Page 1 植物研究雜誌 J. Jpn. Bot. 76: 339-343 (2001) Delineation Of
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植物研究雑誌 J. J. Jpn. Bo t. 76: 76: 339-343 (200 1) Delineation Delineation of Schoenoplectus sect. Malacogeton (Cyperaceae) , New Combination ,and Distinctions of Species a b S. S. Galen SMITH and Eisuke HA YASAKA aDepartment aDepartment of Biology ,University of Wisconsin-Whitewater , Whitewater ,Wisconsin , 53190 U.S. A.; bBotanical bBotanical Garden ,Graduate School of Science ,Tohoku University ,Sendai , 980-0862 JAPAN (Received (Received on 乱1ay 14 , 2001) The new combination ,Schoenoplectus sec t. Malacogeton (Ohwi) S. G. Smith & Hayasaka , is made and a description of the section is provided. Schoenoplectus nipponicus ,Sc h. etuberculatus ,Sch. torreyi and Sch. subterminalis 訂 e included in sec t. Malacogeton , and a key to these is given. We recognize the sections Schoenoplectus , Actaeogeton , Malacogeton and Supini in the genus Schoenoplectus , and a key to the sec- tions tions and a list of North American and East Asian species included in each section are provided. provided. Key words: Cyperaceae , new combination , Schoenoplectus sec t. Malacogeton , taxonomy taxonomy The polymorphic and polyphyletic Scirpus 2000 , Koyama et al. 2000 , Smith and L. L. s. la t. in recent ye 紅 s has been subdivided Yatskievych 1996 ,Wilson 1981). Of these into into about 17 genera (Bruhl 1995 , segregate genera Schoenoplectus is comp 紅 a- Goetghebeur 1998 , Wilson 1981 , 1989). tively large , with species distributed world- Scirpus Scirpus as stric t1 y defined includes about 30 wide , the number of which is v紅 iously species species occurring mainly in the temperate re- estimated at 50 spp. (Goetghebeur 1998) ,60 gions gions of the Northem Hemisphere , with S. spp. (Wilson 1981) ,80 spp. (Hooper 1985) , sylvaticus sylvaticus L. as the type species (Wilson or over 130 spp. (Koyama et al. 2000). For 1989). 1989). The rest of Scirpus s. la t. has been North America and East Asia ,34 species are placed placed in Actinoscirpus (Ohwi) R. W. Haines here recognized (Appendix 11). Following & Lye , Amphiscirpus Oteng ・.Yeboah , Goetghebeur and Simpson (1 991) ,we here Bolboschoenus (Asch.) Palla , Blysmopsis define Schoenoplectus as excluding Oteng- Yeboah ,Blysmus Schul t., Eleogiton Bolboschoenus and the monotypic Link , Ficinia Schrad. , Isolepis R.Br. , Actinoscirpus ,which are sometimes included O 砂 ca ηum Nees ,Phylloscirpus C.B.Clarke , in it (Lye 1995 , Strong 1994). Pseudoschoenus (C.B.Clarke) Oteng- Schoenoplectus as currently delimited is Yeboah , Schoenoplectus (Rchb.) Palla , commonly characterized by its pseudolateral Scirpoides Scirpoides Seg. , Sumatroscirpus Oteng- inflorescences ,i. e. , the proximal bract usu- Yeboah ,Trichophorum Pers. ,and Websteria ally erect and resembling the culm (Wilson S.H.Wright S.H.Wright by recent cyperologists (Bruhl 1981). Close examination shows that this is 1995 , Goetghebeur and Simpson 1991 , a superficial character ,however , because the Goetghebeur 1998 ,Hayasaka and Ohashi proximal involucI 叫 bracts of many species -339- 340 植物研究雑誌第76 巻第6号 平成13 年12 月 now included in Schoenoplectus usually characters such as long leaf blades with un- closely closely resemble the foliage leaf blades and usual architecture , weak rhizomes often 訂 e commonly divergen t. Thus other charac- terminated by tubers , and often aquatic habi- ters ters are now used ,especially embryo form tat and flaccid habi t. Close examination (van (van der Veken 1965 , Wilson 1981) ,stolon shows that these four species all belong to presence , tuber presence and form ,ligule Scirpus sec t.乱 1alacogeton Ohwi [= presence ,culm branching ,leaf position , leaf Schoenoplectus subg. 孔1alacogeton (Ohwi) structure ,spikelet a町 angement ,floral scale Oteng-Yeboah]. The required new combina- indument , and culm anatomy. tion is provided as follows. Species Species relationships within Schoeno ・ plectus 紅 e not clear because of large mor- Schoenoplectus (Rchb.) Palla in Verh. K. phological phological variation in the characters of the K. Zoo l. -Bo t. Ges. Wien 38 (Sitzungsber.): rhizomes ,leaves ,inflorescences ,achenes , 49 (1888) , nom. cons. floral floral bristles and other structures that have Sec t. Malacogeton (Ohwi) S.G.Smith & been been used to define the infrageneric taxa. To Hayasaka , comb. nov. clarify clarify the species relationships within the Scirpus L. subgen. Schoenoplectus Rchb. genus ,a modern infrageneric system should sec t. Malacogeton Ohwi in Mem. Coll. Sci. be be established. This was attempted by Kyoto Imp. Univ. se r. B , 18: 97 (1944). Oteng-Yeboah Oteng-Yeboah (1974) and Raynal (1976b) , Scirpus sec t. Bolboschoenus Asch. ser. but but neither of their treatments 征 e satisfac- Malacogeton (Ohwi) T. Koyama in J. Fac. tory tory (Wilson 1981). In his brief paper , Sci. Univ. Tokyo ,sec t. 3 ,7: 288 (1 958). Oteng-Yeboah Oteng-Yeboah (1 974) recognized and pro 四 Schoenoplectus subgen. Malacogeton vided vided a key to the three subgenera (Ohwi) Oteng-Yeboah in Notes Roy. Bo t. Schoenoplectus , Actaeogeton and Malacoge- Gard. Edinb. 