J. Jpn. Bot. 89: 1–11 (2014)

A New System of (Leguminosae Tribe )

a, b Hiroyoshi Ohashi * and Tomoyuki Nemoto

aHerbarium TUS, Botanical Garden, Tohoku University, Sendai, 980-0862 JAPAN; bDepartment of Biological Sciences, Faculty of Science and Engineering, Ishinomaki Senshu University, Ishinomaki, 986-8580 JAPAN *Corresponding author: [email protected]

(Accepted on August 29, 2013)

The genus Lespedeza (Leguminosae tribe Desmodieae) is distinct from Campylotropis and Kummerowia. Lespedeza includes 44 species in this paper. A new infrageneric system of classification of Lespedeza is proposed. The genus is divided into two subgenera: subgenus Lespedeza confined to North America, and subgenus Macrolespedeza to Asia, mainly East Asia. These subgenera differ only in seedling morphology, but the distinction is strongly supported by their distribution and results of current molecular analyses. The subgenus Lespedeza is subdivided into two sections, Lespedeza and Lespedezariae, and subgenus Macrolespedeza into two sections, Macrolespedeza and Junceae. A revised description of the genus, a key to the infrageneric taxa, an enumeration of the infrageneric taxa with their synonyms, the type of each name, and all the species included in each section are provided.

Key words: Desmodieae, , Junceae, Leguminosae, Lespedeza, Lespedezariae, Macrolespedeza, new classification, sections, subgenera.

The genus Lespedeza belongs to the subtribe of Maximowicz (1873) is broadest, but was Lespedezinae of the tribe Desmodieae with split by Schindler (1913) into three genera: Campylotropis and Kummerowia (Ohashi Lespedeza, Campylotropis and Kummerowia 2005). The genus has a controversial history which correspond to Maximowicz’s subgenera of circumscription in relation to the latter two Lespedeza, Campylotropis and Microlespedeza, genera. Lespedeza was established by Michaux respectively. However, Campylotropis was (1803) based on the North American species, L. included in Lespedeza (Gagnepain 1920, Nakai sessiliflora Michx. (the type selected by Britton 1927, Ohashi 1971) and Kummerowia was also and Brown 1913). The genus was expanded by regarded as Lespedeza (Isely 1948, Fernald Bentham by inclusion of Campylotropis Bunge 1950, Gleason 1963, Clewell 1966a, 1966b, (Bentham 1852) and Hedysarum striatum Ohashi 1982). The three genera in the sense (Thunb.) DC. (Bentham 1861). Maximowicz of Schindler (1913) have been accepted by (1873) first systematized Lespedeza and Hutchinson (1964), Ohashi et al. (1981) and included species of Campylotropis and Ohashi (2005). They have also been adopted Hedysarum striatum as members of subgenus in the floras of China (Lee 1995, Huang et al. Campylotropis and subgenus Microlespedeza, 2010), Indochina (Thuan 1987), Korea (Choi respectively. The genus Lespedeza in the sense 2007) and Taiwan (Huang and Ohashi 1977,

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Table 1. History of Lespedeza with its infrageneric taxa, Campylotropis and Kummerowia Maximowicz (1873) Schindler (1913) Nakai (1927, 1939)* Isely (1948), Fernald (1950), Hutchinson (1964) Clewell (1966), Ohashi (1982)** Lespedeza Lespedeza Lespedeza Lespedeza Lespedeza subg. Lespedeza sect. Eulespedeza [incl. Kummerowia] sect. Eu-Lespedeza sect. Macrolespedeza sect. Macrolespedeza ser. Junceae sect. Eulespedeza ser. Lespedezariae sect. Heterolespedeza ser. Pilosae ser. Violaceae sect. Macro-Lespedeza subg. Campylotropis Campylotropis sect. Campylotropis Campylotropis subg. Microlespedeza Kummerowia Kummerowia * Nakai separated Microlespedeza (= Kummerowia) from Lespedeza (cf. Nakai 1918). ** These authors did not treat Campylotropis. *** Ohashi regarded Campylotropis as distinct (cf. Iokawa and Ohashi 1997).

1993). Kummerowia is currently recognized as was supported by pollen morphology (Xu et al. distinct from Lespedeza in the floras of North 2011). America (Isely 1990, 1998, Haines 2011) (Table Lespedeza is regarded as a subject interesting 1). in molecular phylogenetic studies because of The first infrageneric taxon of Lespedeza its remarkable disjunctive distribution between was proposed by Torrey and Gray (1840) eastern North America and Asia, and the who distinguished a section Lespedezariae (as diversity in opinions for its taxonomic concept ‘Lespedezaria’) among the North American as outlined above. Nemoto et al. (1995) first representatives of the genus. Maximowicz clarified phylogenetic relationships ofLespedeza (1873) divided world Lespedeza (as subgen. with Campylotropis and Kummerowia based Lespedeza) into two sections: Eu-Lespedeza on morphological and anatomical studies and Macro-Lespedeza. He separated section and analyses of RFLPs of cpDNA. Recently, Lespedeza (as Eu-Lespedeza) into four series: phylogenetic relationships among the species of Junceae, Pilosae, Violaceae and Lespedezariae, the genus were intensively analyzed in Japan, of which Violaceae included the type of the Korea and China (Nemoto et al. 2010, Han et al. section and Lespedezariae expanded to include 2010, Xu et al. 2012). These molecular analyses some Asian representatives. Schindler (1913) showed Lespedeza as sister to Campylotropis adopted Maximowicz’s sectional divisions and Kummerowia and the three genera are and recognized two sections: Lespedeza in distinct from each other. Consequently, the Asia and North America and Macrolespedeza results support recent taxonomic recognition of in Asia (Table 1). Nakai (1939) described the three genera (Table 1). a new section Heterolespedeza separated The current molecular analyses indicate, from Macrolespedeza. Akiyama and Ohba however, discrepancy from previous infrageneric (1988) divided section Macrolespedeza into classification systems of Lespedeza. The series Macrolespedeza, Heterolespedeza and analyses suggested that “earlier divergence had Formosae. Ohashi (2001) and Ohashi et al. occurred in Asia resulting into two lineages in (2009a, b) raised the sections Lespedeza and Lespedeza, and then one of them remained in Macrolespedeza in the sense of Schindler to Asia and the other migrated to North America. the rank of subgenera. The subgeneric division After the isolation of both lineages speciation February 2014 The Journal of Japanese Botany Vol. 89 No. 1 3

