植物研究雑誌 J. J. Jpn. Bo t. 79: 79: 101-139(2004) Taxonomy and Distribution of Desmodium and Related Genera (Leguminosae) (Leguminosae) in Malesia (1) Hiroyoshi Hiroyoshi OHASHI Botanical Botanical Garden ，Graduate School of Science ，Tohoku University ，Sendai ， 980-0862 JAPAN (Received (Received on November 8，2003) A taxonomic and phytogeographic account of Desmodium and its related genera in Malesia Malesia is presented as a precursory of treatment these genera for the Fl ora Malesiana. Sixty-four Sixty-four species in 13 genera 訂 'e recognized: Aphyllodium (3 spp よ Codariocalyx (3 spp よDendrolobium (8 spp よDesmodiastrum (1 sp よDesmodium (35 spp. of which 27 native native to Malesia and 8 introduced from America) ， Hanslia (2 spp よHegnera (l sp よ Hylodesmum (4 spp よ Monarthrocarpus (1 sp よ Ohwia (1 sp よ Phyllodium (3 spp よ Tadehagi Tadehagi (l sp.) ， and Trifidacanthus (1 sp.). They 訂 e enumerated with keys to genera ， species species and infraspecific taxa ，selected synonyms and bibliography related to the Malesian Malesian flora ，representative specimens ， and distribution. Of the 56 species native to Malesia Malesia eight (14 %)訂 e endemic to the region; 29 (52 %)訂 e distributed in Malesia and continental continental Asia but not in Australia; 11 (20 %) are common in Malesia and Australia but not not in continental Asia; and eight (14 %) are found in all these areas. Based on the dis- tribution tribution analysis ，a differentiation pattem of the species in these genera in Malesia is presumed. presumed. This This paper is divided into two parts. Aphyllodium ，Codariocalyx ，Dendrolobium ， Desmodiastrum ， and Desmodium are treated in the first p訂t. Following new combina- tions tions or new status are proposed: Codarioc α lyx microphyllus (Thunb.) H. Ohashi ， Desmodiastrum parviflorum (Dalz.) H. Ohashi ，Desmodium nemorosum F. Muell. ex Benth. Benth. subsp. whitfordii (Schind l.) H. Ohashi ， and Desmodium strigillosum Schind l. subsp. subsp. celebicum (Schind l.) H. Ohashi. Key words: Codariocalyx microphyllus ，Desmodium incanum ，Desmodium intortum ， Malesia ， taxonomy. Asian Desmodium and its related genera Huang 1995) ，Taiwan (Huang and Ohashi have been well studied in recent ye 紅 s. 1993) ，Japan (Ohashi 200 1)， and Siberia to Comprehensive lists or floras of the species Russia Far East (Yakovlev et al. 1996). in in these genera have been compiled for West Malesia is the largest 紅 'ea that has not been Asia (Lock and Simpson 1991) ， Pakistan (as treated since Meeuwen (1 961 ， 1962) and West Pakistan; Ali 1977) ， India (Sa 吋appa Ohashi (1971 ， 1973). It needs a recent revi- 1992) ， Nepal (Ohashi 1979) ， Bhutan slO n. (Grierson (Grierson and Long 1987) ， Sri Lanka Malesia was proposed by Zollinger in (Pedley (Pedley 1996) ，South Asia (Kumar and Sane 1857 as a floristic region (Johns 1995) ，and 2003) ，Indo-China (Dy Phon et al. 1994) ， is commonly known as the area for the Flora Indo-China including Thailand (Lock and Malesiana (Steenis 1950). It extends east- Heald 1994) ，China (Ohashi 1995 ，Yang and ward from Sumatera (Sumatra) ， through the -101 ー 102 102 植物研究雑誌第79 巻第2号 平成 16 年4月 Malay Peninsula ， Bomeo ， Sulawesi removing subtribe Bryinae from the previous (Celebes) ，Maluku (Moluccas) ，New Guinea Desmodieae to the aeschynomenoid group as to to the Bismarck Islands. To the north it is de- indicated by the analysis of the chloroplast limited limited by Luzon and to the south by gene rbcL (Bailey et al. 1997 ， Doyle et al. Christmas Christmas Island. This demarcation of 2000). 恥1alesia is adopted here ， although a recent However ，“ in practice ， the main problem redefinition redefinition of Malesia (Brummitt 2001) di- has been the circumscription of Desmodium vides vides it into Malesia in the na 町 ow sense and (Ohashi et al. 1981)" for the system of tribe Papuasia Papuasia by excluding New Guinea and the Desmodieae ， because “there are overall simi- Bismarck Bismarck Ar chipelago. larity among the Desmodium and its allied This This paper aims to enumerate all the genera (Verdcourt 1979)". Both narrower known species and infraspecific taxa of and broader circumscriptions of Desmodium Desmodium and related genera in Malesia as have so far been proposed. Bentham (1 852 ， a precursor to revisions of these genera for 1865b) faced the problem of circumscription Flora Flora Malesiana ， and to analyze distribution of Desmodium. In 1852 he published his first pattems pattems of the native Malesian taxa ， because generic system on Desmodium and its allied no such works on distribution have so far genera. His Desmodium was narrower than been been done for these genera. that of Candolle (1825b ， 1826) and Wight The paper is divided into two parts. and Arn ott (1834). He recognized six satel- Circumscription Circumscription of Desmodium and its rela- lite genera of Desmodium ，i. e. ， Catenaria tives ，a taxonomic history of these genera in Benth. ， Dendrolobiw η(Wight & Arn.) Malesia Malesia and an enumeration including Benth. ， Dicerma DC. ， Ougeinia Benth. ， Aphyllodium ，Codariocalyx ，Dendrolobium ， Phyllodium