J. Jpn. Bot. 88: 166–175 (2013)

New Combinations in North American (Leguminosae: Tribe Desmodieae)

Hiroyoshi Ohashi

Herbarium, Botanical Garden, Tohoku University, Sendai, 980-0862 JAPAN E-mail: [email protected]

(Accepted on May 11, 2013)

Three species groups in Desmodium of North America: D. ciliare group, D. paniculatum group and D. procumbens group are revised for Flora of North America. Each of them is treated as a single species containing one or two varieties. The following four new combinations are published: D. marilandicum var. ciliare (Willd.) H. Ohashi, D. marilandicum var. lancifolium (Fernald & B. G. Schub.) H. Ohashi, D. paniculatum var. fernaldii (B. G. Schub.) H. Ohashi, and D. procumbens var. neomexicanum (A. Gray) H. Ohashi. Desmodium obtusum (Willd.) DC. is regarded to be a synonym of D. marilandicum var. lancifolium; D. glabellum (Michx.) DC. and D. perplexum B.G. Schub. to be synonyms of D. paniculatum; and D. procumbens var. exiguum (A. Gray) B. G. Schub. to be a synonym of D. procumbens var. procumbens.

Key words: Desmodium, , Leguminosae, new combinations, North America.

Recently, Desmodium of North America can at least identify it with the group.” Under has been the subject of much study, notably by this framework, Isely recognized D. ciliare Schubert (1940, 1950a, 1950b, 1970) and Isely (Willd.) DC., D. paniculatum (L.) DC. and D. (1983, 1990, 1998). “Desmodium traditionally procumbens (Mill.) Hitchc., along with each is considered a ‘difficult’ genus” (Isely 1990, p. of their related taxa, as species groups. The 163). Isely (1990) further stated that Desmodium difficulty in classifying these groups is, however, contains “groups of species whose not satisfactorily solved by this approach. Taxa is presently ambiguous or among which the attributed to these groups appear to have been differentiation of kinds is unsatisfactory or recognized by over-weighting such variable overlapping. In such cases, confidently keying characters as type and degree of density of a to a name is often difficult. Thus, I indumentum on the stems and leaves and by the have borrowed a device from Flora Europaea size and shapes of the leaves. The problems in (Tutin et al. 1968, p. xv): ‘Where it is difficult the North American species groups are similar to distinguish between a number of closely to those in the Asian species of Desmodium. similar species in a genus, an ad hoc group For example, Desmodium heterocarpon (L.) has been made, and these groups ... are keyed DC. includes many synonyms that were initially out in the main species key.’ ... For example, if distinguished by over-emphasizing superficially the identity of a specimen in the Desmodium discontinuous characters (Ohashi 1973, 1991). paniculatum (L.) DC. group is uncertain, one To correctly distinguish natural taxa, further

