J. Jpn. Bot. 93(2): 104–120 (2018)

Grona, a Separated from (Leguminosae Tribe )

a,* b Hiroyoshi Ohashi and Kazuaki Ohashi

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

(Accepted on November 4, 2017)

The genus Desmodium is polyphyletic as suggested by recent molecular analyses. Desmodium sections Nicolsonia and Sagotia constitute a monophyletic group distinct from that which includes Desmodium scorpiurus, the type of the genus. The oldest generic name applicable to the monophyletic group is Lour. It was established on the basis of Grona repens Lour. (= Desmodium heterocarpon (L.) DC.) in 1790, but the name is rejected under the conserved name Desmodium Desv. published in 1813. Desmodium and Grona in a new sense are separated from each other based on morphological and palynological characters and evidence from molecular analyses. Grona is, therefore, resurrected in order to accommodate the species belonging to Desmodium sections Nicolsonia and Sagotia under ICN Art. 14.6. The present paper provides 41 new specific combinations with 13 new infraspecific combinations underGrona.

Key words: Desmodium, Desmodium heterocarpon, Desmodium triflorum, , Grona, Loureiro, Nicolsonia, Nogra, Sagotia, tribe Desmodieae.

The circumscription of the genus Desmodium Akschindlium (Ohashi 2003), Hanslia (Ohashi has long been controversial. It is ‘the main 2004b), Monarthrocarpus (Ohashi 2004b), problem’ in the tribe Desmodieae (Ohashi et al. (Ohashi 2005), Ototropis (Ohashi and 1981, Ohashi 2004a). Bentham’s generic concept Ohashi 2012a) and Verdesmum (Ohashi and of Desmodium (Bentham 1865) had widely Ohashi 2012b) are resurrected or produced. The been accepted, but Schindler (1928), Hutchinson taxonomic history of Desmodium and its allied (1964), Ohashi (1973) and Ohashi and Ohashi genera was reviewed by Pedley (1999) and Lima (2012a, b) split the genus into Desmodium and et al. (2014). its satellite genera. The broadest circumscription The tribe Desmodieae contains 32 genera of the genus was published by Bentham (1865), (Ohashi 2005, Ohashi and Ohashi 2012a, b) while the narrowest by Schindler (1928) (Ohashi classified into the group (three 2004a). Ohashi’s circumscription of the genus genera), the group (12 genera) and (1973) was intermediate between these two, but the Desmodium group (17 genera), based mainly it has been modified since 1996. Trifidacanthus on results of an analysis of rbcL by Kajita et (Ohashi et al. 1996), Ohwia (Ohashi 1999), al. (2001). Desmodium is the core genus in the (Ohashi and Mill 2000), Desmodium group. Polyphyly of Desmodium

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Fig. 1. Calyxes (opened view from inside) of Grona and Desmodium sect. Heteroloma. a. Grona barbata (Sauliére 506, K). b. G. triflora (Kanai & H. Ohashi 25747, TI). c. G. heterophylla (Tuyama in 1935, TI). d. G. rubra var. rubra (Katsumata s.n. in 1910, TI). e. Desmodium concinnum (Stainton & al. 8863, TI). f. D. oblongum (Iwatsuki & Fukuoka T.3211, TI). Adaxil lobes (connate lobe) are drawn on the left side. Scale bar: 2 mm. Reproduced with modification from figs. 52 and 53 in H. Ohashi, Ginkgoana 1 (1973) [ H. Ohashi]. was presented by Kajita et al. (1996, [2010 at molecular analyses indicates that D. scorpiurus 5th Legume Conference]), though the results is placed in a clade different from that composed were unpublished. Currently, the polyphyly of of D. heterocarpon, D. heterophyllum and Desmodium was demonstrated by Jabbour et other species belong to the subgenus Sagotia. al. (2018) in molecular phylogenetic analysis of A new classification system of Desmodium is, chloroplast (rbcL, psbA-trnH) and nuclear (ITS- therefore, needed for Desmodium based on the 1) DNA sequences. Desmodium was shown monophyletic group previously included in as polyphyletic separated into three clades: Desmodium. Desmodium heterocarpon-heterophyllum, D. The subgenus Sagotia is the most velutinum, and D. adscendens-intortum clades complicated group in the genus Desmodium. in their phylogenetic tree of cpDNA (Jabbour et Ohashi (1973) characterized the subgenus al. 2018, figs. 2, 3) and chronogram of cpDNA approximately by having terminal racemes (Jabbour et al. 2018, fig. 4). Moreover,Hegnera , composed of 2–4-flowered fascicles, small crinita, Leptodesmia, , flowers lacking bracteoles at the calyx-base, (only in fig. 2), (only in fig. 4- or 5-lobed calyx, diadelphous stamens, 2), Melliniella, and were presented mostly sessile ovary without disk around the between the three Desmodium clades in their base, and rather small-sized articles of the phylogenetic trees in figs. 2 and 4 (Jabbour et al. loments. He classified the subgenus into six 2018). sections: Chalarioidea, Oxytes, Heteroloma, During our systematic studies of Desmodium Nicolsonia, Renifolia and Sagotia. The sections on the basis of morphology, palynology and Chalarioidea and Oxytes are monotypic and molecular analyses, we obtained more detailed differ from others in having asymmetrical results of polyphyly of the genus (Ohashi et persistent stipules. Renifolia includes two species al. in press). Desmodium scorpiurus, the type distinct from others by their unifoliolate leaves species of the genus, was not included in the with a reniform leaflet and calyxes with lobes result of Jabbour et al. (2018), but our result of shorter than the tube (Ohashi 1973, Dy Phon et 106 植物研究雑誌 第 93 巻 第 2 号 2018 年 4 月 al. 1994). examined by Bentham (1865). He considered The remaining three sections are similar in the genus as similar to Kennedia and Galactia having symmetrical stipules, calyxes with lobes of the tribe . Merrill (1935) corrected longer than the tube. Sections Nicolsonia and the misapplication of Grona sensu Bentham to Sagotia are separated from sect. Heteroloma in Phaseoleae and accommodated the species of having 5- or 4-lobed calyx of which the adaxial Grona other than G. repens then known to his lobes are bifid at the apex and lacking secondary new genus Nogra. Merrill recognized Grona as bracts, whereas in Heteroloma the adaxial a valid genus and suggested it to be rejected for lobes are entire or slightly bifid (Fig. 1) and the Desmodium. Schindler (1928) determined the secondary bracts are present or sometimes absent holotype of Grona repens Lour. as Desmodium (Ohashi 1973). The sculpture of pollen grains siliquosum DC., which is a synonym of D. is rugulate-verrucate or fine rugulate, rarely heterocarpon subsp. heterocarpon var. strigosum fine rugulate to microreticulate in Nicolsonia van Meeuwen (Ohashi 1973). After examining and Sagotia (Fig. 2), but it is reticulate in the holotype on the image scanned by BM Heteroloma. (BM000839659), we confirm the as D. Our molecular analyses support that sections heterocarpon subsp. heterocarpon. Nicolsonia and Sagotia constitute a single monophyletic group, but Heteroloma is not. Taxonomic treatment The Nicolsonia and Sagotia group is clearly Grona Lour., Fl. Cochinch. 2: 459 (1790). separated from that including Desmodium Type: Grona helerocarpa (L.) H. Ohashi & K. scorpiurus. After examining eight species of Ohashi (= Grona repens Lour.). the subgenus Sagotia sect. Nicolsonia and Perrottetia DC. in Ann. Sci. Nat. 4: 95 (Jan. sect. Sagotia including D. heterocarpon and 1825), nom. illeg., non Kunth (1824). Type: D. heterophyllum (Ohashi et al. in press), it is Perrottetia barbata (L.) DC. strongly supported that the two sections should Nicolsonia DC., Prodr. 2: 325 (mid Nov. be treated as one group. 1825) [substitute name for Perrottetia DC. The oldest generic name for the Nicolsonia (1825), non Kunth (1824)]; Mem. Leg. 7: 311, t. and Sagotia group is the genus Grona. It was 51 (1826); Schindl. in Repert. Spec. Nov. Regni created by Loureiro (1790) based on Grona Veg. 23: 357, in adnot. (1927). Type: Nicolsonia repens Lour. (Holotype BM000839659) from barbata (L.) DC. (lectotypified by Schindler Vietnam in 1790. The species is a synonym 1927). of Desmodium heterocarpon (L.) DC., which Nicholsonia Span., Companion Bot. Mag. 1: belongs to the section Nicolsonia (DC.) Benth. 346 (1836), sphalm. ut ‘Nicholsonia oxalidifolia in the subgenus Sagotia in Desmodium (Ohashi Span.’ 1973). Nomenclatue of the generic name Sagotia Duchass. & Walp. in Linnaea Grona Lour. was fully discussed by Merrill 23: 737 (1851) [Type: Sagotia triflora (L.) (1935). Although the generic name Grona was Duchass. & Walp.], nom. rej. vs. Sagotia Baill. rejected under the conserved name Desmodium [] in Adansonia 1: 53 (1860), Desv. published in 1813 (Brummitt 1978, ICN nom. cons. App. III), the name can revive here under ICN Desmodium sect. Sagotia (Duchass. & Art. 14.6 (McNeill et al. 2012). We propose, Walp.) Benth., Miq., Pl. Jungh. 2: 222, adnot. therefore, resurrection of the genus Grona Lour. (1852); Benth. in Benth. & Hook. f., Gen. Pl. in order to accommodate Desmodium sections 1: 520 (1865); H. Ohashi in Ginkgoana 1: 233 Nicolsonia and Sagotia. (1973); Pedley in Austrobaileya 5: 227 (1999). – The holotype of Grona repens Lour. was [Meibomia sect. Sagotia (Walp.) Kuntze in Post, April 2018 The Journal of Japanese Botany Vol. 93 No. 2 107

