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Taxonomy and Distribution of Desmodium and Related Related Genera (Leguminosae) in Malesia (11)

Hiroyoshi Hiroyoshi OHASHI

Botanical Botanical Garden ， Graduate School of Science ，Tohoku University ，Sendai ，980 ・0862 JAPAN

(Received on Novernber 8， 2003)

This This second part of my revision of Malesian Desmodium and its relatives includes taxonomic taxonomic treatments of the following eight genera ， Hanslia (including two new combi- nations) ，Hegnera ，Hylodesmum ，Monarthrocarpus ，Ohwia ，Phyllodium ，Tadehagi ， and Trifidacanthus ， and phytogeographic considerations for the taxa of Desmodium and its relatives relatives in Malesia. Thirteen genera and 64 species are recognized as Desmodium and its relatives relatives in Malesia. Their distribution patterns 訂 'e diverse. Of 64 species recognized in this this study 56 訂 'e native to Malesia and 訂 e divided by their distribution patterns into the following following four groups: Malesia (including 8 species) ， Asia (29 spp よAustralia (11 spp よ and and Asia-Australia (8 spp.) groups. All species of the genera native to Jawa ，New Guinea and and the Philippines are discussed. Each 訂 ea has almost equal numbers of native species ， 31 ，32 ， and 29 ，respectively. This fact suggest insufficient collections in New Guinea and the the Philippines. The present -day composition and distribution of taxa in Desmodium and related related genera in Malesia 紅 e considered to be derived from various spatially and tempo- rally rally diverse ancestors.

Key words: Desmodium relatives ，distribution patterns ，Hanslia ，Jawa ，New Guinea.

HANSLIA Key to the species of Hanslia Hanslia Schind l. in Repert. Spec. Nov. 1. Leaves 3-foliolate ， rarely mixed with 1- Regni Veg. 20: 276 (1924) [Type: foliolate; leaflets acute at base … H. hentyi Hansli α adhaerens Schind l. (= Hedysarum 1. Leaves 1- foliolate; leaflets round at base adhaerens Poir. ，non Vah l.)]. .... H. ormocarpoides Desmodium subg. Hanslia (Schind l.) H. Ohashi in Ginkgoana 1: 112 (1 973); Pedley Hanslia hentyi (Verdc.) H. Ohashi ，comb. in in Austrobaileya 5: 226 (1999). nov. (Fig. 5) Hanslia Hanslia is similar to Ohwia in having a Desmodium hen ηi Verdc. in Kew Bull. sc 紅 cely developed rim-aril around the hilum 32: 249 (1 977) [Type: Papua New Guinea. of seeds ， uncinate hairs on the loments ， long Morobe Distric t: ne 訂 Lae ， Atzera Range ， 0 0 articles ，and a coarser ，reticulate sculpture on 6 45"S. ，147 00"E. 210 m. 1 M 訂. 1960. pollen pollen grains. They 紅 e considered to be re- Shrub ， standard reddish purple ， keel and lated lated to each othe r. Ohwia and Hanslia wings pale pink ，pod purple ， adhesive. Henty ormocarpoides have white to creamy flow- in NGF11922 (LAE holo. ， K-photo)] & ers ， but H. hen ηi has purple to pink flowers. Verdc. ，Man. New Guinea Leg.: 398 ，fig. 94- The genus comprises two species and both o (1979). are are found in 恥1alesia. Distr.: 島falesia (New Guinea and new to

-155- 156 156 植物研究雑誌第79 巻第3号 平成16 年6月

Fig. Fig. 5. Hanslia hentyi (Verdc.) H. Ohash i. Bomeo. M t. Ki nabalu. Chew & al. 1444 (K).

Bomeo). Bomeo). plan t. This species occurs under forest and Specimens Specimens examined: Borneo. M t. Ki nabalu. Ulu such habitat is supposed to be similar with Liwagu and Ulu Mesilau ，6 0 N. ， about 116 0 35/E. that of Hylodesmum laxum and H. leptopus. Kundasang. Kundasang. Wings pu 中le ， keels pink. Riverside Fores t. 3 Sep t. 1961 Chew ，Comer & Stainton 1444 (K). (K). New Guinea. Atzera Range near Lae ， al t. 700 ft. Hanslia ormocarpoides (DC.) H. Ohashi ， Henty Henty 11922 (LAE holo.; photo K); Kokoda. Herb ，ca. comb. nov. 2 ft. tall. N.G. Carr 16300 ， 28 Mar. 1936 (L); Conn & Hedys αrum adhaerens Poir. in Lam. ， l. al. 35 (LAE 232235; photo TUS). Encyc l. Supp l. 5: 15 (1817) ， non Vah l. Leaves Leaves of this species have three leaflets (1 791). [Type: Java. Labillardi とre (P ， n. v.)]. or or rarely with single leaflets on the same [Hedysarum ormocarpum Desv. ex Poir. June June 2004 Journal of Japanese Botany Vo l. 79 No. 3 157 in in Lam. ，Encyc l. Supp l. 5: 15 (1 817) ， pro The loment is reticulate-veined ， although it

syn.]. syn.]. was described as not reticulate 田 veined in Desmodium ormocarpoides DC. ，Prodr. 2: Ohashi (1 973) based on young loments. 327 327 (1 825) [Type: 'Cette plante a ete decouverte decouverte a l' ile de ava par M. de HEGNERA Labi11ardiere' Labi11ardiere' (P ?) (fide Pedley 1999)] ; Hegnera Schind l. in Repert. Spec. N ov. Me 町. in Philipp. J. Sci. 14: 409 (1 919) & Regni Veg. 20: 284 (1 924) [Type: Hegnera Enum. Philipp. Flow. Pl. 2: 287 (1923); obcordata (Miq.) Schind l. (= Uraria Meeuwen in Reinwardtia 6: 96 (1961) & 255 obcordata Miq.)]; H. Ohashi in Ginkgoana (1962); (1962); H. Ohashi in Ginkgoana 1: 113 1: 257 (1 973); Dy Phon in Dy Phon & al.， Fl. (1 973); Verdc. ，Man. New Guinea Leg.: 403 Camb. Laos Vietn. 27: 60 (1 994). (1979); (1979); Pedley in Austrobaileya 5: 231 The genus is monotypic. (1999). (1999). D. D. dependens Blume ex Miq. ， F l. Ned. Hegnera obcordata (Miq.) Schind l. in Ind. Ind. 1(1): 248 (1855) [Type: Molucc. Un- Repert. Spec. Nov. Regni Veg. 20: 285 known co11ector. (L holo. 908.115-521)]. (1 924); H. Ohashi in Ginkgoana 1: 257 D. D. pendulum F. Mue11. in Campbe11 ，A (1 973). Year Year New Hebrides: 9 (1 873) [Type: Tana. Uraria obcordata Miq. ， F l. Ned. Ind. Campbe11 Campbe11 (?)]; F. Mue11. ， Fragm. 8: 225 Eerste B 討v. 1: 305 (1 860) [Type: Sumatra (1 874). orien t. Prov. Palembang. prope Mu 紅 aduwa. Hanslia Hanslia adhaerens Schind l. in Repert. Teysmann (L holo.; iso. K)]. Spec. Spec. Nov. Regni Veg. 20: 277 (1924). Desmodium obcordatum (Miq.) Kurz in J. Distr.: Distr.: Malesia ，Australia (N. Queensland) Asia t. Soc. Bengal ， P t. 2，Na t. His t. 42: 229 and and Pacific (New Hebrides ，Solomon Is.: W. (1 874): Meeuwen in Reinwardtia 6: 255 Guadalcanal) Guadalcanal) . (1 962); Backer & Bakh. f. ， F l. Java 1: 604 Specimens Specimens examined: Lesser Sunda Is. Timor. (1 963). Steenis Steenis 18165 (BO). Maluku. Ambon. Teysmann s. n. Distr.: Myanmar ，Thailand ，Indo-China ， in in anno 1907 (BO) ， Robinson 556 (A ，BM ，BO ，K ， and Malesia. NY ， US). Buru. Van Balgooy 4684 (K ， L). New Guinea. Guinea. Mafulu. Brass 5268 (BO ，NY ， US); Kassam- Specimens examined: Jawa. H. Zollinger 1628 (K); Water Rice Road. Brass 32316 (A ，NY ，PNH ， US); W. Maduera Island. Backer 20727 (BO ， L). Malaya: Sec t. Papua. Tributary of Laloki River 2 miles E of Kedah. Henderson s.n. in anno 1933 (BO) ，Researow Rouna. Rouna. T. G. Hartley 10715 (TI). Philippines. Jolo. s. n. Dec. 1933 (K). Sumatera. Palemb. Teysmann 3909413 (BO iso. of Uraria obcordata Miq.?). Ramos & Edano Bur. Sci. 44391 (BM ，BO ， US). Sulawes i. Boton ，SE of Sulawesi. Coode s. n. (K); Uraria obcordata Miq. was recognized Gorontalo. Gorontalo. Ri edel s. n. (K); G. Konkeonkea. van first by Bentham (1 865a) as belonging to Vuuren 750 (BO). Desmodium. He classified the species as a Desmodium ormocarpoides is recorded by member of subsec t. Nephromeria Benth. in Merri11 Merri11 (1910) from Luzon (Whitford 865) ， Desmodium sec t. Heteroloma Benth. ， but did Mindoro Mindoro (Merri11 6223) ， Samar (Me 凶 11 not publish a new combination. 5201) ，Cebu (McGregor Bur. Sci. 1731) and Mindanao (Mrs Clemens 632) ， but these HYLODESMUM specimens 訂 e a11 referable to D. zonatum Hylodesmum H. Ohashi & R. R. Mi11 in Miq. Miq. Edinb. J. Bo t. 57: 173 (2000) [Type: H. The ovary of this species is densely cov- podocarpum (DC.) H. Ohashi & R. R. Mi11 ered ered with white ，straight hairs. These hairs (= Desmodium podocarpum DC.)]; H. become hooked when the Ioments mature. Ohashi [in Sci. Rep. Tohoku Univ. ser. 4， 158 158 植物研究雑誌第79 巻第3号 平成16 年6月

Bio l. 40: 235 (1 999) ，nom. nud.] in K. Iwats. Key to the species of Hylodesmum & al.， Fl. Jap. IIb: 256 (2001). in Malesia Desmodium sec t. Podoc α rpium Benth. in 1. Calyx-1obes not shorter than the tube; pri- Miq. ， P l. Jungh.: 226 (1 852) ， p. p. ，exc l. D. mary bracts ovate to broad1y ovate; axillare axillare & D. securiforme [Type: Desmo- pedicels 15-30( -4 0) mm long; flowers dium podocarpum DC.]; Isely in Brittonia 7: 10nger than 8 mm; stipules patent or 185 185 (1 951). reflexed ， 10-25 mm 1ong ，2-3 mm wide; Papilionopsis Papilionopsis van Steenis in Nova Guinea ， leaflets slightly undulate along the margin;

Bo t. 3: 17 (1 960) [Type: Papilionopsis corolla orange to red ....・ H ・... H. repandum stylidioides stylidioides van Steenis] ，nom. rej. prop. (cf. 1. Calyx-lobes much shorter than the tube; Ohashi Ohashi and Mill in Taxon 53: in press prim 紅 y bracts narrowly ovate or narrowly (2004)). (2004)). triangular ， less than 2 mm wide; pedicels Desmodium sec t. Heteroloma subsec t. 2-20 mm long; flowers less than 10 mm Podocarpia Podocarpia Benth. in Benth. & Hoo k. f.， long; stipules as cending or erect ，less than Gen. Gen. Pl. 1: 520 (1 865). 10 mm long and 2 mm wide;

Desmodium subgen. Podocarpium leafletsentire; corolla pale pink ...・ H ・..… ..2 (Benth.) (Benth.) H. Ohashi in Ginkgoana 1: 120 2. Principal lateral nerves reaching the m 紅ー (1 973) p. p. ，exc l. sec t. Monarthrocarpus gin; fruit stipes usually less than 7(-10)

(Me 町.) H. Ohashi ， p. p. exc l. D. dolabri- mm long …...・ H ・....・ H ・-…... H. podocarpum forme Benth. 2. Principal lateral nerves looped within the Podocarpium (Benth.) Y. C. Yang & P. H. m 訂 gin; fruit stipes usually more than 10

Huang in Bull. Bo t. Lab. North-Eas t. Fores t. mm long …...・ H ・.....・ H ・..…・ H ・H ・...… H ・H ・...3 Ins t. 4: 4 (1979) & in Fl. Reipub l. Popu l. Sin. 3. Art icles 12-18 mm long; leaflets with 41: 41: 47 (1 995) ，nom. illegi t.， non A. Braun ex white spots and indistinct nerves. …- Stizenb. ， Uebers. Verstein. Baden: 90 (1851; … H. leptopus fossil fossil Leguminosae) nec Unger in C. G. F. 3. Art icles 9-10 mm long; leaflets pale green Hochs t.， Pa1aonto l. Neu-See1and ，Geo l. Theil and with distinct nerves beneath …- 1 (2): 13 (post Nov. 1864; fossil Podocarpa- …...H. laxum ceae). ceae). Hylodesmum H. Ohashi & R. R. Mill is Hylodesmum laxum (DC.) H. Ohashi & composed mostly of Desmodium subgenus R. R. Mill in Edinb. J. Bo t. 57: 178 (2000). Podocarpium. Podocarpium. It is morphologically characte- (Fig. 6) rized rized by Ohashi (1 973) and Ohashi and 乱1ill Desmodium laxum DC. in Ann. Sci. Na t. (2000). (2000). It s herbaceous habit and preference (Paris) 4: 102 (1 825) [Type: Nepa l. Wallich for for forest habitats are other characteristic (G-DC)]; Me 町.， Enum. Philipp. Flow. Pl. 2: features features for the genus. It was considered by 287 (1923) ， p. ，exc l. syn. D. gardneri et D. Ohashi Ohashi (1973) to be one of the most ad- leptopus; Meeuwen in Reinwardtia 6: 253 vanced genera in Desmodium and its rela- (1 962) ， p. p. ，exc l. syn. D. leptopus ， D. tives tives and to be a group evolved from japonicum Miq.; Backer & B 法h. f. ， F l. Java Desmodium subgenus Dollinera (Ohashi 1: 604 (1 963); H. Ohashi in Ginkgoana 1: 1971 ， Kajita and Ohashi 1994). 134 & 139 (1 973) ， as subsp. laxum; Verdc. ， The genus comprises 14 species of which Man. New Guinea Leg.: 401 (1 979). four four are found in Malesia. D. gα rdneri Benth. in Miq. ， P l. Jungh.: 226 (1852) ， p. p. ，inc l. specim. cit “ In Ceylona ， Gardner 220 (K syn.)". D. D. elongatum Zoll. & Moritzi in Na t. en June June 2004 Journal of Japanese Botany Vo l. 79 No. 3 159

Geneesk ，Ar ch. Neer l. Ind. 3: 58 (1 846). Elmer 11216 (CAL) and 13820 (CAL) ，were D. D. bambusetorum Miq. ， F l. Ned. Ind. 1(1): cited by Ohashi (1 973) as Desmodium laxum 256 (1 855) [Type: Jawa. Tusschen de subsp. leptopus ， but they were erroneous for Bamboes-struiken Bamboes-struiken op de hellingen van Hylodesmum laxum. Amprong b討Gebok Kl akka. Herb. Zollinger 2537 (L908.115-349)]. Hylodesmum leptopus (A. Gray ex