—166— June 2013 The Journal of Japanese Botany Vol. 88 No. 3 167 analyses of morphological characters and commonly stout ...... D. ciliare genetic features of the groups are necessary. 4. Stems and leaves with various combinations Until that time when such detailed studies can of characters of D. ciliare and D. be carried out, I have decided to regard each of marilandicum, e.g., stems and leaflets the “groups” as single species, D. paniculatum, glabrate but petioles 1–3 mm long or stems D. marilandicum, and D. procumbens, in my pilose but petioles well developed; pedicels treatment of Desmodium for the Flora of North various in length ...... America. Their so-called related species are ...... D. ciliare × D. marilandicum treated as synonymous or as varieties of the recognized species. Isely (1998) noted, however, that “many populations of Desmodium ciliare are of, or 1. The Desmodium ciliare Group include, with various features of D. Plants described in this group as separate marilandicum” (p. 538). “Some plants of D. species are characterized by having similar 2 or marilandicum throughout its range exhibit 3 segmented loments. They include Desmodium various combinations of the characters of D. ciliare (Willd.) DC., D. marilandicum (L.) DC., marilandicum and D. ciliare and are relegated and D. obtusum (Willd.) DC. with D. ciliare × to postulated D. ciliare × D. marilandicum D. marilandicum and D. ciliare × D. obtusum category” (p. 546). This presumed hybrid (Isely 1990, 1998). Isely (1998), Krings (2004), includes “(1) plants with the aspect of D. ciliare, and Raveill (2006), each presented keys to the D. but which are glabrate or have some pubescence; ciliare group, all based on vegetative characters, (2) those with the aspect of D. ciliare, including of which Isely’s is as follows: pubescence, but leaves well petioled; and (3) those with the aspect of D. marilandicum but 1. Leaflets –3 5.3 times as long as wide ...... somewhat pilose” (p. 558). Desmodium ciliare ...... D. ciliare × ? × D. obtusum is presumed to be “plants with 1. Leaflets 1.2–3.5 times as long as wide ...... 2 foliage intermediate between Desmodium ciliare 2. Terminal leaflet generally distinctly longer and D. obtusum and with a mixture of uncinate and narrower than others; medial stems and pilose pubescence (p. 558). sparsely to densely uncinate-pubescent ...... Isely’s (1998) comments indicate that D...... D. obtusum ciliare, D. marilandicum and D. obtusum are 2. Terminal leaflet similar to lateral ones; indistinguishable without recognizing the medial stems glabrous to pilose or also with intermediate forms between them as presumed uncinate pubescence. hybrids. No evidence of hybrid origin of these 3. Stems glabrous(-sparsely uncinate- intermediate forms was presented. I recognize puberulent); petioles 1–2.5 cm long; leaflets this group as a single polymorphic species, D. glabrous or with a few hairs; pedicels marilandicum, with three varieties, although the filiform, generally 0.8–1.5 cm long ...... distinction between the vaieties is sometimes ...... D. marilandicum ambiguous. A key to the varieties is as follows: 3. Stems generally pilose; petioles and pedicels 1. Terminal leaflet elliptic-ovate to narrowly shorter than above or plants lacking the ovate, 5–7.5 × 2–3.3 cm, apex acute to obtuse above combination of characters ...... 4 ...... var. lancifolium 4. Stems conspicuously or thinly pilose(- 1. Terminal leaflet elliptic, ovate, ovate-rhombic uncinate-puberulent); petioles 1–3 mm long, or broadly ovate, 0.9–4 × 0.6–1.7 cm, apex shorter than rachis; leaves subappressed- obtuse ...... 2 pubescent(-glabrate); pedicels 3–8 mm long, 2. Leaves nearly sessile, petioles 1–15 mm 168 植物研究雑誌 第 88 巻 第 3 号 2013 年 6 月

Fig. 1. Holotype of Desmodium ciliare var. lancifolium Fernald & B. G. Schub. (GH). June 2013 The Journal of Japanese Botany Vol. 88 No. 3 169

