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Investigation of Sectional Relationships in the Genus Rhododendron (Ericaceae) Sased on Matk Sequences

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Investigation of Sectional Relationships in the Genus Rhododendron (Ericaceae) Sased on Matk Sequences 植物研究雑誌 J. J. Jpn. Bo t. 73: 73: 143-154(1998) Investigation Investigation of Sectional Relationships in the Genus Rhododendron (Ericaceae) sased on matK Sequences Y 吋iKURASHIGE a，Masaki MINE bペNobuo KOBA Y ASHl bぺ Takashi HANDA b，Kenji TAKAYANAGl b and Tomohisa YUKAWA c aLaboratory aLaboratory of Botany ，Ak agi Nature Park ， Yuhikami Yuhikami 892 ，Minami- Ak agisan ，Ak agi-mura ，Seta-gun ，Gunma ， 379-1113 JAPAN; blnstitute blnstitute of Agriculture and Forestry ，University of Tsukuba ， Tennodai Tennodai 1-1-1 ，Tsukuba ，Ibaraki ， 305-0005 JAPAN; CTsukuba CTsukuba Botanical Garden ，National Science Museum ，Amakubo 4-1-1 ，Tsukuba ，Ibaraki ， 305-0005 JAPAN; dl wate Biotechnology Research Center ，Narita 22-174-4 ，Ki takami ，Iwate ， 024-0003 JAPAN; 巴Tatebayashi Azalea Research Station ，Hanayama 3258 ，Tatebayashi ，Gunma ， 374-0005 JAPAN (Received (Received on October 20 ， 1997) Owing to the enormous number of species and conflicting ideas of classification proposed proposed by many taxonomists ， the genus Rhododendron poses systematic problems at various various infrageneric levels. Sequence data derived from matK and trnK introns were used to to examine relationships among all 8 subgenera and 12 sections of Rhododendron ，and additional additional genera Ledum ，Menziesia ， and all Elliottia. The major results from this study are are as follows: (1) Menziesia and Ledum are nested within the genus Rhododendron. (2) Thus Rhododendron is paraphyletic. (3) R. camtschaticum forms a basallineage of tribe Rhodoreae. Rhodoreae. (4) Subgenus Tsutsusi is monophyletic; however ， R. tashiroi makes both sections sections Tsutsusi and Brachycalyx para-/polyphyletic. (5) R. tsusiophyllum is a member ofthe ofthe subgenus Tsutsusi section Tsutsusi. (6) Subgenus Rhododendron is monophyletic. (7) (7) Subgenus Pentanthera is polyphyletic. (8) Subgenus Pentanthera section Sciadorhodion is is polyphyletic. (9) Subgenus Azaleastrum is polyphyletic. In In the mountain flora of East to Southeast nus in various ways. On the basis of these Asia ， the genus Rhododendron L. (Ericaceae) earlier works ，Sleumer (1949 ， 1980) proposed holds holds the essential position. The diversity of a framework system of Rhododendron with a this this group is represented by the immense main emphasis upon relative positions of flower number of species (e. g. over 1 ，000: Chamber- and leafbuds ，characteristics oflepidote scales lain lain et al. 1996) as well as variation in their on the abaxial surface of leaves ，and persist- habita t. Consequently ， this group presents in- ence or deciduousness of leaves as primary tricate tricate taxonomic problems ， in particularthose diagnostic characters. He recognized the fol- on infrageneric circumscr ヤtion. lowing eight subgenera: Azaleastrum ， Since Linnaeus (1 753) established Rhodo- Hymenanthes ，Pentanthera ，Pseudazalea ， dendron ，taxonomists such as Don (1 834) ， Pseudorhodorastrum ， Rhododendron ， Planchon (1854) ，Maximowicz (1870) ，Wilson Rhodorastrum ，and Tsutsusi (Table 1). and Rehder (1 921) ，and Copeland (1 943) have W orkers of the Royal Botanic Garden ，Ed- attempted over the years to subdivide the ge- inburgh ，subsequently started to revise 一一 143 一一 144 144 植物研究雑誌第73 巻第3号 平成10 年6月 Table 1. Comparison oftwo subdivisions ofthe genus Rhododendron Sleumer (1980) Chamberlain et al. (1 996) Genus Rhododendron Genus Rhododendron Subgen.Rhododendron Sec t. Pogonanthum Sec t. Pogonanthum Subgen. Rhododendron Sec t. Rhododendron Sec t. Rhododendron Sec t. Vireya Sec t. Vireya Subgen. Subgen. Pseudazalea Sec t. Pseudazalea Subgen. Subgen. Pseudorhodorastrum Sec t. Rhabdorhodion Sec t. Rhodobotrys Sec t. Trachyrhodion Subgen. Subgen. Rhodorastrum Sec t. Rhodorastrum Subgen. Subgen. Hymenanthes Sec t. Hymenanthes Sec t. Ponticum Subgen. Hymenanthes Subgen. Subgen. Tsutsusi Sec t. Brachycalyx Sec t. Brachycalyx Subgen. Tsutsusi Sec t. Tsusiopsis Sec t. Tsutsusi Sec t. Tsutsusi Subgen. Subgen. Pentanthera Sec t. Pentanthera Sec t. Pentanthera Subgen. Pentanthera Sec t. Rhodora Sec t. Rhodora Sec t. Sciadorhodion Sec t. Viscidula Sec t. Viscidula Subgen. Subgen. Azaleastrum Sec t. Azaleastrum Sec t. Azaleastrum Subgen. Azaleastrum Sec t. Choniastrum Sec t. Choniastrum Sec t. Mumeazalea Subgen. Mumeazalea Sec t. Candidastrum Subgen. Candidastrum Genus Therorhodio l1 Subgen. Therorhodion Genus TSltsiophylllll l1 Sleumer' Sleumer' s system. Cullen (1980) stressed the (1990) united the monotypic section Tsusiopsis taxonomic taxonomic importance of lepidote scales; he sensu Sleumer with section Tsutsusi ，and they thus thus united all the lepidote (scaly) rhododen- also reduced the monotypic genus drons