全 文 ：植 物 分 类 学 报 43 (6): 557–560(2005) doi:10.1360/aps040072
Acta Phytotaxonomica Sinica http://www.plantsystematics.com
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Received: 27 July 2004 Accepted: 19 October 2004
*Author for correspondence. E-mail: xiangyunzhu@ibcas.ac.cn; Tel.: 86-10-62836111
Taxonomic relationship between Desmodium diffusum DC.
and D. laxiflorum DC. (Fabaceae: Papilionoideae)
1Hiroyoshi OHASHI 2ZHU Xiang-Yun*
1(Botanical Garden, Tohoku University, Sendai 980-0862, Japan)
2(Institute of Botany, the Chinese Academy of Sciences, Beijing 100093, China)
Abstract Desmodium diffusum DC. has usually been included in D. laxiflorum DC., but they
are two distinct species. The former is widely distributed in China, while the latter is found
only in southern China, i.e., Guangdong, Guangxi, south Yunnan, and rarely in Taiwan. D.
unibotryosum C. Chen & X. J. Cui was once applied to the former as a new name, but this
name is illegitimate. The taxonomic history of Desmodium diffusum and D. laxiflorum, key to
these species, their bibliographies, synonyms and distributions are clarified. The lectotype of
D. diffusum is designated.
Key words Desmodium diffusum DC., Desmodium laxiflorum DC., Desmodium
unibotryosum C. Chen & X. J. Cui, Fabaceae, China.
Desmodium diffusum DC. has usually been included in D. laxiflorum DC. since the
treatment of Bentham (1852), but is recognized as a distinct species by Chen and Ohashi
(1983), Ohashi (1985, 1994, 1995) and Cui et al. (1987). This paper aims to clarify their
taxonomic relationship.
The two species were described by de Candolle in January 1825 (de Candolle, 1825a).
He described Desmodium diffusum from East India as“caule fruticoso erectiusculo, racemis
terminalibus elongatis, pedicellis calyce duplo longioribus”, while D. laxiflorum from Nepal
as “caule frutescente erecto, racemis terminalibus laxis, pedicellis hirsutis filiformibus calyce
multo longioribus”. These two species were kept distinct in his succeeding work in November
1825 (de Candolle, 1825b). The type material of D. laxiflorum and of D. diffusum was
examined in de Candolle: Prodromi Herbarium in G-DC (microfiche TUS) in this study.
There are four separate specimens of D. laxiflorum, but all bear the same label, “Glycine.
Napaul. Mr. Wallich 1821.” As pointed out by Chen and Ohashi (1983), on the other hand,
three separate sheets of D. diffusum in Prodromi Herbarium in G-DC were examined, of
which two sheets were respectively mounted with two plants and one sheet with one plant.
These five plants have different labels: “Inde orient Mr. Lambert 1816”and “Inde Roxburgh
Mr. Lambert 1816” on one sheet; “Inde Mr. Lambert 1816” on the other; “Glycine.
Napaul. Mr. Wallich 1821” and unknown (probably the same with Napaul. Mr. Wallich
1821) on another one. De Candolle, however, doubted distinctness of both species by giving a
short note for D. laxiflorum in the original publication as “An forte prioris (=D. diffusum)
varietas” (de Candolle, 1825a) and again for D. diffusum (No. 88 in Prodromus) after the
description of D. laxiflorum (No. 87 in Prodromus) as “an satis a sequente differt?” (de
Candolle, 1825b). Bentham (1852) first regarded D. diffusum as identical with D. laxiflorum.
This treatment was adopted by Baker (1876) for the Flora of British India in which both
species were recorded. This concept for D. laxiflorum was established and has been accepted
Acta Phytotaxonomica Sinica Vol. 43 558
by subsequent taxonomists (Ohashi, 1973).
Chen and Ohashi (1983), however, recognized two forms in Desmodium laxiflorum.
They distinguished the two forms by the differences in the terminal leaflets, primary bracts,
pedicels, flowers, calyces, pods and seeds, and treated them as two subspecies within the
species, i.e., ssp. laxiflorum and ssp. parvifolium H. Ohashi & T. T. Chen. Cui et al. (1987)
considered that D. laxiflorum ssp. parvifolium H. Ohashi & T. T. Chen is a distinct species
from D. laxiflorum and proposed a new name, D. unibotryosum C. Chen & X. J. Cui, pointing
out that their species is identical with D. diffusum (Roxb.) DC. and that this name is a later
homonym of D. diffusum (Willd.) DC. However, they appear to be unaware that Ohashi
(1985) has reached the same conclusion that D. laxiflorum ssp. parvifolium is a distinct
species from D. laxiflorum DC., although his nomenclatural treatment is different from that of
Cui et al. (1987).