Fig. 2. Pollen sculpture patterns of Grona barbata [India. Sauliére 506 K] (A–C). G. heterocarpa subsp. angustifolia [Thailand. Tagawa & al. 2456, TUS] (D–F). G. triflora [Taiwan. Taipei. Ohashi & al. 8809, TUS] (G–I) and G. auricoma [Thailand. Put 478, TUS] (J–L). A, D, G, J. Mesocolpium in equatorial view. B, E, H, K. Sculpture of mesocolpium. C, F, I, L. Aperture in equatorial view. Scale bars: 1 µm (B, E, H, K), 5 µm (A, C, D, F, G, I, J, L). 108 植物研究雑誌 第 93 巻 第 2 号 2018 年 4 月

Lex. Gen. Phan. 357 (1903), nom. nud.]. sometimes elliptic-oblong, abaxial suture slightly Desmodium sect. Nicolsonia (DC.) Benth. or distinctly constricted, hocked-hairy on lateral in Miq., Pl. Jungh. 2: 222 (1852); Benth. in surfaces. Seeds compressed, mostly broadly Benth. & Hook. f., Gen. Pl. 520 (1865); H. elliptic-oblong or almost orbicular in lateral Ohashi in Ginkgoana 1: 196 (1973); Pedley in view, distinctly rim-arillate around the hilum. Austrobaileya 5: 227 (1999). – [Meibomia sect. Nicolsonia (DC.) Kuntze in Post, Lex. Gen. Enummeration of species and Phan. 356 (1903), nom. nud.]. infraspecific taxa ofGrona Desmodium subgen. Sagotia (Duchass. & Grona adscendens (Sw.) H. Ohashi & K. Walp.) Baker in Hook. f., Fl. Brit. India 2: 173 Ohashi, comb. nov. (1876); H. Ohashi in Ginkgoana 1: 166 (1973); Hedysarum adscendens Sw., Prodr. 106 Pedley in Austrobaileya 5: 226 (1999). (1788), Fl. Ind. Occ. 3: 1263 (1806) [Type: Ind. Occ. Swartz (S–holotype)]. – Desmodium Description of the genus: Subshrubs or adscendens (Sw.) DC., Pror. 2: 332 (1825); B. herbs, sometimes shrubs. Leaves 1-, 3- or G. Schub. in Fl. Trop. E. Afr. Legum. 3, Papil. 1–3-foliolate, petiolate, stipulate; stipules free, 1: 461 (1971); H. Ohashi in Ginkgoana 1: 199 nearly symmetric; leaflets pulvinate, stipelate, (1973); Pedley in Dassanayake, Rev. Handb. Fl. chartaceous or subcoriaceous, margin entire, Ceylon 10: 180 (1996); B. G. Schub. in Ann. lateral nerves usually not reaching the margin; Missouri Bot. Gard. 67: 627 (1980); Du Puy stipels subulate. Inflorescences terminal or & Labat, Leg. Madag. 611 (2002); Lima & al. terminal and axillary or very rarely leaf-opposed, in Phytotaxa 169(1): 17 (2014). – Meibomia mostly raceme or panicle, rarely fascicle. adscendens (Sw.) Kuntze, Revis. Gen. Pl. 1: 195 Flowers lax or dense, usually 2–4-flowered per (1891). node. Bracts usually early deciduous, scariose, var. adscendens striate; secondary bracts absent. Bracteoles Distribution: America (West Indies and absent. Calyx campanulate, 4- or 5-lobed, Central and South America), introduced into Sri bilabiate; upper two lobes free or connate in Lanka, India, Thailand, Malaysia, Philippines, various degree with 2-toothed apex, abaxial Indonesia, New Guinea, Melanesia, and widely lobe usually longer than lateral ones, acuminate. in tropical Africa, Madagascar. Standard usually broadly obovate to nearly var. robusta (B. G. Schub.) H. Ohashi & K. orbicular, rounded or often emarginate at the Ohashi, comb. nov. apex, tapering or often suddenly narrowed Desmodium adscendens var. robustum B. to the base, not auriculate; wings not longer G. Schub. in Bull. Jardin Bot. Bruxelles 22: 290 than keel-petals, usually incurved, rounded or (1952) [Type: Congo. Kivu, Mokoto Lakes. obtuse at apex; keel-petals almost incurved, Ghesquière 4994 (BR–holotype)]; Verdcourt, apex obtuse, rarely rounded or subacute, not Man. New Guinea Leg.: 392 (1979). beaked, with claw longer than that of wings. Distribution: Congo, New Guinea. Stamens diadelphous, vexillary stamen free; anthers uniform. Ovary mostly sessile, without Grona appressipila (B. G. Schub.) H. Ohashi disk around the base; stigma terminal, slightly & K. Ohashi, comb. nov. capitate. Loments sessile or stipitate, compressed Desmodium appressipilum B. G. Schub. in or rarely slightly turgid, more than 3-jointed, Kew Bull. 25: 61 (1971), & Fl. Trop. E. Afr. adaxial suture more or less thickened, continuous Legum. 3, Papil. 1: 458 (1971) [Type: Tanzania. or slightly constricted between articles; articles Elton Plateau. Richards 14184 (K–holotype; A– small-sized, semiorbicular to quadrangular, isotype)]. April 2018 The Journal of Japanese Botany Vol. 93 No. 2 109

