A MONOGRAPH OF THE SUBGENUS DOLLINERA
OF THE GENUS DESMODIUM
(LEGUMINOSAE)
by
Hiroyoshi OHASHI
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Dollinera is an Asiatic subgenus of the genus Desmodium. The subgenus is characterized largely in having monadelphous stamens and many-jointed indehiscent pods. Most of the species of the subgenus are distributed in the subtropical and warm regions from the Himalayas eastwards through Burma, N. Thailand to S. W. China (mostly in Yunnan and Szechuan). Several species, D. megapfiyllum, D. multiflorum and D. sequax, extend their geographical areas from the Himalaya China region southwards to Malaysia and/or eastwards to Taiwan. Dollinera was established by Endlicher as a distinct genus in 1840 distinguished from the genus Desmodium mainly by the monadelphous stamens, but he did not express what species were included in his new genus. In 1842, however, Walpers adopted Endlicher's genus citing two species, Dollinera, rufescens Endl. and Dollinera sambuensis Endl. Before the founding of Dollinera as a genus these two species had been treated generally under the genus Desmodium. Bentham in 1852 merged the Endlicher's genus with Desmodium as a section and this view of the status of Dollinera had usually been accepted by the subsequent taxonomists (Bentham 1865, Baker 1876, Taubert 1894, Dalla Torre & Harms 1900-1907, Merrill 1910). The study by Schindler (1926) is the only revision of Dollinera of the genus Desmodium. He elevated it to subgeneric status and divided it into three sections, i.e. sect. Tiliifoliae, Floribundae and Khasianae, based on the presence or absence of the bracts and bracteoles. He described a number of new species and consequently recognized 21 species within his subgenus, of which one species, D. elegans (as D. tiliifolium), was subdivided into three varieties and five forms. In the present paper the subgenus Dollinera is delimited newly to include several species hitherto not placed properly in the genus Desmodium, and is classified in four sections, i.e. sect. Dollinera, Kingianae, Sequax and Siamensia. The section Kingianae and the section Siamensia were not recognized as members of the subgenus by Schindler. The only representative of the section Siamensia, D. siamense, was described by Schindler under the genus Phyllodium as P. siamense, but the species is considered to be better placed in the subgenus. The section Kingianae comprising two species is a rather isolated group in the structure of pods and the geographical preference, but seems to be natural to include in this subgenus due to the similarity in the vegetative, floral and pollen morphology. Among the subgenus the monotypic section Sequax is apparently distinct mainly in the structure of flower and pollen grains. The section Dollinera, the largest and the most complicated group comprising nine species, is, furthermore, split to three subsections, i.e. subsect, Dollinera, Khasianae and Tiliifoliae. The present study aims to provide a basic revision of the subgenus Dollinera as a part of my study of the Asiatic Desmodium. All species treated in the revision have detailed descriptions of their external and pollen morphology together with a number of illustrations of the specimens, petals, pods, seeds and pollen grains. The seeds and pollen grains are described in this paper mostly for the first time. This revision is based mainly upon studies on the specimens of the following herbaria: Arnold Arboretum, Harvard University, Cambridge, Massachusetts, U.S.A. (A); Botanical Section, Plant Industry Division, Department of Agri culture, Bangkok, Thailand (BK); The Forest Herbarium, Royal Forest Department, Bangkok, Thailand (BKF); British Museum (Natural History), London, Great Britain (BM); Central National Herbarium, Botanical Survey of India, Calcutta, India (CAL); Royal Botanic Gardens, Edinburgh, Great Britain (E); Gray Herbarium, Harvard University, Cambridge, Massachusetts, U.S.A. (GH); The Herbarium and Library, Royal Botanic Gardens, Kew, Great Britain (K); Department of Botany, Faculty of Science, Kyoto University, Kyoto, Japan (KYO); The New York Botanical Garden, New York, U.S.A. (NY); Mus éum National d'Histoire Naturelle, Laboratoire de Phanérogamie, Paris, France (P); Department of Botany, Faculty of Science, University of Tokyo, Tokyo, Japan (TI); The National Science Museum, Tokyo, Japan (TNS); Herbarium of the University of California, Department of Botany, University of California, Berkeley, California, U.S.A. (UC). Abbreviations of the herbarium names are those of the Index Herbariorum, Part I (Lanjouw and Stafleu,ed. 5, 1964). I wish to express my sincere thanks to the Directors of all these Institutions for their kind cooperation. I particularly indebted to Professor Hiroshi Hara, Depart ment of Botany, University of Tokyo, for his valuable suggestions and criticisms. Desmodium subgen. DOLLINERA (Endl.) Schindl. in Fedde, Rep. 22: 262 (1926). Shrubs or occasionally small trees, usually much branched, young branches pubescent and often angular, old ones usually glabrescent and terete. Leaves 3- foliolate or rarely 3- and 1-foliolate, petiolate, stipulate; when 3-foliolate terminal leaflets petiolulate and 2-stipelate, lateral leaflets sessile (but the leaflet-pulvini evident, usually more densely pubescent) and 1-stipelate; when 1-foliolate leaves slightly petiolulate but stipelate. Stipules often early deciduous, usually falcate, scariose, striate, free, pubescent and often minutely puberulent on the outside, glabrous but usually sparsely puberulent at the base of the inside, pubescent along the margin; stipeis scariose, striate, those of the terminal leaflets often