Ohashi (1985) considered that Desmodium diffusum DC. in Prodr. 2: 335 (1825) is a
correct name and different from D. diffusum (Willd.) DC. in Prodr. 2: 336 (1825). The type of
D. diffusum DC. is Lambert (G-DC), although Ohashi (1994) designated its type to be
Roxburgh, Cat. 57 (BM) that is the type of Hedysarum diffusum Roxb., non Willd. (Ohashi,
1994). According to the IDC microfiche/Prodromi Herbarium, as aforementioned, there are
three sheets of D. diffusum DC., i.e., two of Lambert 1816 and one of Wallich 1821. In total
five individuals are mounted on the three sheets and all are referable to D. diffusum DC. The
two Lambert sheets (three individuals) must be the syntypes of D. diffusum DC. Desmodium
diffusum (Roxb.) DC. is not published name, although Cui et al. (1987) cited it. de Candolle
(1825a) quoted “Hedysarum diffusum Roxb., non Willd.”only as its synonym in the original
description of Desmodium diffusum, and actually H. diffusum Roxb. in Hortus Bengalensis
(1814) is an invalid nude name. Hedysarum diffusum (DC.) Roxb. was correctly published in
1832 based on Desmodium diffusum DC., but is a later homonym of Hedysarum diffusum
Willd. published in 1802.
Desmodium diffusum (Willd.) DC. (1825) based on Hedysarum diffusum Willd. is an
entirely different species from D. diffusum DC. This is a synonym of D. dichotomum (Willd.)
DC. based on H. dichotomum Willd. published in 1802. In China Desmodium dichotomum is
known only in Yunnan (Ohashi, 1995; Yang & Huang, 1995).
A new illustration of Desmodium diffusum DC. is provided on the plate 123 (on page
258) in Flora of Taiwan (Huang & Ohashi, 1993) as D. laxiflorum. The plate 3: 4–11 (on
page 18) in Flora Reipublicae Popularis Sinicae 41 (Yang & Huang, 1995) is similar to D.
diffusum. Illustration of true D. laxiflorum DC. is presented on the plate 18 (on page 89) in
Flore du Cambodge du Laos et du Vietnam (Ohashi, 1994). The figure 2623 of D. laxiflorum
in Iconographia Cormophytorum Sinicorum 2: 447 (1972), which is same with figure 473 in
Flora Illustralis Plantarum Primarum Sinicarum: 484 (1955), must be drawn on the basis of
the two species. The inflorescences, primary bract and calyx apparently are those of D.
laxiflorum, but the pod is D. diffusum. Unfortunately, we could not find voucher specimens of
the figure 2623 in Iconographia Cormophytorum Sinicorum in PE.
In summary, Desmodium diffusum DC. and D. laxiflorum DC. are distinguished as
follows:
1. Terminal leaflet 2.5–11 cm long, apex obtuse or acute, lateral veins 5–9 pairs; primary bract usually less
than 4 times as long as broad; corolla blue-violet or violet; pods smooth to more or less rough on lateral
surfaces, obliquely jointed, both sutures constricted at nodes; hilum about 0.15 mm long; prostrate or
ascending herbs or subshrubs ........................................................................................................ D. diffusum
1. Terminal leaflet (5.5–)8–20 cm long, apex acute or acuminate, lateral veins 8–11 pairs; primary bract more
than 4 times as long as broad; corolla white or pale violet; pods striate-veined on lateral surfaces, vertically
jointed, both sutures almost straight at nodes; hilum about 0.3 mm long; erect or ascending shrubs or
No. 6 H. OHASHI et al.: Taxonomic relationship between Desmodium diffusum and D. laxiflorum 559
subshrubs......................................................................................................................................D. laxiflorum
Desmodium diffusum DC. in Ann. Sci. Nat. 4: 100. 1825; et Prodr. 2: 335, no. 88. 1825;
Ohashi in Hara, Origin Evolution Divers. Pl. & Pl. Commun. 186. 1985; Ohashi et al. in Sci.