long; stems and petioles uncinate-puberulent D. marilandicum var. lancifolium (Fernald and mostly pilose; pedicels 3–8 mm long ...... & B. G. Schub.) H. Ohashi, comb. nov...... var. ciliare D. ciliare var. lancifolium Fernald & B. G. 2. Leaves petiolate, petioles 12–30 mm long; Schub., Rhodora 40: 437 (1938). [Fig. 1] stems essentially glabrous, petioles sparsely Hedysarum obtusum Willd., Sp. Pl. 3(2): uncinate-puberulent; pedicels (6–)8–19 mm 1190 (1802). long ...... var. marilandicum H. rigidum Elliott, Sketch Bot. S. Carolina 2: 215 (1823). Desmodium marilandicum (L.) DC., Prodr. D. obtusum (Willd.) DC., Prodr. 2: 329 2: 328 (1825); Schub. in Gray’s Man. ed. 8: 920 (1825); Isely, Native Natur. Leg. US: 547 (1950); Schub. in Man. Vasc. Pl. Texas: 864 (1998); Krings in Vulpia 3: 144, fig. 9 (2004). (1970); Isely, Vasc. Fl. SE US 3(2): 175 (1990); D. rigidum (Elliott) DC., Prodr. 2: 330 Isely, Native Natur. Leg. US: 546 (1998); Krings (1825). in Vulpia 3: 145, fig. 8 (2004); Raveill in Vulpia Meibomia obtusa (Willd.) Vail in Bull. 5: 20 (2006). Torrey Bot. Club. 19: 115 (1892); Schind. in Hedysarum marilandicum L., Sp. Pl. 2: 748 Repert. Spec. Nov. Regni Veg. 23: 357 (1927). (1753). Meibomia rigida (Elliott) Kuntze, Revis. Meibomia marilandica (L.) Kuntze, Revis. Gen. Pl. 1: 198 (1891). Gen. Pl. 1: 198 (1891). Stems and petioles sparsely to densely var. marilandicum uncinate-puberulent. Leaves shortly petiolate; Stems essentially glabrous. Leaves mostly petioles sparsely to densely uncinate-puberulent, petiolate; petioles sparsely uncinate-puberulent, 2–20 mm long; terminal leaflet elliptic-ovate to 12–30 mm long; terminal leaflet ovate, ovate- narrowly ovate, 5–7.5 × 2–3.3 cm, apex acute to rhombic or suborbicular, 1.5–2.5(–4) × 1–1.7 cm, obtuse. Pedicels 4–10 mm long. apex obtuse. Pedicels (6–)8–19 mm long. 2. The Desmodium paniculatum Group D. marilandicum var. ciliare (Willd.) H. Schubert (1950a, 1950b) treated Desmodium Ohashi, comb. nov. paniculatum (L.) DC. and its related species Hedysarum ciliare Willd., Sp. Pl. 3(2): 1196 and clarified the classification of the group by (1802). recognizing D. glabellum (Michx.) DC. and Desmodium ciliare (Willd.) DC., Prodr. 2: describing D. perplexum B. G. Schubert. Isely 329 (1825); Isely, Native Natur. Leg. US: 538 (1983) surveyed the taxonomic relations among (1998); Krings in Vulpia 3: 145, fig. 6 (2004). these species in detail and recognized them as Meibomia ciliaris (Willd.) S. F. Blake in Bot. distinct within his Desmodium paniculatum Gaz. 78: 275 (1924). group. These species are characterized by having M. ciliaris var. acutifoliola Schindl. in straight loments with 3–5, angled articles. Repert. Spec. Nov. Regni Veg. 23: 357 (1927). Morphologically, however, as pointed out by M. ciliaris var. albiflora Schindl. in Repert. Isely (1990) “Desmodium paniculatum fades Spec. Nov. Regni Veg. 23: 357 (1927). into both D perplexum and D glabellum, and Stems uncinate-puberulent and patent long- authors have interpreted the species variously.” pilose. Leaves nearly sessile, petioles uncinate- Isely (1998) considered the relationships puberulent and patent long-pilose, 1–15 mm among these species as “a continuum” with long; terminal leaflet elliptic, ovate or ovate- variation between the members of the group rhombic, 0.9–4 × 0.6–1.7 cm, apex obtuse. suggesting introgression through various hybrid Pedicels 3–8 mm long. combinations: “D. paniculatum × D. perplexum; 170 植物研究雑誌 第 88 巻 第 3 号 2013 年 6 月

Fig. 2. Holotype of Desmodium perplexum B. G. Schub. Fig. 3. Holotype of Desmodium fernaldii B. G. Schub. (GH). (GH).

D. paniculatum × D. glabellum; and D. fruits. I consider that D. fernaldii should be glabellum × D. perplexum.” recognized as distinct from D. paniculatum, Krings (2004) characterized the abaxial but at the rank of variety, even though they leaf surface of the D. paniculatum group in are distinguished by somewhat continuous or North Carolina as follows: D. glabellum: overlapping characteristics: abaxial midvein with dense, long, spreading, 1. Abaxial leaf surface sparsely strigillose, straight trichomes, and uncinate trichomes uncinate-puberulent on veins; stems and lacking or nearly so; D. fernaldii: abaxial petioles glabrous or uncinate-pubescent ...... midvein with dense uncinate trichomes, and ...... var. fernaldii scattered appressed, straight trichomes; D. 1. Abaxial leaf surface slightly strigose to paniculatum: abaxial midvein with appressed, conspicuously subappressed-villous and straight trichomes of ca. 0.2 mm long; and D. sometimes uncinate-pubescent; stems and perplexum: abaxial midvein with long, straight, petioles glabrescent to conspicuously pilose spreading or appressed trichomes and shorter or uncinate-pubescent ...... var. paniculatum uncinate trichomes. Through the wide range of morphological Desmodium paniculatum (L.) DC., Prodr. variations in the D. paniculatum group, I 2: 329 (1825); B. G. Schub. in Rhodora 52: 152 recognize the members merely as forms of (1950); in Gray’s Manual ed. 8: 922 (1950); a single polymorphic species, because these Isely, Vasc. Fl. SE US 3(2): 173 (1990); Isely, species are basically uniform in flowers and Native Natur. Leg. US: 550 (1998); Krings in June 2013 The Journal of Japanese Botany Vol. 88 No. 3 171