D. D. podocarpum var. Z似 um (DC.) Baker in Benth.) H. Ohashi & R. R. Mill in Edinb. J. Hoo k. f. ， F l. Bri t. India 2: 165 (1 876). Bo t. 57: 179 (2000). Podoc α rpium 1αxum (DC.) Y. C. Yang & Desmodium leptopus A. Gray ex Benth. in P. P. H. Huang in Bull. Bo t. Lab. N. E. Fores t. 乱1iq. ， Pl. Jungh.: 226 (1852) [Type: Ins t.， Harbin 4: 7 (1 979) ，nom. illegi t. Philippines. Luzon. Exped. Explor. Dist r.: Sri Lanka ，India ，Nepal ，Thailand ， Americanas. (K lecto.; isolecto. GH ， US)]; Indo-China ，島'I alesia ，Solomon Is. (Guadal- Miq. ， F l. Ned. Ind. 1(1): 255 (1 855). (Fig. 7) canal) ，China ，and Japan. D. gardneri Benth. in Miq. ， P l. Jungh.: Specimens Specimens examined: Borneo. Sarawak. Y. P. 226 (1852) [Type: Ceylon. Walker (K Ching Ching S. 44685 (K). Jawa. Hasskarl s. n. (L 908.115- lecto.)]; Miq. ， F l. Ned. Ind. 1(1): 255 (1 855). 349) ， Backer 37187 (L). Bali. Soenda. Steenis 8086 D. laxum DC. subsp. leptopus (A. Gray ex (L); (L); Slooten 8086 (A). Lesser Sunda Is. Sumbawa. Benth.) Benth.) H. Ohashi in Ginkgoana 1: 141 t. M t. Batulanteh ， moist Dipterocarpus retusus forest ， 500-700 m al t. Kostermans 18759 (BO ， K ， L). (1 973); Verdc. ，Man. New Guinea Leg.: 402

Maluku. Buru: Wae Kose ，prim 訂 y forest ，on a river- (1 979). side ，0- 50 m al t. Mogea & Ismail JPM-5203 (BO ， K). Podocarpium leptopus (A. Gray ex New Guinea. Schlechter 14106 (BM ， K); Schlechter Benth.) Y. C. Yang & P. H. Huang in Bull. 19009 19009 (NY); Vogelkop Peninsula. Aifat river valley ， Bo t. Lab. N. E. Fores t. Ins t.， Harbin 4: 6 t. al t. 750 m. P. van Royen & H. Sleumer 6914 (A ，K ， L); (1979) ，nom. illegi t. 乱1orobe distric t. Kauli Lake ， al t. 4500 ft.， alongside creek creek in Castanopsis fores t. A. Kairo 45433 (A ， K); W. Desmodium tashiroi Matsum. in It o & of of Bulolo. Frodin & Hill (NY); Path at top of Atzera Matsum. ，J. Coll. Sci. Imp. Univ. Tokyo 12 range ， al t. 600 ft. Verdcourt & al. 5100 (K). (Ten t. Fl. Lutchu よ 415 (1899) [Type: Philippines. Philippines. Luzon. M t. Province ， Banaue ， M t. Ryukyu: insula Okinawa. Y. Tashiro anno Taggute. Taggute. Dr. Cinklin & del Rosario PNH72413 (A ，K ， 1887 (TI holo.)]. L ，PNH); Elmer 22372 (A ，BM ，L ，NY). Mindanao. Elmer 11216 (A ，BM ，CAL ，E ，L ，NY; photo .T US) ， D. laxum auc t. non DC.: Rid l.， Fl. Malay Elmer 13820 (A ，BM ，CAL ， L; photo TUS). Negros. Penins. 1: 608 (1 922). Elmer 10029 (A ，BM ，E ，L ，NY); Gutierrez & al. PNH Distr.: Indo-China ， Malesia ， S. China ， 108894 108894 (PNH). Sarawa k. Proposed G. Murud Taiwan ，and S. Japan. National National Park ， 4th Division ，mixed Dipterocarp forest Specimens Specimens examined: Borneo. M. S. Clemens at at 1200 m. Yii Puan Ching S-44685 (K). Sulawes i. 10901 10901 (A). Sarawak. Proposed G. Murud National Sulawesi Sulawesi Utara ，220 m. de Vogel & Vermeulen 6561 P 紅 k， 4th Division ，mixed Dipterocarp forest at 1200 (L) ， Burley & al. 3750 (A). m. m. Y. P. Ching S-44685 (K). Jawa. Zollinger 2537 (BM) ， Kostermans 161 (A ，BO ，K ， L); Bakh. v/d. Hylodesmum laxum shows a much varia- Brink 4386 (K ， L); N.Wirawan 114 (BO ，K ， L). Bali. tion tion in the shape of leaflets in the Soenda. Steenis 8086 (L). Lesser Sunda Is. Sumbawa. Philippines. Philippines. Elmer 10029 (A) and others M t. Batulanteh ， moist Dipterocarpus retusus forest ， have a narrowly elliptic terminalleaflet ， and 500-700 m al t. Kostermans 18759 (BO ，K ，L) ， Kostermans Kostermans 18395 (L). Malaya. Pahang. Ridley s. n. ， Elmer 13820 (A) has an ovate one but (BM ， SING). Selangor. Ridley s. n. (SING); Burkill Elmer 22372 (A) has a narrowly elliptic one 8851 (SING). Perak. Henderson 10821 (SING). in in the lower leaves and an ovate one in the Maluku. Romassi 1467 (L); Buru. Mogea & Ismail upper upper leaves. 5203 (BO ，K) ，Mogea 5404 (BO ，K) ，Nooteboom 5117 The specimens from the Philippines ， (BO) ， van Balgooy 4783 (K ， L). New Guinea. D. 160 160 植物研究雑誌第79 巻第3号 平成16 年6月

Fig. Fig. 6. Hylodesmum 1似 um (DC.) H. Ohashi & R. R. Mill subsp. laxum. a: Philippines. Luzon. Dr. Cinklin & del Rosario PNH72413 (A). b: Inflorescence showing characteristic long stipe in young loments (en- larged larged from fig. a).

Sayers Sayers NGF 24245 (A). Philippines. Luzon. Exped. Hylodesmum laxum and H. leptopus 紅 e Explor. Explor. Americanas. (K lecto.; isolecto. GH ， US); found found in New Guinea ， but the latter appe 紅 S Angilog ， al t. 1200 m. A. Loher 5916 (K); Biliran. R. C. to to be r紅 e. McGregor Bu r. Sci. 18729 (K); Camiguin Is. E. Fenix Bur.Sci .4 132 (K); Catanduanes. Ramos & Edano Bur.Sci.75287 Bur.Sci.75287 (NY ，UC); Leyte. Frohne PNH 35687 Hylodesmum podocarpum (DC.) H. (L ，PNH). Sulawes i. Yusuf & Wahyono 78 (BO ， L). Ohashi & R. R. Mill in Edinb. J. Bo t. 57: Sumatera. Sumatera. R. S. Boeea 10676 (A ， L); Lampung. M. 181 (2000). Jacobs Jacobs 8163 (K). Desmodium podocarpum DC. in Ann. Sci. Na t. (P 紅 is) 4: 102 (1 825) [Type: Nepa l. June June 2004 Joumal of Japanese Botany Vo l. 79 No. 3 161

Wallich Wallich (G ・DC)]; Me 町. in Philipp. J. Sci. ， C. i. e. ， M t. Santo Tomas in Benguet Ramos & 5: 5: 83 (1910) & Enum. Philipp. Flow. Pl. 2: Edano Bur. Sci. 45098 (K ，NY ，SING ，US) ， 287 (1 923); H. Ohashi in Ginkgoana 1: 142 and loc. ci t. Merrill 11655 (SING ， US). (1973). (1973). Merrill (1 910) reported Desmodium Hedysarum podocarpum (DC.) Spreng. ， podocarpum DC. from Luzon based on two Sys t. Veg. 3: 317 (1826). specimens ，i. e. ， Williams 1398 and Merrill Podocarpium podocarpum (DC.) Y. C. 4356 ， although Merrill 4356 in K and NY Yang & P. H. Huang in Bull. Bo t. Lab. N.E. (Fig. 8) was erroneously determined by him Fores t. Ins t.， Harbin 4: 7 (1979) ，nom. illegi t. as D. sca1l フe DC. ， a synonym of H. repandum. repandum. These two specimens are without Key to the subspecies of doubt H. podocarpum subsp. podocarpum. 均，lodesmum podocarpum The occu 町 ence of the species was confirmed 1. 1. Terminal leaflets broadly obovate to or- in the Philippines by our recent collection ， bicular ， broadest at or above the middle ， Nemoto & al. 10750 (TUS).

apex apex rounded with an acute tip …...・ H ・..…・ Meeuwen (1 962) recorded Desmodium ...・.. .subsp. podocarpum racemosum (Thunb.) DC. from the 1. 1. Terminal leaflets elliptic-ovate ， broadest Philippines based on Saran & Party 30147. 1 below the middle ， apex acute to acuminate could not find this specimen in L and other … subsp. oxyphyllum herbaria. A specimen ， Dr. Cinklin & del Rosario Rosario Phi l. Na t. Herb.72413 (L) ，was de- subsp. subsp. podocarpum: H. Ohashi & R. R. termined by Meeuwen as D. racemosum. Mill Mill in Edinb. J. Bo t. 57: 181 (2000). This specimen (as D. podocarpum) is kept in

Desmodium podocarpum DC. subsp. several herbaria ， but all 訂 e Hylodesmum podocarpum: H. Ohashi in Ginkgoana 1: 154 laxum (DC.) H. Ohashi & R. R. Mill subsp. (1 973); Huang & H. Ohashi in Fl. Taiwan laxum (Fig. 6). Meeuwen (1 962) included D. ed. ed. 2， 3: 262 ，t. 126 (1 993). podocarpum DC. ， D. gardneri Benth. and D. D. D. racemosum auc t. non (Thunb.) DC.: fallax Schind l. in her D. racemosum as syno- Meeuwen in Reinwardtia 6: 257 (1 962) ， nyms. She circumscribed D. racemosum p. p. p. ，inc l. syn. D. podocarpum. most broadly. Excepting Desmodium Distr. Distr. (subsp. podocarpum): India ，Nepal ， gardneri ， other three species she regarded as Malesia Malesia (Philippines: Luzon ， rare) ，China ， conspecific are distinguished as subspecies Taiwan (rare) ， Japan and Korea. of Hylodesmum podoca ψum (DC.) H. Specimens Specimens examined: Philippines. N. Luzon. Prov. Ohashi & R. R. Mill ，i. e. ， subsp. fallax Bengue t: M t. Santo Tomas. 4 f t. high ， flowers greenish (Schind l.) H. Ohashi & R. R. Mill ， subsp. pu 中le. 12 Oc t. 1904. Williams 1398 (NY); Bagino to oxyphyllum (DC.) H. Ohashi & R. R. Mill (= Ambuklao ， Prov. Bengue t. Merrill 4356 (NY); typical D. racemosum) ，and subsp. podo- Mountain Prov. ，Bauko Town ， above Kamatagan. 16 0 56'N. 120 0 54'E. al t. ca. 1450 m. T. Nemoto & al. carpum. Desmodium gardneri comprises H. 10750 10750 (TUS). laxum and H. leptopus (A.Gray ex Benth.) Distribution Distribution of Hylodesmum podocarpum H. Ohashi & R. R. Mill. (DC.) (DC.) H. Ohashi & R. R. Mill subsp. podocarpum is confined to Luzon. Occurr- subsp. oxyphyllum (DC.) H. Ohashi & R. ence ence of the species was ，however ，uncertain R. Mill in Edinb. 1. Bo t. 57: 182 (2000). in in the Philippines. Specimens kept as D. Hedysarum racemosum Thunb. in Mu 町ay ， podocarpum DC. from the Philippines so f紅 Sys t. Veg. ed. 14: 675 (May 1784) & as as 1 have examined 紅 'e all referable to H. Thunb. ， F l. Jap.: 285 (Aug. 1784) [Type: repandum (Vahl) H. Ohashi & R. R. Mill ， Japan. Thunberg (UPS)] ，non Aub l. (1755). 162 162 植物研究雑誌第79 巻第3号 平成16 年6月

Fig.7. Fig.7. Hylodesmum leptopus (A. Gray ex Ben 出.) H. Ohashi & R. R. Mill. a: Philippines. Luzon. Lectotype (K). (K). b: P 訂 t of the lectotype with label (enlarged frorn fig. a). c: Lornent (enlarged frorn fig. a).

Desmodium oxyphyllum DC. in Ann. Sci. D. racemosum (Thunb.) DC. var. Na t. (Paris) 4: 102 (Jan. 1825) [Type: Nepa l. pubescens F. P. Metcalf in Lingnan Sci. J. Wallich Wallich (G- DC)]. 19: 605 (1 940) [Type: China. “Fukien. near D. D. racemosum (Thunb.) DC. ， Prodr. 2: Chekiang border ，600-1200 m. viii 1924. 337 337 (Nov. 1825) ，nom. illegi t.; :品 1eeuwen in Ching 2513 (US holo.)]. Reinwardtia Reinwardtia 6: 257 (1962) ， p. p. ，exc l. syn. D. podocarpum DC. subsp. oxyphyllum t. ci t. D. podocarpum ， D. gardneri ， et D. (DC.) H. Ohashi in H. Hara ， F l. E. Hima l. fallax. fallax. 2: 65 (1 971) & in Ginkgoana 1: 150 (1973). D. D. japonicum Miq. ， Ann. Mus. Bo t. Podocarpium podoc αrpum (DC.) Y. C. Lugduno ・Batavi 3: 46 (1867) ，p. p・ Yang & P. H. Huang var. oxyphyllum (DC.) D. D. oxyphyllum DC. v征 japonicum Y. C. Yang & P. H. Huang in Bull. Bo t. Lab. Matsum. in J. Coll. Sci. Imp. Univ. Tokyo 12 N.E. Fores t. Ins t.， Harbin 4: 9 (1 979) ，nom. (Ten t. F l. Lutchu.): 415 (1899) [Type: Japan. illegi t. insula “insula Oshima". ix 1887. Y. Tashiro s. n. P. podocarpum v紅 . japonicum (Matsum.) (TI (TI holo.)]. P. H. Huang in Bull. Bo t. Res. ， Harbin 18: 6 June June 2004 Joumal of Japanese Botany Vo l. 79 No. 3 163

輪、ー

J

、、)'1 戸

a 裂l:l.1O;:. ~r ))護側出

Fig. Fig. 8. Hylodesmum podoca ψum (DC.) H. Ohashi & R. R. M i11 subsp. podoca 中 um. a: Philippines. Luzon. Me 凶 11 4356 (NY). b: Loments (enlarged from fig. a).