Rep. Tohoku Univ. ser. 4, Biol. 39: 215. 1988; Ohashi in Dy Phon et al., Fl. Camb. Laos
Vietn. 27: 86. 1994, excl. syn. cit. H. recurvatum Roxb. et D. recurvatum (Roxb.) Wight &
Arn.; Ohashi in J. Jpn. Bot. 70: 113. 1995; T. C. Huang & Ohashi in Fl. Taiwan, ed. 2, 3: 252,
plate 123 ut D. laxiflorum (1993). (Lectotype: India orient. Lambert (G-DC), n.v. fide
microfiche Candolle: Prodromi Herbarium in TUS)
Hedysarum diffusum Roxb., (Hort. Beng. no. 57 (1814), nom. nud.) Fl Ind. 3: 357. 1832;
non Willd. 1802. Type: India orient. Roxburgh, Cat. no. 57 (BM), n.v., nom. illegitimate.
D. laxiflorum DC. var. formosense Ohwi in J. Jpn. Bot. 26: 235. 1951. Type: Formosa.
Fujieda in Kizangun. leg. S. Okamoto (KYO).
Desmodium laxiflorum ssp. parvifolium H. Ohashi & T. T. Chen in J. Jpn. Bot. 58: 268,
figs. 1, 2. 1983. Type: Taiwan. Pingtung Co. Mutan, alt. 350–400 m. In open evergreen forest,
among grass, along a path. Ohashi et al. 13486 (TUS holo., iso. TAI, TI, TUS); Ohashi et al.,
Sci. Rep. Tohoku Univ. ser. 4, 38: 305. 1984.
Desmodium unibotryosum C. Chen & X. J. Cui, nom. nov. in Acta Bot. Yunnan. 9: 307.
1987.
Desmodium laxiflorum auct. non DC.: Benth. in Miq., Pl. Jungh. 228. 1852, p.p.; Baker
in Hook. f., Fl. Brit. Ind. 2: 164. 1876, p.p.; Forbes & Hemsley in J. Linn. Soc. Bot. 23: 173.
1887; Gagnep., Fl. Gen. Indo-Chine 2: 591. 1920, p.p.; Merr., Enum. Philip. Pl. 2: 286. 1923;
van Meeuwen, Reinwardtia 6: 252. 1962, p.p.; Ohashi in Ginkgoana 1: 101. 1973, p. p., incl.
plate 12a et fig. 25: 1–4, 26: 4; T. C. Huang & Ohashi, Fl. Taiwan 3: 262. 1977, p. maj. p.;
Ohashi in Hara & Williams, Enum. Flow. Pl. Nepal 2: 118. 1979, p. p.; Verdc., Man. New
Guin. Leg. 401. 1979, p.p.; Yang & Huang in Fl. Reip. Pop. Sin. 41: 20. 1995, p.p. excl. syn.
cit. D. diffusum (Willd.) DC.
Distribution: India, Nepal, Sikkim, Bhutan, Myanmar, Thailand, Indochina, Malesia,
China (Guangdong, Guangxi, Guizhou, Sichuan, Taiwan, Yunnan).
Habitat: Margin of forests, roadsides, in grasslands; alt. 50–2000 m.
Desmodium laxiflorum DC. in Ann. Sci. Nat. 4: 100. 1825et Prodr. 2: 335, no. 88. 1825;
Meeuwen in Reinwardtia 6: 252. 1962, p.p., excl. syn. cit. D. diffusum DC., H. roxburghii
Spreng. p.p.; Ohashi in Ginkgoana 1: 101. 1973, p.p., excl. syn. cit. D. diffusum DC.,
Hedysarum roxburghii Spreng. p.p., D. laxiflorum var. formosense Ohwi; Ohashi in Hara,
Origin Evolution Divers. Pl. et Pl. Commun. 178–182. 1985; Grierson & Long, Fl. Bhutan 1:
678. 1987; Ohashi et al. in Sci. Rep. Tohoku Univ. ser. 4, Biol. 40: 19. 1991; Ohashi in Dy
Phon et al., Fl. Camb. Laos Vietn. 27: 87, pl. 18: 1–3. 1994; T. C. Huang & Ohashi in Fl.