somewhat smaller than those of the laterals, ciliate and pubescent outside, glabrous inside. Inflorescences usually terminal and axillary but rarely only axillary (in sect. Siamensia), the terminals on young shoots and branches usually paniculate or occasionally racemose (in sect. Sequax, sect. Siamensia, and D. khasianum), when paniculate varying greatly in form, the axillary ones from the axils of upper leaves usually racemose but sometimes paniculate (in spiciform). Flowers usually fascicu late but sometimes umbellate, usually binate or ternate but often appearing simul taneously 1-3(-4) flower-bud (especially in D. elegans, D. megaphyIlum, D.yunnan ense & D. Rockii), subtended by the bracts. Bracts often early deciduous, scariose, striate, usually dimorphic, i.e. primary and secondary bracts, pubescent and often minutely puberulent outside, more densely pubescent along the margin, glabrous inside; the primary bract 1, subtending the secondary bracts which subtending the fascicles of flowers; secondary ones present or rarely absent, usually 2 but often appearing 1-4 poorly developed smaller ones which subtending the flower-buds. Bracteoles 2 or reduced to none, usually at or near the base of the calyx but very rarely on midway of the pedicel, when present usually early deciduous, scariose, striate, subulate or narrowly triangular or narrowly ovate, pubescent outside and along the margin, glabrous inside. Calyx narrowly campanulatc, pubescent, 4- lobed, the lobes equal or unequal in length, the upper (=adaxial or dorsal) lobe entire or minutely 2-toothed at the apex (only in sect. Sequax rather distinctly 2- toothed), the laterals usually triangular, often shorter than. the others, the lower (=abaxial or ventral) one often longer than the others. Standard not auricled, shortly clawed or cuneately tapering to the point, wings inflexed or deflexed, usually more or less auriculate at the upper base of lamina, obtuse at the apex; wings and keel-petals distinctly clawed, thinly membranous, ofte faintly adherent to each other; keel-petals folded down the base of lamina or the upper of the claw when anthesis, hence usually distinctly shorter than the wings, inflexed, usually swollen to the outside near the middle of lamina, often with a linear appendage on the outside. Stamens monadelphous, the vexillary one free usually above the middle but occasionally below the half (in sect. Sequax), the other united most of their length, free part of the filaments unequal in length, glabrous. Pistils slender, usually pubescent; style bent at right angles to the ovary or rarely not so bent (occasionally in sect. Sequax) and exserted from the androecium, not thickened in the upper part, glabrous; stigma minute, capitate. Pods (Figs. 29, 40 & 48) narrowly oblong, indehiscent, sessile or stalked, more than 5-jointed, usually compressed but rarely nearly moniliform (only in sect. Sequax), glabrous or pubescent, the upper (=adaxial) suture shallowly undulate or nearly straight, the lower (=abaxial) suture usually undulate but occasionally not or slightly undulate (in sect. Siamensia and sect. Kingianae), calyx persistent at the base; joints nearly once or twice as long as broad, transversely to transversely broadly oblong or elliptic or nearly quadrangular. Seeds (Figs. 29, 40 & 48) transversely elliptic or depressed obovate, hilum with thickened ring-like margin, i.e. rim-arillate. Seedlings: Cotyledons epi gaeous, elliptic, sessile, opposite at the first node; the first and second leaves simple and opposite at the second node, stipulate, broadly ovate; usually succeeding leaves 3-foliolate, alternate, but occasionally the third and the forth opposite and succeedings alternate. Pollen grains (Pls. 13-14 & Fig. 41) tricolporate; colpi long, narrow or opened, with or without margin, often constricted and protruding at the equator, remaining parts usually bordered by the endexine thickening, the membrane granulated; pores rounded or elliptic, when elliptic usually equatorially or rarely meridionally elongated; usually protruding or occasionally flattened, slightly marginate, the membrane granulated or not; exine mostly entire or rarely undulate (in sect. Sequax), mostly fine reticulate or very rarely fine reticulate mixed partly with verrucate or rugulate sculptures (in sect. Sequax), when reticulate lumina 1 µ or below in diameter, tectate, 1-2 µ thick, the ektexine thicker than the endexine, muri simplibaculate, columellae short.
Key to the sections of the subgenus Dollinera.
1. Sect. DollineraShrubs or small trees. Leaves usually 3-foliolate but rarely mixed with 1-foliolate leaves (only in D. yunnanense); stipules usually narrowly ovate or narrowly triangular. Inflorescences usually terminal and axillary but occasionally only terminal (in D. yunnanense and D. Rockii). Flowers usually fasciculate but sometimes umbellate. Bracts dimorphic or rarely trimorphic (i.e. primary, secondary and third bracts); secondary bracts present or absent (in D. khasianum); third bracts appearing only in umbellet, very small in size. Calyx 4-lobed, the upper lobe entire or minutely 2-toothed at the apex. Wings longer than the keel-petals, auriculate or not; keel petals auriculate or not (in subsect. Dollinera and subsect. Khasianae). Pods sessile or shortly stalked, usually pubescent with straight, appressed hairs but rarely (in D. megaphyllum} both straight and hooked hairy; lower sutures undulate or rather deeply indented (only in D. confertum); joints longer than. broad, more than 3 mm wide. Type species: D. multiflorum DC. Key to the subsections of the section Dollinera.
1-(1). Subsect. Dollinera Key to the species of subsection Dollinera.