Table 1. Continued Ohashi (1971) Isely (1990, Nemoto et al. Ohashi (2001)*** Ohashi (2005) Present work 1998)** (1995)

Lespedeza Lespedeza Lespedeza Lespedeza Lespedeza Lespedeza [incl. Campylotropis] subg. Lespedeza subg. Lespedeza subg. Macrolespedeza sect. Lespedeza sect. Macrolespedeza sect. Lespedezariae sect. Heteroloespedeza subg. Macrolespedeza sect. Macrolespedeza sect. Junceae

Campylotropis Campylotropis Campylotropis Kummerowia Kummerowia Kummerowia Kummerowia Kummerowia Kummerowia

had occurred independently in Asia and North basis of their distichously arranged winter bud America” (Nemoto et al. 2010). Therefore, the scales. However, in this study we exclude the genus Lespedeza should be divided first into two species from Macrolespedeza circumscribed two groups: Asian species and North American by Ohashi et al. (2009b) in accordance with the species. results of Xu et al. (2012). The remaining species The North American species group is clearly of Macrolespedeza is distinct from members separated by the analyses into two subgroups of Lespedeza of the Asian species group in that correspond to the sections Lespedeza and morphology and molecular analyses. Lespedezariae (Torrey and Gray 1840), which Members of subgenus Lespedeza of the are traditionally distinguished by floral color and Asian species group appear to form at least two shape in floras of North America. clades, but they are still insufficiently related The Asian species group is revised from the in morphology and molecular analyses. For previous interpretation classifying the group example, “clade C” obtained from cpDNA in into two subgenera Lespedeza and subgen. Nemoto et al. (2010) consisting of Lespedeza Macrolespedeza (Ohashi 2001, Table 1). chinensis, L. floribunda and L. virgata is similar Members of subgenus Macrolespedeza to a clade obtained from ITS including these constitute a distinct clade among the group in three species and L. tomentosa in Han et al. the results of Nemoto et al. (2010) and Han et al. (2010, Fig. 3). Xu et al. (2012, Fig. 1) obtained (2010), but Xu et al. (2012) showed two separate a similar clade, called “clade D”, composed of clades among Macrolespedeza, of which one these four species with L. dunnii, L. fordii and L. included many species common with other fasciculiflora var. hengduanshanensis from ITS, analyses and another includes Lespedeza dunnii but the last variety was excluded by cpDNA Schindl. and L. fordii Schindl. These two species from the “clade D”. Lespedeza fasciculiflora endemic to China were analyzed only by Xu (corresponding to var. fasciculiflora) was not et al. (2012) and their clade was included in a included in the “clade C” and the position was clade with some members of subgen. Lespedeza. uncertain in Nemoto et al. (2010). Xu et al. Ohashi et al. (2009b) examined herbarium (2012) placed L. fasciculiflora var. fasciculiflora specimens of these species in detail. Both in a different clade with L. caraganae, L. cuneata species are placed in subgen. Macrolespedeza and L. lichiyuniae based on ITS. Lespedeza in lacking cleistogamous flowers and are tomentosa was placed in a clade by cpDNA with accommodated in sect. Heterolespedeza on the L. cuneata, L. daurica, L. forrestii, L. inschanica 4 植物研究雑誌 第 89 巻 第 1 号 2014 年 2 月 and L. juncea in Nemoto et al. (2010), Han E), while those of subgenus Macrolespedeza et al. (2010) and Xu et al. (2012). Han et al. are characterized by first two leaves opposite (2010) found a similar clade using ITS with L. (Figs. 1F–J, 2). These two patterns of seedlings caraganae, L. lichiyuniae and L. pilosa but not correspond to the disjunctive distribution and L. tomentosa. Xu et al. (2012) also obtained the separation indicated by the molecular analyses similar clade of these species in addition with L. between the subgenera. The subgenus Lespedeza fasciculiflora var. fasciculiflora, L. jiangxiensis, is subdivided into two sections: Lespedeza and L. hispida and L. potaninii from analysis of ITS Lespedezariae, and the subgenus Macrolespedea sequence data. Diversification of Asian species into two sections: Macrolespedea and Junceae. of subgenus Lespedeza is more conspicuous in A revised description of the genus Lespedeza, molecular phylogeny than those of subgenus a key to the infrageneric taxa, an enumeration of Macrlespedeza, but critical relationships are the infrageneric taxa with their synonyms and still uncertain because of the incongruence the type of each name, and all the species with between the cpDNA and ITS phylogenies. their subspecies included in each section are Here, we accommodated all members of provided. subgenus Lespedeza of the Asian species group in a same section, Junceae, distinct from those The genus Lespedeza with its bibliography Macrolespedeza. Lespedeza dunnii and L. fordii and a description are included in section Junceae. Lespedeza Michx., Fl. Bor.-Amer. 2: 70 Species distributed in Indian Subcontinent (1803); Benth. in Benth. & Hook. f., Gen. Pl. and Western Asia (Afghanistan), belonging 1: 524 (1865), p. p., excl. gen. Campylotropis; to the subgenus Lespedeza, have not yet been Maxim. in Trudy Imp. S.-Peterburgsk. Bot. Sada investigated molecular phylogenetically. 2: 345 (1873), p. p., excl. subg. Campylotropis They are Lespedeza aitchisonii Ricker, L. & subg. Microlespedeza; Britton in Trans. N.Y. elegans Camb., L. juncea f. pseudovariegata Acad. Sci. 12: 57 (1893); Britton & Brown, Pramanik & Thoth., L. kanaoriensis Camb., L. Ill. Fl. N. U.S. ed. 2. 2: 402 (1913); Schindl. nuristanica Rech. f. and L. variegata Camb. in Bot. Jahr. Syst. 49(5): 570 (1913); Nakai, Morphologically these species are regarded as Lespedeza Jap. Korea: 1 (1927), p. p., excl. sect. similar to L. juncea in having cleistogamous Campylotropis; Fernald, Gray’s Manual Bot. flowers and habit of subshrubs or herbs. We ed. 8: 923 (1950); Ohwi, Fl. Jap.: 677 (1953); also accommodated them in the section Junceae Hutch., Gen. Fl. Pl. 1: 487 (1964); Clewell in proposed below in this paper. Rhodora 68: 365 (1966); H. Ohashi, Polhill Thus, the previous infrageneric systems & Schubert in Polhill & Raven, Adv. Leg. needs to be revised in harmony with the results Syst. 1: 300 (1981); Isely, Vasc. Fl. S.E. U.S., of the molecular analyses. We propose a new Leguminosae: 57 (1990); T. Nemoto & H. infrageneric classification system of the genus Ohashi in J. Pl. Res. 106: 121 (1993) [seedlings]; Lespedeza in this paper. We recognize here T. C. Huang & H. Ohashi in Fl. Taiwan ed. 2, 3: the North American group of the species as 317 (1993); Li & Chen in Fl. Reip. Pop. Sin. 41: subgenus Lespedeza and the Asian one as 131 (1995); Isely, Native Natural. Legum. U.S.: subgenus Macrolespedea. The distinction 624 (1998); H. Ohashi in Sci. Rep. Tohoku Univ. between the two subgenera is in seedling ser. 4 (Biol.), 40(3): 238 (1999); H. Ohashi in K. morphology (Nemoto and Ohashi 1993, Iwats. & al., Fl. Jap. IIb: 259 (2001); H. Ohashi Nemoto et al. 2010). Seedlings of subgenus in Lewis & al., Leg. World: 435 (2005); P. H. Lespedeza are characterized by first two leaves Huang, H. Ohashi & T. Nemoto in Fl. China 10: subsequent to cotyledons alternate (Fig. 1A– 302 (2010). February 2014 The Journal of Japanese Botany Vol. 89 No. 1 5