Distribution: Tanzania. var. barbata Distribution: Africa and tropical America. Grona auricoma (Grah. ex Benth.) H. var. argyrea (Baker) H. Ohashi & K. Ohashi, Ohashi & K. Ohashi, comb. nov. comb. nov. Desmodium auricomum Grah. ex Benth. in Desmodium dimorphum var. argyreum Baker Miq., Pl. Jungh. 2: 223, in adnota (1852) [Type: in Oliver & al., Fl. Trop. Afr. 2: 161 (1871) Myanmar. Tavoy. Wallich 5704 (K000858859– [Type: Angola. Huila. Welwitsch 2161 (LISU– holotype; K000858860–isotype)]; H. Ohashi in holotype, fide Schubert 1971)]. – Nicolsonia Ginkgoana 1: 234 (1973), & in Dy Phon & al., barbata var. argyrea (Baker) Schindl. in Repert. Fl. Cambodge Laos & Vietnam 27: 106 (1994). Spec. Nov. Regni Veg. 23: 359 (1927). – var. auricoma Desmodium barbatum var. argyreum (Baker) Distribution: India, Myanmar, Thailand, B. G. Schub. in Bull. Jard. Bot. État Bruxelles Cambodia, Laos, Vietnam, Indonesia (Java). 22: 298 (1952) & in Fl. Trop. E. Afr. Legum. 3, var. pseudoauricoma (H. Ohashi) H. Ohashi Papil. 1: 479 (1971). & K. Ohashi, comb. nov. Distribution: Tropical Africa, Madagascar. Desmodium auricomum v a r . var. dimorphum (Welw. ex Baker) H. Ohashi pseudoauricomum H. Ohashi in Dy Phon & al., & K. Ohashi, comb. nov. Fl. Cambodge Laos & Vietnam 27: 106 (1994), Desmodium dimorphum Welw. ex Baker ut ‘pseudo-auricomum’ [Type: Cambodia. in Oliver & al., Fl. Trop. Afr. 2: 161 (1871). E. Poilane 13923 (P–holotype MNHN- – Nicolsonia barbata var. dimorpha (Bak.) P-P02142443)]. Schindl. in Repert. Spec. Nov. Regni Veg. 23: Distribution: Cambodia, Vietnam. 359 (1927). – D. barbatum var. dimorphum (Welw. ex Baker) B. G. Schub. in Bull. Jard. Bot. Grona barbata (L.) H. Ohashi & K. Ohashi, État Bruxelles 22: 298 (1952), & in Fl. Congo comb. nov. Belge et Ruanda-Urundi 5: 205 (1954); Hepper Hedysarum barbatum L., Syst. Nat., ed. in Fl. West Trop. Afr. ed. 2. 1(2): 584 (1958); B. 10, 2: 1170 (1759) [Lectotype (designated by G. Schub. in Fl. Trop. E. Afr. Legum. 3, Papil. 1: Schubert 1971): [“Habitat in Jamaica.”] Herb. 478 (1971). Linn. No. 921.48 (LINN)]; Jarvis, Ord. Chaos Distribution: Tropical Africa, south Africa, 558 (2007). – Perrottetia barbata (L.) DC. in Madagascar. Ann. Sci. Nat. (Paris) 4: 96 (1825). – Nicolsonia var. procumbens (Baker) H. Ohashi & K. barbata (L.) DC., Prodr. 2: 325 (1825) & Mem. Ohashi, comb. nov. Leg. 7: 313 (1826); Schindl. in Repert. Spec. Desmodium barbatum var. procumbens B. Nov. Regni Veg. 23: 359 (1927). – Uraria G. Schub. in Bull. Jard. Bot. État Bruxelles 22: barbata (L.) Desv., Mém Soc. Linn. Paris 4: 309 297 (1952) [Type: Congo. W. of Lake Albert, (1826). – Desmodium barbatum (L.) Benth. in between Bunia and Gety. Lebrun 4028 (BR– Miq., Pl. Jungh. 2: 224 (1852); B. G. Schub. in holotype)], & in Fl. Trop. E. Afr. Legum. 3, Fl. Trop. E. Afr. Legum. 3, Papil. 1: 477 (1971), Papil. 1: 478 (1971). & in Ann. Missouri Bot. Gard. 67: 634 (1980); Distribution: Tropical Africa, Madagascar. McVaugh, Fl. Novo-Galiciana 5: 459 (1987); Du subsp. saulierei (Schindl.) H. Ohashi & K. Puy & Labat, Leg. Madag. 611 (2002); Lima & Ohashi, comb. nov. al. in Phytotaxa 169(1): 30 (2014). – Meibomia Desmodium barbatum var. saulierei Schind. barbata (L.) Kuntze, Revis. Gen. Pl. 1: 195 in Repert. Spec. Nov. Regni Veg. 23: 359 (1927) (1891). [Type: India. Madras, Pulney Hills. Saulière 506 subsp. barbata (K–holotype)]. – D. barbatum subsp. saulierei 110 植物研究雑誌 第 93 巻 第 2 号 2018 年 4 月

(Schindl.) H. Ohashi in Ginkgoana 1: 203 Saisorn 264 (KKU–holotype; AAU, BK, BKF– (1973). isotypes)]. Distribution: Southern India. Distribution: Thailand.

Grona birmanica (Watt ex Prain) H. Ohashi Grona brownii (Schindl.) H. Ohashi & K. & K. Ohashi, comb. nov. Ohashi, comb. nov. Desmodium birmanicum Watt ex Prain in Desmodium brownii Schindl. in Notes J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 66: 401 Roy. Bot. Gard. Edinburgh 25: 13 (1926), & in (1897) [Lectotype (designated by Ohashi 1973; Repert. Spec. Nov. Regni Veg. 22: 256 (1926) cf. pl. 28b): Myanmar. Pegu, Yomah. S. Kurz [Type (syntypes): Australia. North Coast. Brown 1677 (CAL)]. – Desmodium heterocarpon (L.) s.n. (E?); Brown 4188 BM, K, (fide Pedley DC. var. birmanicum (Watt & Prain) H. Ohashi 1999)]; Pedley in Austrobaileya 5: 239 (1999). in Ginkgoana 1: 213 (1973). Desmodium neurocarpum Benth. var. gracile Distribution: Myanmar. Benth., Fl. Austr. 2: 234 (1864) [Type: Australia. North Coast. Brown (K–holotype)]. Grona brachystachya (Kuntze) H. Ohashi & Distribution: Australia (Western Australia, K. Ohashi, comb. nov. Northern Territory, Queensland). Desmodium brachystachyum Grah. ex Benth. in Miq., Pl. Jungh. 223, in adnot. (1852), Grona capitata (Burm. f.) H. Ohashi & K. non Walpers (1842) [Type: India. Nathpur. Ohashi, comb. nov. Hamilton. ‘Hed retusum Hb. Ham … Herb. Hedysarum capitatum Burm. f., Fl. Ind.: Bentham. 1854’ (K–holotype)]; Baker in Hook. 167, t. 54. fig. 1 (1768) [Original material: f., Fl. Brit. India 2: 171 (1876). – Meibomia Burman, Fl. Ind. tab. 54, fig. 1 (1768) and brachystachya Kuntze, Revis. Gen. Pl. 1: 197 Plukenet, Amaltheum Botanicum 161. t. 433, (1891). – Nicolsonia brachystachya Schindl. in fig. 3 (1705)]. – Desmodium capitatum (Burm. Repert. Spec. Nov. Regni Veg. 23: 359 (1927). f.) DC., Prodr. 2: 336 (1825); Rugayah in – Desmodium benthamii Balakr. in J. Bombay Floribunda 1(8): 31, fig. 1 (1988); H. Ohashi in Nat. Hist. Soc. 63: 328 (1964); H. Ohashi in J. Jpn. Bot. 79(2): 120 (2004). – Ginkgoana 1: 249 (1973). capitata (Burm. f.) Hassk., Cat. Hort. Bog. 281 Distribution: Southern India. (1844), & Pl. Jav. Rar. 390 (1848). – Meibomia Kuntze (1891) published [Meibomia] capitata (Burm. f.) Kuntze, Revis. Gen. Pl. brachystachya [Bkr. “Grah.”] as a new 1: 195 (1891). – D. styracifolium (Osbeck) combination based on D e s m o d i u m Merr. sensu H. Ohashi in Ginkgoana 1: 224 brachystachyum Grah. ex Benth. (1852), (1973), p.p., incl. syn. Hedysarum capitatum, non Walpers (1842), but the name, not as a Desmodium capitatum, Pseudarthria capitata, combination, has priority from 1891 (ICN Art. Meibomia capitata; & in Dy Phon & al., Fl. 58.1, 2012). Desmodium benthamii Balakr. is Cambodge Laos & Vietnam 27: 90 (1994), p.p. superflous underGrona. Distribution: India, Thailand, Malaysia, Indonesia, Phillipines, Micronesia (Yap). Grona brevipedicellata (W. Saisorn, Chantar. & Balslev) H. Ohashi & K. Ohashi, comb. nov. Grona ferruginea (Wall. ex Thwaites) H. Desmodium brevipedicellatum W. Saisorn, Ohashi & K. Ohashi, comb. nov. Chantar. & Balsslev in Phytotaxa 263(2): 154, Desmodium ferrugineum Wall. [Numer. List fig. 2 (2016) [Type: Thailand. Sakon Nakhon: n. 5732 (1831–32), nom. nud.] ex Thwaites, Phu Phan National Park. 19 Oct. 2014. W. Enum. Pl. Zeyl. 87 (1859) [Type: Southern April 2018 The Journal of Japanese Botany Vol. 93 No. 2 111