1) Desmodium amethystinum Dunn in Gard. Chron. ser. 3, 32: 210 (1902) -Craib, Fl. Siam. Enum. 1: 4-03 (1928). [Fig. 27a]
A shrub, usually up to 2 m high; young branches angular and very densely softly white or brown-velutinous. Stipules rather persistent, narrowly ovate with an acuminate apex, 12-15 mm by 2.5-3.5 mm in size, very densely appressed velutinous outside. Petioles (1.5-)3-5 cm long, very densely velutinous like the young branches. Leaves 3-foliolate; leaflets subcoriaceous, elliptic or ovate, acute or mucronate at the apex, nearly entire along the margin, the upper surface densely velutinous but becoming sparsely hairy with age, the lower surface very densely uniformly white or brown-velutinous, lateral nerves 8-10 on each side of the midrib and reaching the margin, reticulate-veins inconspicuous; stipels narrowly triangular, 3-6 mm long, 1-2 mm wide at the base, rather sparsely sericeous outside; terminal leaflets usually cuneately obtuse at the base, (5-)7-12 cm long, (3.5-)4-6(-6.5) cm wide; petiolules 1-2 cm long, pubescent like the petioles; lateral leaflets obliquely rotund at the base, (4-)6-8(-14) cm long, (2.5-)3-5 cm wide. Inflorescences terminal and axillary; terminals paniculate, copiously branched, 15-25 cm long; axillary ones racemose; rachis very densely velutinous like the young branch but decreasing gradually the degree of pubescence (usually in rachilla densely spreading velutin ous) with age. Flowers mostly 2- but occasionally 3-flowered; flowering pedicels 4-6 mm long, fruiting pedicels 8-10 mm long, rather densely very minutely puberulent. Primary bracts broadly ovate with an acute apex (6-8 mm by 3.5- 5 mm in size) to narrowly ovate with an acuminate apex (5-7 mm by 1.5-2 mm in size), very densely velutinous outside; secondary bracts narrowly ovate, acuminate at the apex, 3-4 mm by 0.5-1 mm in size. Bracteoles absent. Calyx rather densely very minutely puberulent and mixed with very sparse longer hairs, 4-lobed; tube about 1 mm long; the upper lobe broadly triangular, entire or minutely 2-toothed at the apex, about 1.5 mm long, 1.5-2 mm wide, laterals triangular, 1.2-1.5 mm long, 1-1.2 mm wide at the base, the lower broadly triangular, 1.5-2 mm long, 1 mm wide. Corolla (Fig. 28: b) amethyst-coloured (ex Dunn); standard obovate with an emarginate apex, tapering to the base, 8.5-10 mm long and 5.5-6 mm wide; wings transversely elliptic, 10.5-11.5 mm long (including 1.5-2 mm long claw), 3-3.5 mm wide, auriculate at the lamina-base; keel-petals 8-10.5 mm long (including 1.5-3 mm long claw), 2.5-3 mm wide, acute at the apex. Stamens 8.5- 9 mm long. Pistils 9-10 mm long (including the 4-4.5 mm long style), sparsely to moderately very minute puberulent (0.05 mm long) except for the glabrous style; style thickened slightly above the middle. Pods (Fig. 29: a) sessile, compressed, 6-7-jointed, glabrous, swollen on the seed, both sutures not thickened but appar ently marginate, the upper suture nearly straight or slightly undulate, the lower indented, isthmus usually about 2/3 as broad as the pod, scarsely reticulate-veined; joints transversely elliptic-oblong, 3-4 mm by 4-7 mm in size. Seeds (Fig. 29: b) transversely broadly elliptic, reddish brown in colour, 2.5×2.5-3 mm in size. Pollen grains (Pl. 13: a-d) tricolporate, subprolate, or prolate spheroidal, 37-40× 30-38 (average 38.7×33.3) µ in size, rhomboidal in the equatorial view; colpi long, narrow, the marginal thickening distinct, slightly intruding, with or without con striction at the equator, the membrane granulated; pores large, about 5×10 µ in size, equatorially elongated, slightly protruding; exine evenly fine reticulate, lumina about 1 µ in diameter, tectate, about 2 µ thick, the ektexine thinner than the endexine, muri simplibaculate. Distr. Thailand and China. In vegetative structure and fruit this species is very similar to D. multiflorum which is usually known as D.floribundum or D. sambuense, but differs consistently in the size of stipules and stipels, the degree of indumentum on pods and leaves, shape of styles, and the equatorial shape of pollen grains. However in floral structure the species is closely related to D. confertum especially in the venation pattern of the wings and the shape of styles. 2) Desmodium confertum DC. in Ann. Sci. Nat. 4: 101 (Jan. 1825); Prodr. 2: 335 (Nov. 1825)-Benth. in Miq., Pl. Jungh. 225 (1852), in adnota-Baker in Hook. f., Fl. Brit, Ind. 2: 167 (1876)-Prain in Journ. Asiat. Soc. Beng. 66 (2): 396 (1897)-Ohashi in Hara, Fl, E, Himal. 149 (1966). [Fig. 27b] Hedysarum dioicum Buch.-Ham. ex D. Don, Prodr. Fl. Nepal. 