Type (designated by Britton and Brown in proximal fascicles, bracteolate, enclosed in 1913): Lespedeza sessiliflora Michaux, nom. calyx; corolla reduced in size, stamens reduced illeg. (Medicago virginica L., L. virginica (L.) in size and number. Fruits loments, compressed, Britton). unilocular, 1-seeded, indehiscent, pubescent, Perennial herbs or shrubs, unarmed, without villose or glabrescent; chasmogamous loments uncinate indumentum. Rootstocks woody. (oriented from chasmogamous flowers) Stems erect, ascending or prostrate, often usually stipitate, usually elliptic-ovate or woody near base in herbs, usually branched. suborbicular, style straight; cleistogamous Leaves alternate, pinnately trifoliolate, petiolate. loments (oriented from cleistogamous flowers) Stipules persistent, free, subulate or narrowly usually crowded at base of peduncle, sessile, deltate, acuminate, striate-nerved, ciliate. obovate to suborbicular, slightly smaller than Petioles pulvinate, distinct in medial leaves but chasmogamous one, with a curved short style. decreasing gradually to nearly sessile in distal Seeds 1, asymmetrical, generally elliptic or leaves. Leaflets estipellate, ovate to elliptic, oblong, rim-arillate, chasmogamous seeds narrowly elliptic, narrowly oblong, narrowly slightly longer or similar to cleistogamous one. ovate or narrowly obovate, entire, apex usually Seedlings: cotyledons 2, epigeous, opposite, apiculate, abaxial surface generally pale green, green, sessile, estipulate, oblong, 5–13 mm principle lateral veins running upward within long, glabrous, apex rounded, base rounded; margin, ciliate; terminal leaflet petiolulate, leaves subsequent cotyledons 1-foliolate, single pulvinate, usually larger than lateral ones in or paired at the first node, stipulate, petiolate, size and shape; lateral leaflets pulvinate, sessile pulvinate at both ends of petiole, leaf-blade or subsessile, often more or less oblique. reticulate nerved, lower surface puberulent; Inflorescences racemes (strictly pseudoracemes), leaves at and after the second node 3-foliolate rarely capitate; axillary, sometimes seemingly or rarely 1-foliolate, alternate. Chromosome terminal in compound racemes (panicles) numbers x = 9, 11; 2n = 18, 22, 36, 40, 42, and by reduction of their subtending leaves; ca. 44. primary bract 1, subtending (1–)2 flowers; Species included: 44, listed under each secondary ones 2, subtending a flower. Flowers section below. Ca. 43 natural hybrids (33 hybrids chasmogamous or cleistogamous, pedicellate in subgen. Lespedeza and ca. 10 in subgen. or subsessile; bracteolate. Chasmogamous Macrolespedeza). flowers papilionaceous; calyx campanulate, Distribution: Asia (mainly China to Japan, 5-lobed, lobes longer than tube, adaxial 2 free few extending to India to Afghanistan and to or proximally more or less connate with apex Malesia) and eastern N. America (U.S.A. to 2-toothed; corolla pink-purple or white to pale Canada). Subgenus Lespedeza is confined in yellow; standard broadly obovate to orbicular, North America, while subgenus Macrolespedeza proximally short clawed or cuneate, with in Asia and Malesia. inflexed auricles, darker purple marks (nectar guides) at throat of adaxial surface of lamina; Key to the infrageneric taxa of Lespedeza wings and keel petals long clawed, lamina 1. native to North America; first two elliptic-oblong, proximally rounded. Stamens leaves subsequent to cotyledons alternate in diadelphous, anthers uniform. Disk present seedling ...... 2 [subgen. Lespedeza] around base of ovary. Ovary 1-ovulate, stipitate 1. Plants native to Asia; first two leaves (less than 1 mm, rarely to 2 mm); style adaxially subsequent to cotyledons opposite in incurved, slightly exserted from stamens; stigma seedling ...... 3 [subgen. Macrolespedeza] minute, terminal. Cleistogamous flowers often 2. Corolla pink-purple; calyx 4-lobed (adaxial 6 植物研究雑誌 第 89 巻 第 1 号 2014 年 2 月

Fig. 1. Seedlings of Lespedeza subgen. Lespedeza sect. Lespedeza (A–C), sect. Lespedezariae (D, E) and subgen. Macrolespedeza sect. Macrolespedeza (F–J). A. L. frutescens. B. L. procumbens. C. L. violacea. D. L. capitata. E. L. hirta. F. L. bicolor. G. L. buergeri. H. L. cyrtobotrya. I. L. formosa subsp. formosa. J. L. formosa subsp. patens. Horizontal lines (g) under cotyledons show ground level. co, cotyledon; co’, scar of fallen cotyledon; l1, first unifoliolate leaf (A–E) and fi rst two unifoliolate leaves opposite (F–J) at fi rst node subsequent to cotyledons;l 2, second leaf subsequent to fi rst one (A–E) at second node; l3, third leaf subsequent to fi rst two opposite ones (F–J) at second node. Scale bar = 1 cm. B, E and I reproduced from Nemoto and Ohashi (1993) with permission from The Botanical Society of Japan and Springer Japan. Drawing by T. Nemoto. February 2014 The Journal of Japanese Botany Vol. 89 No. 1 7

Fig. 2. Seedlings of Lespedeza subgen. Macrolespedeza sect. Junceae. A. L. caraganae. B. L. chinensis. C. L. cuneata. D. L. daurica. E. L. fl oribunda. F. L. inschanica. G. L. juncea. H. L. pilosa. I. L. tomentosa. J. L. virgate. Horizontal lines (g) under cotyledons show ground level. co, cotyledon; co’, scar of fallen cotyledon; l1, fi rst two unifoliolate leaves opposite at fi rst node subsequent to cotyledons; l3, third leaf subsequent to fi rst two opposite ones at second node. Scale bar = 1 cm. E and F reproduced from Nemoto and Ohashi (1993) with permission from The Botanical Society of Japan and Springer Japan. Drawing by T. Nemoto. 8 植物研究雑誌 第 89 巻 第 1 号 2014 年 2 月