drons into a single subgenus Rhododendron ， Tsusiophyllum to section Tsutsusi. Judd and while while Sleumer (1 980) scattered them among Kron (1995) moved parts of sections four four subgenera: Rhododendron ，Pseudazalea ， Brachycalyx (of subgenus Tsutsusi) and Pseudorhodorastrum ，and Rhodorastrum. Rhodora (of subgenus Pentanthera) to section Philipson Philipson and Philipson (1986) assigned sec- Sciadorhodion (of subgenus Pentanthera). tions tions Mumeazalea and Candidastrum (of Chamberlain et al. (1996) compiled these re- subgenus subgenus Azaleastrum sensu Sleumer) to a sults and recognized the following eight subgeneric subgeneric rank ，and they reduced genus subgenera: Azaleastrum ，Candidastrum ， Therorhodion Therorhodion to a subgeneric rank of the ge- Hymenanthes ，Mumeazalea ，Pentanthera ，Rho- nus nus Rhododendron. Chamberlain and Rae dodendron ，Therorhodion ，and Tsutsusi (Ta- June June 1998 Journal of Japanese Botany Vo l. 73 No. 3 145 ble ble 1). We will use this system as the reference ， recognized by Chamberlain et al. (1 996) were because because (1) it covers the whole genus ，and (2) sampled; since tribe Rhodoreae (of subfamily it it took account of the results from recent stud- Rhododendroideae) comprises three genera ， les. les. Rhododendron ，Ledum ，and Menziesia ，we There are many studies that intended to chose one species each from the latter two clarify clarify the phylogenetic relationships within genera. A single species of Elliottia (sub- Rhododendron ，e.g. ， in terms of secondary family Rhododendroideae tribe Clado- substances substances (Harborne 1980 ，Harborne and thamneae) was also included. A single species Williams Williams 1971 ，King 1980) ，interspecific cross of Cassiope (subfamily Vaccioideae tribe compatibility compatibility (Williams et al. 1985 ，Yamaguchi Cassiopeae) was selected as the outgroup based et et al. 1985) ， ontogeny (Palser et al. 1985 ， on the results of an analysis of rbcL (Kr on and Philipson Philipson 1980) ，and micromorphology (Seith Chase 1993) and 18s rDNA (Kr on 1996) se- 1980). 1980). However ， these results have been of quences. Table 2 shows the materials exam- limited limited value in determining the phylogeny of ined and the sources where the plants were Rhododendron. Rhododendron. Therefore ，it is clear that we grown. All the voucher specimens are depos- require require another approach to resolve ited in TNS. m 仕ageneric systematic problems of Rhodo- Total DNA was extracted from fresh tissue dendron. dendron. following the methods of Kobayashi et al. Comparison of DNA sequences of matK ， (1995). Sequences were determined with PCR- the the maturase-encoding gene located in an intron amplified the matK gene and its flanking trnK of of the chloroplast gene trnK ， the former of introns from a total DNA extract by use ofthe which has evolved approximately three times primers shown in Fig. 1 and Table 3. Single- faster faster than the chloroplast gene rbcL (Johnson stranded DNA for dideoxy sequencing was and So It is 1994) ， has proven to be a powerful produced in a second round of amplification tool tool for phy logenetic reconstruction within using the double-stranded product as a tem- angiosperm families and genera (Johnson and plate. Both the forward and reverse strands Soltis Soltis 1994 ，Steele and Vilgalys 1994 ，Soltis et were sequenced for all taxa. l. al. 1996). In this study ，we compared matK and All parsimony analyses were conducted trnK introns sequences to investigate with PAUP ，Phylogenetic Analysis U Par- sing subgeneric subgeneric and sectional relationships in Rho- simony ， Version 3.1 (Swofford 1993). The dodendron. dodendron. heuristic search option with 100 random repli- cates cates (Maddison 1991) was used to perform Materials Materials and Methods Fitch parsimony analyses (Fitch 1971). Branch Twenty-two species representing all eight lengths for trees were calculated by ACCTRAN subgenera subgenera and 12 sections of Rhododendron optimization (Swofford and Maddison 1987). trnK trnK 3914F trnK 1MF matK 462F matK 3MF ーー... -.. ーー ... ーー ... m o. tK matK 174R m o. tK1MR matK 1848R trnK 2R 司咽ー- ，.._ -時一 4 ‘一ー Fig. Fig. 1. Relative position of the PCR amplification and sequencing primers used for matK and trnK introns. introns. Arrows indicate the direction of strand synthesis. Boxed areas represent coding regions. 146 146 植物研究雑誌第73 巻第3号 平成 10 年6月 Table Table 2. Species of Rhododendron and its related genera used for matK sequencing. Subdivision of Rhododendron is is based on Chamberlain et al. (1 996). Classification of all other taxa is based on Stevens (1 971) GenBank Species Species Subgenus Section Voucher access lO n Subfamily Subfamily Rhododendroideae Tribe Tribe Rhodoreae Rhododendron ovatum (Lind l.) Maxim Aza1eastrum Aza1eastrum Wi1son 1391 AB012729 Rhododendron stamineum Franch. Aza1eastrum Choniastrum Akagi N ature Park 94/0084 AB012730 Rhododendron albiflorum
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