Desv. and Pteroloma Desv. ， Desmodiastrum ， and Desmodium are treated based on species previously included in in in this par t. Other genera (Hanslia ，Hegnera ， Desmodium ， although he included Nicolso ・ Hylodesmum ， Monarthrocarpus ， Ohwia ， nia recognized by Candolle (1825b ) in Phyllodium ，Tadehagi ， and Trifidacanthus) ， Desmodium. This concept was 1ater strength- phytogeographic phytogeographic considerations and refer- ened by Schindler (1 924a ，1924b ，1928). He ences ences will be presented in the next part. recognized 15 genera as closely related to Desmodium. Schindler' s core Desmodium Circumscription Circumscription of Desmodium (1 928) is almost confined to the Old W orld and related genera species excluding almost all the New World Desmodium and related genera belong to species as Meibomia. This is the na 町 owest the the tribe Desmodieae in the subfamily circumscription of Desmodium ever pro- Papilionoideae. Papilionoideae. Desmodieae 紅 e considered posed. to to be one of the most highly advanced tribes In 1865 ，Bentham (1865b) adopted a new of of the subfamily Papilionoideae (Polhill concept of Desmodium changing his previ- 1981 ， Ohashi et al. 1981). “ There are high ous system of Desmodium and its allied gen- levels levels of specialisation and diversity with era created in 1852. With the exception of good correlation in a notable range of fea- Ougeini αhe united the five remaining genera tures tures to give a rather precise stepwise pro- into Desmodium ，recognizing them as sec- gression gression through series ，section ，subgenus ， tions within Desmodium. Pedley (1 999) ex- genus genus and generic group" (Ohashi et al. plained why Bentham (1 865a) changed his 1981). 1981). The tribe is divided into two concept of Desmodium in ‘ Genera Plantarum' . subtribes ，i. e. ， Desmodiinae and Lespedez- This is the broadest circumscription of inae inae (Ohashi in Lewis et al. in prep.) ，by Desmodium and this concept was followed Apri12004 Joumal of Japanese Botany Vo l. 79 No. 2 103 by Baker (1 876) ， Taubert (1 894) ， Merrill are here separated as distinct genera (Ohashi (1910 ，1923) ， Schubert (1940 ，1971 ，1980) ， in Lewis et a1. in prep ふ Meeuwen (1 962) ， Fosberg (1 966) ， Verdcourt In the present paper ，Desmodium and re- (1979) ， Sanjappa (1 992) ， and others. lated genera in Malesia 紅 e grouped as com 回 Naturally ，Desmodium in the broad sense posing of the following 13 genera ，i. e. ， contains contains variously diverse species ， and the Aphyllodium ，Codariocalyx ，Dendrolobium ， boundary boundary between Desmodium in the Desmodi αstrum ， Desmodium ， Hanslia ， broader broader sense and other genera of Desmodi- Hegnera ， Hylodesmum ，Monarthrocarpus ， inae inae becomes obscure. Ohwia ， Phyllodium ， Tadehagi and Trifi- A concept intermediate between the dacanthus. Excepting Desmodiastrum and broadest broadest Desmodium of Bentham (1865b) Tr がdacanthus ， the other 11 genera were and and the narrowest circumscription of treated as belonging to the members of Schindler Schindler (1 928) was proposed by Ohashi Desmodium and related genera in Ohashi (1973) (1973) and was used in Ohashi et a1. (1 981) ， (1 973). Dy Phon et a1. (1 994) ， Ohashi (1 995) ，Yang and and Huang (1995) and Pedley (1 996 ，1999). History of taxonomic studies on However ， in this system “it has the disadvan- Desmodium and related genera tages tages that a small number of species are 住an- in Malesia sitional sitional to some degree ， and that the The first important contribution to the tax- difference difference between some subgenera of onomy of Desmodium and re1ated genera in Desmodium and the segregates are m 紅 ginal Ma1esia was made by Bentham (1 852). He so so that stability for the classification is not classified 21 species from Jawa and assured" assured" (Ohashi et a1. 1981 on page 293). Sumatera in the Junghuhn herbarium as fo1- All All 19 genera in the subtribe Desmodiinae lows: Desmodium with 15 species ，Dendro- recognized recognized in Ohashi et a1. (1 981) form a lobium with two species ， and one species monophy letic group marked by the loss of each in Catenaria ，Dicerma ， Phyllodium and the the chloroplast rpl2 intron (Bailey et a1. Pteroloma. Merrill (1 923) adopting the 1997). 1997). However ， previous concepts for broadest concept of Desmodium (= Desmodium proposed by Bentham (1 852 ， Desmodium s. 1.) proposed by Bentham 1965b) ， Ohashi (1 973) ， Ohashi et a1. (1 981) (1865b) recorded 31 species from the are are considered to be p訂 aphyletic suggested Phi1ippines in which 28 are indigenous and by results of analyses of the chloroplast gene three are alien introductions from America ， rbcL (Kajita et a1. 1996 and in prep.). Also ， but excluded Desmodium securiforme Benth. Meibomia proposed by Schind1er (1 924a) is as a separate genus Monarthrocarpus and monophyletic monophyletic with Desmodium. The broad- described a new genus Trifidacanthus based est est Desmodium is morphologically indistin 輔 on T. unifoliolatus Merr. Backer and guishable guishable from the close genera Alysicarpus ， Bakhuizen van den Brink Jr.