lobes proximally connate), less than half as Lespedeza ser. Lespedezariae (Torr. & A. long as flower and loment; bracteoles shorter Gray) Maxim. in Trudy Imp. S.-Peterburgsk. than calyx-tube; racemes 4–14-flowered.sect. Bot. Sada 2: 346 & 372 (1873), p.p., incl. L. Lespedeza capitata, L. hirta, cet. excl. 2. Corolla white to pale yellow; calyx 5-lobed, Species included (4 spp.): Lespedeza nearly equaling or exceeding corolla and angustifolia (Pursh) Elliott, L. capitata Michx., loment; bracteoles longer than calyx-tube; L. hirta (L.) Hornem. (subsp. hirta) [with L. hirta racemes 10–40-flowered subsp. curtissii Clewell] and L. leptostachya ...... sect. Lespedezariae Engelm. 3. Flowers only chasmogamous, usually 10–16 mm long; loments stipitate; shrubs or herbs, erect or 2. Subgen. Macrolespedeza (Maxim.) H. ascending, to 3 m tall...... sect. Macrolespedeza Ohashi in J. Jap. Bot. 57: 29 (1982); H. Ohashi 3. Flowers cleistogamous and chasmogamous in K. Iwats. & al., Fl. Jap. IIb: 260 (2001); H. (except L. dunnii, L. fordii and L. forrestii Ohashi, T. Nemoto & K. Ohashi in J. Jpn. Bot. with only chasmogamous flowers), usually 84(4): 199 (2009); P. H. Huang, H. Ohashi & T. to 10 mm long (except L. gerardiana 11-13 Nemoto, Fl. China 10: 302 (2010). mm long, fide Ohashi et al. 2009); loments 2-1. sect. Macrolespedeza Maxim. in Trudy sessile or subsessile; subshrubs or herbs, Imp. S.-Peterburgsk. Bot. Sada 2: 346 & 359 erect, ascending or prostrate, to 1.5 m tall ..... (1873), as “Macro-Lespedeza”; Schindl. in Bot...... sect. Junceae Jahrb. Syst. 49(5): 574 (1913); Nakai, Lespedeza Jap. Kor.: 7 (1927), ut “sect. Macrolespedeza Enumeration of the infrageneric taxa and Nakai”; Nakai in J. Jap. Bot. 15(9): 531 (1939), species (with subspecies) of the genus ut “sect. Macrolespedeza (Maxim.) Nakai”; Lespedeza Akiyama in Univ. Mus. Univ. Tokyo Bull. no. 1. Subgen. Lespedeza [autonym newly 33: 91 (1988); Li & Chen in Fl. Reip. Pop. Sin. circumscribed here incuding only American 41: 131 (1995). Type (chosen by Nakai, l. c. species] 15(9): 531. 1939): Lespedeza bicolor Turcz. 1-1. sect. Lespedeza. Type: Lespedeza Lespedeza sect. Heterolespedeza Nakai in J. virginica (L.) Britt. Jap. Bot. 15(9): 531 (1939) in nota; H. Ohashi, Lespedeza ser. Violaceae Maxim. in Trudy Nemoto & K. Ohashi in J. Jpn. Bot. 84(4): 200 Imp. S.-Peterburgsk. Bot. Sada 2: 346 & (2009). Type: Lespedeza buergeri Miq. 359 (1873), p.p., incl. Lespedeza repens, L. L. ser. Heterolespedeza (Nakai) S. Akiyama reticulata, L. stuevei (as “stuvei”), L. violacea, & H. Ohba in Univ. Mus. Univ. Tokyo Bull. no. cet. excl. Type (designated here): Lespedeza 33: 142 (1988). violacea (L.) Pers. L. ser. Formosae S. Akiyama & H. Ohba Species included (7 spp.): Lespedeza in Univ. Mus. Univ. Tokyo Bull. no. 33: 116 frutescens (L.) Hornem., L. procumbens Michx., (1988). Type: Lespedeza formosa (Vogel) L. repens (L.) W. P. C. Barton, L. stuevei Nutt., Koehne. L. texana Britt., L. violacea (L.) Pers. and L. Species included (9 spp.): Lespedeza bicolor virginica (L.) Britt. Turcz., L. buergeri Miq., L. cyrtobotrya Miq., L. davidii Franch., L. homoloba Nakai, L. maritima 1-2. sect. Lespedezariae Torr. & A. Gray, Nakai, L. maximowiczii C. K. Schneid., L. Fl. N. Amer. 1(3): 368 (1840), “Lespedezaria”. melanantha Nakai and L. thunbergii (DC.) Type (designated here): Lespedeza hirta (L.) Nakai [with subsp. elliptica (Benth. ex Maxim.) Hornem. H. Ohashi, subsp. formosa (Vogel) H. Ohashi, February 2014 The Journal of Japanese Botany Vol. 89 No. 1 9 subsp. patens (Nakai) H. Ohashi and subsp. Plantae Junghuhnianae, pp. 205–269. Sythoff. Leiden. satsumensis (Nakai) H. Ohashi]. Bentham G. 1861. Flora Hongkongensis. li+482 pp. Reeve & Co., London. Britton N. L. and Brown A. 1913. An Illustrated Flora 2-2. sect. Junceae (Maxim.) H. Ohashi & T. of the Northern United States, Canada and British Nemoto, stat. nov. Possessions. ed. 2. rev. and enl. C. Scribner, New York. Lespedeza ser. Junceae Maxim. in Trudy Choi B. H. 2007. Fabaceae. In: Flora of Korea Editorial Imp. S.