244 (Feb. 1825). A much branched shrub; usually up to 2 m high; young shoots and branches terete but often very slightly angular, finely to densely whitish or yellowish hairy especially along the ridges, afterwards mostly glabrescent. Stipules early deciduous, triangular, acuminate at the apex, 5-7 mm long, 2-4 mm wide at the base, seri ceous outside. Petioles 1.5-5 cm long, rather densely to sparsely tomentose. Leaves 3-foliolate; leaflets subcoriaceous to coriaceous, elliptic or obovate, acute or often obtuse or occasionally apiculate at the apex, entire along the margin, the upper surface densely appressed tomentose when young and afterwards becoming gla brous, the lower surface densely appressed sericeous or tomentose, especially on the nerves, and usually glaucous, lateral nerves usually 5-9 pairs and reaching the margin, reticulate-veins prominent, leaflet-pulvini very densely silky hairy; stipels narrowly triangular or filiform, 2-3 mm long; terminal leaflets usually cunate at the base, (3-)6-13(-17) cm long, (1.5-)2.5-7(-10) cm wide; petiolules (1-)1.5- 3 (-4-) cm long, tomentose like the petioles; lateral leaflets oblique and rotund to obtuse at the base, (2.5-)4.5-8(-11) cm long, (1.5-)2-4(-6) cm wide. Inflorescences terminal paniculate and axillary racemose, when young cone-like with overlapping large primary bracts, often seemingly subsecund especially in fruit, usually 10-20 cm long; rachis densely to rather sparsely spreading pilose and densely very mi nutely tomentose. Flowers mostly binate; pedicels 3-5 mm long, minutely spreading pubescent (0.2-0.3 mm long) and densely very rninute puberulent (below 0.1 mm long). Primary bracts ovate with an acuminate apex, 9-14 mm by 3-5 mm in size, ciliate white hairy (about 1 mm long) and very minutely puberulent outside; secondary bracts occasionally early deciduous, narrowly ovate or narrowly elliptic with an acute apex, 1.5-2.5 mm by 0.7-1 mm in size, ciliate. Bracteoles usually absent but rarely present and very early deciduous, when present 0.8-1.5 mm long about 0.2 mm wide, linear or subulate, puberulent outside. Calyx 3-5 mm long very minutely puberulent and often mixed with longer hairs at the very base and the margin of the lobes, the tube 2-3 mm long, 4-lobed; the upper lobe broadly triangular, entire and obtuse or 2-toothed at the apex, (1.5-)2-2.5 mm long and 2-3 mm wide at the base, the lateral one triangular, nearly same or slightly longer than the upper one, the lower lobe triangular, acute or slightly acuminate at the apex, usually slightly longer than the lateral ones, (2-) 2.5-3 mm long. Corolla (Fig. 28: c) mauve to purple; standard normally to broadly obovate with a rounded or slightly emarginate apex, (9-) 10-14 mm by 5-9.5 mm in size; wings 10-16 mm long (including the 2-2.5 mm long claw), 3-4(-4.5) mm wide, obtuse or rounded at the apex, transversely narrowly elliptic, auriculate at the adaxial base of the lamina, usually almost straight or often slightly deflexed; keel-petals 9-13 mm long (including 4-5 mm long claw), 2.5-3.5 mm wide, acute at the apex. Stamens about 10 mm long, glabrous. Pistils 10.5-12 (-13) mm long (including the 5-6 mm long style), inflexed at a right angle near the middle of the style, densely appressed puberulent except the glabrous style. Pods (Fig. 29: c) compressed, shortly stalked (2-5 mm long), (2-)3-4-jointed, very densely spreading pale brown or grayish sericeous on both sutures and densely appressed so on the lateral surfaces, the upper suture more or less undulate, the lower suture deeply incised, isthmus 1/2-1/4 as broad as the pod, not reticulate-veined, the calyx usually persistent at the base; joints depressed obovate or transversely elliptic, 8-11×4-5 mm in size. Pollen grains tricolporate, suboblate, oblate spheroidal, prolate spheroidal or subprolate, 30-41× 30-35 (average 34.9×33.0) µ in the equatorial view, often almost hexagonal in the outline at the polar view when oblate spheroidal grains; colpi long, narrow, slightly marginate, intruding, the membrane granulated; pores large, about 8×15 µ in size, equatorially elongated, strongly protruding, without constriction at the equator; exine evenly fine reticulate, lumina about 1 µ in size, decreasing the size of lamina towards tile polar area, tectate, about 2 µ thick, the ektexine thinner than, the endexine, muri simplibaculate, columellae short. Distr. Himalaya (Nepal, Sikkim, Bhutan) and E. India (Assam). Repres. specim. exam.: Nepal. Kathmandu: Pouwa (Hara et al, 6301468 TI); near Kusma, alt. 3000 ft. Gully side in Shorea forest (Stainton, Sykes & Williams 9228 BM, TI); Galchi. (Bailey's collectors s.n. BM); Sim Chautara-Thare, Trisuli Valley, alt. 1000 m. Along sunny road side in Shorea forest (Hara, K-urosawa & OhasbiJune 2, 1969 TI); Phusri-Dhara Pani (Hara et al. 6301432 