India. Wallich Cat. n. 5732 (K-W)]; H. (CANB–holotype; BRI, DNA, K, L–isotypes)]. Ohashi in Ginkgoana 1: 204 (1973); Pedley in Distribution: Australia (Western Australia, Dassanayake, Rev. Handb. Fl. Ceylon 10: 186 Northern Territory). (1996). Distribution: Sri Lanka, India, Micronesia Grona griffithiana (Benth.) H. Ohashi & K. (Yap). Ohashi, comb. nov. Desmodium griffithianum Benth. in Miq., Grona filiformis (Zoll. & Moritzi) H. Ohashi Pl. Jungh. 222 (1852), in adnota [Lectotype & K. Ohashi, comb. nov. (designated here): India. Khasiya. Griffith s.n. Desmodium filiforme Zoll. & Moritzi in (K; Fig. 3); Griffith 352 (K–isolectotype), Natuur- Geneesk. Arch. Ned.-Indië 3: 77 (1846) Griffith 11 (CAL–isolectotype)]; Baker in Hook. [Type: Java. Sandy coast near Poeger. Zolinger f., Fl. Brit. India 2: 171 (1876); H. Ohashi in 2738 (BM–holotype, BO–isotype. fig. 3 in H. Ginkgoana 1: 207 (1973), & in Dy Phon & al., Ohashi 2004)]; Pedley in Austrobaileya 5: 238 Fl. Cambodge Laos & Vietnam 27: 132 (1994); (1999); H. Ohashi in J. Jpn. Bot. 79(2): 122, fig. P. H. Huang & H. Ohashi in Fl. China 10: 273 3 (2004). (2010). – Meibomia griffithiana (Benth.) Kuntze, Distribution: Indonesia, New Guinea, Revis. Gen. Pl. 1: 198 (1891). Australia (Western Australia, Northern Territory, Distribution: India, Myanmar, Thailand, Queensland). Laos, Vietnam, China (Guizhou, Sichuan, Yunnan). Grona flagellaris (Benth.) H. Ohashi & K. Bentham (1852) cited two specimens as ‘in Ohashi, comb. nov. Assam et Khasiya’ in the protologue. There Desmodium flagellare Benth., Fl. Austr. 2: are two sheets of ‘Khasiya Griffith’ stamped 233 (1864) [Type: Australia. Beagle Valley. ‘Herbarium Benthamianum 1854’ in K, but no Mueller s.n. in 1855 (K–holotype)]; Schindl. in specimen labeled ‘in Assam’. One of ‘Khasiya Repert. Spec. Nov. Regni Veg. 22: 255 (1926); Griffith’ bears label numbered ‘352’ and another Pedley in Austrobaileya 5: 240 (1999). without number, but these seem to be duplicate. Distribution: Australia (Western Australia, Baker (1876) cited two specimens as ‘Khasia, Northern Territory). alt. 4–6000 ft., Griffith, Hook. fil. & Thomson’, but a specimen labeled ‘Hook. fil. & Thomson’ Grona fulvescens (B. G. Schub.) H. Ohashi was not found in K. The ‘Khasia, alt. 4–6000 ft., & K. Ohashi, comb. nov. Griffith’ must be the same as ‘Khasiya Griffith’. Desmodium fulvescens B. G. Schub. in Bull. We designate here one of ‘Khasiya. Griffith’ Jard. Bot. État Bruxelles 22: 296 (1952) [Type: as the lectotype, not holotype as annotated by Congo. Haut-Katanga. Shinsenda. Ringoet 7 Ohashi in 1988, and another is isolectotype in May 1912 (BR–holotype)], & in Fl. Congo numbered 352 on the label (Fig. 3). There is Belge et Ruanda-Urundi 5: 203 (1954). another duplicate in CAL, but this bears the label Distribution: Congo. Griffith 11 (Ohashi 1973).

Grona glareosa (Pedley) H. Ohashi & K. Grona hannii (Schindl.) H. Ohashi & K. Ohashi, comb. nov. Ohashi, comb. nov. Desmodium glareosum Pedley in Desmodium hannii Schindl. in Repert. Austrobaileya 5: 242 (1999) [Type: Australia. Spec. Nov. Regni Veg. 21: 4 (1925) [Neotype Northern Territory: ca. 8 miles NNE of Edith (designated by Pedley 1999): Australia. River Siding. M. Lazarides & L. G. Adams 115 [Carpentaria. F. C. Hann (B†)]; Queensland. 112 植物研究雑誌 第 93 巻 第 2 号 2018 年 4 月

Cook District. J. R. Clarkson 4780 (BRI); Grona repens Lour., Fl. Cochinch. 2: CANB, K, MBA, QRS–isoneotypes]; Pedley in 459 (1790) [Type: Vietnam. In collihus Austrobaileya 5: 241 (1999). agrestibus Cochinchinae. J. de Loureiro s.n. Distribution: Australia (Western Australia, (BM000839659–holotype)]. Northern Territory, Queensland). Hedysarum polycarpon Poir. in Lam., Encycl. Meth. 6: 413 (1805) [Type: Indes Grona harmsii (Schindl.) H. Ohashi & K. orientales in herb. Lam (P–holotype MNHN- Ohashi, comb. nov. P-P00296540)]. – Desmodium polycarpum Desmodium harmsii Schindl. in Bot. Jahrb. (Poir.) DC., Prodr. 2: 334 (1825). Syst. 54: 54 (1916) [Type: Cochinchina. Desmodium trichocaulum DC. in Ann. In sabulosis ad caput Ti wan praefecturae Sci. Nat. 4: 101 (1805) [Type: Napaulia (v.s. Baria. L. Pierre 5821 (P–holotype HNHN- comm. a cl. Wallich)]. – D. polycarpum var. P-P02142446)]; H. Ohashi in Ginkgoana 1: 209 trichocaulon (DC.) Baker in Hook. f., Fl. Indica (1973), & in Dy Phon & al., Fl. Cambodge Laos 2: 172 (1876). & Vietnam 27: 123 (1994). Distribution: Sri Lanka, India, Nepal, Bhutan, Distribution: Vietnam. Myanmar, Thailand, Malaysia, Cambodia, Laos, Vietnam, Indonesia, Philippines, China, Taiwan, Grona helenae (Buscal. & Muschl.) H. and southern Japan, NE Australia, Pacific Ohashi & K. Ohashi, comb. nov. islands, Africa. Desmodium helenae Buscal. & Muschl. in subsp. heterocarpa Bot. Jahrb. Syst. 49: 472 (1913) [Type: Zambia. var. heterocarpa Steppe am Banguelo-See. 21 May 1910. Distribution: Sri Lanka, India, Nepal, Bhutan, ([Buscalioni & Muschler] 961. Herb. BR)]; B. Myanmar, Thailand, Malaysia, Cambodia, Laos, G. Schub. in Fl. Congo Belge et Ruanda-Urundi Vietnam, Indonesia, Philippines, China, Taiwan, 5: 183 (1954). and southern Japan, New Guinea, NE Australia, Distribution: Congo, Rhodesia. Pacific islands, Africa. var. gymnocarpa (Schindl.) H. Ohashi & K. Grona heterocarpa (L.) H. Ohashi & K. Ohashi, comb. nov. Ohashi, comb. nov. Desmodium heterocarpon var. gymnocarpum Hedysarum heterocarpon L., Sp. Pl. 2: 747 Schindl. in Repert. Spec. Nov. Regni Veg. 21: (1753) [Lectotype (designated by Pedley 1996): 5 (1925) [Type: Sri Lanka. Thwaites (B†)]; H. Ceylon. Herb. Hermann 2: 32, No. 294, left-hand Ohashi in Ginkgoana 1: 213 (1973); Pedley in specimen (BM)]; Jurvis, Ord. Chaos 560 (2007). Dassanayake, Rev. Handb. Fl. Ceylon 10: 189 – Desmodium heterocarpon (L.) DC., Prodr. 2: (1996). 337 (1825), ut heterocarpum; Merr. in Trans. Distribution: Sri Lanka, southern India. Amer. Philos. Soc. ser. 2, 24(2): 200 (1935). – B. var. strigosa (van Meeuwen) H. Ohashi & K. G. Schub. in Fl. Trop. E. Afr. Legum. 3, Papil. Ohashi, comb. nov. 1: 462 (1971); H. Ohashi in Ginkgoana 1: 210 Hedysarum siliquosum Burm. f., Fl. Ind. 169, (1973), & in J. Jpn. Bot. 66: 17 (1991), & in Dy t. 55, fig. 2 (1768). – Desmodium siliquosum Phon & al., Fl. Cambodge Laos & Vietnam 27: (Burm. f.) DC., Prodr. 2: 336 (1825). 134 (1994); Pedley in Dassanayake, Rev. Handb. Desmodium heterocarpon var. strigosum van Fl. Ceylon 10: 187 (1996), & in Austrobaileya 5: Meeuwen in Reinwardtia 6: 95 (1961) [Type: 244 (1999); H. Ohashi in J. Jpn. Bot. 79(2): 124 New Guinea. Hollandia Hamadi. C. Kalkman (2004). – Meibomia heterocarpa (L.) Kuntze, BW3596 (L–holotype)] & 6(3): 251 (1962); H. Revis. Gen. Pl. 1: 196 (1891). Ohashi in J. Jpn. Bot. 79(2): 125 (2004). April 2018 The Journal of Japanese Botany Vol. 93 No. 2 113