-Peterburgsk. Bot. Sada 2: 346 & 367 Committee, The Genera of Vascular Plants of Korea, pp. 585–622. Academy Publishing Co., Seoul. (1873). Type (designated here): Lespedeza Clewell A. F. 1966a. Native North American species of juncea (L. f.) Pers. Lespedeza (Leguminosae). Rhodora 68: 359–405. L. ser. Pilosa Maxim. in Trudy Imp. S.- Clewell A. F. 1966b. I. Identification of the of Peterburgsk. Bot. Sada 2: 346 & 381 (1873). North America. II. Selected bibliography of Lespedeza. Bull. Tall Timbers Res. Sta. No. 7. 29 pp. Type: Lespedeza pilosa (L. f.) Pers. Fernald M. L. 1950. Lespedeza. In: Fernald M. L., Gray's L. sect. Lespedeza, p. p., incl. Asian spp., Manual of Botany, ed. 8, pp. 923–927. American Book excl. American spp.: Nakai, Lespedeza Jap. Kor. Company, New York. 73 (1927), “sect. Eulespedeza Nakai”; Li & Gagnepain F. 1920. Papilionées. In: Lecomte H. (ed.), Chen, Fl. Reip. Pop. Sin. 41: 145 (1995). Flore Générale de lʼIndo-Chine 2: 505–613. Mus. Nat. d'Hist. Natur. Phanérogamie, Paris. L. subgen. Lespedeza, p. p., incl. Asian spp: Haines A. 2011. New England Wildflower Society's Flora H. Ohashi in Iwatsuki & al., Fl. Jap. IIb: 259 in Novae Angliae. 973 pp. Yale University Press, New key & 263 (2001); H. Ohashi, T. Nemoto & K. Haven. Ohashi in J. Jpn. Bot. 84(4): 145 (2009); P. H. Han J.-E., Chung K.-H., Nemoto T. and Choi B.-H. 2010. Phylogenetic analysis of eastern Asian and eastern Huang, H. Ohashi & T. Nemoto, Fl. China 10: North American disjunct Lespedeza. Bot. J. Linn. Soc. 303 (2010). 164: 221–235. Species belonging (24 spp.): Lespedeza Huang P. H., Ohashi H. and Nemoto T. 2010. Lespedeza. aitchisonii Ricker, L. caraganae Bunge, L. In: Wu Z. Y., Raven P. H. and Hong D. Y. (eds.), Flora chinensis G. Don, L. cuneata (Dum.Cours.) G. of China 10, Fabaceae, pp. 302–311. Science Press, Beijing and Missouri Botanical Garden Press, St. Don, L. daurica (Laxm.) Schindl., L. dunnii Louis. Schindl., L. elegans Cambess., L. fasciculiflora Huang T. C. and Ohashi H. 1977. Leguminosae. In: Franch., L. floribunda Bunge, L. fordii Schindl., Editorial Committee of the Flora of Taiwan (ed.), Flora L. forrestii Schindl., L. gerardiana Wall. ex of Taiwan 3: 148–421. Epoch Publishing Co., Ltd., Taipei. Maxim., L. hisauchii T. Nemoto & H. Ohashi, Huang T. C. and Ohashi H. 1993. Leguminosae. In: L. hispida (Franch.) T. Nemoto & H. Ohashi, L. Editorial Committee of the Flora of Taiwan, Second inschanica (Maxim.) Schindl., L. jiangxiensis Bo Edition (ed.), Flora of Taiwan, ed. 2, 3: 160–396. Xu, X. F. Gao & Li Bing Zhang, L. juncea (L. Department of Botany, National Taiwan University, f.) Pers., L. kanaoriensis Camb., L. lichiyuniae Taipei. Hutchinson J. 1964. Leguminosae. The Genera of T. Nemoto, H. Ohashi & T. Itoh, L. nuristanica Flowering Plants 1: 221–489. Oxford University Press, Rech. f., L. pilosa (Thunb.) Siebold & Zucc., London. L. tomentosa (Thunb.) Siebold ex Maxim., L. Iokawa Y. and Ohashi H. 1997. Two new taxa of variegate Camb. and L. virgata (Thunb.) DC. Campylotropis (Leguminosae) from China. J. Jpn. Bot. 72: 139–143. Isely D. 1948. Lespedeza striata and L. stipulacea. Rhodora We thank Dr. Kazuaki Ohashi of Iwate 50: 21–27. Medical University for his reading and Isely D. 1990. Vascular Flora of the Southeastern United comments on a preliminary manuscript. States. 3(2) Leguminosae (Fabaceae). The University of North Carolina Press, Chapel Hill. Isely D. 1998. Native and Naturalized Leguminosae References (Fabaceae) of the United States exclusive of Alaska Bentham G. 1852. Leguminosae. In: Miquel F. A. W., and Hawaii. Monte L. Bean Life Science Museum, 10 植物研究雑誌 第 89 巻 第 1 号 2014 年 2 月