TI), Dhankuta, alt. 1200 m (Hara et al. 6301457, 6301474 TI), Dumhan (Hara et al. 6301458 TI), Garhi Danra-Tuwa (Hara et al. 6301459 TI), Khebang, alt. 1400 m (Hara et al. 6301460 TI), Khebang-Bharomdin (Hara et al. 6301461 TI), Ranga Pani-Ghorwa (Hara et al. 6301462 TI). Sikkim. Darjeeling: Runjeet (C. B. Clarke 26300A CAL); Pashok, alt. 1500ft. (Gamble 7356 CAL), (Prain's collector 164 CAL), Testa (T. Anderson s.n. CAL), (Kari 463 TI). Bhutan. (Griffith 1458 CAL), (G. King s.n. CAL); Mungur (R. E. Cooper 4665 BM), Lhuntzi Dzong, alt. 5000 ft. (Ludlow et al. 20930 BM); Khoma, alt. 5000 ft. Dry hillside (Ludlow, Sherriff & Hicks 21240 BM, TI); Demri Chu, Monyul, alt. 3000 ft. Along river (Ludlow, Sherriff & Taylor 7241 BM, TI). India. Assam: Khasia (Griffith 1438 CAL), (Oldham s.n. CAL), (C. B. Clarke 6233 CAL); Cherapunji (K. Biswas 3950 CAL), Saleri-But, alt. 5000 ft. In oak and pine forest (Rankin & Pretzlik 82 BM). This Himalayan species approaches D. multiflorum in vegetative characters, but differs distinctly in flowers and legumes. The venation of wing-petals of D. con fertum and D. amethystinum differs consistently from that of wings of D. multiflorum as illustrated in Fig. 28: c. The venation of the wings of D. confertum shows princi pally the same pattern of that of the keel-petals of the species, i.e. most of the veins branch off at or near the base of lamina and seemingly extend in rather parallel. While in D. multiflorum the venation pattern is apparently different from that of the keel-petals of the species and, also of D. confertum, i.e. most of the veins spring from a few central veins rather pinnately and often branch off near the margin of lamina. The latter type of the venation of wings is a general pattern in the subgenus Dollinera. The style is always slightly constricted at the base in D. confertum, but is not so in D. multiflorum. The pods of D. confertum are also different greatly from those of D. multiflorum as shown in Fig. 29: c. Moreover, D. confertum is usually distin guishable from the latter by the less sharply angular young branches, the lethery leaflets, and the larger bracts. The hairiness on pods in D. confertum varies rather greatly in age from blackish brown in colour when young gradually to pale or yellowish brown when mature. Among the specimens of the species examined a gathering from Bhutan (Ludlow, Sherriff & Hicks 21240) shows an extraordinary variation in the size of flower and the presence of secondary bracts and bracteoles. The standard petals of the collec tion are 12-14 mm by 8-9.5 mm in size, while many collections from Nepal, Sikkim and Assam are 9-12 mm by 4.5-7.5 mm in size. Also the wings and keel-petals of the Bhutanese specimen are distinctly larger than the others (wings: 13.5-16 mm by 3.8-4.5 mm against 10.5-12.5 mm by 2.8-3.6 mm in size; keel-petals: 11.5-13 mm by 3-3.5 mm against 9-10.5 mm by 2.4-3 mm in size). The pistils of the Ludlow et al. 21240 are 11-12 mm long but those of the others 10.5-11.5 mm long. However the calyx of the Bhutanese specimen tends to show a different variation from the specimens from other regions. In the former specimen calyx-tubes are not shorter than the lower calyx-lobes and the lateral lobes are nearly same or slightly shorter than the upper lobes, i.e. the tube 2.5 mm long, the upper and the lateral lobes 2.2 mm long, the lower lobe 2.5 mm long (all are average length when 20 samples are measured). Whereas in the later specimens calyx-tubes are distinctly shorter than the lower calyx-lobes and the lateral lobes are shorter than the upper lobes, i.e. the tube 2.2 mm long, the upper lobe 2.3 mm long, the lateral lobe 2.0 mm long and the lower lobe 2.8 mm long (all are average length when 41 samples are ex-amined). Schindler (1926) placed this species in his sect. Floribundae of the subgen. Dollinera, which was characterized in having no secondary bracts and bracteoles. However, the secondary bracts occur in several specimens, and bracteoles are found only on one sheet from the Bhutanese specimen. 3) Desmodium kulhaitense C. B. Clarke ex Prain in Journ. Asiat. Soc. Beng.66 (2): 395 (1897)-Schindl. in Fedde, Rep. 22: 268 (1926); in Fedde, Rep. Beih. 49: 281 (1928)-Ohashi in Journ. Jap. Bot. 40: 364 (1965); in Hara, Fl. E.Himal. 