Fig. 3. Lectotype of Desmodium griffithianum Benth. K (TUS, photo). 114 植物研究雑誌 第 93 巻 第 2 号 2018 年 4 月

Distribution: India, Nepal, Myanmar, 3(2): 1201 (1802) [Type: ‘Sri Lanka (Ceylon).’ Thailand, Malaysia, Cambodia, Laos, Vietnam, Herb. Willdenow 13832 (B-W microfiche). Indonesia, S. China, Taiwan, Philippines, New Based on Hedysarum triflorum var. β L. (fide Guinea, N & NE Australia, New Caledonia, Pedley 1996)]. – Desmodium heterophyllum Pacific islands, Africa. (Willd.) DC., Prodr. 2: 334 (1825); H. Ohashi subsp. angustifolia (H. Ohashi) H. Ohashi & in Ginkgoana 1: 239 (1973), & in Dy Phon & K. Ohashi, comb. nov. al., Fl. Cambodge Laos & Vietnam 27: 110 Desmodium heterocarpon subsp. (1994); Pedley in Dassanayake, Rev. Handb. angustifolium H. Ohashi in Ginkgoana 1: 212 Fl. Ceylon 10: 179 (1996), & in Austrobaileya (1973), ‘subsp. angustifolium (Craib) Ohashi’, 5: 236 (1999); Du Puy & Labat, Leg. Madag. & in J. Jpn. Bot. 66(1): 17 (1991), & in Dy 615 (2002); P. H. Huang & H. Ohashi in Fl. Phon & al., Fl. Cambodge Laos & Vietnam 27: China 10: 275 (2010). – Meibomia heterophylla 140 (1994), & in J. Jpn. Bot. 79(2): 125 (2004). (Willd.) Kuntze, Revis. Gen. Pl. 1: 196 (1891). – D. reticulatum Champ. ex Benth. in Hook., Desmodium triflorum [var.] α. majus Wight Kew J. 4: 46 (1852) [Type: China. Hong Kong. & Arn., Prodr. Fl. Ind. Orient. 1: 229 (1834) Champion 254 (K–holotype)]; P. H. Huang & H. [Type: India. ‘Wight, Cat. n. 779 (partly), 781’]. Ohashi in Fl. China 10: 274 (2010). Distribution: Sri Lanka, India, Nepal, Distribution: Myanmar, Thailand, Cambodia, Myanmar, Thailand, Cambodia, Laos, Vietnam, Laos, Vietnam, S and SW China, Malaysia. S & SE China, Taiwan, Japan (Ryukyus, subsp. ovalifolia (Prain) H. Ohashi & K. Ogasawara), Pacific islands, Philippines, Ohashi, comb. nov. Indonesia, New Guinea, Australia (introduced), Desmodium polycarpum DC. var. ovalifolium New Caledonia (introduced), Madagascar (rare). Prain in J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 66: 141 (1897) [Lectotype (designated by Ohashi Grona hirta (Guill. & Perr.) H. Ohashi & K. 1994): Penang. Wallich 5730a (K-WALL)]. Ohashi, comb. nov. – D. ovalifolium Wall. [Numer. List n. 5730 Desmodium hirtum Guill. & Perr., Fl. Seneg. (1831–32), n.n.] ex Merr. in Philipp. J. Sci. C. Tent. 209 (1832) [Type: Senegal. Perrottet 247 5: 85 (1910). – D. heterocarpon var. ovalifolium (BM–isosyntype, fide Schubert 1971)]; B. G. (Wall.) Rugayah in Reinwardtia 10: 382 (1987). Schub. in Fl. Congo Belge et Ruanda-Urundi – D. heterocarpon subsp. ovalifolium (Prain) H. 5: 186 (1954); Hepper in Fl. West Trop. Afr. ed. Ohashi in J. Jpn. Bot. 66: 21 (1991), & in Dy 2. 1(2): 585 (1958); B. G. Schub. in Fl. Trop. E. Phon & al., Fl. Cambodge Laos & Vietnam 27: Afr. Legum. 3, Papil. 1: 457 (1971); Du Puy & 139 (1994), & in J. Jpn. Bot. 79(2): 126 (2004). Labat, Leg. Madag. 615 (2002). Distribution: Thailand, Cambodia, Laos, Distribution: Tropical Africa, Madagascar. Vietnam, S and SW China, Malaysia. Grona jucunda (Thwaites) H. Ohashi & K. Grona heterophylla (Willd.) H. Ohashi & K. Ohashi, comb. nov. Ohashi, comb. nov. Desmodium jucundum Thwaites, Enum. Pl. Hedysarum triflorum [var.] β L., Sp. Pl. 2: Zeyl. 411 (1864) [Type: Sri Lanka (Ceylon). 749 (1753) [Type: β from Sri Lanka (fide Pedley Matelle District, Lakkaigalla. A. Oswald 1996)]. Brodie 3778 (K000858872–holotype)]; H. Hedysarum triflorum [var.] γ L., Sp. Pl. 2: Ohashi in Ginkgoana 1: 216 (1973); Pedley in 750 (1753) [Type: γ from India (fide Pedley Dassanayake, Rev. Handb. Fl. Ceylon 10: 185 1996)]. (1996). – Meibomia jucunda (Thwaites) Kuntze, Hedysarum heterophyllum Willd., Sp. Pl. Revis. Gen. Pl. 1: 198 (1891). April 2018 The Journal of Japanese Botany Vol. 93 No. 2 115