Brigham Young University, Provo. Michx. und ihre nächsten Verwandten. Bot. Jahrb. Syst. Maximowicz C. J. 1873. Synopsis Generis Lespedezae, 49: 570–658. Michaux. Trudy Imp. S.-Peterburgsk. Bot. Sada 2: Thuân N. van 1987. Kummerowia, Lespedeza, 329–388. Campylotropis. In: Thuân N. van, Dy Phon P., Michaux A. 1803. Flora Boreali-Americana 1. Paris, Niyomdham C. et Vidal Y. 1987. Flore du Cambodge Strasbourg. du Laos et du Viêtnam 23, Légumineuses– Nakai T. 1918. Report on the Vegetation of Diamond Papilionoidées, pp. 138–148. Muséum National Mountains, Corea. The Government of Chosen, Seoul. d'Histoire Naturelle Laboratoire du Phanérogamie. Nakai T. 1927. Lespedeza of Japan and Korea. The Forest Paris. Experiment Station of Government General of Chosen, Torrey J. and Gray A. 1840. A Flora of North America 1. Keijo. Wiley & Putnam, New York. Nakai T. 1939. Notulae ad plantas Asiae orientalis (IX). J. Xu B., Gao X. F., Wu N. and Zhang L. B. 2011. Pollen Jap. Bot. 15: 523–541. diversity and its systematic implications in Lespedeza Nemoto T. and Ohashi H. 1993. Seedling morphology of (Fabaceae). Syst. Bot. 36(2): 352–361. Lespedeza (Leguminosae). J. Pl. Res. 106: 121–128. Xu B., Gao X. F. and Zhang L. B. 2013. Lespedeza Nemoto T., Ohashi H. and Tamate H. 1995. Phylogeny jiangxiensis, sp. nov. (Fabaceae) from China based on on Lespedeza and its allied genera (Desmodieae- molecular and morphological data. Syst. Bot. 38(1): Lespedezinae). In: Crisp M. and Doyle J. J. (eds.), 118–126. Advances in Legume Systematics 7, Phylogeny, pp. Xu B., Wu N., Gao X. F. and Zhang L. B. 2012. Analysis 351–358. Royal Botanic Gardens, Kew. of DNA sequences of six chloroplast and nuclear genes Nemoto T., Yokoyama J., Fukuda T., Iokawa Y. and Ohashi suggests incongruence, introgression, and incomplete H. 2010. Phylogeny of Lespedeza (Leguminosae) lineage sorting in the evolution of Lespedeza based on chloroplast trnL-trn F sequences. J. Jpn. Bot. (Fabaceae). Mol. Phyl. Evol. 62: 346–358. 85: 213–229. Ohashi H. 1971. A taxonomic study of the tribe Coronilleae Appendix. (Leguminosae), with a special reference to pollen Voucher specimens (all in TUS) of seedlings in Figs. 1 morphology. J. Fac. Sci. Univ. Tokyo Sect. III (Bot.) and 2. 11: 25–92. Subgen. Lespedeza sect. Lespedeza Ohashi H. 1982. Leguminosae. In: Satake Y., Ohwi J., Lespedeza frutescens (L. ) Hornem.: T. Nemoto 5873, seeds Kitamura S., Watari S. and Tominari T. (eds.), Wild from the cultivated in Tohoku Univ. grown from Flowers of Japan, Herbaceous Plants (including Dwarf seed from the specimen (U.S.A., Indiana, Brown Co., A. F. Subshrubs) 2: 186–212, pls. 179–200. Heibonsha Ltd., Clewell 1570, TUS). Publishers. Tokyo (in Japanese). L. procumbens Michx.: T. Nemoto 8590, seeds from Ozark Ohashi H. 2001. Leguminosae. In: Iwatsuki K., Boufford National Forest, Johnson Co., Arkansass, U.S.A. D. E. and Ohba H. (eds.), Flora of Japan IIb: 213–279. L violacea (L.) Pers.: T. Nemoto 5872, seeds from the Kodansha Ltd., Tokyo. plant cultivated in Tohoku Univ. grown from seed from the Ohashi H. 2005. Tribe Desmodieae. In: Lewis G. P., Schrire specimen (U.S.A., Indiana, Brown Co., A. F. Clewell s.n., B., Mackinder B. and Lock M. (eds.), Legumes of the 28 Sep. 1959, TUS). World, pp. 433–445. Royal Botanic Gardens, Kew. Ohashi H., Nemoto T. and Ohashi K. 2009a. A revision Subgen. Lespedeza sect. Lespedezariae of Lespedeza subgenus Lespedeza (Leguminosae) of Lespedeza capitata Michx.: T. Nemoto 5869, seeds from China. J. Jpn. Bot. 84: 143–166. the plant cultivated in Tohoku Univ. grown from seeds Ohashi H., Nemoto T. and Ohashi K. 2009b. A revision of from Ontario, Canada. Lespedeza subgenus Macrolespedeza (Leguminosae) L. hirta (L.) Hornem. subsp. hirta: T. Nemoto 8589, seeds in China. J. Jpn. Bot. 84: 197–223. from Western Lake, Bay Co., Florida, U.S.A. Ohashi H., Polhill R. M. and Schubert B. G. 1981. Desmodieae. In: Polhill R. M. and Raven P. H. (eds.), Subgen. Macrolespedeza sect. Macrolespedeza Advances in Legume Systematics 1: 292–300. Royal Lespedeza bicolor Turcz.: Seeds collected by T. Nemoto Botanic Gardens, Kew. from Shiroishi-shi, Miyagi Pref., Japan (Nov. 29, 1984). Schindler A. K. 1912a. Kummerowia Schindler novum L. buergeri Miq.: T. Nemoto 2414, seeds from Hayama- genus Leguminosarum. Repert. Spec. Nov. Regni Veg. machi, Kanagawa Pref., Japan. 10: 403–404. L. cyrtobotrya Miq.: T. Nemoto 2419, seeds from Shiroishi- Schindler A. K. 1912b. Das Genus Campylotropis. Repert. shi, Miyagi Pref., Japan. Spec. Nov. Regni Veg. 11: 338–347; 424–431. L. thunbergii (DC.) Nakai subsp. formosa (Vogel) H. Schindler A. K. 1913. Einige Bemerkungen über Lespedeza Ohashi: T. Nemoto 5852, seeds from Lishan, Taichung Co., February 2014 The Journal of Japanese Botany Vol. 89 No. 1 11