33: 315 (1 974). ton , but he neither gave descriptions for the Perennials , erect or flaccid , 40-200 cm subgenera subgenera nor the included species except high. Rhizomes long ,weak , very soft , often for for the type species. Raynal (1 976a ,b , 1977) terminated by fleshy tubers to ca. 3 mm recognized recognized the four sections Schoenoplectus , thic k. Culms trigonous or terete , ca. 0.5-5 Actaeogeton ,Supini , and Pterolepis; with mm thic k. Leaves basal , ca. 3 to more than emphasis emphasis on sec t. Supini , in which he recog- 20; sheaths often free from culm and resem- nized nized 21 species , mainly from Africa. His bling blades; sheath fronts hyaline ,splitting detailed detailed study of Sch. sec t. Supini should be early ,usually pinnate- fibrillose; blades 2 to extended extended to the rest of the species to provide ca. 20 per culm ,some species forming emer- a more complete classification for Schoeno- gent ,erect blades an d/ or submerged ,flaccid plectus. plectus. The present paper , which empha- blades ,cross-section mostly trigonous or lat 同 sizes sizes East Asian and North American erally flattened , apices often blunt and ec- species ,contributes to the establishment of a centric; distal bl 争de longer than sheath , ca. Ill odern in 仕ageneric system for the genus. It 0.2-10 mm wide. Inflorescences of one also also provides the name Schoenoplectus sec t. spikelet or capitate or 1x ーto 3x ー branched; 乱1alacogeton and other information needed proximal bract erect ,structure like foliage for for the senior author' s 位eatment of the genus leaf blades ,ca. 1-32 cm long. Spikelets 1 to in in an upcoming volume of the Flora of N orth 20 per inflorescence , 5-25 x 3-6 mm. Floral America America (Flora of North America Editorial scales narrowly oblong-ovate , 4-7 mm long , Committee Committee 2001). smooth or awn sparsely spinulose ,some- In In Schoenoplectus three species in eastern times sp 訂 sely ciliolate distally ,flanks each N orth America and one in East Asia share usually with 2 to ca. 10 prominent veins (at December 2001 Journa1 of Japanese Botany Vo 1. 76 No. 6 341 least least in proximal part of spikelet) , apices ing achene; eastern North American subacute ,entire or minutely notched ,awn coastal plains …・・ - (extension (extension of midrib) ca. 0.1-0.5 mm. …...・ H ・.Schoenoplectus etuberculatus Perianth Perianth bristles stout , from ca. 1/2 of 同 to 3. Leaf blades ca. 1-2 mm wide; floral twice twice achene length ,retrorsely to antrorsely scales awned , the apical midrib exten- spinulose. spinulose. Anthers ca. 2-3.5 mm long. Styles sions 0.2 -0 .5 mm; inflorescence of 1 to 4 3-fid 3-fid or 2-fid. Achenes yellow- to dark spikelets; not forming submerged mead- brown when ripe , ovoid to obovoid ,promi- ows; widespread in eastern North nently nently beaked , compressed-trigonous or America ...・ H ・...・ H ・.Schoenoplectus torreyi biconvex ,2 -4 .5 mm , smooth. Flowers ab- 3. Leaf blades 1 mm or less wide; floral sent sent from basal sheaths. Fresh-water lakes , scales mucronate , the apical midrib ex- ponds ,streams ,ditches , often deeply emer- tensions to 0.1 mm; inflorescence of one gent gent or submerged aquatic , in deep water erect spikelet; forming submerged mead- flaccid flaccid and sometimes entirely vegetative. ows of vegetative plants in deep or flow- Chromosome numbers reported: 2n = 42 , ing water; widespread in N orth America , 70 ,74 , 76. mostly eastern... …・ Type species: Schoenoplectus n伊'/)onicus …. .Schoenoplectus subterminalis (Makino) (Makino) Sojak. The type species is Sch. nipponicus , not Koyama (1 958 , under Scirpus) confused Sch. Sch. etuberculatus as stated by Oteng- the species of Sch. sec t. Malacogeton with Yeboah (1 974) , because Ohwi (1 944) , in his those of Bolboschoenus , which is now key key to species of Scirpus subg. treated as a distinct genus even when Scirpus Schoenoplectus , established Scirpus sec t. is treated in a broader sense including Malacogeton , and included only Scirpus Schoenoplectus (Kozhevnikov 1988). Oteng- nipponicus. nipponicus. Koyama (1 958) reduced Scirpus Yeboah (1 974) reg 紅 ded Sc h. sec t. nipponicus nipponicus to S. etuberculatus subsp. Malacogeton as intermediate between n伊ponicus , and Oteng-Yeboah (1 974) ap- Schoenoplectus and Bolboschoenus but parently parently adopted Koyama's treatment of S. closer to the former in having a 'net-like nipponicus nipponicus and gave Sch. etuberculatus as ground tissue' and other characters. the the type species of Sch. subg. Malacogeton. Schoenoplectus sec t. Malacogeton is here We think the diagnostic characters of Sch. considered to be distinguishable from the sec t. Malacogeton 紅 'e not sufficient to 回 at other sections as indicated in the key to the it it as a subgenus , but still great enough to rec- sections below (Appendix 1). A close rela- ognize ognize it as a distinct section. tionship between Sc h. sec t. Malacogeton and Sc h. sec t. Schoenoplectus was suggested by Key to species of sect. Malacogeton van der Veken (1965) based on embryo mor- 1.