Distribution: Sri Lanka. Desmodium nemorosum var. simplex subvar. whitfordii Schindl. in Repert. Spec. Nov. Regni Grona juruenensis (Hoehne) H. Ohashi & Veg. 21: 10 (1925) [Lectotype (designated by K. Ohashi, comb. nov. Ohashi 2004): Phillippines. Luzon. Prov. Bataan, Desmodium juruenense Hoehne, Relat. Mt. Mariveles, Lamo River. H. N. Whitford 227 Commiss. Linhas Telegr. Estratég. Matto Grosso (K); CAL, NY, US–isotypes]. – D. nemorosum Amazonas Annexo 5, Bot. viii. 73, t. 148, f. 1 var. whitfordii (Schindl.) H. Ohashi in Ginkgoana (1919); Lima & al. in Phytotaxa 169(1): 55, fig. 1: 220 (1973). – D. whitfordii (Schindl.) Pedley 18 (2014) [Lectotype (designated by Lima et al. in Austrobaileya 5: 247 (1999). – D. nemorosum 2014): Brazil. Mato Grosso: Salto Augusto. Jan. subsp. whitfordii (Schindl.) H. Ohashi in J. Jpn. 1915. J. K. Kuhlmann 460 (R)]. – Meibomia Bot. 79(2): 128 (2004). juruenense (Hoehne) Hoehne in Anexos Mem. Distribution: Philippines. Inst. Butantan, Secc. Bot. 1, fasc. 1: 33, pl. 11 (1921). – Nicolsonia juruenense (Hoehne) Grona orinocensis (DC.) H. Ohashi & K. Schindl. in Repert. Spec. Nov. Regni Veg. 23: Ohashi, comb. nov. 359 (1928). Desmodium adscendens orinocense Distribution: Brazil. DC., Prodr. 2: 332 (1825) [Type: Humboldt, Bonpland & Kunth, Nov. Gen. Sp. 6: tab. 597 Grona muelleri (Benth.) H. Ohashi & K. (1823)]. – Nicolsonia orinocensis (DC.) Schindl. Ohashi, comb. nov. in Repert. Spec. Nov. Regni Veg. 23: 358 (1927). Desmodium muelleri Benth., Fl. Austr. 2: Distribution: Venezuela. 235 (1864) [Type: Australia. Upper Victoria River. Mueller s.n. (K–holotype)]; Pedley in Grona pullenii (Pedley) H. Ohashi & K. Austrobaileya 5: 241 (1999). – Meibomia Ohashi, comb. nov. muelleri (Benth.) Kuntze, Revis. Gen. Pl. 1: 198 Desmodium pullenii Pedley in Austrobaileya (1891). 5: 243 (1999) [Type: Australia. Northern Distribution: Australia (Western Australia, Territory, 12 km S of Hayes Creek. April 1988. Northern Territory, Queensland). R. Pullen & Spottswood 11169 (CANB– holotype; BRI, DNA, NSW, PERTH–isotypes)]. Grona nemorosa (F. Muell. ex Benth.) H. Distribution: Indonesia (Lesser Sunda Ohashi & K. Ohashi, comb. nov. Islands) and Australia (Western Australia, Desmodium nemorosum F. Muell. ex Northern Territory, Queensland). Benth., Fl. Austr. 2: 234 (1864) [Syntypes in K, fide Pedley (1999): Australia. Brisbane Grona pycnotricha (Pedley) H. Ohashi & K. River. Mueller; Pine River. Fitzalan; sine loc. Ohashi, comb. nov. Leichhardt]; Pedley in Austrobaileya 5: 247 Desmodium pycnotrichum Pedley in (1999); H. Ohashi in J. Jpn. Bot. 79(2): 127 Austrobaileya 5: 236 (1999) [Type: Australia. (2004). – Meibomia nemorosa (F. Muell. ex Northern Territory. Coastal Plains Research Benth.) Kuntze, Revis. Gen. Pl. 1: 198 (1891), Station. M. Lazarides 6828 (CANB–holotype; p.p. BRI, K, MEL–isotypes)]. subsp. nemorosa Distribution: Australia (Western Australia, Distribution: New Guinea, Australia Northern Territory). (Queensland, New South Wales). subsp. whitfordii (Schindl.) H. Ohashi & K. Grona ramosissima (G. Don) H. Ohashi & Ohashi, comb. nov. K. Ohashi, comb. nov. 116 植物研究雑誌 第 93 巻 第 2 号 2018 年 4 月

Desmodium ramosissimum G. Don, Gen. in Dy Phon & al., Fl. Cambodge Laos & Syst. 2: 294 (Oct. 1832) [Type: W. Trop. Africa. Vietnam 27: 128 (1994). St. Tomas (S. Tomè). G. Don (BM–holotype, Distribution: Cambodia, Vietnam. fide Schubert 1963, 1971)]; B. G. Schub. in J. Arn. Arb. 44: 293 (1963), & in Fl. Trop. E. Grona schweinfurthii (Schindl.) H. Ohashi Afr. Legum. 3, Papil. 1: 464 (1971); Du Puy & & K. Ohashi, comb. nov. Labat, Leg. Madag. 618 (2002). Desmodium schweinfurthii Schindl. in Bot. Distribution: Tropical Africa (Uganda, Jahrb. Syst. 54: 57 (1916) [Syntypes: Im Lande Kenya, Tanganyika, Zambia), Madagascar, der Djur, gr. Seriba Ghattas. Schweinfurth n. II. Mascarene Is. 48 (B†), etc.]; B. G. Schub. in Fl. Congo Belge et Ruanda-Urundi 5: 184 (1954). Grona rubra (Lour.) H. Ohashi & K. Ohashi, Distribution: Congo. comb. nov. Ornithopus ruber Loureiro, Fl. Cochinch. Grona setigera (E. Mey.) H. Ohashi & K. 452 (1790) [Type: Cochinchina. Loureiro s.n. Ohashi, comb. nov. (n.v.)]. – Desmodium rubrum (Lour.) DC., Prodr. Nicolsonia setigera E. Mey., Comm. Pl. 2: 327 (1825); H. Ohashi in Dy Phon & al., Fl. Afr. Austr. (Meyer) 1: 124 (1836) [Type: Cambodge Laos & Vietnam 27: 124 (1994), & South Africa, Cape Province, Umzimvubu in J. Jpn. Bot. 79(2): 128 (2004). – Meibomia [Omsamwubo] R., Drege [446] (W, ?iso.), fide rubra Kuntze, Revis. Gen. Pl. 1: 198 (1891). Schubert (1971)]. – Desmodium setigerum (E. var. rubra Mey.) Harv. in Fl. Cap. 2: 229 (1862); B. G. Distribution: Cambodia, Vietnam, China Schub. in Fl. Congo Belge et Ruanda-Urundi (Guangdong, Hainan). 5: 187, fig. 11/E (1954), & in Fl. Trop. E. Afr. var. macrocarpa (H. Ohashi) H. Ohashi & Legum. 3, Papil. 1: 460 (1971). K. Ohashi, comb. nov. Distribution: Cameroun, Congo, Ghana, Desmodium rubrum var. macrocarpum H. Kenya, Liberia, Malawi, Mozambique, Nigeria, Ohashi in Ginkgoana 1: 222 (1973) [Type: Rhodesia, South Africa, Tanganyika, Uganda, Vietnam. Tourane. J. & M. S. Clemens 3096 (P– Zambia. holotype)], & in Dy Phon & al., Fl. Cambodge Laos & Vietnam 27: 128 (1994). Grona stolzii (Schindl.) H. Ohashi & K. Distribution: Vietnam. Ohashi, comb. nov. var. uncinata (H. Ohashi) H. Ohashi & K. Desmodium stolzii Schindl. in Repert. Spec. Ohashi, comb. nov. Nov. Regni Veg. 21: 9 (1925) [Lectotype Desmodium rubrum var. uncinatum H. (designated by Schubert 1971): Tanganyika. Ohashi in Dy Phon & al., Fl. Cambodge Laos & Masukulu Forest. Stolz 745 (G)]; B. G. Schub. Vietnam 27: 128 (1994) [Type: Vietnam. Thuân in Fl. Trop. E. Afr. Legum. 3, Papil. 1: 463 Hai, Ca Na. Poilane 9101 (P–holotype)]. (1971). Distribution: Vietnam. Distribution: Tropical Africa (Burundi, Congo, Malawi, Tanzania, Zambia, Zaire, Grona schubertiae (H. Ohashi) H. Ohashi & Zimbabwe). K. Ohashi, comb. nov. Desmodium schubertiae H. Ohashi in Grona styracifolia (Osbeck) H. Ohashi & K. J. Arnold Arbor. 71(3): 381 (1990) [Type: Ohashi, comb. nov. Vietnam. Saigon, November 1864. Poilane Hedysarum styracifolium Osbeck, Dagbok 32489 (P–holotype; TUS–isotype)]; H. Ohashi Ostind. Resa: 247 (1757) [Type: From China. April 2018 The Journal of Japanese Botany Vol. 93 No. 2 117

