, Zone. 29 Aug. 1998. Noshiro & al. 9810179 (TI, TUS); 3–5 mm, acuminate; petiole 3–8 cm long; Chitwan Distr. 18 Jun. 1996. Mikage & al. 9614173 (TUS). stipels small, subulate; rachis 1–3 cm long, THAILAND. Chanburi. 26 Nov. 1956. Smitinand 3643 hirsute; leaflets thinly chartaceous; terminal (BKF); Dong Lan Forest. 28 Nov. 1958. Sørensen & al. leaflet ovate to broadly ovate, 4.5–10 × 3–6.5 6107 (TI); Chieng Mai, Doi Chieng Dao. 8 Sep. 1959. Bunchaui 1009 (BKF); Sai Yok. Larsen 8754 (BKF); N. cm, both surfaces sparsely appressed hirsute, Chiang Mai. 1 Dec. 1984. Koyama & al. T-48821 (TUS). basal veins 3, lateral veins 4 or 5 on each side, CHINA. Hainan. Paoting Hsien. 6 Jan. 1934. H. Y. Liang slender, base rounded or nearly truncate, rarely 64056 (PE, ISBC). broadly cuneate, apex acuminate; lateral leaflets Sa and Gilbert (2010) adopted Shuteria obliquely ovate, smaller. Racemes axillary, ferruginea (Kurz) Baker as the correct name 42 植物研究雑誌 第 92 巻 第 1 号 2017 年 2 月 for S. hirsuta Baker (1876) based on their sequences: A maximum likelihood approach. J. Molec. recognition “that name [= S. hirsuta Baker] Evol. 17: 368–376. Felsenstein J. 1985. Confidence limits on phylogenies: An was published three years later than Pueraria approach using the bootstrap. Evolution 39: 783–791. ferruginea [Kurz]”. They regarded P. ferruginea Hu J. M., Lavin M., Wojciechowski M. F. and Sanderson was published by Kurz (1874) as a new species, M. J. 2000. Phylogenetic systematics of the tribe and cited it as the basionym for Shuteria Millettieae (Leguminosae) based on chloroplast trnK/ ferruginea. 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a a b 大橋一晶 ，那谷耕司 ，大橋広好 ：マメ科マメ亜科イ ンゲンマメ連の新属 Harashuteria マメ科マメ亜科インゲンマメ連に属する Shuteria を行い，その結果に基づいて系統樹 (Figs. 2, 3) を作成 は 4–5 種からなり，スリランカ，インド，ヒマラヤ した．Shuteria hirsuta は Shuteria (S. vestita) のクレード 東部（ネパール・シッキム），アッサムから東南アジ と離れて，意外にも中南米の Cologania に類縁があった． アおよび中国南部に分布するとされている (Schrire 形態上は S. hirsuta は Shuteria の他の種とは線形の豆果 2005)．Shuteria は ヤ ブ マ メ 属 Amphicarpaea， ノ サ サ に多数の種子を入れ，種子の間には隔膜があることで異 ゲ属 Dumasia，Cologania に近縁とされている (Lackey なっていた．しかし，S. hirsuta と Cologania は花，豆果， 1981)．Shuteria の 1 種 S. hirsuta Baker (Fig. 1) は 分 類 染色体数の違いが明瞭であり，同属と見なすことはでき 上の位置が不安定で，クズ属 Pueraria とされたり，中 ない．Cologania はメキシコから中南米と南米に分布域 国 で は Amphicarpaea lineata と し て も 記 載 さ れ て い があり，インド〜中国の Shuteria とは分布上も著しく る．最近の Flora of China Vol. 10 Fabaceae でも Sa and 隔離している．このため，S. hirsuta に基づいて，新属 Gilbert (2010) は S. hirsuta の正名として誤った学名 S. Harashuteria K. Ohashi & H. Ohashi を設立した．また， ferruginea と A. lineata とを Shuteria として認めている． 新組み合わせ Harashuteria hirsuta (Baker) K. Ohashi & 一方，Ohashi and Ohashi (2016) は花粉形態に基づいて H. Ohashi を提案した．属名は原 寛先生 (1911–1986) S. hirsuta を Shuteria と 認 め，A. lineata を S. hirsuta の を記念した． 異名とした． （a岩手医科大学薬学部， 本研究は Shuteria hirsuta の分類上の不安定な位置を b東北大学植物園津田記念館） 検討するために S. hirsuta とその近縁種の分子系統解析