151 (1966). [Fig. 30a] An erect shrub, 60-150 cm high; young branches slightly angular, densely spreading grayish velutinous or woolly (1-2 mm long) as are the petioles, the petio lules, and the lower part of the inflorescence-rachides. Stipules narrowly elliptic ovate with an acuminate apex, (8-) 10-15 mm by 1.5-2.5 mm in size, densely velu tinous along the margin and partly on the back. Petioles 3-5 cm long. Leaves 3 foliolate; leaflets subcoriaceous, narrowly elliptic with an acuminate apex, glabrous or very sparsely silky hairy along the midrib above, densely appressed whitish seri-ceous beneath, reticulate-veins not prominent, lateral nerves 5-8 on each side of the midrib and reaching the margin; stipels narrowly ovate, 2-3 mm by 0.5-0.7 mm in size, slightly pubescent outside; terminal leaflets obtuse at the base, (5-)9-13 (-15) cm by (1.5-)3-4(-5) cm in size; petiolules 1.5-2.5 cm long; lateral leaflets obliquely obtuse or rotund at the base, (4-) 7-10 (-11) cm by (1.5-)3-4 cm in size. Inflorescences terminal and axillary, the terminal one much branched paniculate or rarely simple racemose, the axillary one racemose, 15-25 cm long; the lower part of rachis pubescent like the young branch, the upper part glabrous except for the base of bracts. Flowers binate; pedicels slender filiform, 7-10 mm. long when an-thesis, afterwards 11-15 mm long in fruit, very sparsely pilose and very minute hooked hairy. Primary bracts scariose, striate, narrowly triangular or narrowly ovate, acuminate to cuspidate at the apex, 8-10 mm by 1.2-1.5 mm in size,densely sericeous along the margin and rather densely sericeous to nearly glabrous on the back; secondary bracts absent. Bracteoles absent. Calyx 4-5 mm long, rather densely very minute uncinate hairy, 4-lobed; tube 2-3 mm long, the upper lobe broadly triangular, minutely 2-toothed or entire with an obtuse apex, 1-1.5 mm long, 1.5-2 mm wide at the base, the lateral ones triangular, acute at the apex, 1.5-2 mm long, the lower lobes narrowly triangular, acuminate at the apex, slightly longer than the others, 2-2.5 mm long, about 1 mm wide at the base. Corolla (Fig. 28: d) purple-rose; standard obovate, rounded or slightly emarginate at the apex, shorter than the other petals, 8-11 mm by 5-6 mm in size; wings 11.5-14 mm long including the 1.5-2 mm long claw, 3.5-4 mm wide, straight or slightly incurved, obtuse at the apex; keel 10-11 mm long, 2.7-3 mm wide, acute or apicu-late at the apex. Stamens 7.5-8.5 mm long, Pistils 8.5-9.5 mm long. Pods (Fig. 29: k) sessile, (6-)8-9(-10)-jointed, glabrous, the upper suture nearly straight or slightly undulate, the lower one constricted, isthmus usually about 2/3 as wide as the pod, swollen on seeds, reticulate-veined; joints about 5 × 3.5-4 mm in size. Pollen grains tricolporate, subprolate or prolate, 29-31×22-27 (average 31.2×24.3) p. and el-lipsoidal in the equatorial view; colpi long narrow, marginate, intruding, the membrane granulated, constricted and protruding at the equator; pores elliptic, about 6 × 10 µ in size, elongated equatorially, sometimes constricted at the equator, slightly proturding, no marginal thickening; exine evenly fine reticulate, lumina about 1 µ in diameter, tectate, 2 µ thick, the ektexine thicker than the endexine, muri simplibaculate, columellae short. Distr. Endemic to E. Himalaya (Sikkim). Specim. exam.: Sikkim. Hee, alt. 4000 ft. (C. B. Clarke 13109K Oct. 12, 1870 CAL-Isotype); Rungli, alt. 3000 ft. (W. W. Smith 4501 CAL); Bok, alt. 7000 ft. (G. H. Cave s.n. Lloyd Bot. Gard., Darjeeling); above Dentam, alt. 5000 ft. (Ribu & Rhomoo 4913 CAL); Penlong La, near Gangtok, alt. 2100 m (H. Hara & K. C. Pradhan 6301481 Sept. 13, 1964 TI). This endemic Sikkimese species is one of the less variable species of Desmodium and appears to be closely related to D. multiflorum especially in the structure of flower. From D. multiflorum, D. kulhaitense is distinctly separable by the many jointed legumes, the narrowly elliptic, acuminate leaflets, the patent velutinous or woolly young branches, petioles and inflorescence-rachis, and the longer stipules. 4) Desmodium multiflorum DC. in Ann. Sci. Nat. 4: 101 (Jan. 1825); Prodr. 2: 335 (Nov. 1825)-Wall., Cat. no. 5705 A, B, C pro parte (1831-32)-Benth. in Miq., Pl.Jungh. 225 (1852), in adnota. Desmodium angulatum DC., l.c. 101 (Jan. 1825); l.c. 335 (Nov. 1825). A much branched erect shrub, usually 1-2 m high; branches of the old years terete, glabrescent, dark brown with scattered elliptic lenticels, young shoots sharply angular, especially along the ridges densely ascending or often appressed or oc-casionally spreading pale-yellowish to whitish pubescent. Stipules rather persistent, narrowly ovate to ovate with an acuminate apex, (4-) 5-11 mm long and 1.5-2 (-2.5) mm wide. Petioles 1.5-5 cm long, sparsely to densely tomentose. Leaves 3 foliolate; leaflets subcoriaceous, elliptic or obovate, usually obtuse or acute but rarely acuminate at the apex, the upper surface rather sparsely to densely appressed sericeous but afterwards glabrescent, the lower surface densely appressed or ascending sericeous especially on principal veins and mostly more or