Dane’s island, near Whampoa. Osbeck s.n. 25 Encycl. 4(2): 451 (1798). – Pleurolobus triflorus Oct. 1851 (P), fide Merrill (1916)]. – Nicolsonia (L.) J. St.-Hil., Nouv. Bull. Soc. Philom. 3: 192 styracifolia (Osb.) Desv. in Ann. Sci. Nat. (Paris) (1812). – Desmodium triflorum (L.) DC., Prodr. 9: 418 (1826). – Desmodium styracifolium 2: 334 (1825); B. G. Schub. in Fl. Congo Belge (Osbeck) Merr. in Am. J. Bot. 3: 580 (1916); 5: 187 (1954), & in Fl. Trop. E. Afr. Legum. 3, Enum. Philipp. Fl. Pl. 2: 289 (1923); H. Ohashi Papil. 1: 459 (1971); H. Ohashi in Ginkgoana 1: in Ginkgoana 1: 224 (1973), p.p., excl. syn. 245 (1973), & in Dy Phon & al., Fl. Cambodge Hedysarum capitatum, Desmodium capitatum, Laos & Vietnam 27: 111 (1994); Pedley in Pseudarthria capitata, Meibomia capitata, et Austrobaileya 5: 235 (1999); B. G. Schub. in D. celebicum; H. Ohashi in Dy Phon & al., Fl. Ann. Missouri Bot. Gard. 67: 660 (1980); H. Cambodge Laos & Vietnam 27: 90 (1994); Ohashi in J. Jpn. Bot. 79(2): 131 (2004); Du Puy Pedley in Dassanayake, Rev. Handb. Fl. Ceylon & Labat, Leg. Madag. 620 (2002); Lima & al. in 10: 181 (1996); H. Ohashi in J. Jpn. Bot. 79(2): Phytotaxa 169(1): 91, fig. 33 (2014). – Sagotia 130 (2004). triflora (L.) Duchass. & Walp. in Linnaea 23: Distribution: Sri Lanka, India, Myanmar, 738 (1850). – Nicolsonia triflora (L.) Griseb. Thailand, Malaysia, Cambodge, Laos, Vietnam, in Abh. Königl. Ges. Wiss. Göttingen. 7: 202 S. China, Philippines, Indonesia (Sulawesi). (1857). – Meibomia triflora (L.) Kuntze, Revis. Gen. Pl. 1: 197 (1891) with many infraspecific Grona tiwiensis (Pedley) H. Ohashi & K. taxa (cf. B. G. Schubert 1980). Ohashi, comb. nov. Hedysarum stipulaceum Burm. f., Fl. Desmodium tiwiense Pedley in Austrobaileya Ind. 168, t. 54, fig. 2 (1768). – Desmodium 5: 248 (1999) [Type: Australia. Northern stipulaceum (Burm. f.) Hassk., Cat. Pl. Hort. Territory: Bathurst Island, Runku. I. D. Cowie Bot. Bogor.: 274 (1844). 5424 (DNA–holotype; BRI, K, etc.–isotypes)]. Desmodium triflorum var. β minus Wight & Distribution: Australia (Northern Territory: Arn., Prodr. Fl. Ind. Orient. 1: 229 (1834) [Type: Bathurst Island). India. ‘Wight, Cat. n. 777, 779 (partly)’], & var. γ villosum Wight & Arn., l.c. 1: 229 (1834) Grona trichostachya (Benth.) H. Ohashi & [Type: India. ‘Wight, Cat. n. 778’]. K. Ohashi, comb. nov. Distribution: Worldwide in tropics. Native in Desmodium trichostachyum Benth., Fl. tropical Asia and Africa. Austr. 2: 234 (1864) [Lectotype (designated by Pedley 1999): Australia. Port Essington. Grona wittei (B. G. Schub.) H. Ohashi & K. Armstrong (K)]; H. Ohashi in Ginkgoana 1: 244 Ohashi, comb. nov. (1973); Pedley in Austrobaileya 5: 237 (1999). Desmodium wittei B. G. Schub. in Bull. Distribution: New Guinea, Australia (Western Jard. Bot. État Bruxelles 22: 292 (1952) [Type: Australia, Northern Territory, Queensland). Congo. Upper Katanga. Parc Nat. Upemba, Lupiala. G. de Witte 2918 in Oct. 1947 (BR– Grona triflora (L.) H. Ohashi & K. Ohashi, holotype)]. comb. nov. Distribution: Congo. Hedysarum triflorum L., Sp. Pl. 2: 749 (1753), p.p., excl. var. β & var. γ [Lectotype Grona wynaadensis (Bedd. ex Gamble) H. (designated by Fawcett and Rendle 1920): Ohashi & K. Ohashi, comb. nov. India. Herb. Hermann 1: 21, No. 297 Desmodium wynaadense Bedd. ex Gamble, (BM000594452)]; Jarvis, Ord. Chaos 562 Fl. Madras 2: 343 (key) and 346 (1918) & (2007). – Aeschynomene triflora (L.) Poir., in Kew Bull. 1919: 223 (1919) [Lectotype 118 植物研究雑誌 第 93 巻 第 2 号 2018 年 4 月

Fig. 4. Lectotype of Desmodium wynaadense Bedd. ex Gamble. K000858771 (©The Board of Trustees of the Royal Botanic Gardens, Kew). April 2018 The Journal of Japanese Botany Vol. 93 No. 2 119