(1998) ，nom. illegi t. desmum podocarpum subsp. oxyphyllum Distr. Distr. (subsp. oxyphyllum): India ，Nepal ， (DC.) H. Ohashi & R. R. Mill. The two taxa Sikkim ，Bhutan ，Myanm 紅， Laos ，Vietnam ， are mounted on the same herbarium sheet in China ，Taiwan ，Japan ，Korea ，and Russian A. A duplicate specimen of Zollinger 1905 is Far Far Eas t. Probably e町 oneous from Malesia kept in BM ， but this composes purely O. (Jawa). (Jawa). caudata. Hylodesmum podoca ψum subsp. Specimen Specimen examined: Jawa. Zollinger 1905 ，p. p. 0 砂phyllum mounted on the sheet in A may (A). (A). be erroneously mixed in Zollinger 1905. The specimen ， Zollinger 1905 cited above ， This specimen must be brought from outside is is labeled as “ planta J avanica a c l. Zollingero of Jawa ， because H. podocarpum subsp. lecta" ， but is composed of two plants; Ohwia 0 砂'P hyllum has not been known in Jawa. caudat α(Thunb.)， H. Ohashi and Hylo- A specimen ， Lorzing 11737 (BO) from 164 164 植物研究雑誌第79 巻第3号 平成16 年6月

North Sumatera ， is named Desmodium Podocarpium repandum (Vahl) Y. C. oxyphyllum oxyphyllum (= Hylodesmum podocarpum Yang & P. H. Huang in Bull. Bo t. Lab. N.E. subsp. subsp. oxyphyllum) ， but is D. elegans DC. Fores t. Ins t.， Harbin 4: 13 (1979) ，nom. This This specimen was collected from a culti- il1 egi t. vated vated plant and D. elegans is not indigenous Distr.: Africa and Asia. Asia: North in in Malesia. Yemen ， Sri Lanka ， India ， Myanmar ，

Thailand ，Indo 由 China ，Malesia ，and China Hylodesmum repandum (Vahl) H. (Yunnan). No records from Australia. Ohashi & R. R. Mi l1 in Edinb. J. Bo t. 57: Specimens examined: Borneo. M t. Ki nabalu. M. 185 185 (2000). Togashi s. n. 16.9.1966 (TI) ， Upper Ki nabalu. J. & M. Hedysarum repandum Vahl ，Symb. Bo t. S. Clemens 34170 (BM ， NY). Jawa. Backer 9558 (K); Blume Blume (NY); o. Kuntze 24.5.75 (NY). Lesser Sunda [Type: [Type: Yemen. Forsskal 2: 2: 82 (1791) 1763. Is. Lombok. Elbert 837 (K). Forbes Timor. 3496 (BM). (holo. (holo. C-Vahl)]. Malaya. Pahang. Ridley 13553 (BM). Maluku. Aeschinomene remota Poir. in Lam. ， Mittel-Ceram. E. Stresemann 99 (L). New Guinea. Encyc. Encyc. 4: 452 (1 798). Brass 4141 (NY); Morobe Prov.: Bulldong Road ， al t. Desmodium repandum (Vahl) DC. ，Prod r. 2650 m ，montane fores t. M. Fallen 519 (L); Star Mts. ， Sibil-valley ， riverside. C. Kalkman 4046 (L). 2: 2: 334 (1 825); 恥1eeuwen in Reinwardtia 6: Philippines: Philippines: Luzon: Prov. Bengue t. Merrill 4330 (K ， 258 (1 962); Backer & Bakh. f.， Fl. Java 1: L， NY). Mindanao. M t. Katanglad. M. D. Sulit 604 (1 963); H. Ohashi in Ginkgoana 1: 160 PNH10019 (L). Sulawes i. Johansson & al. 308 (K); (1973); (1973); Verdc. ，Man. New Guinea Leg.: 404 Sumatera. W. & C. Bangham (K); East Coas t. Yates (1979). (1979). 1401 (NY). D. scα かe DC. ， Prodr. 2: 334 (1825) Hylodesmum repandum is often very simi- [Type: [Type: Mauritius. “ in insula Borbonia et lar to H. podoc αrpum subsp. podocarpum es- forsan forsan Madagascar" (G-DC: two specimens ， pecially in leaflets and loments. Sometimes both both app 紅白t1 y from Mauritius)]; Rid l.， Fl. both. were misidentified each other as noted Malay Penins. 1: 608 (1922); Me 町.， Enum. under the latter. Philipp. Philipp. Flow. Pl. 2: 289 (1 923). Verdcourt (1979) doubted the indigenous D. D. strangulatum Wight & Arn.， Prodr. Fl. status of Hylodesmum repandum (as Ind. Ind. Orien t.: 228 (1 834) [Type: Peninsula Desmodium repandum) in New Guinea. Ind. Ind. orientalis. Wight 774 (K lecto.; isolecto. Distribution pattem of this species differs en- E)]. E)]. tirely from those of other members of the D. D. cq 伊um Eckl. & Zeyh. ，Enum. Pl. genus. Afric. Afric. Austra l. 251 (Jan. 1836) [Type: South Africa. “ In locis humidis in hiatu montis MONARTHROCARPUS sylvoso sylvoso ad flumen ‘Makasanirivier' Monarthrocarpus Merr. in Philipp. J. (Kaffe r1 and) ， Maio". Ecklon & Zeyher Sci. ， C. 5: 88 (1910) [Type: Monarthro ・ (MEL? holo.; iso. K)]. carpus securiforme (Benth.) Merr. (= D. D. schimperi Hochs t. ex A. Rich. ，Ten t. Desmodium securiforme Benth.)]; Hutch. ， l. Fl. Abyss. 1: 205 (1847). Gen. Fl. Pl. 1: 480 (1 964). Papilionopsis Papilionopsis sη lidioides Steenis in Nova Desmofischera Holthuis in Blumea 5: 188 Guinea ，Bo t. 3: 17 (1960) [Type: The apical (1 942) [Type: Desmofischera monosperma part part of an inflorescence of Hylodesmum Holthuis]. repandum (Fabaceae) ， D. Bergman 287 (S) ， Desmodium subgen. Podocarpium sec t. which also comprises foliage ， roots and a Monarthrocarpus (Me 町.) H. Ohashi ，Gink- young scape of Burmannia disticha L. goana 1: 125 (1 973). (B urmanniaceae )]. M onarthrocarpus Me 町. is distinct from June June 2004 Journal of Japanese Botany Vo l. 79 No. 3 165

Fig.9. Fig.9. Monarthrocarpus securiforme (Benth.) Me 町. a: Philippines. Luzon. Clemens 16730 (UC). Reproduced from Ginkgoana 1， p l. 15a (1 973). b: Infructescence (enlarged from fig. a).

Desmodium in having falcate ，トseeded Ohashi in Ginkgoana 1: 128 (1 973); Verdc. ， loments ，sc 紅白ly rim-arillate ，flat seeds ， and Man. New Guinea Leg.: 407 (1979). diadelphous diadelphous stamens ， although it was in- Monarthrocarpus securiforme (Benth.)

cluded cluded in Desmodium subgenus Podo 目 Me 町. var. monophylla Me 町. in Philipp. J. carpium carpium (Ohashi 1973). Sci. ， C. 5: 90 (1 910) [Type: Philippines. The genus is monotypic. Mindanao ，District of Davao ， Catalonan. Copeland Copeland 937. Apr. 1904. in forests ，altitude Monarthrocarpus securiforme (Benth.) 125 m (herb. ?)] & Enum. Philipp. Fl. Pl. 2: Me 町. in Philipp. 1. Sci. ， C. 5: 88 (1 910) & 291 (1923). Enum. Philipp. Flow. Pl. 2: 291 (1 923). (Fig. Desmofischera monosperma Holthuis in 9). 9). Blumea 5: 189 ，fig. 5 (1 942) [Type: Taland Desmodium securiforme Benth. in Miq. ， is l.， Karakelong ， O. v. Beo. 26 Apr. 1926. H. l. Pl. Jungh.: 226 (1 852) [Type: Philippines. J. Lam 2637 (L holo.; iso. BO)]. Luzon. Luzon. Cuming 576 (K holo.; iso. BM ， K)]; Distr.: Endemic to Malesia: Maluku ，New Miq. ， Fl. Ned. Ind. 1(1): 255 (1 855); Guinea ，Philippines ， and Sulawesi. Meeuwen in Reinwardtia 6: 258 (1 962); H. Specimens examined: Maluku: M t. Permatang 500 166 166 植物研究雑誌第79 巻第3号 平成 16 年6月 m. Kostermans 958 (A ，BO ，K ， L) M t. Permatang ， (1 964) ， but it has been widely recognized as along along Sabgawi river ， al t. 200 m. Kostermans 908 (L) a member of Desmodium. Although O. t. M t. Totodohoe 30 m. Kostermans 761 & 1455 (L); caudata has remarkable characters in com- Morotai. Morotai. Kostermans 7856 (L); Morotai. H. J. Lam 3645 3645 (L); Mortai. Lam 3681 (K); Halmahera Island. mon with Desmodium ，i. e. ，jointed pods Taylor Taylor P-319 (A); Maluku. Yoshida 2429 (L); Obi Is l. with hooked hairs ， the differences between Anggai ， al t. 200 m. E.F. de Vogel 4195 (L). New Ohwia and Desmodium are described by Guinea: Guinea: Misoo l. near Waima. Shrub ，1 m high ，fls Ohashi (1 973) for D. caudatum in morpho- white. white. D. R. Pleyta 1046 (A ，K ， L); Mori Ri ver ，Abau logical and palynological characters. It is re- sub-district ，Papua ， al t. 50 m. Sayers NGF19660 (L); Ohwia it Usino ， al t. 400 ft. Henty NGF28020 (L); Sorong ， al t. vived to a genus ， because is distinct 20 m. Plezte 629 (L). Philippines: Luzon. Cuming 576 from Desmodium in having a disk around the (K holo.; iso. BM ，K) ，lrosin. Elmer 15907 (NY ，TI) ， base of the pistil and seeds with a scarcely Isabela ， Calamangan-Mariano. M.S. Clemens 16729 developed hilum lacking a rim-ari l. (UC); (UC); Isabela Prov. ， rather open forest trail above Moreover ， Ohwia differs from most of Calamangan ，M t. Moises. Aug. 1926. M. S. Clemens Desmodium in having distinctly veined 16730 16730 (UC). Sulawesi. Taland is l.， Karakelong. Clealed Clealed old folest ， rather frequent ， al t. 90 m. Small chartaceous petals (against slightly veined undershrub ，p 訂 tly prostrate ， cololla and filaments membranaceous petals in most of Desmo ・ white ，仕. green. Lam 2637 (L ，BO types); Taland is l. dium) ， equally indented linear pods and Salebabae ， top G. Ajambana. 20 May 1926. H. J. Lam coarsely reticulate pollen grains (Ohashi 3057 3057 (L); weg Oedjoeng Lamoeroe- Takalala. C. 1999). Monod de Froideville 421 (L). The genus comprises two species of which one one is found in Malesia. OHWIA Ohwia H. Ohashi in Sci. Rep. Tohoku Ohwia caudata (Thunb.) H. Ohashi in Univ. Univ. ser. 4，Bio l. 40: 243 (1 999) [Type: Sci. Sci. Rep. Tohoku Univ. ser. 4，Bio l. 40: 243 Ohwia caudata (Thunb.) H. Ohashi (= (1 999); H. Ohashi in K. Iwats. & al.， Fl. Jap. Hedysarum caudatum Thunb.)]. IIb: IIb: 254 (2001); Kumar & Sane ， Leg. S. Catenaria Catenaria Benth. in 乱1iq. ， Pl. Jungh.: 217 Asia: Asia: 212 (2003). & 220 (1 852) [Type: Catenaria laburnifolia Hedysarum caud α，tum Thunb. in Mu 町 ay ， (Poir.) (Poir.) Benth. (Hedysarum laburnifolium Sys t. Veg. ed. 14 ， 675 (1 784); Thunb. ， F l. Poir.)] ，non Roussel ， F l. Calvados ed. 2: 85 Jap.: Jap.: 286 (1 784) [Type: Japan. Thunberg (1806) ， nec. Stemberg ， Versuch Fl. Vorwelt (UPS)]. (UPS)]. (Tentamen): 1 (Tentamen): 25: t. 52 ，f. 1 (1 825); Schind l. H. H. laburnifolium Poir. in Lam. ，Encyc l. 6: in in Repert. Spec. Nov. Regni Veg. 20: 275 422 (1 805) [Type: from Java. (P) ，n. v.]. (1 924); Hutch. ， Gen. Fl. Pl. 1: 480 (1 964). Desmodium caudatum (Thunb.) DC. ， Desmodium sec t. Catenaria (Benth.) Prodr. Prodr. 2: 337 (1825); Meeuwen in Benth. Benth. in Benth. & Hoo k. f. ， Gen. Pl. 1: 520 Reinwardtia Reinwardtia 6: 248 (1 962); Backer & Bakh. (1 865); Taub. in Engler & Prantl ，Na t. f. ， F l. Java 1: 605 (1 963); Ohwi ， F l. Jap.: ed. Pflanzenfam. Pflanzenfam. 111 ，3: 328 (1894). Eng l.: 561 (1 965); H. Ohashi in Ginkgoana Desmodium subgen. Caten α ria (Benth.) 1: 1: 91 (1 973); Huang & H. Ohashi in Fl. Baker Baker in Hoo k. f.， Fl. Bri t. Ind. 2: 163 Taiwan 3: 256 ， p l. 572 (1 977); H. Ohashi in (1876); (1876); H. Ohashi in Ginkgoana 1: 91 Enum Fl. Pl. Nepal 2: 115 (1979); Grierson (1 973). & Long ， F l. Bhutan 1(3): 677 (1 987); Ohwia H. Ohashi is named instead of Sanjappa ， Leg. India: 150 (1992); Huang & Catenaria Catenaria (Ohashi 1999). Catenaria was H. H. Ohashi in Fl. Taiwan ed. 2 ，3: 252 (1 993); created created by Bentham (1852) and maintained H. H. Ohashi in Dy Phon & al.， Fl. Camb. Laos by Schindler (1924b ) and Hutchinson Vietn. Vietn. 27: 78 (1994); Yang & Huang in Fl. June June 2004 Journal of Japanese Botany Vo l. 79 No. 3 167

Reipub l. Popu l. Sin. 41: 19 (1 995); Pedley ， Baker in Hoo k. f. ， F l. Bri t. lnd. 2: 162 Rev. Rev. Hand. Fl. Ceylon 10: 175 (1996). (1876). D. D. laburnifolium (Poi r.) DC. ， Prodr. 2: The genus comprises eight species of 337 (1825); Rid l.， Fl. Malay Penins. 1: 608 which two are found in Malesia. (1922). (1922). Catenaria Catenaria laburnifolia (Poir.) Benth. in Key to the species of Phyllodium Miq. ， P l. Jungh.: 220 (1 852); Miq. ， F l. Ned. in 島lalesia lnd. lnd. 1(1): 257 (1855). 1. Lateral leaflets similar to the terminal one C. C. caudata (Thunb.) Schind l. in Repert. and distinct from primary bracts; flowers

Spec. Spec. Nov. Regni Veg. 20: 275 (1 924); more than 10 mm long ...・ H ・... P. vestitum Hutch. ，Gen. Fl. Pl. 1: 480 (1964). 1. Lateral leaflets smaller (ca. 2 times) than Distr.: Distr.: Sri Lanka ， E. lndia ，Myanm 飢 the terminal one and similar to primary lndo-China ，Malesia ，China ，Taiwan ，and bracts; flowers less than 7 mm long …… 2 Japan. Japan. 2. Pods usually 2-jointed ，sp 訂 sely pubescent Specimens Specimens examined: Jawa. Reinwardt (L); on lateral surfaces ， reticulate-veined; Zollinger Zollinger 1905 (BM; A ， p. p. ，exc l. Desmodium stipules narrowly triangular ，6-8 mm long. podocarpum subsp. 0巧'phyllum) ，Junghuhn 169 (K ，L) ， Lowest calyx 国 lobe almost equal to the lat 田 Backer Backer 2363 (K ，L); Tjisaroea-South (WNW slope of t. M t. Gede) ， al t. 1200 m ， in moist fores t. S. 1. van eral one. Bracteal leaflets uniformly Ooststroom Ooststroom 12737 (PNH). Many specimens collected appressed pubescent on nerves on adaxial

in in Jawa are kept in BO. Malaya. Peninsula Malaysia. surfaces... ・H ・..…...・ H ・.....・ H ・..P. pulchellum Johor. Johor. Ri dley 15434 (BM). Sumatera. A りeh ，in dense 2. Pods usually 3-4オ ointed; stipules broadly evergeen evergeen fores t. Iwatsuki & al. 812 (BO ，KYO ，L); triangular ，3-5 mm long. Lowest calyx- near near Lake Tawar. Bangham 803 (A ， K); Korinchi. lobe lobe longer than the lateral one. Bracteal Robinson Robinson & Kl oss 20 (BM). lea f1 ets densely hairy on nerves on adaxial

PHYLLODIUM surface ...・ H ・-…...・ H ・.....・ H ・....・ H ・.P. elegans Phyllodium Desv. ，J. Bo t. Agric. 1: 123 ， tab. tab. 5， fig. 24 (1 813) ，exc l. Phyllodium Phyllodium elegans (Lour.) Desv. in lutescens lutescens Desv. [Type: Phyllodium Mem. Soc. Linn. Paris 4: 324 (1 825) [Type: pulchellum pulchellum (L.) Desv. (= Hedysarum China. Canton-Macao (Gaudichaud 147 P pulchellum pulchellum L.)]; H. Ohashi in Ginkgoana 1: neo. designated by Dy Phon in F l. Camb. 260 (1973); H. Ohashi & a l.， Adv. Leg. Sys t. Laos Vietn. 27: 38 ， 1994)]; H. Ohashi in 1: 1: 299 (1 981); Allen & Allen ， Leg.: 516 Ginkgoana 1: 269 (1 973); Dy Phon in Dy (1981); (1981); Huang & H. Ohashi ， F l. Taiwan ed. Phon & a l.， Fl. Camb. Laos Vietn. 27: 37 2， 3: 350 (1993); Dy Phon in Dy Phon & al.， (1 994). l. Fl. Camb. Laos Vietn. 27: 28 (1 994); Y. C. V紅. javanicum Schind l. in Repert. Spec. Yang & P. H. Huang in F l. Reipub l. Pop. Nov. Regni Veg. 20: 270 (1924) [Syntypes: Sin. Sin. 41: 8 (1 995); Pedley ， Rev. Hand. Fl. e Java & Madura. (BO ，CAL ，etc.)]; H. Ceylon 10: 164 (1 996); H. Ohashi in Sc i. Ohashi in 1. Jpn. Bo t. 72: 350 (1 997). Rep. Rep. Tohoku Univ. ser. 4，Bio l. 40: 245 Phyllodium vestitum Miq. ，non DC. ， F l. (1999); (1999); Pedley in Austrobaileya 5: 255 Ned. Ind. 1(1): 261 (1855) ，exc l. syn. ci t. (1999). (1999). Desmodium blandum van Meeuwen in Desmodium sec t. Phyllodium (Desv.) Reinwardtia 6: 247 (1962) ， p. p. ， inc l. Benth. Benth. in Benth. & Hook. f. ，Gen. Pl. 1: 519 specim. eJ awa; Backer & B 紘 h. f. ， F l. Java (1865). (1865). 1: 603 (1 963) ， ut D. blandum Knaap. Desmodium subgen. Phyllodium (Desv.) Distr. (var. javanicum): Malesia (Jawa) and Thailand. 168 168 植物研究雑誌第79 巻第3号 平成16 年6月

Specimins Specimins examined: Jawa. Teyjesmann 1868 (K ， Phyllodium vestitum Benth. in Miq. ， P l. NY) ， Backer s. n. July 1904 (K ，L) ，Miquel s. n. (K) ， Jungh.: 217 (1 852) [Type: “Dicerma Horsfield Horsfield 60 (BM) & Horsfield 81 (GH ，K ，L) ， vestitum vestitum Wall. ，Numer. List n. 5739. ln Horsfield Horsfield 585 (K) ， Junghuhin 117 (K) ， Winterbottom s.