Taiwan. L. floribunda Bunge: T. Nemoto 2413, seeds from Beijing L. thunbergii subsp. patens (Nakai) H. Ohashi: T. Nemoto Botanical Garden, Institute of Botany, CAS, China. 4234, seeds from Itoigawa-shi, Niigata Pref., Japan. L. inschanica (Maxim.) Schindl.: T. Nemoto 8587, seeds from plant cultivated in Tohoku Univ., originated from Subgen. Macrolespedeza sect. Junceae Tong-myeon, Chonwon-gun, Chungchongnam Prov., Lespedeza caraganae Bunge: T. Nemoto 2408, seeds from Korea. Beijing Botanical Garden, Institute of Botany, CAS, China. L. juncea (L. f.) Pers.: T. Nemoto 2400, seeds from Sendai- L. chinensis G. Don: T. Nemoto 2404, seeds from the plant shi, Miyagi Pref., Japan. cultivated in Tohoku Univ., originated from Tailukohsia, L. pilosa (Thunb.) Siebold & Zucc.: T. Nemoto 2405, seeds Hualien Co., Taiwan. from Sendai-shi, Miyagi Pref., Japan. L. cuneata (Dum.Cours.) G. Don: T. Nemoto 2399, seeds L. tomentosa (Thunb.) Siebold ex Maxim.: T. Nemoto from Sendai-shi, Miyagi Pref., Japan. 2411, seeds from Kitaazumi-gun, Nagano Pref., Japan. L. daurica (Laxm.) Schindl.: T. Nemoto 2401, seeds from L. virgata (Thunb.) DC.: T. Nemoto 2406, seeds from Beijing Botanical Garden, Institute of Botany, CAS, China. Takahashi-shi, Okayama Pref., Japan.