(designated here): India. Wynaad. Beddome Plantae Junghuhnianae. pp. 205–269. Sythoff. Leiden. 3/80 [Nilgiri hills: Carcoor Ghat (March 1880)] Bentham G. 1865. Leguminosae. In: Bentham G. and Hooker J. D., Genera Plantarum 1: 434–600. Reeve & (K000858771)]; Pedley in Dassanayake, Co., London. Rev. Handb. Fl. Ceylon 10: 187 (1996). – Brummitt R. K. 1978. Report of the committee for D. ferrugineum Wall. ex Thwaites subsp. Spermatophyta, 20. Taxon 27(2/3): 285–314. wynaadense (Bedd. ex Gamble) H. Ohashi in Candolle A. P. de 1825. Ordo LXIII. Leguminosae Juss. In: Ginkgoana 1: 206 (1973). Prodromus Systematis Naturalis Regni Vegetabilis 2: 93–524. Treuttel et Würtz, Paris. Distribution: Southern India (W. Ghats, Dy Phon P., Ohashi H. and Vidal J. E. 1994. Leguminosae Nilgiri, Anamalai, Travancore Hills). (Fabaceae), Papilionoideae, Desmodieae. In: Morat Gamble (1918) published Desmodium Ph., Flore du Cambodge du Laos et du Viêtnam. 27. wynaadense Bedd. with diagnosis of the species Muséum National d’Histoire Naturelle Laboratoire du Phanérogamie. Paris. in the key to the Desmodium species and a note Gamble J. S. 1918. Flora of the Presidency of Madras. under the name together with four regions of its 1(2). Celastraceae to Leguminosae-Papilionatae. West distribution, but did not cite specimens in the Newman & Co., Adlard & Son, London. protologue. The species was published again Hutchinson J. 1964. Genera of Flowering with description with six specimens under the (Angiospermae) based principally on the Genera Plantarum of G. Bentham and J. D. Hooker. vol. 1. regions in the following year. These specimens Oxford University Press, London. correspond to the regions cited in the protologue. Jabbour F., Gaudeul M., Lambourdière J. Ramstein They are, therefore, regarded as the original G., Hassanin A., Labat J-N. and Sarthou C. 2018. material as indicated in ICN 9.3 (MacNeill et Phylogeny, biogeography and character evolution in the tribe Desmodieae (Fabaceae: Papilionoideae), al. 2012). We select ‘Nilgiri hills: Carcoor Ghat with special emphasis on the New Caledonian endemic (March 1880), Beddome (K) as the lectotype genera. Mol. Phylogenet. Evol. 118: 108–121. of Desmodium wynaadense Bedd. ex Gamble. Kajita T., Ohashi H., Nemoto T., Bailey C. D. and Doyle (Fig. 4). J. J. 1996. Polyphyly of the legume tribe Desmodieae: evidence from the chloroplast genome. Amer. J. Bot. 83: 165–166 (abstract). Grona zenkeri (Schindl.) H. Ohashi & K. Kajita T., Ohashi H., Tateishi Y., Bailey C. D. and Doyle J. Ohashi, comb. nov. J. 2001. rbcL and legume phylogeny, with particular Desmodium zenkeri Schindl. in Repert. reference to Phaseoleae, Millettieae and allies. Syst. Spec. Nov. Regni Veg. 21: 8 (1925) [Type: Bot. 26: 515–536. Lima L. C. P., Queiroz L. P. de, Tozzi A. M. G. A. and Cameroons. Zenker 1615, 4787, Lotz 5 (B†– Lewis G. P. 2014. A taxonomic revision of Desmodium syntypes)]. (Leguminosae, Papilionoideae) in Brazil. Phytotaxa Distribution: Cameroons. 169(1): 1–119. Loureiro J. de 1790. Flora Cochinchinensis: sistens plantas in regno Cochinchina nascentes. Ulyssipone. We wish to express our thanks to the curators McNeill J., Barrie F. R., Buck W. R., Demoulin V., Gtriyrt and directors of the following herbaria: A, GH, W., Hawksworth D. L., Herendeen P. S., Knapp S., BM, BO, CAL, E, K, KYO, L, TI and TUS Marhold K., Prado J., Prud’homme van Reine, W. for allowing us to examine their collection. F., Smith G. F., Wiersema J. H. and Turland N. J. We thank Professor T. Nemoto of Ishinomaki (eds.) 2012. International Code of Nomenclature for algae, fungi, and plants (Melbourne Code). Regnum Shenshu University for his help on our studies Vegetabile 154. Koeltz Scientific Books, Königstein. and comments to the manuscript and Dr. B. Ye Merrill E. D. 1935. A comentary on Loureiro’s “Flora of Tohoku University for providing her pictures Cochinchinensis.” Trans. Amer. Philos. Soc. n. ser. of pollen grains used in this paper. 24(2): 1–445. Ohashi H. 1973. The Asiatic Species of Desmodium and Its Allied Genera (Leguminosae). Ginkgoana 1. Academia References Scientific Book, Inc., Tokyo. Bentham G. 1852. Leguminosae. In: Miquel F. A. W., Ohashi H. 1999. The genera, tribes and subfamilies of 120 植物研究雑誌 第 93 巻 第 2 号 2018 年 4 月

Japanese Leguminosae. Sci. Rep. Tohoku Univ., 4th Ohashi H., Nemoto T. and Wu T. L. 1996. The taxonomic ser. (Biology) 40: 186–269. position of Trifidacanthus (Leguminosae). J. Jpn. Bot. Ohashi H. 2003. A new circumscription of and a 71(2): 57–66. new genus Akschindlium (Leguminosae). J. Jpn. Bot. Ohashi H. and Ohashi K. 2012a. Ototropis, a genus 78: 269–294. separated from Desmodium (Leguminosae). J. Jpn. Bot. Ohashi H. 2004a. and distribution of 87(2): 108–118. Desmodium and related genera (Leguminosae) in Ohashi H. and Ohashi K. 2012b. Verdesmum, a new genus Malesia (I). J. Jpn. Bot. 79: 155–185. of Leguminosae: tribe Desmodieae. J. Jpn. Bot. 87(5): Ohashi H. 2004b. Taxonomy and distribution of 299–306. Desmodium and related genera (Leguminosae) in Schindler A. K. 1928. Die Desmodiinen in der botanischen Malesia (II). J. Jpn. Bot. 79: 155–185. Literatur nach Linne. Repert. Spec. Nov. Regni Veg. Ohashi H. 2005. Desmodieae. In: Lewis G., Schrire B., Beih. 49: 1–371. Mackinder B. and Lock M. (eds.), Legumes of the Schubert B. G. 1980. Desmodium. In: Dwyer J. D. and World. pp. 433–446. Royal Botanic Gardens, Kew. collaborators. Flora of Panama, part 5, Fam. 83 Ohashi H. and Mill R. R. 2000. Hylodesmum, a new name Leguminosae subfam. Papilionoideae. Ann. Missouri for Podocarpium (Leguminosae). Edinburgh J. Bot. 57: Bot. Gard. 67(3): 622–662. 171–188.

a b 大橋広好 ，大橋一晶 ：マメ科シバハギ属の新学名 マメ科ヌスビトハギ連 Desmodieae は 32 属から構成 ギである．このため，Grona の和名をシバハギ属とし， される．Ohashi (2005) はヌスビトハギ連を Lespedeza Desmodium の和名をアコウマイハギ属に再度変更す 群（3 属），Phyllodium 群（12 属）および Desmodium る．和名アコウマイハギは台湾の阿猴（現屏東県屏東 群（17 属）の 3 群に分けた．しかし，分子系統学的解 市）で発見され，早田文蔵によって Desmodium akoense 析からは Desmodium 群内は単系統ではない属を含むこ Hayata に付けられた．この学名は D. scorpiurus (Sw.) とが明らかにされている（Kajita et al. 1996，［2010 未 Desv. の異名であるが，この種は属名 Desmodium のタ 発表］，Jabbour et al. 2018）．われわれは分子系統学的解 イプであるため，アコウマイハギ属は Desmodium の 析をさらに進めた結果，Desmodium 群のうち属として 安定した和名となる．アコウマイハギ D. scorpiurus Desmodium にまとめられていた属内分類群の多くが多 (Sw.) Desv. は日本では初島住彦『琉球植物誌』(1971) 系統であることを確認した (Ohashi et al. in press). これ に伊江島への帰化が記録されている．本論文では 41 までに旧ヌスビトハギ属からヌスビトハギなどを別属 種 13 種内分類群を新たにシバハギ属と認め，新組み Hylodesmum に移し，その和名もヌスビトハギ属とし， 合わせの学名を与えた．その中で日本産種はシバハギ 元の Desmodium をシバハギ属に改めた（大橋広好，本 Grona heterocarpa (L.) H. Ohashi & K. Ohashi， カ ワ リ 誌 77(1): 59–60, 2002）．しかし，シバハギ Desmodium バマキエハギ Grona heterophylla (Willd.) H. Ohashi & K. heterocarpon (L.) DC. を含む Desmodium sect. Nicolsonia Ohashi，ハイマキエハギ Grona triflora (L.) H. Ohashi & とカワリバマキエハギ D. heterophyllum Willd. を含む K. Ohashi の 3 種である． Desmodium sect. Sagotia とは分子系統学的解析の結果 （a東北大学植物園津田記念館， から Desmodium から除かれ，属として Grona にまと b岩手医科大学薬学部） まることが明らかとなった．Grona のタイプはシバハ