Taiwan, ed. 2, 3: 256. 1993, excl. pl. 123. (Type: Nepal. Wallich 1821 (G-DC, four syntypes),
n.v. fide Candolle: Prodromi Herbarium microfiche in TUS)
D. recurvatum (Roxb.) [Grah. ex Wall., Cat. no. 5717, 1831–1832, comb. nud.] Wight &
Arn. in Prodr. 226. 1834; Wight, Icon. t. 374, 1838; Hosokawa, J. Soc. Trop. Agric. 4: 201.
1932.
Distribution: Bhutan, China (Guangdong, Guangxi, Taiwan, Yunnan), Indonesia, Laos,
Myanmar, Nepal New Guinea, Philippines, Sikkim, Thailand, Vietnam.
Habitat: In open grasslands and forests, margin of forests; alt. 100–1500 m.
Acknowledgements We thank Ms. GAO Xin-Fen, Chengdu Institute of Biology, the
Chinese Academy of Sciences, for her comments on the manuscript.
Acta Phytotaxonomica Sinica Vol. 43 560
References
Baker J G. 1876. Desmodium Desv. In: Hooker J D ed. The Flora of British India. Kent: Reeve & Co. 2:
161–175.
Bentham G. 1852. Leguminosae. In: Miquel F A W ed. Plantae Junghuhnianae. Leiden: Sythoff. 205–269.
Chen T-T, Ohashi H. 1983. Two forms in Desmodium laxiflorum DC. (Leguminosae). The Journal of Japanese
Botany 58: 263–269.
Cui X-J (崔现举), Chen C (陈介), Wu C-Y (吴征镒). 1987. New taxa of the genus Podocarpium and a new
name of Desmodium. Acta Botanica Yunnanica 9: 305–308.
De Candolle A P 1825a. Notice sur quelques genres et espèces nouvelles de lègumineuses, extraite de divers
Mémoires présentés à la Société dHistoire naturelle de Genève, pendant le cours des années 1823 et
1824. Annales des Sciences Naturelles 4: 90–103.
De Candolle A P 1825b. Ordo LXIII. Leguminosae Juss. In: de Candolle A P ed.: Prodromus Systematis
Naturalis Regni Vegetabilis. Paris: Treuttel et Würtz. 2: 93–524.
Huang T-C, Ohashi H. 1993. Leguminosae. In : Flora of Taiwan. 2nd ed. Taipei: National Taiwan University.
3: 160–396.
Ohashi H. 1973. The Asiatic species of Desmodium and its allied Genera (Leguminosae). Ginkgoana 1: 1–318.
Ohashi H. 1985. Specific diversity in Desmodium subgenus Desmodium (Leguminosae). In: Hara H. ed.
Origin and Evolution of Diversity in Plant and Plant Communities. Tokyo: Academic Scientific Book Inc.
176–187.
Ohashi H. 1994. Desmodium Desv. In: Dy Phon P, Ohashi H, Vidal J E ed. Flore du Cambodge du Laos et du
Vietnam 27. Legumineuses-Desmodiees. Paris: Museum National d’Histoire Naturelle, Laboratoire de
Phanerogamie. 62–142.
Ohashi H. 1995. An enumeration of Chinese Desmodium and its allied genera (Leguminosae). The Journal of
Japanese Botany 70: 111–117.
Yang Y-C (杨衔晋), Huang P-H (黄普华). 1995. Desmodieae. In: Flora Reipublicae Popularis Sinicae (中国
植物志). Beijing: Science Press. 41: 1–161.
弥散山蚂蝗与大叶拿身草(豆科:蝶形花亚科)
的分类学关系
1大桥广好 2朱相云*
1(东北大学植物园 仙台市 980-0862 日本)
2(中国科学院植物研究所 北京 100093)
摘要 弥散山蚂蝗Desmodium diffusum(新拟)不同于大叶拿身草D. laxiflorum,然而前者在中国却不被承
认。弥散山蚂蝗广布于中国,与大叶拿身草相较更为普遍,而大叶拿身草仅分布于中国的广东、广西、台
湾及云南南部。崔现举等将弥散山蚂蝗置于单序山蚂蝗D. unibotryosum种中。根据国际命名法规,单序
山蚂蝗为一非法名称,属弥散山蚂蝗的异名。本文介绍了弥散山蚂蝗和大叶拿身草的分类历史、种的检
索表、种的文献、异名及分布。同时,指定了弥散山蚂蝗的后选模式。
关键词 弥散山蚂蝗; 大叶拿身草; 单序山蚂蝗; 豆科; 中国