a b 大橋広好 ，根本智行 ：マメ科ハギ属の新分類体系 ハ ギ 属 は 1803 年 に Michaux (1803) に よ っ て 北 は両地域で独自に分化したと推測された．ハギ属の系統 ア メ リ カ 産 の 種 に 基 づ い て 設 立 さ れ，1873 年 に 関係から見ると，これまでハギ属をアジアのヤマハギ亜 Maximowicz (1873) に よ っ て 初 め て 体 系 化 さ れ た． 属とアジア・北アメリカのハギ亜属に分けていたが，こ Maximowicz は今日の分類では別属とされるハナハギ の分類体系は系統を反映していないことが明らかとな 属とヤハズソウ属もハギ属に含めて，ハギ属を広く範 った． 囲づけた．その後 Maximowicz のハギ属の範囲付け そこで本研究ではハギ属の記載を改めると共に，最 について Schindler (1913), Nakai (1927), Isley (1948), 近の系統関係を反映した次のような新しい分類体系を Hutchinson (1964), Ohashi (1971, 2001, 2005), Ohashi 提案した．ハギ属は現在のところ 44 種よりなる．また et al. (1981) などによる意見の変遷があり，今日ではハ 約 43 雑種が記録されている．まずハギ属を北アメリカ ギ属（狭義），ハナハギ属，ヤハズソウ属に分けられる 産のハギ亜属 subgenus Lespedeza とアジア産のヤマハ ようになった（Table 1）．これらの属はヌスビトハギ連 ギ亜属 subgenus Macrolespedeza (Maxim.) H. Ohashi ハギ亜連としてまとめられている． に分けた．ハギ亜属は芽生えの第 1 節で葉が互生（Fig. ハギ亜連では属の範囲付けに異説があることから、分 1A–E）するが，一方のヤマハギ亜属は対生する（Fig. 子系統解析研究の格好のテーマと考えられた．Nemoto 1F–J お よ び Fig. 2）．次にハギ亜属をハギ節 section et al. (1995) はハギ属，ハナハギ属，ヤハズソウ属の系 Lespedeza（ 7 種）と section Lespedezariae Torr. & A. 統関係を葉緑体 DNA の解析によって初めて明らかにし Gray（ 4 種）とに細分した．ヤマハギ亜属はヤマハギ節 た．さらに，最近ではハギ属はアジアと北アメリカに section Macrolespedeza Maxim.（ 9 種）と新節シベリ 隔離分布することで種間の系統関係が注目された．そ アメドハギ節 section Junceae (Maxim.) H. Ohashi & T. の成果が日本 (Nemoto et al. 2010)，韓国・日本 (Han et Nemoto（ 24 種）の 2 節に分けた．括弧内は各節に属す al. 2010)，中国 (Xu et al. 2012) でそれぞれ発表された． る種数を示した．それらの種名と亜種名とは本文中に挙 その結果からハギ属，ハナハギ属，ヤハズソウ属はそれ げた． ぞれ独立属とする見解が支持された．また，ハギ属はア （a 東北大学植物園津田記念館， ジアに起源し，その一部が北アメリカに分布し，その後 b 石巻専修大学理工学部生物科学科）