Vulpia 3: 144, fig. 2 (2004); Raveill in Vulpia 5: circumscribed the species including D. exiguum 19 (2006). A. Gray. Desmodium procumbens is divided Hedysarum paniculatum L., Sp. Pl. 2: 749 into four varieties based on differences in the (1753). leaves; one variety is in North America north of H. glabellum Michx., Fl. Bor. Amer. 2: 73 Mexico. The North American representative of (1803). the species, regarded as var. exiguum (A. Gray) Desmodium glabellum (Michx.) DC., Prodr. B. G. Schub., has broad, unifoliolate lower 2: 329 (1825); Isely, Native Natur. Leg. US: 541 leaves and linear to narrowly ovate leaflets (1998); Krings in Vulpia 3: 146, fig. 13 (2004). on the upper leaves. Isely (1998) commented D. dillenii Darl., Fl. Cestr. ed. 2: 414 (1837). on D. procumbens as follows: “[Desmodium Meibomia paniculata (L.) Kuntze, Revis. procumbens] var. exiguum, which extends north Gen. Pl. 1: 198 (1891). to the United States, differs from the others in M. glabella (Michx.) Kuntze, Revis. Gen. Pl. the extraordinary development of the broad 1: 198 (1891). unifoliolate lower leaves, which contrast with D. perplexum B.G. Schub., Rhodora 52: the narrow leaflets of the upper stem leaves. 154 (1950); Isely, Native Natur. Leg. US: 551 Although D. procumbens appears well-defined (1998); Krings in Vulpia 3: 144, fig. 20 (2004). in the United States, this may not be true in [Fig. 2] Mexico and southward (McVaugh 1987).” D. paniculatum var. epetiolatum B. G. McVaugh (1987) reports D. procumbens to be Schub. in Rhodora 52: 153 (1950). “a variable species with respect to leaf-form and D. paniculatum var. dillenii (Darl.) Isely, indument as well as to plant-habit.” Amer. Midl. Naturalist 49: 927 (1953). The habit of Desmodium procumbens varies D. dichromum Shinners, Spring Fl. Dallas- locally. It is described as an erect or ascending Fort Worth Area: 409 (1958). perennial in North America (Isely 1998), or as var. paniculatum an erect or procumbent annual (Schubert 1940, 1980; McVaugh 1987). The epithet, procumbens D. paniculatum var. fernaldii (B. G. Schub.) is derived from Hedysarum procumbens Mill. H. Ohashi, comb. nov. described from Jamaica. D. fernaldii B. G. Schub. in Rhodora 52: Desmodium procumbens var. exiguum was 147 (1950); Isely, Native Natur. Leg. US: 540 distinguished from var. procumbens by having (1998); Krings in Vulpia 3: 146, fig. 12 (2004). variously shaped leaflets versus uniform leaflets [Fig. 3] in the latter (Schubert 1940), but the leaflets of var. exiguum are polymorphic, exhibiting 3. The Desmodium procumbens Group continuous variation with those of var. Desmodium procumbens (Mill.) Hitchc. procumbens. occurs from SE Arizona to Trans-Pecos Texas Isely (1998) grouped Desmodium in the southwestern United States, southward procumbens with D. neomexicanum A. Gray through Mexico, Central America to the and D. rosei B. G. Schub. in the D. procumbens northern half of South America and in the West group. Desmodium neomexicanum is united with Indies. It is introduced and naturalized in the D. procumbens by having twisted loments, but Philippines, Maluku (Ambon) and tropical is recognized at the rank of variety, distinguished Africa. The species is characterized by having as follows: twisted loments, which are strongly so when 1. Margin of loments not involute; articles young. Desmodium procumbens became known glabrous, abaxial margin rounded .....D. rosei in North America when Schubert (1940) newly 1. Margin of loments involute; articles 172 植物研究雑誌 第 88 巻 第 3 号 2013 年 6 月

Fig. 4. Holotype of Desmodium exiguum A. Gray (GH). June 2013 The Journal of Japanese Botany Vol. 88 No. 3 173