Moulmega" (K 町 W)]; Ohashi in Ginkgoana n. n. July 1846 (K) ，Waitz s. n. (L); Tjis 訂 oa. Moler on 17/4 17/4 1897 (A). 1: 279 (1 973); Dy Phon in Dy Phon & al.， Fl. Phyllodium Phyllodium elegans var. javanicum is dis- Camb. Laos Vietn. 27: 34 (1994). tinguished tinguished from v紅 . elegans by its grayish Desmodium vestitum (Benth.) Benth. ex hairy hairy leaves ， broadly ovate terminal leaflet Baker in Hoo k. f.， Fl. Birt. Ind. 2: 162 with with rounded base and distinct lateral and (1 876); Rid l.， Fl. Malay Penins. 1: 607 third third (between lateral nerves) nerves ，wrin- (1 922). kled kled prim 訂 y leafy bracts with conspicuous Distr.: Myanm 紅， Thailand ，Indo-China ， third third nerves and 4- 5 mm long fruiting and Malesia. pedicels. pedicels. Specimen examined: Malaya. Peninsula Malaysia. Phyllodium Phyllodium eleg α ns var. eleg α ns is not Kehah. Rawei island. Ridley 15980 (BM). found found in Malesia ， and is confined to Thailand ， Indo-China and S. China (Ohashi TADEHAGI 1997c). 1997c). Tadehagi H. Ohashi ，Ginkgoana 1: 280 (1 973) ， p. p. ， exc l. subgen. Godefroya Phyllodium pulchellum (L.) Desv. ， J. (Schind l.) H. Ohashi [Type: Tadeh α gi Bo t. Agric. 1: 124 (1813) & in Mem. Soc. t1 匂uetrum (L.) H. Ohashi (= Hedysarum Linn. Linn. Paris 4: 324 (1825); Benth. in Miq. ， P l. triquetrum L.)] & in 1. Jpn. Bo t. 78: 276 Jungh.: Jungh.: 217 (1852); H. Ohashi in Ginkgoana (2003). 1: 1: 276 (1 973); Dy Phon in Dy Phon & al.， Fl. The genus comprises six species of which Camb. Laos Vietn. 27: 40 (1994); Pedley in one is found in Malesia. Me 町 ill (1923) re- Austrobaileya Austrobaileya 5: 257 (1 999). corded Tadehagi auriculatum (DC.) H. Hedysarum pulchellum L. ， Sp. Pl.: 747 Ohashi (as Desmodium auriculatum DC.) (1753) (1753) [Type: Herb. Linn. No. 92 1. 24 from Luzon ， but it was misidentification of (LINN dlecto. ， see Pedley in Turland & T. triquetrum (L.) H. Ohashi nothosubsp. arvis Jarvis 1997)]. phil 伊pinense H. Ohashi var. phil 伊pinense Dicerma pulchellum (L.) DC. in Ann. Sci. H.Ohashi. Na t. (Paris) 4: 236 (1 825) & in Prodr. 2: 239 (1825) (1825) & Mem. Leg.: 328 (1826). Tadehagi triquetrum (L.) H. Ohashi in Desmodium pulchellum (L.) Benth. ， F l. Ginkgoana 1: 290 (1 973) ， p. p. ，inc l. subspp. Hongkong.: Hongkong.: 83 (1 861); Rid l.， Fl. Malay triquetrum & pseudotriquetrum ， cet exc l. & Penins. Penins. 1: 607 (1922); Merr. in Philipp. 1. in 1. Jpn. Bo t. 78: 284 (2003). Sci. ， C. 5: 79 (1910) & Enum. Philipp. Flow. Hedysarum t1 匂uetrum L. ， Sp. Pl.: 746 l. Pl. 2: 288 (1 923); Meeuwen in Reinwardtia (1753) [Type: Herb. Hermann 3: 39 & 4: 18 6: 6: 276 (1 962); Backer & B はh. f. ， F l. Java (BM) ， fide Pedley 1996]. 1: 1: 603 (1 963); Verdc. ，Man. New Guinea Leg.: Leg.: 404 (1 979). Key to the subspecies and their hybrids Distr.: Distr.: Asia ， Australia and Pacific 。， f Tadeh αεi 釘iquetrum (Solomon Is ふ Asia: Sri Lanka ， lndia ， 1. Loments densely to sparsely pubescent on

Myanmar ，Thailand ，Indo-China ，Malesia lateral surfaces... ・H ・H ・H ・.....・ H ・.....・ H ・H ・H ・.2 (throughout) ，China ，Taiwan ，and S. Japan 1. Loments glabrous on lateral surfaces …・ .3 (Ryukyus). (Ryukyus). 2. Leaf-blade usually more than 3 times longer longer than broad ，linear to narrowly ob- June June 2004 Joumal of Japanese Botany Vo l. 79 No. 3 169

long-ovate long-ovate or narrowly ovate.. 44688 (A ，K ，NY ， US). Sulawes i. Rachmat 24 (L). …… subsp. triquetrum Sumatera. Bacon 121 (K) ， Lorzing 13087 (A ，BO ，K) ， 2. 2. Leaf-blade less than 3 times longer than Yates 2236 (NY). broad ， usually ovate or narrowly ovate... .. subsp. subsp. pseudotriquetrum (DC.) H. …...・ H ・.. .nothosubsp. philippinense var. var. palawanense Ohashi in Ginkgoana 1: 295 (1 973) in 1. Jpn. 3. 3. Leaf-blade less than 3 times longer than Bo t. 78: 287 ，fig. 10 (2003). broad ， usually ovate or narrowly ovate. Desmodium pseudotriquetrum DC. in An n. Sci. Na t. (Paris) 4: 100 (1825) [Type: …… subsp. pseudotriquetrum Nepa l. Wallich (G holo. ，A iso. partly)] ，& 3. 3. Leaf-blade usually mo 問 than 3 times longer longer than broad ， linear to narrowly ob- Mem. Leg. 320 (1 826); Me 町. in Philipp. 1. long-ovate long-ovate or narrowly ovate. Sci. ， C. 5: 87 (1910) ，Enum. Philipp. Flow. l. Pl. 2: 288 (1 923) ，& in Trans. Amer. Phi l. ..nothosubsp. …..nothosubsp. phil 伊'P inense Soc. Soc. 24: 201 (1935). ...・ ...var. phil 伊'P inense Desmodium triquetrum subsp. pseudo- triquetrum triquetrum (DC.) Prain in 1. Asia t. Soc. subsp. subsp. triquetrum: H. Ohashi in Gin- Bengal ， P t. 2，Na t. His t. 66: 390 (1 897): kgoana 1: 296 ， p l. 42a (1 973) & in 1. Jpn. Meeuwen in Reinwardtia 6: 263 (1 962); Bo t. 78: 286 ，figs. 8-9 (2003). Verdc. ，Man. New Guinea Leg.: 410 (1 979). Desmodium triquetrum (L.) DC. ， Prodr. 2: Distr. Distr. (subsp. pseudotriquetrum): India ， 326 (1825); Prain in 1. Asia t. Soc. Bengal ， Nepal ，Bhutan ，Malesia (rare in New Guinea t. Pt. 2，Na t. His t. 66: 390 (1 897) ， ut subsp. and Philippines) ， S. & SW. China ，and genuina genuina Prain; Rid l.， Fl. Malay Penins. 1: Taiwan. Taiwan. 611 (1 922); Me 町.， Enum. Philipp. Flow. Pl. 2: 2: 290 (1 923); Meeuwen in Reinwardtia 6: Specimens examined: New Guinea. Milne Bay Distric t. Brass 24360 (K ，L ， US). Philippines. Luzon. 262 (1962) ， ut subsp. triquetrum; Backer & Lepanto. Merrill 4477 (K ，NY ， US). Bakh. f. ， F l. Java 1: 604 (1 963); Verdc. ， Man. New Guinea Leg.: 410 (1 979) ， ut nothosubsp. philippinense H. Ohashi in J. subsp. subsp. t1 匂uetrum. Jpn. Bo t. 78: 291 (2003) ，as ‘'p hili} フ'P inensis" . Pteroloma Pteroloma triquetrum (L.) Desv. ex Benth. 二 Tadehagi triquetrum subsp. triquetrum in in Miq. ， P l. Jungh.: 220 (1 852). xTadehagi triquetrum subsp. pseudotriquet- Distr. Distr. (subsp. triquetrum): Asia ， N. rum (DC.) H. Ohashi Australia Australia (?)， and Pacific. Sri Lanka ，India ， var. philippinense H. Ohashi in J. Jpn. Bangladesh ， Myanm 征， Thailand ， Indo- Bo t. 78: 293 ， figs. 11-12 (2003) ， as china ， Malesia (throughout) ， S. & SW. ‘'phil 伊'P inensis" . China ，and Taiwan (rare). Pteroloma pseudotriquetrum x? triquet- Specimens Specimens examined: Borneo. About 3 miles from rum: Schind l. in Repert. Spec. Nov. Regni Tamu Dara t. In dense bracken and lalang in full sun ， Veg. 20: 273 (1924). by path. Bright pinkish- mauve flowers ，no scen t. Damton 78 (BM ， L); Ki nabalu. 1. & M. S. Clemens Desmodium auriculatum auc t. non DC.: 26795 26795 (K ， L). Jawa. Bl ume (A) ，Horsfield L66 (K); Me 汀.， Enum. Philipp. Flow. Pl. 2: 284 Timor. Timor. Unknown collector (L ，G-DC as Desmodium (1 923). auriculatum auriculatum DC. in Candolle: Prodromi Herbarium ， Dist r. Malesia. Endemic to the Philippines IDC micro-edition sheet no. 381). Maluku. Amboina. (Luzon). Robinson Robinson 557 (GH ，K ， US); de Wiljes-Hissink 99 (A). New Guinea. Papua. Milne Bay Distric t. Brass 24360 Specimens examined: Philippines. Luzon. Prov. (A ，K ，L ， US); Goragatabu Creek area. Pullen 6766 (A ， Rizal ， Bosoboso. July 1906. Ramos Bu r. Sci. no. 1036 ， ， l. K ， L). Philippines. Luzon. Ramos & Edano Bur. Sci. (GH-holotype NY US); Prov. Riza Ramos Bur. Sci. 22685 22685 (GH); Riza l. Ramos & Edano Bu r. Sci. 29494 170 170 植物研究雑誌第79 巻第3号 平成16 年6月

(K ， US); Manila. Vidal 2610 (A ， K). Trifidacanthus unifoliolatus Merr. in This This hybrid has loments glabrous on the Philipp. J. Sci. ， C. 12: 269 (1917) [Type: lateral lateral surface as subsp. pseudotriquetrum. Luzon. M. Ramos Bur.Sci. 27196 (K holo.)]; Merrill Merrill (1923) determined this hybrid as Me 江， Enum. Philipp. Flow. Pl. 2: 292 Desmodium auriculatum based on “Ramos (1 923); H. Ohashi et al. in J. Jpn. Bo t. 71: 63 B. B. S. 1036 ，Ramos B. S. 22685 and Merrill (1 996). 7394 ぺalthough 1 have not examine the last Desmodium horridum Steen. in Blumea one. one. See the differences between Tadehagi 24: 482 (1 978) [Type: Lombo k. De Voogd auriculatum auriculatum and T. triquetrum (nothosubsp. 2130 (L holo.; iso. A ， K)] & in Reinwardtia phil 伊']J inense var. phil 伊']J inense) in Ohashi 10: 24 (1 982). in in J. Jpn. 78: 279 (2003). Desmodium unifoliolatum (Me 町.) Steen. V 征・ palawanense H. Ohashi in J. Jpn. in Reinwardtia 10: 24 (1 982). Bo t. 78: 293 ，fig. 13 (2003) ， as ''p alawa- Distr.: Malesia (Lesser Sunda Is: Lombok nensis" nensis" and Flores and the Philippines: Luzon) ， S. Distr. Distr. Malesia. Philippines and New Vietnam and S. China (Hainan). Guinea. Guinea. Specimens examined: Lesser Sunda Is. Lombok ， Specimens Specimens examined: Philippines. Palawan. Edano Pr 司ja. De Voogd 2130 (L holo. of D. horridum; iso. 4 Mar. 195 1. PNH14138 (A-holotype ， L). New K). Philippines. Luzon. M. Ramos Bur. Sci. 27196 (K Guinea. Guinea. Fergusson Is. Brass 27226 (A ，K ，L ， US). holo. of T. unifoliolatus; iso. A); Il ocos Norte. Ramos This This hybrid has loments pubescent on the Bu r. Sci. 32925 (BM ，NY). lateral lateral surface. Phytogeographic Phytogeographic considerations TRIFIDACANTHUS 1. Distribution of genera in Malesia Trifidacanthus Trifidacanthus Merr. in Philipp. J. Sci. ， The boundary between Malesia and Indo- C. C. 12: 269 (1917) [Type: Trifidacanthus China floristic regions lies slightly to the unifoliolatus unifoliolatus Merr.]; H. Ohashi & al.， Adv. north of the political border of Malaysia Leg. Leg. Sys t. 1: 299 (1 981); Dy Phon in Dy (Johns 1995). Continental Asia in this paper Phon & a l.， Fl. Camb. Laos Vietn. 23: 94 means an 訂 ea excluding the southem half of (1987); (1987); Y. C. Yang & P. H. Huang in Fl. Malay Peninsula. Malesia has much diversity Reipub l. Pop. Sin. 41: 2 (1995). in distribution of organisms. In cu 町 ent Trifidacanthus Trifidacanthus Me 町. was first established phytogeographical studies ， Malesia is di- and placed ，by Merri l1 (1 917) ， close to vided into three floristic regions: Westem Arthroclianthus Arthroclianthus Baill. endemic to New Malesia (Sunda Shel f) including Bomeo ， Caledonia ， but was included in Desmodium Jawa ，southem half of Malay Peninsula ， by Steenis (1 982). It was again recognized as Palawan ， and Sumatera ， Central Malesia in- distinct distinct from Desmodium and was consid- cluding the Lesser Sunda Islands ，Maluku ， ered ered one of the genera nearest to Desmodium the Philippines (excluding Palawan) ， and

(Ohashi (Ohashi et al. 1996). Trifidacanthus is ch 訂- Sulawesi ， and Eastem Malesia or Papuasia acterized acterized in having lignified 佐ifid spines (de- including New Guinea and the Bismarck rived rived from an inflorescence axis and two Archipelago (Johns 1995). The boundary be- lateral lateral pedicels) ， the m 征 ginal venation pat- tween these floristic regions seems to co 町 e- tem of the leaflets ， sympodial terminal spond to a line previously proposed on the inflorescences ，and loments without hooked basis of distribution of remarkable animals hairs. hairs. or plants in Malesia. Various faunal or The genus is monotypic. floristic lines have been proposed in 恥1alesia: Haxley's Haxley's or Merrill's line ， Lydekker's line ， June June 2004 Journal of Japanese Botany Vo l. 79 No. 3 171

Wallace's Wallace's line ， Weber's line ，Zollinger's floristic region and southern China. line ，etc. Some of the lines agree entirely or Hylodesmum has four species in Malesia: H. p征 tly with the eastern or western boundaries laxum ， H. leptopus and H. repandum are of of the Central floristic region and others run common and widely distributed in Malesia ， within within the floristic region. The Lydekker' s but H. podocarpum is rare and known only faunal faunal line seems to correspond with the in Luzon ， the Philippines. floristic floristic boundary between the Eastern and The remaining six genera (Aphyllodium ， Central Central floristic regions. Codariocalyx ，Dendrolobium ，Desmodium ， The distribution patterns of the 13 genera Phyllodium ， and Tadeh α gi) contain species of of Desmodium and relatives are diverse in distributed widely in continental Asia Malesia. Malesia. The monotypic Monarthrocarpus is through Malesia to Australia ， but their distri- endemic endemic to Malesia and is apparently con- bution patterns 紅 e individually characteris- fined fined to the Central and Eastern Malesia tic. Aphyllodium comprises eight species floristic floristic regions. It is known in 乱1aluku ，New including four endemic to Australia (Pedley Guinea ， Luzon in the Philippines and 1999) ， one each endemic to New Guinea (A. Sulawesi. Sulawesi. Hanslia is also found in the novoguineense) and Myanm 紅 (A. hispidum Central Central and Eastern Malesia floristic regions (Schind l.) H. Ohashi; Ohashi 1997b) ， one and and Australia but not in continental Asia. confined to N ew Guinea and Australia (A.