less glaucous, lateral nerves conspicuous, usually 6-8 on each side of tlie midrib and reaching margin, entire along the margin; stipels narrowly triangular, 1-3 mm long and 0.3-0.8 mm wide at the base; terminal leaflets usually cuneate and obtuse or rarely rotund at the base, 3-10(-13) cm by 2-5(-6) cm in size; petiolules 0.7-1.5 cm long; lateral leaflets oblique and rotund or rarely obtuse at the base, 2.5-7 (-9) cm by 1.5-3.5(-4) cm in size. Inflorescences terminal and axillary, terminals mostly paniculate but occasionally racemose, axillary ones racemose, usually 6-9 cm but occasionally up to 20 cm long, when young cone-like with overlapping bracts; rachis densely ascending serious and minute uncinate hairy. Flowers binate; pedicels (3-)4-7(-10) mm long, rather sparsely to densely spreading hooked or straight hairy. Primary bracts narrowly ovate to broadly ovate, entire and acuminate or acute at the apex, 7-11 mm by 2-3 mm in size, sparsely to densely sericeous outside; secondary bracts ovate, sericeous along the margin, (1.2-) 1.5-3.5 mm by 0.8-1.5 mm in size. Bracteoles absent or very rarely present at the base of the calyx (paired, filiform, 0.4-0.6 mm long, sparsely pubescent). Calyx densely uncinate minute hairy and sparsely to scarcely sericeous, 3-5(-5.5) mm long, 4-lobed, the tube 2-2.5 mm long, the upper lobe broadly triangular, entire or minutely 2-toothed at the apex, 1.5-2 (-2.5) mm long, lateral ones triangular, acute at the apex,almost same length to the upper one, the lower triangular, slightly longer than the others, 2-2.5(-4) mm long, acuminate at the apex. Corolla (Fig. 28: e & f) mauve, pale pink or pink to purple; standard normally to broadly elliptic or obovate, 7.5-11.5 mm by 4-7.5 mm in size, more or less emarginate at the apex, cuneate at the base; wings narrowly elliptic, slightly inflexed, 8-14(-14.5) mm (including the 1.5-2.5 mm long claw) by 3-4.5 (-5) mm in size, obtuse at the apex; keel distinctly smaller than the wings, 7-10(-13) mm long (including the 3-4(-5) mm long claw), 2-3.5 mm wide, acute or obtuse or slightly apiculate at the apex. Stamens 6-7 mm long. Pistils 8.5-9 mm long, sparsely or rather densely appressed pubescent except the glabrous style. Pods (Fig. 29: d) sessile, (4-) 5-7 (-8) -jointed, densely appressed brown-sericeous, the upper suture nearly straight or slightly undulate, the lower indented, isthmus usually about 2/3 as broad as the pod, swollen on seeds, reticulate-vdned, calyx often persistent at the base; joints depressed obovate, 3-4 mm×3.5 mm in size. Seeds (Fig. 29: e) transversely elliptic, 1.5-2×2.5-3 mm in size. Pollen grains (Pl. 13: e-h) tricolporate, oblate spheroidal, prolate spheroidal, subporiate or prolate, 33-47 ×26-39 (average 39.3×31.7) µ in size; colpi long, narrow, distinctly marginate, slightly intruding; pores almost rounded or elliptic, 6-9×x 10-15 µ in size, equatorially elongated, slightly constricted and protruding at the equator; exine evenly fine reticulate, lumina about l µ or slightly larger than l µ in diameter, tectate, about l µ thick, the ektexine thinner than the endexine, muri simpli-baculate, columellae short. Distr. Himalaya (N.W. & N.E. India, Nepal, Sikkim, Bhutan), Burma, Thailand, Laos, and China. Repres. Specim. exam.: India. Kumaon: (Hume s.n. CAL), Bachgawan, alt. 4500 ft. (N. Gill 54 TI); Simla: (Gamble 4535E, 6408B CAL), (Schlich in 1888 s.n. CAL), Kasauli, alt. 6000 ft. (J. W. Chiddell 43 BM); Ghurwol (G. King s.n. CAL), (R. S. Hole 791 CAL); near Musoourie (G. King s.n CAL); Assam: Khasia Hills (Griffith 685 CAL), (J. D. Hooker & Thomson s.n. CAL), Shillong, alt. 5000 ft. In pine woods (C. B. Clarke 44582E CAL), (F. Kingdon-Ward 18637 BM, A), Naga Hill, Konema (D. Prain s.n. CAL), Jaintia Hills, Maolamang alt. about 4500 ft. (Burkill & Banerjee 313 CAL). Nepal. Maina, alt. ca. 8000 ft. On earthy ledges among rockes (Polunin, Sykes & Williams 450 BM), alt. 7000 ft. On old cultivation terraces and earth banks (Polunin, Sykes & Williams 481 BM, TI); Tamur Valley, Thapabu Khola, N. of Taplejung, alt 6000 ft. On open slopes (J. Stainton 1191 BM, TI); near Jagat, alt. 8500 ft. On dry south slope (Stainton, Sykes & Williams 3379 BM, TI); Annapurna Himal: Seti Khola, alt. 6500 ft. (Stainton, Sykes & Williams 6714 BM, TI); Kathmandu: Thankot (K. de B. Codrington 219, 243 BM), Godavari-Phulchauki, alt. 2000 m. Margin of thickets, sunny place (Ohashi June 23. 