Fig. 5. Holotype of Desmodium bigelovii A. Gray (GH). 174 植物研究雑誌 第 88 巻 第 3 号 2013 年 6 月

uncinate-puberulent (terminal one sometimes D. procumbens var. neomexicanum (A. glabrous), abaxial margin angled or Gray) H. Ohashi, comb. nov. [based on autonym sometimes rounded at maturity ...... D. neomexicanum A. Gray var. neomexicanum ...... D. procumbens (1879)]. 2. Loments distinctly spirally twisted, stipitate Desmodium neomexicanum A. Gray, (stipe 1.5–3.5 mm long); primary bracts Smithsonian Contr. Knowl. 3(5): 53 (1852), as 1.5–2.5 mm long; leaves 1- and 3-foliolate ... “Neo-Mexicanum”; Schub. in Cont. Gray Herb...... var. procumbens 129: 21, pl.1, fig. C, 1–9 (1940); McVaugh, Fl. 2. Loments slightly spirally twisted, subsessile Novo-Galic. 5: 475 (1987); Isely, Native Natur. or stipitate (stipe 0.3–2 mm long); primary Leg. US: 546 (1998). bracts 2.5–5.5 mm long; leaves mostly Desmodium bigelovii A. Gray, Smithsonian 3-foliolate ...... var. neomexicanum Contr. Knowl. 5(6): 47 (1853). [Fig. 5] D. neomexicanum var. bigelovii (A. Gray) Desmodium rosei B. G. Schub., Contr. Gray S. Watson, Rep. US Geogr. Surv., Wheeler 6 Herb. 129: 22, pl. 1, fig. A. 1940. (Botany): 101 (1879). Meibomia neomexicana (A. Gray) Kuntze, Desmodium procumbens (Mill.) Hitchc. in Revis. Gen. Pl. 1: 198 (1891). Ann. Rep. Missouri Bot. Gard. 4: 76 (1893); M. bigelovii (A. Gray) Kuntze, Revis. Gen. Schub., Contr. Gray Herb. 129: 5, pl. 1, fig. B1–7 Pl. 1: 197 (1891). (1940) as var. typicum; Schub. in Ann. Missouri Herbs, annual. Stems inconspicuously Bot. Gard. 67: 651 (1980); McVaugh, Fl. Novo- uncinate-pubescent or glabrescent. Leaves Galic. 5: 481 (1987). mostly 3-foliolate, apex obtuse, both surfaces Hedysarum procumbens Mill., Gard. Dict. uncinate-puberulent. Primary bracts usually ed. 8, Hedysarum no. 10 (1768) [Type. Jamaica. persistent, 2.5–5.5 mm long. Loments slightly Houstoun, 1730 (BM; photo GH). fide Schubert spirally twisted, subsessile or stipitate; stipe (1980)]. 0.3–2 mm long; articles (1–)2–5. D. exiguum A. Gray, Smithsonian Contr. Knowl. 5(6): 46 (1853). [Fig. 4] This study was mainly based on specimens Meibomia exigua (A.Gray) Kuntze, Revis. of Desmodium in the Harvard University Gen. Pl. 1: 198 (1891). Herbaria (A, GH, NEBC), which had long M. procumbens (Mill.) Schind. in Repert. been studied by Dr. Bernice G. Schubert Spec. Nov. Regni Veg. 20: 151 (1924). (1913–2000) who carefully annotated almost D. procumbens var. exiguum (A. Gray) B. all of them collected in North America. I would G. Schub. in Contr. Gray Herb. 129: 12, pl. 1, like to thank Dr. D. E. Boufford and Ms. E. fig. B8–9 (1940); Schub. in Ann. Missouri Bot. W. Wood (Harvard University Herbaria) for Gard. 67: 651.1980; Isely, Native Natur. Leg. various help when I worked on the treatment US: 552 (1998). for FNA at Harvard in 2007, 2010 and 2012. var. procumbens I am grateful to Drs. T. Azuma (Hokkaido Herbs, perennial. Stems uncinate-puberulent University), Y. Endo (Ibaraki University), Y. and sparsely pubescent. Leaves 1- and 3-foliolate, Iokawa (Jyoetsu Kyoiku University), T. Kajita apex acute or acuminate, both surfaces uncinate- (Chiba University), T. Nemoto (Ishinomaki puberulent and villose. Primary bracts caducous Senshu University), R. E. Spangler (Harvard or partly persistent, 1.5–2.5 mm long. Loments University), M. Suzuki (Tohoku University), Y. distinctly spirally twisted, stipitate, stipe 1.5–3.5 Tateishi (Ryukyu University), and J. Yokoyama mm long; articles 2–3. (Yamagata University) for their collaboration June 2013 The Journal of Japanese Botany Vol. 88 No. 3 175 in my field work in the United States in 1995, Its Allied Genera (Leguminosae). 318 pp., 76 pls. 1996 and 1997. I thank Dr. J. Raveill (University Ginkgoana 1. Academia Scientific Book, Inc., Tokyo. Ohashi H. 1991. Taxonomic studies in Desmodium of Central Missouri) for his help in editing the heterocarpon (L.) DC. (Leguminosae). J. Jap. Bot. 66: original manuscript of Desmodium for Flora 14–25. of North America, which includes the new Raveill J. A. 2006. Identification of Missouri species of combinations proposed here. I am also grateful the tribe Desmodieae (Fabaceae) using vegetative to Dr. Boufford for comments on the original characters. Vulpia 5: 14–22. Schubert B. G. 1940. Desmodium: Preliminary studies-I. manuscript. Contr. Gray Herb. 129: 3–31. Schubert B. G. 1950a. Desmodium: Preliminary studies III. References Rhodora 52: 135–155. Isely D. 1983. The Desmodium paniculatum (L.) DC. Schubert B. G. 1950b. Desmodium. In: Fernald M. (Fabaceae) complex revisited. Sida 10: 142–158. L., Gray’s Manual of Botany, 8th ed. pp. 915–923. Isely D. 1990. Vascular Flora of the southeastern United American Book. Co., New York. States. 3(2) Leguminosae (Fabaceae). The University Schubert B. G. 1970. Desmodium. In: Correll D. S. and of North Carolina Press, Chapel Hill and London. Johnston M. C., Manual Vasc. Pl. Texas. pp. 855–869. Isely D. 1998. Native and Naturalized Leguminosae Texas Research Foundation, Renner, Texas. (Fabaceae) of the United States exclusive of Alaska and Schubert B. G. 1980. Desmodium. In: Dwyer J. D. and Hawaii. Monte L. Bean Life Science Museum, Brigham collaborators. Flora of Panama, part 5, Fam. 83 Young University, Provo, Utah. Leguminosae subfam. Papilionoideae. Ann. Missouri Krings A. 2004. Abaxial foliar vestiture of Desmodium Bot. Gard. 67(3): 622–662. Desv. (Fabaceae) in North Carolina and vegetative Woods M. 2008. The genera Desmodium and Hylodesmum recognition of the species. Vulpia 3: 140–172. (Fabaceae) in Alabama. Castanea 73(1): 46–69. Ohashi H. 1973. The Asiatic Species of Desmodium and