Five Five genera 訂 'e distributed in continental schindleri) ， and only A. biarticulatum dis- Asia Asia and Malesia: Desmodiastrum ，Hegnera ， tributed widely in Sri Lanka ，India ，Myan- Hylodesmum ，Ohwia ， and Trifidacanthus ， mar ， Indo-China ， Hainan ， Western and but but their distribution pattems differ from Central Malesia and Australia. Thus there each each other within these regions. With the ex- 紅 e three species in Malesia ， but there is a ception ception of Hylodesmum these genera are rep- distant gap in the distribution of A. resented resented by only a single species each in biarticulatum between JawalSulawesi and Malesia. Malesia. All species of Desmodiastrum (4 Australia. This disjunction straddles the spp.) 紅 e confined to India except Desmo ・ range of the two East Malesian species ， A. diastrum diastrum parviflorum ， which extends to East novoguineense (New Guinea) and A. Jawa in the Western Malesia floristic region ， schindleri (New Guinealnorthern Australia). but but is not found elsewhere in Indo-China. Dendrolobium comprises 19 species and is Hegnera is distributed in Myanmar ， abundant in Indo-China (9 spp.) and Malesia Thailand ，Indo-China ， and in Jawa ，Malay to Australia (7 spp.) (Ohashi in Lewis et al. Peninsula ， and Sumatera in the Western in prep ふIt includes four species endemic to Malesia Malesia floristic region. Ohwia is widely dis- Malesia of which three are from the tributed tributed in India ， Sri Lanka ，Nepal ，Sikkim ， Philippines. Dendrolobium papuacola is en- Bhutan ，Myanmar ，Vietnam ，Laos ， and E. demic to N ew Guinea and close to D. Asia ， but in Malesia is confined to the arbuscul αwhich is distributed in New Western Western Malesia floristic region. Hegnera Guinea ， northern Australia and F討i. and and Ohwia are considered to be derived in Dendrolobium rugosum is app 訂 ently part of Malesia Malesia from Indo-China ， but it is possible the Indo-Chinese floristic element and is that that Desmodiastrum migrated to Malesia di- confined to Malay Peninsula in Malesia. It rectly ， rather than through Indo-China. seems to be derived from continental Asia. Trifidacanthus Trifidacanthus is monotypic and rare ， being Dendrolobium tri α ngulare and D. umbel- found found sporadically in Hainan ， south latum are widespread species ， but the former Vietnam ，Luzon and Lombo k. This suggests occurs on hills or low altitude montane sides ， a connection between the Central Malesia while the latter is coasta l. 172 172 植物研究雑誌第79 巻第3号 平成16 年6月

Desmodium includes one species (D. three to each of Aphyllodium ， Codariocalyx bolsteri) bolsteri) endemic to the Philippines ， 15 spe- and Phyllodium ，two to Hanslia ， and one cies cies common in Asia ，eight in Australia and each in Desmodiastrum ，Hegnera ，Monar- only only three covering both continents. Of the throcarpus ，Ohwia ，Tadehagi ， and Trifida- 15 15 species distributed in continental Asia canthus. The native Malesian species 訂 e only only five are found in N ew Guinea ， while all diverse in distribution pattem ，and 紅 e pri-

the the eight of the Australian species 紅 e dis 噛 marily divided into four groups: 1) species tributed tributed in New Guinea. This pattem of dis- endemic to Malesia ， 2) species distributed in tribution tribution shows that Desmodium of New Malesia and continental Asia ， 3) species dis- Guinea Guinea and Australia appe 紅 s to be closely tributed in Malesia and Australia ， and 4) spe- related related than that between New Guinea and cies distributed in the whole area from the the regions west of New Guinea treated in continental Asia through Malesia to this this paper (Central and Westem Malesia Australia. These groups are referred to here floristic floristic regions and continental Asia). as Malesia group ， Asia group ，Australia Phyllodium Phyllodium appears to have undergone a group and Asia-Australia group ，respectively m 司jor radiation in Indo-China where six of (Table 1). eight eight species of the genus occur. Four of these these six species 紅 e almost endemic to the (i) Malesia group region. region. In Malesia ， Phyllodium pulchellum is The eight following species (14 % of the widely widely distributed ， but P. vestitum is con- 56 native Malesian species) 訂 e distributed as fined fined to the Malay Peninsula and P. elegans indicated in parentheses after each name: V紅 . javanicum to Java (var. elegans is con- Aphyllodium novoguineense (New Guinea) ， fined fined to Indo-China and southem China). Dendrolobium cumingianum (Luzon) ，Den- Tadehagi Tadehagi is represented by T. triquetrum in drolobium geesinkii (Luzon) ，Dendrolobium Malesia. Malesia. It shows unique variation in the papuacola (New Guinea) ， Dendrolobium Philippines ， Lesser Sunda islands and N ew quinquepetalum (Luzon and rarely in Guinea Guinea (Ohashi 2003). The genus is rare in Mindoro) ， Desmodium bolsteri (Luzon) ， Australia: Australia: T. triquetrum is uncertain in Hanslia hen ηi (New Guinea and newly in Australia Australia and only one endemic T. robustum Borneo) and Monarthrocarpus securiforme is is known in north Westem Australia (Pedley (Luzon and Isabela islands in the 1999). 1999). Philippines ，Maluku ， and New Guinea). The following two subspecies are also en- 2. 2. Distribution of species in Malesia demic to Malesia: Desmodium nemorosum Species Species of Desmodium and its related gen- subsp. whi ザordii is endemic to Luzon and era era may be considered difficult to interpret in Desmodium strigillosum subsp. celebicum is terms terms of distribution due to their weedy na- confined to Sulawesi. Tadeh α gi triquetrum ture ture and remarkable ability for dispersal of produces hybrids (nothosubsp. philippinense) seeds. seeds. However ，many species have definite between subsp. triquetrum and subsp. distribution distribution pattems. Those represented by pseudotriquetrum in Luzon ，Palawan ， and introduced introduced and widespread species are ex- New Guinea. The hybrid is divided into two

ceptiona l. Of the sixty 四 four species 1 recog- varieties: Var' philippinense in Luzon and nize nize in the group in Malesia ，eight species of va r. palawanense in Palawan and New Desmodium are introductions from tropical Guinea. These subspecies and notho- or or subtropical America. Fifty-six are native subspecies belong actually to the Malesia species species of which 27 belong to Desmodium ， group ， but they are not treated in this eight eight to Dendrolobium ， four to Hylodesmum ， phytogeographical consideration as sep 紅 ate 」吉ロ oNCChF Table Table 1. Distribution pattems of species in Desmodium and relatives in Malesia

Malesia Malesia group Asia group Australia group Asia-Australia group

Aphyllodium Aphyllodium novoguineense Codariocalyx gyroides Aphyllodium schindleri Aphyllodium biarticulatum

Dendrolobium cumingianum C. motorius Dendrolobium arbuscula Cod α， riocalyx microphyllus D. geesinkii Dendrolobium rugosum Desmodium brachypodium Dendrolobium umbellatum D. papuacola D. triangulare D.filiforme Desmodium gangeticum D. quinquepetalum Desmodi α strum parviflorum D. gunnii D. heterocarpon Desmodium bolsteri Desmodium auricomum D. nemorosum D. tl 併orum Hanslia Hanslia hentyi D. capU αtum D. pullenii Phyllodium pulchellum 旬。ロ円ロ Monarthrocarpus Monarthrocarpus securiforme D. dichotomum D. rhytidophyllum Tadeh α gi triquetrum [5 [5 gen. 8 spp.] D. diffusum D. tenax [6 gen. 8 spp.] mw-o D. D. heterophyllum D. trichostachyum 同』 D. D. laxiflorum Hanslia ormoc α rpoides 8820 D. D. megaphyllum [4 gen. 11 spp.] D. D. renifolium D. D. rubrum 回。 S D. sequ 似 ロ〕『〈。 D. D. strigillosum D. D. s ηracifolium --13Z0 D. teres D. D. velutinum ・ω D. D. zonatum Hegnera obcordata

Hylodesmum 1似 um H. H. leptopus H. H. podocarpum H. H. repandum Ohwia caudata Phyllodium Phyllodium elegans P. P. vestitum Trifidacanthus Trifidacanthus unifoliolatus [9 [9 gen. 29 spp.]

目、吐い 174 174 植物研究雑誌第79 巻第3号 平成16 年6月 from its mother species. gion. The latter subgroup contains Dendro-

Among the Malesia group ， five species lobium rugosum ，Desmodiastrum parviflo ・ (and (and one subspecies and one nothovariety) rum ，Desmodium αuricomum ， D. renifolium ， are are found in Luzon island in the Philippines ， D. rubrum ， D. teres ，Hegnera obcordata ， and four species (and one nothovariety) 紅 e Hylodesmum podocarpum ， .Ohwia caud αta ， found found in New Guinea. In addition ，one sub- Phyllodium elegans ，and P. vestitum. species species of an Asian group is endemic to Dendrolobium rugosum ， Desmodium Sulawesi ，Desmodium strigillosum subsp. rubrum ， and Phyllodium vestitum 紅 e note- celebicum. celebicum. With the exception of worthy as they are confined only to the Monarthroc α rpus securiforme ， which is Malay Peninsula. Infraspecific taxa of D. found found widely in Malesia (Maluku ，New heterocarpon (i ム subsp. angustifolium and Guinea ，Philippines and Sulawesi) ， other en- subsp. ovalifolium) and D. strigillosum (i ム demic species occur in locally limited habi- subsp. strigillosum) are also limited to the re- tats. tats. gion. Desmodium heterocarpon subsp. A disjunction between the Westem and angustifolium is confined to the Malay Eastem Malesia floristic regions is app 紅 ent Peninsula ， but subsp. heterocarpon is widely in in the distribution pattem of Hanslia hentyi distributed ， belonging to the Asia-Australia and Tadehagi triquetrum nothosubsp. group. Hylodesmum podocarpum and its philippinense philippinense v 紅 . palawanense. infraspecific taxa 紅 e COmrrlon in continental Asia ， but is very r紅 e in Malesia: subsp. (ii) (ii) Asia group podocarpum is known rarely from Luzon in

Twenty 四 nine species (52 % of the 56 na 園 the Philippines (Fig. 8) and subsp. tive tive species) are distributed in Malesia and oxyphyllum is probably not found in Malesia. continental continental Asia (excluding southem half of The taxa of the West Malesia subgroup 紅 e Malay Peninsula) ， but not in Australia. They considered to have come into the area from belong belong to nine genera as shown in Table 1. continental Asia. Desmodium heterophyllum was recorded Of the 29 species belonging to the Asia from Australia by Whele & Frances [in Bo t. group ，De ωS幻modi ωα strum Bull. Bull. Queensland 22: 14 (1 920)] but the spe- only example 血tha 抗 tis 副dis 司dunc 凶tivel 砂 y distributed cies cies is said to be recent introduction to in India and J awa. Desmodium dichotomum Australia Australia (Pedley 1999). Desmodium has a similar distribution pattem ， occurring strigillosum strigillosum and D. velutinum were also re- in India ， China (Yunnan) and Malesia (J awa ， garded garded as recent introductions (Pedley 1999). Lesser Sunda Islands ，and Sulawesi). They Desmodium ren{ 戸Jlium was recorded from are not in Indo-China or Thailand. Australia Australia (Meeuwen 1962 ， Ohashi 1973) ， but Trifidacanthus unifoliolatus shows a is is not included in Pedley's work (1 999). I unique distribution pattem in the Asia group. have have not examine specimens of the species It is a rare species and is found only in one from Australia. or a few localities in Hainan ，southem The Asian group is divided into two sub- Vietnam (Phu Kh anh) ，Luzon ，Lombok and groups groups according to their distribution pat- Flores. It s principal areas of distribution 紅 e tems; tems; a general subgroup and a West around the South China Sea ，and disjunc- Malesia Malesia subgroup. Species of the general tively Lesser Sunda Islands. The species is subgroup subgroup include 18 species and are found in not known from Indo-China except for the two or three floristic regions in Malesia ， but one locality in southem Vietnam and the the the others include 11 species and are distrib- West Malesia floristic region in Malesia. uted uted only in the West Malesia floristic re- Desmodium sequax is widely distributed June June 2004 Joumal of Japanese Botany Vo l. 79 No. 3 175 in in montane areas in Northern India ，Nepal ， (iv) Asia-Australia group Sikkim ， Bhutan ， Myanmar ， Thailand ， Eight species (1 4 % of the native Malesian Malesia ，Laos ，Vietnam ，China ， and Taiwan. species) occur across Malesia ，continental It It usually occurs above 600 m altitude ，al- Asia ， and Australia (Table 1). Tadehagi though though it has been collected between 200 m tr 匂uetrum was recorded from Australia by (in (in Vietnam) and 2850 m (in China) above Meeuwen (1 961) ，but ， according to Pedley sea sea leve l. It is distributed sporadically in (1 999) ，its occu 町'ence seems dubious. Malesia Malesia (Maluku ，New Guinea ，Philippines ， Several other species need to be confirmed Sulawesi ， and Sumatera) ， where it is appar- as native to Australia or no t. Codariocalyx ently ently confined to high altitudes. This distri- motorius was first recorded from Australia bution bution pattern suggests that the species may by Holtze in 1892 (Chapman 1991) ， spread spread by birds and may be imported from Desmodium renifolium by Bailey in 1900 continental continental Asia. The distribution pattern is (Chapman 1991) and D. repandum by similar similar to that of Ohwia caudl αta. Meeuwen (1962) ， but Pedley (1 996 ， 1999) denied denied their occu 町 ence in Australia. 1 have (iii) (iii) Australia group not found specimens of these species from The following eleven species (20 % of the Australia and therefore include them in the 56 native Malesian species) 紅 e found in Asia group in this study.