1967 TI); Taplejung-Garhi Danra (Hara et al. 6301438 TI-Syntype of D. nepalense Ohashi); Garhi Danra-Tuwa (Hara et al. 6301484 TI-Holotype of D. nepalense Ohashi). Sikkim. Darjeeling (J. S. Gamble 8351 CAL); Gangtok, alt. 4000 ft. (Ribu & Rhomoo 4004 CAL). Bhutan. (R. E. Cooper 4561, 4579 BM); Dotena Limpu, alt. 8000 ft. (Cooper 2506, 2654 BM); Linchu, alt. 5000 ft. (Cooper 3400 BM); Valley of Kuru Chu, alt. 6500 ft. On dry hillsides, red soil (Cooper 4377 BM); Lhuntzi Dzong, alt. 5000 ft. Amongst shrubs on hillside (Ludlow,Sherriff & Hicks 20930 BM, TI). Burma. Pegu (S. Kurz 1676 CAL); Mogok (A. Rodger 174 CAL); Chin Hills (A. Huk s.n. CAL); Shan State, Taungyi (A. Khalil s.n. CAL); Mindat Sakan, alt. 4500 ft. Thickets & waste places (F. Kingdon-Ward 22545 BM, TI); Ukhrul, alt. 5000 ft. In thickets (Kingdon-Ward 17787 NY), alt. 6000 ft. Common in open places (Kingdon-Ward 17920 NY, BM, A). Thailand. Loei (Dee 232 BKF), Phu Krading (Native collector DE303B A), (D. Bunpheng 681 BKF), alt. ca. 1200 m. Open grassland (T. Shimizu et-al. T.9011 KYO), Phu Luang. On moist, sandy ground in open place (M. Tagawa et al. T.1545 KYO), (K. Bunchuai 176 BKF); Chieng Mai, Doi Sutep (A. F. G. Kerr 782 BK, CAL), (B. Hayata s.n. Oct. 14, 1921 TI), (K. Iwatsuki et al. T.9379 KYO), below the summit of Doi Chiang Dao, alt. 1900-2175 m. Among bush on limestone ridge (T. Shimizu et al. T.10118 KYO). China. Yunnan: (D. J. Anderson in 1868 s.n. CAL), (C. Shneider 2492, 2499 GH), (A. Henry 9802 CAL), (Henry 9802A NY), (H. T. Tsai 58997 A); Szemao (A. Henry 10003B TNS); Ping-pien Hsien (Tsai 61464, 62426 A); Mienning, Poshang, alt. 2350 m. In thickets (T. T. Y-ii 17857 A); Chlh-tse-lo, alt. 2500 m. On road side (Tsai 54175 A), alt. 3200 m. In forest (Tsai 5848 A); Der-la, Champu-tung, alt. 2300 m (C. W. Wang 66831 A); Kiukiang Valley, alt.1400 m. Margin of thickets (Y Ü20389 A); Chengkang, Maliling, alt. 1900 m (YÜ 16846 A); Che-tse-lo, alt. 3200 m. In forest (Tsai 58479 A); Lichiang Range (McLaren's native collectors 91B, 211B BM); Szechuan: Hsl-ching Hsien (YÜ 1224 A); Juei-she Hsien (YÜ 1044 A); Kweichow: Tuhshan, border of Kwansi, alt. 400 m. In open places (Y. Tsiang 6478 A, NY); Kwangsi: YeoMar Shan, N. Hin Yen, alt. 5000ft. In open thickets (R. C. Ching 7167 A, NY); Kwang-tung: Yao-shan, An-Keng, alt. 2500 ft. (S. Sin 11830 NY); Fukien: Shaown and vicinity, Satasun, alt. 4000 ft. (F. H. Niao 9497 A); Hupeh: (E. H. Wilson 2624 NY), (Henry 2502 CAL). Taiwan. (R. Torie s.n. TI); Taichung Hsien: Peikoutashan (U. Mori in 1910 TI); Nantou Hsien: Mt. Mushan (Kawakami & Mori in 1906 TI), Mt. Neng-Kao (M. T. Kao 5581 TI), Jeuai, alt. 1000 m. On sunny grassfield along valley (Ando, Tate-ishi & Watanabe 351 TI). This species, usually known as D. floribundum, resembles D. amethystinum, D. confertum, D. kulhaitense and D. Hayatae. From D. amethystinum, D. confertum, D. kulhaitense and D. Hayatae, however, D. multiflorum differs consistently as noted on each species, respectively. Sometimes the species approaches to D. elegans especially when sterile or often when anthesis. From D. elegans, D. multiflorum is clearly distin-guishable by the sharply angular young branches and inflorescence-rachis, the elliptic or obovate, usually obtuse terminal leaflets, the binate-flowered racemes, persisting bracts subtending the flower-buds on inflorescences, and the usually ebracteolate calyx. D. multiflorum is highly polymorphic in external morphology but is less variable in pollen morphology. In pollen morphology, among the grains examined in this study those from Burma (Kingdon-Ward 22545 TI) show the largest in size, 44-47 × 32-36 (average 45.4×33.5) µ, and those from Nepal (Stainton 1191 TI) are the smallest, 34-38×27-30 (average 35.9×28.5) µ in size. In outer morphology, D. multiflorum constitutes somewhat distinct local assemblages. Collections from the Himalayas especially Nepal are referable to the typical form which has acute, elliptic or obovate, glaucous leaflets, without bracteoles, and sparsely to moderately patent hooked hairy on inflorescence-rachis and petioles. In addition to these features the size of keel-petals of the specimens from Nepal is often remarkably smaller than that of the specimens from other regions. The specimen from Burma (Kingdon-Ward 22545) of which pollen grains mentioned above, tends to have sharply acuminate elliptic leaflets, lax flowered racemes, distinctly longer pedicels and larger flowers (standard 9.5-11.5×4.5-7 mm in size, wings 11.5-14.5×3.5-5 mm and keel-petals 10.5-13×3-3.5 mm in size). Specimens from Thailand, how-ever, have densely velutinous, not glaucous leaflets, densely patent hooked hairy on inflorescence-rachis and pedicels, and often minutely bracteolate near the base of the calyx. It proved impossible, however, to subdivide several forms of D. multiflorum into varieties, for examination of herbarium material showed that no discontinuity of characters could be found. 1-(2). Subsect. Khasianae (Schindl.) Ohashi, stat. nov. |