大橋広好：北アメリカ産シバハギ属（マメ科ヌスビトハ ギ連）の新組み合わせ Flora of North America vol. 11 のマメ科シバハギ属 海道胆振からも報告がある．市街地で見られるもっとも Desmodium を執筆した際に Desmodium marilandicum 普通のシバハギ属植物で，荒れ地のありふれた雑草とな (L.) DC., D. paniculatum (L.) DC. お よ び D. っている．小葉は多型で，狭長卵形が典型的な形である procumbens (Mill.) Hitchc. について種の範囲を新たに が，線形から広卵形，ほぼ円形あるいは菱形までの変異 設定した．これに伴って4 個の新組み合わせを行ったので， が見られる．日本に帰化している形を見ても小葉にはかな これらを本文中で発表した．また， D. marilandicum では りの変異が見られる．D. marilandicum と同種と見なした 近縁種とされていた Desmodium obtusum (Willd.) DC.， Desmodium obtusum (Willd.) DC. も日本に帰化しており， D. paniculatum で は D. glabellum (Michx.) DC. と D. 1940 年大阪府茨木市で採集され，今日ではアメリカヌス perplexum B. G. Schub.， および D. procumbens で は ビトハギと呼ばれている（日本の帰化植物 2003. p. 106. D. procumbens var. exiguum (A. Gray) B. G. Schub. を 平凡社）．この種は D. marilandicum の変種である var. 異名と見なした．Desmodium paniculatum (L.) DC. は lancifolium (Fernald & B. G. Schub.) H. Ohashi に当た 北アメリカ東部に広く分布し，もっとも普通に見られる種 ると考えられる． (Isely 1990) である．日本でアレチヌスビトハギと呼ばれ （東北大学植物園津田記念館） ていて，東北地方南部以西沖縄まで広く帰化していて北