Malesia Malesia and Australia (areas in Malesia 紅 e Cod α， riocalyx microphyllus ， Desmodium shown in parentheses): Aphyllodium heterocarpon ， D. velutinum and Tadehagi schindleri schindleri (New Guinea) ， Dendrolobium triquetrum are included in this group ， but arbuscula arbuscula (New Guinea) ， Desmodium their infraspecific taxa show different pat- brachypodium brachypodium (New Guinea) ， D. filiforme terns of distribution from the type taxa. (Jawa (Jawa and New Guinea) ， D. gunnii (New Codariocalyx microphyllus occurs widely in Guinea) ， D. nemorosum subsp. nemorosum Malesia and continental Asia and less fre- (New Guinea; subsp. whi ゆrdii belongs to quently in northeastern Australia ， but it has a the the Malesia group) ， D. pullenii (Alor in the variety endemic to Assam and a form to Lesser Lesser Sunda Islands and New Guinea) ， D. Nepal (Ohashi 1984). Desmodium hetero- rhytidophyllum rhytidophyllum (New Guinea) ， D. tenax carpon subsp. heteroca ψon belongs to the (New Guinea) ， D. trichostachyum (Maluku Asia-Australia group ， while subsp. and and New Guinea) ， and Hanslia . angustifolium and subsp. oval 扮lium are not ormocarpoides ormocarpoides (many places in Malesia and found in Australia ， so the latter two subspe- New Hebrides) (Table 1). As can be seen ， in cies belong to the Asia group. Desmodium Malesia ，these species 訂 e mainly distributed velutinum is divided into subsp. velutinum in in or even confined to New Guinea. All ex- and subsp. longibracteatum; the latter sub-

cept cept Hanslia ormocarpoides occur more fre 闇 species belongs to the Asia group. Tadehagi quently quently in Australia where they are often triquetrum ， though its distribution in widely widely distributed. Desmodium filiforme is Australia is dubious ， includes subsp. disjunctively disjunctively distributed in Jawa from New pseudotriquetrum of the Asia group. Subsp. Guinea Guinea and northern Australia. It is close to pseudotriquetrum is frequent in continental D. D. pullenii ， which is found in Alor in Asia but r訂 e in Malesia where it is only Malesia Malesia and northern Australia. Direct found sporadically in New Guinea and the dispersals dispersals from northern Australia to Jawa or Philippines. From their distribution patterns Alor ，rather than via New Guinea ， are sup- and infraspecific differentiation patterns posed posed from their distribution patterns. Codariocalyx microphyllus ， Desmodium heterocarpon ， D. velutinum ， and Tadehagi 176 176 植物研究雑誌第79 巻第3号 平成16 年6月 t1 匂uetrum may be regarded mainly as mem- Desmodium and related genera were re- bers bers of the Asia group. corded from Jawa (Backer and Bakhuizen Among other members of the Asia- van den Brink Jr. 1963) ， of which 28 were Australia Australia group ， Aphyllodium biarticulatum ， considered to be native. The three excluded Desmodium gangeticum ， D. triflorum ， and species 訂 e naturalized from America ， Phyllodium Phyllodium pulchellum may also be reg 紅 ded Desmodium adscendens (v 紅. trifoliastrum) ， as as members of the Asia group on the basis of D. tortuosum ， and D. uncinatum. Thirty-one

their their distribution pattems. Only Dendro 幽 species are recognized as native to Jawa in lobium lobium umbellatum is different from the this paper and 紅 'e classified into the follow- other other members of the Asia group. It is a ing 10 genera: Aphyllodium biarticulatum coastal coastal plant and disperses the seeds en- (not included in Backer and Bakh. fよ closed closed in corky pericarp by an ocean cu 町 en t. Codl αriocalyx 3 spp. (as Desmodium With the exception of the last species ， all gyroides ， D. microphyllum ，and D. motorium members of the Asia-Australia group are in Backer and B 紘 h. fふ Dendrolobium 2 considered considered as originally distributed in Asia. spp. (as Desmodium tri α ngulare and D. umbellatum umbellatum in Backer and Bakh. fよ 3. 3. Comparisons of Desmodium and re- Desmodiastrum parviflorum (as Desmodium lated lated genera between Jawa ，New Guinea alysicarpoides in Backer and Bakh. fよ and the Philippines Desmodium 16 spp. (1 5 species in Backer Comprehensive Comprehensive works for Leguminosae and B 北 h. f. ， of which D. lasiocarpum is were provided by Merrill (1 923) for the synonymous; and D. diffusum and D. teres Philippines ， Backer and Bakhuizen van den 紅 e added) ， Hegnera obcordata (as Brink Brink Jr. (1963) for Jawa and Verdcourt Desmodium obcordatum in Backer and (1979) (1979) for New Guinea. Desmodium and re- B 紘 h. f.)， Hylodesmum 3 spp. (as lated lated genera in these regions appe 紅 to be Desmodium laxum and D. repandum in better better known than in other regions in Backer and Bakh. f.; and Hylodesmum Malesia. Malesia. However ，Jawa is estimated as hav- leptopus is added ， but H. podocarpum is du- ing ing been botanized more than 2-4 times bious and excluded) ， Ohwi α1 sp. (as more intensively than the Philippines and Desmodium caudatum in Backer and Bakh. Papua New Guinea based on comparisons of fふ Phyllodium 2 spp. (as Desmodium previous previous collections in each regions blandum and D. pulchellum in Backer and (Campbell (Campbell and Hammond 1989 ， Johns B 紘 h. fよ and Tadehagi triquetrum (as 1995). 1995). The relatively recent discoveries of Desmodium triquetrum in Backer and Bakh. such such distinct species as Desmodium hentyi f.) (Table 2). Verdc. Verdc. from near Lae in Papua New Guinea ， Distribution pattems of these species are Dendrolobium papuacola from Loloki River as follows: 22 species belong to the Asia in in Papua New Guinea and Dendrolobium group (71 % of the 31 native species in geesinkii geesinkii from Luzon may reflect a paucity Jawa) ， one (Desmodium filiforme) to the of of collections of Desmodium and related Australia group (3 % of species in Jawa) ， genera genera from New Guinea and the and eight belong to the Asia-Australia group Philippines. Philippines. In spite of the disparity in the in- (26 % of Javanese natives). All species ex-

ventories ventories among these regions ，1 have at 同 cept Desmodium filiforme relate to continen- tempted tempted a prelimin 紅 y comparison of species tal Asia in distribution. and and distribution pattems for the genera in the (ii) New Guinea: Verdcourt (1 979) re- areas: areas: corded 33 species including one unknown (i) (i) Jawa: 百lirty-one species of from New Guinea ， of which seven were in- June June 2004 Journal of Japa Ii ese Botany Vo l. 79 No. 3 177

Table 2. Distribution of Desmodium and related species in Jawa ， the Philippines and New Guinea with those in in Indo-China and Australia

Indo-China Indo-China 1 Jawa Philippines New Guinea Australia 2 F3--Fδ-附一白九山政邸一倒的問帥伽訪問仇幻附叩O 一ゆn m由一加問一如一戸一ePM ny.Fn回一一如一一間一一dN側一・・・・・・一川一・・…Gune司一一 一 一例励伊一似幻伽一一一一仕一一一時一一中一一一一一世一一巳一一 一一丸一一一 一一一一回一一一 一一…・・・・・・一一...... 一...... 一-・・・・・・… cbhMZfMWJ 仰 D.D.aamm-Mm1OLnm

叫

Jω州問 pc--o・叫んぬ仰mg…m 白島 mwm伽…同一問問伽 仰 D.a 似仰仰仰仰一∞一 及的mM'ZmF α EE 伽…町一氏附幻 一戸一円…湖一伽仰山川一一 一一 一均一一一 d Psr-.m N G-mLU DlLH dN rL-4F-m nm a ew 自 a 向 nr--dn-e-qu-n 戸しw-w e先 iw 。 u U M 一一 一一 一一町一一一一一一一・・・……・・・・・・一一 一 一 一・・・・…一 ω ic--rohmZWN ..... 仇 停 D.n ••• 仏 mM Jωm--itmmmmdα Hm 脆一問一勾畑一明一ω ω me ・ 一 …一一…… …川一刷同一柳川帥初刑制，一町一一江-m一回一一 一 03- O N β 1-ca M5W ωnuea、}ノ 一 …附 一問問問・ 一一一 …・・・・・・一一一 Ummzmwd WM hh 仰 mmω dd幻 msm 仇及 加伊…仰白mMm nu 及 ahω 誌な協同おお州mωおじ間的仰仰リ削んり 6 6 向島崎市川仇仰仇 均 ・・・ Mdmwcmm ••••••• ma0 md印刷 MZU仰 aahh・m 似仇 仰伽m 印刷伽ゎ伽附叫仰 d' MG 叩 hhanuaaa仰仇 ••••••••••••••••••• 仇一mMZammb mZZ 附 Hmm u ・-一例

附 ω 勾仇肋Md一切 砂川

1 Dy Phon ， Ohashi and Vidal (1994) ， 2Pedley (1 999) ，3 su bsp. whi φrdii ， 4 subsp. nemorosum ，5 v紅 . elegans ，6var .javanicum. 178 178 植物研究雑誌第79 巻第3号 平成16 年6月 troduced troduced from America. Thirty-two indige- and Australia groups in these genera seems nous species 訂 'e recognized in this study to agree with that of Malesia and Papuasia. including including Verdcourt' s unknown (= Desmo- Among the Asia-Australia group only dium sequax) (Table 2). They are Aphyllo- Aphyllodium biarticulatum is not found in dium 2 spp. (as Desmodium biarticulatum in N ew Guinea ， although this species is widely Verdcourt ， but is Aphyllodium novoguine- but sporadically distributed in Asia ， Malesia ense; ense; and A. schindleri is added) ，Codario- and Australia. However ， in New Guinea ， calyx calyx 2 spp. (as Desmodium gyroides and D. there is one endemic species of Aphyllodium microphyllum microphyllum in Verdcourt) ，Dendrolobium as well as a member of the Australia group. 3 spp.(as Desmodium umbellatum in Differentiation of the genus in New Guinea Verdcourt ， but Dendrolobium arbuscula and and Australia needs to be studied. D. D. papuacola are added) ，Desmodium 17 (iii) The Philippines: Merrill (1923) enu-

spp. spp. (14 species in Verdcourt; and Desmo ・ merated 32 species of Desmodium and one dium dif.fu sum ， D. pullenii and D. tenax are species of each of Mon α rthrocarpus and added) ， Hanslia 2 spp. (as Desmodium h仰の li Trifidacanthus in the Philippines of which 30 and D. ormocarpoides in Verdcourt) ，Hylo- species 紅 e native to the region (Table 2). desmum 3 spp. (as Desmodium laxum and D. Desmodium malacophyllum ， D. procumbens ， repandum in Verdcourt; Hylodesmum D. scorpiurus and D. trifoliastrum 紅 e ex- leptopus leptopus is added) ， Monarthrocarpus 1 sp. cluded from his list as species introduced (as (as Desmodium securiforme in Ve 吋court) ， from America. Twenty-nine species are rec- Phyllodium Phyllodium 1 sp. (as Desmodium pulchellum ognized in this paper and 訂 e reclassified as in in Verdcourt) ， and Tadeh α gi 1 sp. (as follows: Codariocalyx 3 spp. (as Desmodium Desmodium tf 匂uetrum in Verdcourt). Addi- gyrans ， D. gyroides and D. microphyllum in

tions tions to Verdcourt (1 979) are seven species ， Me 町 ill) ，Dendrolobium 4 spp. (as Desmo ・ i. i. e. ， Aphyllodium schindleri (as D. biarti- dium cumingianum ， D. quinquepetalum and culatum culatum var. australiense in Verdcourt) ， D. umbellatum in Merrill; and Dendrolobium Dendrolobium arbuscula ， D. papuacola ， geesinkii) ，Desmodium 13 spp. (as 14 species

Desmodium dif.fu sum ， D. pullenii ， D. tenax in Me 町 ill in which D. buergeri ， D. lasio ・ (as (as D. rhytidophyllum subsp.α cutifoliolum in carpum and D. ovalifolium are synonymous; Verdcourt) ，and Hylodesmum leptopus (as D. and D. d{ がlsum and D. nemorosum (subsp. laxum subsp. leptopus in Verdcourt). whi ゆrdii) 紅 e added); Hanslia ormocarpoi- Thirty-two Thirty-two species of Desmodium and des (as Desmodium ormocarpoides in relatives relatives in New Guinea comprise four spe- Me 町 ill) ，Hylodesmum 4 spp. (as Desmodium cies cies of the Malesia group including two en- laxum ， D. podocarpum ，and D. scal) フe in demic species (Aphyllodium novoguineense Merrill; and Hylodesmum leptopus is added) ， and Dendrolobium papuacola) ， 10 from the Monarthrocarpus securiforme (as in

Asia Asia group ， 11 from the Australia group ，and Me 町 ill) ， Phyllodium pulchellum (as Desmo ・ seven seven from the Asia-Australia group. All dium pulchellum in Me 町 ill) ， T. α dehagi species species of the Australia group (i. e. 100 %) triquetrum (as Desmodium auriculatum ， D. are are found in New Guinea ， whilst 10 ofthe29 pseudotriquetrum and D. triquetrum in species species of the Asia group known in Malesia Merrill) and Trifidacanthus unifoliolatus (as (34 (34 %) are found in New Guinea. It is appar- in 乱1errill). ent ent that the members of Desmodium and The distribution pattems of these species relatives relatives in New Guinea have s仕onger ties to fit with the Malesia ，Asia ，Australia ，and

Australian Australian taxa than continental Asian ones. Asia 目 Australia groups. The Malesia group The distribution boundary between the Asia comprises five species (1 7 % of native spe- June June 2004 Joumal of Japanese Botany Vo l. 79 No. 3 179 cies cies in the Philippines) ，i. e. ， Dendrolobium Guinea (Table 2). With the exception of cumingianum ， D. geesinkii ， D. quinquepet α-， Codarioc α lyx motorius and Desmodium

lum ，Desmodium bolsteri ， and Monarthro 凶 capitatum the remaining 11 J awanese spe- carpus carpus securiforme. The Asia group includes cies not found in New Guinea are also not the the following 15 species (52 % of species in distributed in the Philippines the the Philippines): Codariocalyx gyroides ， C. Fourteen species from New Guinea (44 % motorius ， Desmodium caph αtum ， D. of New Guinea species) are not found in diffusum ， D. heterophyllum ， D. 1似 iflorum ， Jawa (Table 2). Other than the four species D. D. sequax ，D. のracifolium ， D. velutinum ， D. belonging to the Malesia group ， the remain- zonatum ，Hylodesmum 1ωum ， H. leptopus ， ing species (10 spp.) all belong to the

H. H. podocarpum ， H. repandum ， and Trifid ル， Australia group (Table 1). This indicates that canthus canthus un 約liolatus. The Australia group the species of Desmodium and its relatives in comprises comprises two species (7 % of species in the New Guinea are very close to those of Philippines) ，Desmodium nemorosum (subsp. Australia but those in Jawa 紅 e no t. whi ゆrdii) and Hansli α ormocarpoides. The (v) Comparison between Jawa and the Asia-Australia Asia-Australia group has seven representa- Philippines: Jawa and the Philippines have tives tives in the (24 Philippines % of the 紅 ea's nearly equal number of species in the genera species) ，i. e. ，Cod α， riocalyx microphyllus ， treated in this paper. Nineteen species of Dendrolobium umbellatum ， Desmodium Jawa (61 % of the island's species) are also gangeticum ， D. heterocarpon ， D. triflorum ， found in the Philippines (Table 2) ， but 12 Phyllodium Phyllodium pulchellum and Tadehagi species (the remaining 39 %) are indigenous trzquetrum. trzquetrum. to Jawa (Table 2). The Jawanese species ab- Geographically ， the Philippines lies close sent in the Philippines suggest a close con- to to south China and Taiwan. Most of the spe- nection between Jawa and Indo-China and cies cies belonging to the Asia and Asia-Australia India in distribution ， because they 紅 e found groups 訂 e also found in China an d/ or in Indo-China an d/ or India ， but not (or only Taiwan ， while the two species of the rarely) in China. Desmodiastrum parviflo- Australia Australia groups found in the Philippines do rum ， Desmodium auricomum ， D. teres ， not not occur in China and Taiwan. This indi- Hegnera obcordata ， and Phyllodium elegans cates cates a close distributional relationship in var. javanicum are not distributed in China ， Desmodium and related genera between the Aphyllodium biarticulatum is only known in Philippines Philippines and south China and Taiwan. Hainan and Desmodium dichotomum and D. (iv) (iv) Comparison between Jawa and megaphyllum are found in Yunnan in China. New Guinea: Thirty-one species of Apart from four species endemic to the Desmodium and relatives 紅 e recorded from Philippines ， the following six species of the Jawa and 32 from New Guinea. Such simi- Asia and Asia-Australia groups are not found larity larity in numbers of species may suggest in Jawa: Desmodium nemorosum ， D. sηraci- those those in New Guinea are poorly known in folium ， Hanslia ormocarpoides ，Hylodes- the the light of its larger areas ，richness of envi- mum podocarpum ， Monarthrocarpus securi- ronmental ronmental conditions and rem 泊ning unde- forme ， and Trifidac α nthus unifoliolatus. veloped veloped natural 訂 'eas. It seems probable that Hylodesmum podocarpum is very r訂 e in more new species will be found in New Luzon and is not found in Indo-China but is Guinea Guinea in future. common in continental China and Japan. Its Of the 31 J avanese species ，18 紅 e in com- ranges from NW India ， through Nepal ， mon with New Guinea (58 % of all species China ， Taiwan ， Korea and into Japan. in in Jawa) ， and 13 紅 e not found in N ew Trifidacanthus unifoliolatus is a very r訂 e 180 180 植物研究雑誌第79 巻第3号 平成16 年6月 species species not only in Malesia but also in China pressions of Desmodium from the Pleiocene and Vietnam. These facts may indicate that of Westem Caucasus (Sh 北 η1 1992). These species species in Desmodium and related genera in records suggest that the genera of the the Philippines 紅 e remote from J awa. Three Desmodiinae were rather differentiated endemic endemic Dendrolobium species 訂 e charac- quickly in the Miocene (Tertiary) ， because teristic teristic of these genera in the Philippines. both taxa and localities are diverse. I inter- (vi) (vi) Comparison between New Guinea pret the Malesian taxa of Desmodium and re- and the Philippines: Twenty species from lated genera as derived from ancestral New Guinea 訂 e also found in the species from the Sino- Indian region. There is Philippines Philippines (Table 2). Twelve New Guinean no evidence to indicate the time initiate ex- species ，all of which belong to the Australia pansion of distribution of those genera from group ， are absent from the Philippines (Table Sino- Indian regions to Malesia. It is sup- 3). 3). Desmodium nemorosum is distributed in posed that it occurred probably in the both both regions ， but separated into subsp. Miocene. Immigration must have been done nemorosum in New Guinea and subsp. from time to time since then when geo- whitfordii whitfordii endemic to the Philippines. graphic and climatic corridors were available Codariocalyx Codariocalyx motorius ， Dendrolobium for colonization. Malesian taxa appe 紅 to cumingianum ， D. geesinkii ， D. quinquepeta- have settled and differentiated in several lum ，Desmodium bolsteri ， D. capitatum ， D. major regions following this expansion of

sη racifolium ，Hylodesmum podocarpum and Sino- Indian ancestors. Subsequent immigra 嶋 Tl 切dα canthus unifoliolatus are not found in tion must have stemmed from Australia New Guinea. With the exception of four which appears to be one of the areas of diι endemics ， the remaining Philippine species ferentiation of Desmodium. Subgenus all all belong to the Asia group and show a Sagotia of the genus seems an example that close close distributional association to Indo-China differentiated secondarily in Australia where and and China ， not to New Guinea. 24 species including 9 endemics 紅 'e known (Pedley (Pedley 1999). All species of Desmodium in 4. 4. Diversity and distribution in the Australia group (Table 1) belong to the Desmodium and related genera in Malesia subgenus ， and it is recognized as the most The origin of diversity in genera and spe- advanced group in Desmodium (Ohashi cies cies as well as distribution pattems of 1973). The concentration of endemic species Desmodium and its related genera in Malesia in Luzon and New Guinea seems to be the is is considered here. These genera are most di- most remarkable feature in the evolution of

verse verse in the Sino- Indian region and are con- the Desmodium group in Malesia. It is natu 国 sidered sidered to have primarily originated and ral that the present-day composition and dis- differentiated differentiated in this region (Ohashi et al. tribution of taxa in Desmodium and its 1981). 1981). They are included in the subtribe related genera in Malesia 征 e diverse ，be- Desmodiinae. Desmodiinae. Fossil woods of Ougeinia (= cause they 紅 e derived from various spatially Desmodium in Ohashi 1973 but is recog and temporally diverse ancestors. nized nized as distinct ， cf. Nemoto and Ohashi 2003) 2003) were found from the Miocene of I wish to express my sincere thanks to the Assam ， India (cf. Wheeler and Baas 1992) ， Directors and Curators of A & GH ，B 加I，BO ， leaves leaves of Codariocalyx (as Desmodium cf. CAL ，CAS ，E ，K ，KYO ，L ，LAE ，LD ，NY ， microphyllum) ，Desmodium and Phyllodium PNH ，S ，SING ，TI ，TNS ，TUS ，UC ，UPS ， (as (as D. pulchellum) from the Miocene of and US for allowing me to work in their her- China (Guo and Zhou 1992) and lea f1 et im- baria an d/ or for loans of specimens. I would June June 2004 Journal of Japanese Botany Vo l. 79 No. 3 181 like like to thank F. Adima (L) ， D. E. Boufford Brummitt R. K. 2001. World Geographical Scheme for and and E. W. Wood (Harvard University Recording Plant Distributions. ed. 2. Hunt Institute for for Botanical Documentation ， Carnegie Mellon Herb 訂 ia) ， B. Verdcourt ， R. M. Polhill ， G. University ，Pittsburgh. Lewis ， B. Schrire ， and Lulu Rico- Ar ce (K) ， Campbell D. G. and Hammond H. D. (eds.) 1989. and and H. Wiriadinata and B. Sunarno (BO) for Floristic inventory of tropical countries: The status their their help when 1 had worked in their her- of plant systematics ，collections ， and vegetation ， b紅白. 1 am also very grateful to Y. Endo plus recommendations for the future. New Y ork (lbaragi (lbaragi University) ， Y. lokawa (Jyoetsu Botanical Garden ，New Y or k. Kyoiku University) ， T. Kajita and J. Murata Candolle A. P. de 1825a. Notice sur quelques genres et esp とces nouvelles de legumineuses ，extraite de di- (The (The University of Tokyo) ， T. Nemoto vers Memoires presentes a la Societe d'Histoire (lshinomaki (lshinomaki Senshu University) ， and Y. naturelle de Geneve ， pendant le cours des annees Tateishi Tateishi (Ryukyu University) for their col- 1823 et 1824. Ann. Sci. Na t. (Paris) 4: 90-103. laboration laboration in field and herbarium works and 一一一 1825b. Desmodium D C. Prodromus Systematis Naturalis Naturalis Regni Vegetabilis 2: 325-339. Treuttel et for for providing necess 訂 y references. 1 sin- Wurtz ，Paris. cerely cerely thank to Y. Kadota (TNS) ，1. Murata ， 一一一 1826. Revue de la tribu des Hedysarees. H. H. Nakai (Sapporo lntemational University) ， Memoires sur la famille des Legumineuses. pp. L. L. Rico- Ar ce and E. W. W ood for their help 295-355. A. Belin ，P 紅 is. in in bibliography; to Gregory Kenicer (Edin- Chapman A. D. 199 1. Australian Plant Name Index burgh burgh University) for reading the manu- D- 1. Australian Fl ora and Fauna Series Number script; script; and to lhsan A. Al-Shehbaz (MO) ， D. 13. Australian Government Publishing Service Canberra. Canberra. E. E. Boufford ， Alice Grabovskaya (LE) ， L. Dalzell Dalzell N. A. 185 1. Contributions to the Botany of Rico- Ar ce ， and H. Walter Lack (B) for their Western India. Hook. 1. Bo t. 3: 206-212. help help searching the type of H edysarum Doyle 1. J. ，Chappill J. A.， Bailey C. D. and Kajita T. malacophyllum Link an d/ or Chamisso col- 2000. Towards a comprehensive phylogeny of leg- lections. lections. umes: evidence from rbcL sequences and non- molecular molecular data. In: Herendeen P. S. and Bruneau A. A. (eds よ Advances in Legume Systematics 9: 1- References References 20. Royal Botanic Gardens ，Kew. Ali Ali S. 1. 1977. Flora of West Pakistan. No. 100 Dy Phon ， P. 198 1. Taxons nouveaux pour la flore de Papilionaceae. Papilionaceae. 398 pp. Dep 訂 tment of Botany ， 1' lndochine. Bull. Mus. Natn. His t. Na t. Paris 4 University University of Karachi ， Karachi. (Desmodium pp. ser. ，sec t. B ， Adansonia 1: 111-119. 346-354). 346-354). 一一一， Ohashi H. and Vidal J. E. 1994. Fl ore du Backer Backer C. A. and Bakhuizen van den Brink Jr. R. C. Cambodge du Laos et du Vietnam. 27 Legumino- 1963. 1963. Fl ora of Java. 1: Papilionaceae. pp. 565-645. sae (Fabaceae) ， Papilionoideae ， Desmodieae. N. N. V. P. Noordhoff ， Groningen. Museum National d'Histoire Naturelle Laboratoire Bailey Bailey C. D. ， Doyle J. J. ，K 司ita T. ，Nemoto T. and du Phanerogamie ，Paris. Ohashi Ohashi H. 1997. The chloroplast rpl2 intron and Fosberg F. R. 1966. Critical notes on Pacific Island ORF184 as phylogenetic markers in the legume Plants. 1. Micronesica 2: 143-152. tribe tribe Desmodieae. Systematic Botany 22: 133- 一一一 1968. Critical notes on Pacific Island Plants. 2. 138. 138. Micronesica 4: 255-259. Baker Baker J. G. 1876. Desmodium. In: Hooker J. D. ，The Greuter W. ， McNeill J. ， Barrie F. R.， Burdet H. M. ， Flora Flora of British India 2: 161-175. Ken t. Demoulin V. ，Filgueiras T. S. ， Nicolson D. H. ， Bentham G. 1852. Leguminosae. In: Miquel F. A. W. ， Silva P.C. ，Skog 1. E. ， Trehane P， Turland N. J. ， Plantae Plantae Junghuhnianae. pp. 205-269. Leiden. and Hawksworth D. L. 2000. International Code of 一一一 1865a. Description of some new genera and spe- Botanical Nomenclature (Saint Louis Code). cies cies of tropical Leguminosae. Trans. Li nn. Soc. Koeltz Scientific Books ，Konigstein. 25: 25: 297-320. 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Gardens ，Kew. Philippine Fl owering Plants. 2: 241-323 (Desmo ・ Hollis Hollis S. and Brummitt R. K. 1992. World dium pp. 28 4- 291 ， Monarthrocarpus p. 291 ， Geographical Geographical Scheme for Recording Plant Trifidac α nthus p. 292). Man i1 a (Bureau of Distributions. Distributions. Hunt Institute for Botanical Printing). Documentation ， Carnegie Mellon University ， Nemoto T. and Ohashi H. 2003. Diversity and evolu- Pittsburgh. Pittsburgh. tion of anatomical structure of loments in tribe Holmgren P. K.， Holmgren N. H. and Barnett L. C. Desmodieae (Papilionoideae). In: K1 itgaard B. and 1990. 1990. Index Herbariorum. P紅 t 1: The Herbaria of Bruneau A. (eds よ Advances in Legume the the World. New York Botanical Garden ，New Systematics. Part 10. pp. 395 -4 12. Royal Botanic Yor k. Gardens ，Kew. Huang T. C. and Ohashi H. 1993. Desmodium. In: Nicolson D. H. 1978. Ill egitimate “basionyms ，" impact Huang T. C. et al. (eds.) ，Fl ora of Taiwan Second on priority and author citation or ， the rise of edition. edition. 3: 251-270. Editorial Committee of the Desmodium incanum and fall of D. canum Fl ora of Taiwan ， Second Edition. Taipei ， Taiwan. (Fabaceae). Taxon 27: 365-370. Hutchinson Hutchinson J. 1964. Genera of Fl owering Plants Ohashi H. 197 1. A monograph of the subgenus (Angiospermae) (Angiospermae) based principally on the Genera Dollinera of the genus Desmodium (Leguminosae). Plantarum Plantarum of G. Bentham and J. D. Hooker. vo l. In: Hara H. (ed.) ，Fl ora of East Himalaya 2: 259- 1， 516 pp. Oxford Univ. Press ， London. (Desmo- 320. University of Tokyo Press ， Tokyo. dieae dieae pp. 477 -4 84). 一一1973. The Asiatic Species of Desmodium and It s Johns Johns R. J. 1995. Malesia-An introduction. Curtis Bo t. Allied Genera (Leguminosae). 318 pp. ， 76 pls. Mag. 12(2): 52-62. Ginkgoana 1. Academia Scientific Book Inc. ， Kajita Kajita T. and Ohashi H. 1994. Chloroplast DNA varia- Tokyo. tion tion in Desmodium subgenus Podocarpium 一一一 1979. Leguminosae. In: Hara H. and Williams L. (Leguminosae): (Leguminosae): Infrageneric phylogeny and H. J. (eds よ On Enumeration of the Fl owering infraspecific infraspecific variations. J. Plant Research 107: Plants of Nepal 2: 103-133. British Museum 349-354. 349-354. (Natural History) ， London. 一一一，一一一， Nemoto T. ， Bailey C. D. ， Doyle J. L. and 一一一 1984. Infraspecific taxa of Desmodium Doyle Doyle J. J. 1996. Polyphyly of the legume tribe microphyllum (Thunb.) DC. (Leguminosae). J. Jpn. Desmodieae: Desmodieae: Evidence from the chloroplast Bo t. 59: 42 -4 7. genome. genome. Am. J. Bo t. 83: S165. 一一一 1988. Desmodium (Leguminosae) of the Kumar S. and Sane P. V. 2003. Legumes of South Hawaiian Islands. Sci. Rep. Tohoku Univ. ser. 4， Asia ，a check-lis t. Royal Botanic Gardens ，Kew. Bio l. 39(3): 171-189. Lock J. M. and Heald J. 1994. Legumes of Indo-China ， 一一一 199 1. Taxonomic studies in Desmodium

a check 働 lis t. Royal Botanic Gardens ，Kew. heterocarpon (L.) D C. (Leguminosae). 1. Jpn. Bo t. 一一一 and Simpson K. 1991. Legumes of West Asia ，a 66: 14- 25. check-lis t. Royal Botanic Gardens ，Kew. 一一一 1994. Desmodium Desvaux. In: Dy Phon P. ， Meeuwen M. S. van 196 1. Notes on Desmodium Desv. Ohashi H. et Vidal J. E. ，Fl ore du Cambodge du In: In: M.S. van Meeuwen & al.， Prelimin 訂 y revisions Laos et du Vietnam. 27 Leguminosae (Fabaceae) ， of of some genera of Malaysian Papilionaceae 11. Papilionoideae ， Desmodieae. pp. 62-142. Museum Reinwardtia Reinwardtia 6: 93-102. National d'Histoire Naturelle Laboratoire du Meeuwen M. S. Kn aap-van 1962. Preliminary revi- Phanerogamie. P訂 is. sions sions of some genera of Malaysian Papilionaceae 一一一 1995. An enumeration of Chinese Desmodium V -A census of the genus Desmodium. Reinwardtia and its allied genera (Leguminosae). J. Jpn. Bo t. 6: 6: 239-276. 70: 111-117. 一一一 1963. Desmodium megaphyllum Zoll. ，a new re- 一一一 1997a. Three species of Dendrolobium in cord cord for Australia. In: C. G. G. J. van Steenis ， Malesia ， Australia and Fiji (Leguminosae- Misce Il aneous Botanical Notes XII. Blumea 12: Papilionoideae: Desmodieae). Taiwania 42: 135- 13-17. 13-17. 14 1. Me 町ill E. D. 1910. An enumeration of Philippine 一一一 1997b. Aphyllodium (DC.) Gagnep. instead of Leguminosae ， with keys to the genera and species. Dicerma DC. (Leguminosae-Papilionoideae: Philipp. Philipp. 1. Sci. ， C. 5: 1-136. Desmodieae). Taiwania 42: 142-148. June June 2004 Journal of Japanese Botany Vo l. 79 No. 3 183

一一一 1997c. Taxonomic status of Phyllodium elegans Ri dley H. N. 1922. Leguminosae. The flora of the (Lour.) (Lour.) Desv. v訂 . javanicum Schind l. (Legumi- Malay Peninsula. L. Reeve & Co. ，Lt d. ， London. nosae-Papilionoideae: nosae-Papilionoideae: Desmodieae). J. Jpn. Bo t. Rugayah 1988. Desmodium capitatum dan Desmodium 72: 72: 347-35 1. styracifolium. Fl oribunda 1(8): 29-32.

一一一 1998a. Dendrolobium (Leguminosae-Papilio ‘ Sanjappa M. 1992. Legumes of India. 338 pp. Bishen noideae: noideae: Desmodieae) in the Philippines. J. Jpn. Singh Mahendra Pal Singh. Dehra Dunn ，India. Bo t. 73: 248-258. (Desmodium pp.149-166). 一一一 1998b. The taxonomic position of Desmodium Schindler A. K. 1924a. Desmodium und Meibomia. bolsteri bolsteri (Leguminosae) ，a species endemic to the Repert. Spec. Nov. Regni Veg. 20: 13 6- 155. Philippines. Philippines. J. Jpn. Bo t. 73: 259-263. 一一一 1924b. uber einige k1 eine Gattungen aus der 一一一 1999. The genera ，tribes and subfamilies of Verwandschaft von Desmodium Desv. Repert. Japanese Japanese Le guminosae. Sc i. Rep. Tohoku Univ. Spec. Nov. Regni Veg. 20: 266-286. ser. ser. 4，Bio l. 40: 187-268. 一一一 1925. Desmodii generumque affinium species et 一一一 200 1. Leguminosae. In: Iwatsuki ， K.， Ohba H. combinationes novae. Repert. Spec. Nov. Regni and and Boufford D. E. (eds よFl ora of Japan IIb: 213- Veg. 21: 1-2 1. 279. 279. Kodansha Lt d. ， Tokyo. 一一一 1928. Die Desmodiinen in der botanischen 一一一 2003. A new circumscription of Tadehagi and a Literatur nach Linne. Repert. Spec. Nov. Regni new genus Akschindlium (Leguminosae). J. Jpn. Veg. Beih. 49: 1-37 1. Bo t. 78: 269-294. Schubert B. G. 1940. Desmodium: preliminary studies 一一一 in prep. Tribe Desmoideae. In: Lewis G. P. ， 1. Contr. Gray Herb. 129: 3-3 1. Schrire Schrire B. ， Mackinder B. and Lock M. (作 ed む叫.) s. 一一 1943. Desmodium. pp. 413 -4 39. In: Macbride J.

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一一一， Nemoto T. and Wu T. L. 1996. The taxonomic 3) subfamily Papilionoideae. In: Milre 国 Redhead E. position position of Trifidacanthus (Leguminosae). J. Jpn. and Polh i11 R. M. (eds よFl ora of Tropical East Bo t. 71: 57-66. Africa. Crown Agents for Oversea Govemments 一一一， Polhill R. M. and Schubert B. G. 198 1. and Administrations. Desmodieae. Desmodieae. In: Polhill R. M. and Raven P. H. 一一一 1980. 18. Desmodium. Fl ora of Panama ， part 5， (eds.) ， Advances in Legume Systematics. Part 1. Fam. 83 Leguminosae subfam. Papilionoideae. pp. pp. 292-300. Royal Botanic Gardens ，Kew. Ann. Missouri Bo t. Gard. 67(3): 622-662. 一一一， Ye B. and N emoto T. 1999. Dendrolobium Shakryl A. K. 1992. Leguminosae species from the ter- 。rbuscula (Domin) H. Ohashi ， D. umbellatum (L.) tiary of Abkhazia. In: Herendeen P.H. and Dilcher Benth. Benth. and a related new species (Leguminosae- D. L. (eds.) ， Advances in Legume Systematics. P制 Papilionoideae: Papilionoideae: Desmodieae). J. Jpn. Bo t. 74: 14- 4. The Fossil Record. pp. 189-206. Royal Botanic 24. 24. Gardens ，Kew. Pedley Pedley L. 1996. Fabaceae (Leguminosae) Subfamily Steenis C. G. G. J. van 1950. An Introduction. Fl ora Faboideae Faboideae (Papilionoideae) (continued). Tribe Malesiana Series 1. 1. Desmodieae. Desmodieae. pp. 149-198 (including 11. 一一一 1978. Misce l1 aneous Botanical Notes XXV. Pseudarthria Pseudarthria by V. E. Rudd). In: Dassanayake M. Blumea 24: 479 -4 84. D. D. and Clayton W. D. (eds.) ，A Revised Handbook 一一- 1982. 156. Reduction of Trifidacanthus Meη. to to to the Fl ora of Ceylon 10. A. A. Balkema ， Desmodium (Leguminosae). In: Steenis C. G. G. J. Rotterdam. Rotterdam. van and Veldkamp J. F. ，Misce l1 aneous Botanical 一一一 1999. Desmodium Desv. (Fabaceae) and related Notes XXV I. Reinwardtia 10: 21-26. genera genera in Australia: a taxonomic revision. Taubert P. 1894. Leguminosae. In: Engler A. and Austrobaileya Austrobaileya 5(2): 209-261. Prantl K.， Die natu r1 ichen Pflanzenfamilien. IU ， 3: Polhill Polhill R.島 1. 1981. Papilionoideae. In: Polhill R.乱 1. 70-396. Leipzig. and and Raven P. H. (eds.). Advances in Legume Turland N. J. and Jarvis C. E. 1997. Typification of Systematics. Systematics. Part 1. pp. 191-208. Royal Botanic some Linnaean specific and varietal names in the Gardens ，Kew. Leguminosae (Fabaceae). Taxon 46: 457 -4 85. Pramanik Pramanik A. and Thothathri K. 1986. On the status of Verdcourt B. 1979. A Manual of New Guinea Desmodiastrum Prain (Fabaceae). J. Indian Bo t. Legumes. Office of Forests ， Division of Botany ， Soc. Soc. 65: 373-379. Lae ， Papua New Guinea. 646 pp. (Desmodium pp. 184 184 植物研究雑誌第79 巻第3号 平成16 年6月

385 -4 13). 178-298. Parbury ， Allen & Co. ， London. Wheeler E. and Baas P. 1992. Fossil wood of the Yakovlev G. P. ， Sytin A. K. and Roskov Yu. R. 1996. Leguminosae: Leguminosae: A case study in xylem evolution and Legumes of N orthern Eurasia. A Checklis t. Royal ecological ecological anatomy. In: Herendeen P. H. and Botanic Gardens ，Kew. Dilcher Dilcher D. L. (eds よ Advances in Legume Yang Y. C. and Huang P. H. 1995. Desmodieae. In: Systematics. Systematics. Part 4. The Fossil Record. pp. 281- Lee S. K. (ed よ Fl. Reip. Popu l. Sin. 41. 301. 301. Royal Botanic Gardens ，Kew. Leguminosae (3) Papilionoideae 2. pp. 1-16 1. Wight R. and Arnott G. A. W. 1834. Leguminosae. Science Press ， Beijing (in Chinese). Prodromus Fl orae Peninsula lndiae Orientalis 1:

大橋広好: Malesia のマメ科シバハギ属と近縁属 の分類と分布 (11)

本研究では Malesia のシパハギ属とその近縁属 0 巧Iphyllum (D C.) H. Ohashi & R. R. Mill としては は13 属65 種よりなることを明らにした.前編(本 ハーバード大学標本館に所蔵されるジャワ島産ミ 誌79 巻 2 号)の Aphyllodium から Desmodium まで ソナオシ標本に貼り付けられている断片的な個体 に続いて，この後編では Hanslia から Trifidacan- がある.この標本はジャワで 1905 年に採集された thus までの 8 属14 種について分類学的な整理を行っ ものである.しかし，この標本によってジャワ島 た.また Malesia におけるシパハギ属と近縁属に にヌスピトハギがあるとするには疑問が残る.こ ついて分布を整理し，系統進化的な考察を試みた. のミソナオシ標本の重複品がロンドンの自然史博 Hanslia は Malesia に固有の属で，形態および花 物館にもあるが，それはミソナオシだけが貼り付 粉形質においてミソナオシ属に類似するが，柱頭 けられた標本である.また これまでジャワ島か が花柱の側面につく点で異なる • Hanslia hen かi は らヌスピトハギが記録されたことはなく，一方， 1977 年にニューギニアから 1 点だけの果期の標本 ミソナオシはジャワにふつうである.ハーバード に基づいて記載された種であった.その後やはり 大学標本は他の地域で採集されたヌスピトハギが ニューギニアからの標本が 2 点見付かったが，花 混入されたものである可能性がある.多分 も果実もない標本であった.今回の標本は Kew Malesia 地域にはヌスピトハギは分布しないと思 から未同定のまま Leiden に貸し出されていたも われる.現在のところヌスピトハギは台湾北部が ので，断片的な花と数個の花膏とをつけている 南限である.日本では沖縄本島嘉津宇岳が南限地 (Fig. (Fig. 5). これが 4 枚目の標本であり， Borneo か であり，沖縄における唯一の産地であるが，かつ らの初発見である.本種については完全な状態の ては山原に稀ではなかったときもあったようで， 花は未知であり，この種の正体はまだ分からない 開発などによって生育地が破壊されたという(沖 点がある. 縄県 1996 沖縄県の絶滅の恐れのある野生生物) . ヌスピトハギ属は Malesia に 4 種が分布する. アジア大陸では暖帯から亜寒帯に広く分布する. Hylodesmum repandum はこの属で唯一アフリカに Malsesia 地域内における 13 属の分布についてみ も分布する種であるが，形態的にはマルバヌスピ ると，固有属は Monarthrocarpus のみ， Malesia と トハギに類似し，しばしば誤ってそれと同定され アジア大陸に分布するのは Desmodiastrum ， ている. Verdcourt (1 979) によれば， New Guihea Hegnera ，ヌスピトハギ属 Hylodesmum ， ミソナオ のH. repandum は帰化であろうという.オオバヌ シ属，および Tr 折dacanthus の 5 属であり， スピトハギ Hylodesmum laxum (DC.) H. Ohashi & R. Malesia とオーストラリアに限られるのは Hanslia R. R. Mill とトキワヤブハギ H. leptopus (A. Gray ex のみである.残りの 6 属 (Aphyllodium ，マイハギ Benth.) Benth.) H. Ohashi & R. R. Mill は Malesia に広く分 属，ナハキハギ属， Desmodiastrum ， ウチワツナ 布し， New Guinea まで見られる.一方マルバヌ ギ属，タデハギ属)は Malesia ，アジア大陸，オー スビトハギH. podocarpum (D C.) H. Ohashi & R. R. ストラリアに分布する.なお マレー半島のほぼ Mill Mill subsp. podocarpum は Malesia ではルソン島に 南半分は，植物地理学上の Malesia に含まれるた だけ生育し (Fig. 8) ，ここが分布の南限である. めに，本論文でアジア大陸とよぶ地域からはここ マルパヌスビトハギは日本と中国にはふつうであ を除いている. るが，台湾中部南投県と嘉義県に稀であり，台湾 次に， Malesia に自生する 57 種を対象として種 北部と沖縄を飛んで九州、|やアジア大陸に分布する. の分布について調べてみた.ここではアメリカか Malesia のヌスピトハギ H. podoc αrpum subsp. らの帰化種である 8 種を除いて考察する. Malesia June June 2004 Journal of Japanese Botany Vo l. 79 No. 3 185

固有種， Malesia とアジア大陸に分布する種， た研究が発表されている.したがって標本も多く， Malesia とオーストラリアに分布する種，アジア シパハギ属と近縁属の自生種について他の地域よ 大陸・ Malesia' オーストラリアに分布する種をま りもよく調べることができた. 3 地域を比較して とめ，それぞれを Malesia ，Asia ，Australia ，Asia- みると，ジャワには 10 属31 種，ニューギニアには Australia Australia group と名付ける.種数をみると Malesia 9 属32 種，フィリピンには 9 属29 種が認められた group は 5 属 8 種(全体の 14 %)， Asia group は 9 (Table 2). この結果から，まず， 3 地域がほぼ同 属29 種(全体の 52 %)， Australia group は 4 属 11 数の種数をもつことが注目された. しかし， 3 地 種(全体の 20 %)， Asia-Australia group は 6 属 8 域の聞には面積，生育地環境の多様性，開発程度 種(全体の 14 %)であった (Table 1). 種数の比 の多少などの条件に明らかな差のあることから推 較からみると， Malesia におけるシパハギ属と近 定するに，ニューギニアとフィリピンにはジャワ 縁属はアジア大陸の種との関連が顕著である. よりも多数の種が生育すると予想される.この結 Asia Asia group の種の中では Westem Malesia floristic 果はニューギニアとフィリピンの調査がジャワの region region (マレ一半島南部，スマトラ，ボルネオ， それに比べて不足していることに起因するためで ジャワ，パラワン島よりなる)にのみ分布する種 あろうと考えられる. が11 種あり，アジア大陸との結ぴつきを強く示す. 次に， 3 地域に共通する種および 2 地域間ごと さらに Asia-Australia group の 8 手重の中でナハキハ の共通種と非共通種とをまとめてみた (Table 2). ギ Dendrolobium umbellatum を除く他の 7 種もア ジャワにはインドシナあるいはインドとの共通種 ジア大陸との結びつきを示す種であると思われる. がみられ，オーストラリアとの共通種はない.ニュー なお， Malesia に分布するヌスピトハギ属 4 種は ギニアは Malesia 固有種とオーストラリアとの共 全て Asia group に属する. 通種とが特色であり フィリピンには Malesia 固 一方， Malesia とオーストラリアとの共通種の 有種と中国との共通種がみられることがわかる. 中にはシバハギ属 8 種が含まれるが，これらは高 最後に Malesia においてシパハギ属と近縁属の 度に分化したと考えられる種である.また，固有 種類と分布型の多様性とがどのように生まれてき 種の多くは狭い分布域をもっ . Dendrolobium cumin- たのかについて考察した.シバハギ属と近縁属は gianum ， Dendrolobium geesinkii ，Desmodium bolsteri インドから中国に至るアジア大陸で属レベルの分 はルソン島，Dendrolobium quinquepetalum はルソ 化が見られ，ここで第一次的に分化したと考えら ン島と稀にミンドロ島 ， Aphyllodium novo guineense ， れている (Ohashi et al. 1981). Desmodiinae 亜連

Dendrolobium papuacola はニューギニア ， Hanslia に属するシバハギ属， Ougeinia ， Cod α， riocalyx ， hentyi はニューギニアと新しくボルネオ島で発見 Phyllodium の中新世からの化石はインドと中国か された.ただ 1 種 Monarthrocarpus securiforme は ら発見されている. Malesia においてシパハギ属 ルソン，イサベラ，モルッカ，ニューギニアに知 と近縁属の化石は記録されていないが， Asia られており，広い分布域をもっている. 8 種のう group とAsia-Australia group の種は基本的にアジ ち 5 干重カ宝ルソン島で 4 手重カ宝ニューギニアでみら ア大陸の種に由来して今日 Malesia に分布するに れる.亜種としては Desmodium nemorosum subsp. 至った種であると考えられる.さらに Malesia を whi ゆrdii がルソン島に ， D. strigillosum subsp. 経てオーストラリアに分布を広げたと予測される. celebicum がスラウェシに固有である.雑種 この過程で地方的な種分化がおこり， Malesia 固 Tadehagi Tadehagi triquetrum nothosubsp. philippinense はフィ 有種である Malesia group の種が生まれ，オース リピン，パラワン，ニューギニアでみられる.固 トラリアでも種レベルでの分化がおこり，今日 有種の多くはアジア大陸の種に形態的に最も近く， Malesia にみられる Australia group の手重はオース また固有種がルソン島とニューギニアに集中する トラリアから Malesia に侵入したと想像される. ことは Malesia におけるシパハギ属と近縁属の進 Malesia におけるシパハギ属と近縁属の多様性は 化に関して問題を解く鍵となる現象であると思わ 歴史を経て分布を広げてきた進化の結果であると れる. 考えられる. Malesia の中ではジャワ，ニューギニア，フィ (東北大学大学院理学研究科 リピンがマメ科について他の地域よりもまとまっ 附属植物園津田記念館)