Delineation of Schoenoplectus Sect. Malacogeton (Cyperaceae), New Combination, and Distinctions of Species

5 Pages·2001·0.39 MB·English

by  S. Galen Smith

植物研究雑誌
J. Jpn. Bot.
76: 339-343 (2001)
Delineation of Schoenoplectus sect. Malacogeton (Cyperaceae)，
New Combination，an d Distinctions of Species
a
S. Galen SMITHand Eisuke HAY ASAKAb
aDepartment of Biology，Un iversity of Wisconsin-Whitewater，
Whitewater，W isconsin，5 3190 U.S.A.;
bBotanical Garden，Gr aduate School of Science，To hoku University，Se ndai，98 0-0862 JAPAN
(Received on乱1ay14，2 001)
The new combination，S choenoplectus sect. Malacogeton (Ohwi) S. G. Smith &
Hayasaka，i s made and a description of the section is provided. Schoenoplectus
nipponicus，S ch. etuberculatus，Sc h. torreyi and Sch. subterminalis eincluded in sect.
訂
Malacogeton，a nd ak ey to these is given. Wer ecognize the sections Schoenoplectus，
Actaeogeton，Ma lacogeton and Supini in the genus Schoenoplectus，an d ak ey to the sec-
tions and ali st of North American and East Asian species included in each section are
provided.
Key words: Cyperaceae， new combination， Schoenoplectus sect. Malacogeton，
taxonomy
The polymorphic and polyphyletic Scirpus 2000， Koyama et al. 2000， Smith and
L. s. lat. in recent ye shas been subdivided Yatskievych 1996，W ilson 1981). Of these
紅
into about 17 genera (Bruhl 1995， segregate genera Schoenoplectus is comp a-
紅
Goetghebeur 1998， Wilson 1981， 1989). tively large，w ith species distributed world-
Scirpus as strict1yd efined includes about 30 wide，t he number of which is v iously
紅
species occurring mainly in the temperate re- estimated at 50 spp. (Goetghebeur 1998)，6 0
gions of the Northem Hemisphere，w ith S. spp. (Wilson 1981)，8 0 spp. (Hooper 1985)，
sylvaticus L. as the type species (Wilson or over 130 spp. (Koyama et al. 2000). For
1989). The rest of Scirpus s. lat. has been North America and East Asia，3 4 species are
placed in Actinoscirpus (Ohwi) R. W. Haines here recognized (Appendix 11). Following
& Lye， Amphiscirpus Oteng・.Yeboah， Goetghebeur and Simpson (1991)，w e here
Bolboschoenus (Asch.) Palla，B lysmopsis define Schoenoplectus as excluding
Oteng-Yeboah，B lysmus Schul ，.t Eleogiton Bolboschoenus and the monotypic
Link， Ficinia Schrad.， Isolepis R.Br.， Actinoscirpus，wh ich are sometimes included
O砂caηumNees，P hylloscirpus C.B.Clarke， in it (Lye 1995，S trong 1994).
Pseudoschoenus (C.B.Clarke) Oteng- Schoenoplectus as currently delimited is
Yeboah， Schoenoplectus (Rchb.) Palla， commonly characterized by its pseudolateral
Scirpoides Seg.， Sumatroscirpus Oteng- inflorescences，i. e.，t he proximal bract usu-
Yeboah，T richophorum Pers.，a nd Websteria ally erect and resembling the culm (Wilson
S.H.Wright by recent cyperologists (Bruhl 1981). Close examination shows that this is
1995， Goetghebeur and Simpson 1991， as uperficial character，h owever，b ecause the
Goetghebeur 1998，H ayasaka and Ohashi proximal involucI叫 bractsof 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 resemble the foliage leaf blades and usual architecture， weak rhizomes often
ecommonly divergent. Thus other charac- terminated by tubers，a nd often aquatic habi-
訂
ters are now used，e specially embryo form tat and flaccid habit. Close examination
(van der Veken 1965，W ilson 1981)，s tolon shows that these four species all belong to
presence，t uber presence and form，l igule Scirpus sect.乱1alacogeton Ohwi [=
presence，c ulm branching，le af position，le af Schoenoplectus subg.孔1alacogeton(Ohwi)
structure，s pikelet aangement，floral scale Oteng-Yeboah]. The required new combina-
町
indument，a nd culm anatomy. tion is provided as follows.
Species relationships within Schoeno
・
plectus enot clear because of large mor- Schoenoplectus (Rchb.) Palla in Verh. K.
紅
phological variation in the characters of the K. Zool.-Bot. Ges. Wien 38 (Sitzungsber.):
rhizomes，l eaves，i nflorescences，a chenes， 49 (1888)，n om. cons.
floral bristles and other structures that have Sect. Malacogeton (Ohwi) S.G.Smith &
been used to define the infrageneric taxa. To Hayasaka，c omb. nov.
clarify the species relationships within the Scirpus L. subgen. Schoenoplectus Rchb.
genus，a m odern infrageneric system should sect. Malacogeton Ohwi in Mem. Coll. Sci.
be established. This was attempted by Kyoto Imp. Univ. ser. B，18 : 97 (1944).
Oteng-Yeboah (1974) and Raynal (1976b)， Scirpus sect. Bolboschoenus Asch. ser.
but neither of their treatments esatisfac- Malacogeton (Ohwi) T. Koyama in J. Fac.
征
tory (Wilson 1981). In his brief paper， Sci. Univ. Tokyo，s ect. 3，7: 288 (1958).
Oteng-Yeboah (1974) recognized and pro Schoenoplectus subgen. Malacogeton
四
vided a key to the three subgenera (Ohwi) Oteng-Yeboah in Notes Roy. Bot.
Schoenoplectus，Ac taeogeton and Malacoge- Gard. Edinb. 33: 315 (1974).
ton，b ut he neither gave descriptions for the Perennials，e rect or flaccid，4 0-200 cm
subgenera nor the included species except high. Rhizomes long，w eak，v ery soft，o ften
for the type species. Raynal (1976a，b ，1 977) terminated by fleshy tubers to ca. 3 mm
recognized the four sections Schoenoplectus， thick. Culms trigonous or terete，c a. 0.5-5
Actaeogeton，S upini，a nd Pterolepis; with mmt hick. Leaves basal，ca . 3t o more than
emphasis on sect. Supini，in which he recog- 20; sheaths often free from culm and resem-
nized 21 species，m ainly from Africa. His bling blades; sheath fronts hyaline，s plitting
detailed study of Sch. sect. Supini should be early，u sually pinnate-fibrillose; blades 2t o
extended to the rest of the species to provide ca. 20 per culm，s ome species forming emer-
am ore complete classification for Schoeno- gent，er ect blades and/or submerged，f laccid
plectus. The present paper，w hich empha- blades，cr oss-section mostly trigonous or lat
同
sizes East Asian and North American erally flattened，a pices often blunt and ec-
species，co ntributes to the establishment of a centric; distal bl争delonger than sheath，ca .
Illodern in仕agenericsystem for the genus. It 0.2-10 mm wide. Inflorescences of one
also provides the name Schoenoplectus sect. spikelet or capitate or 1x to3x branched;
ー ー
乱1alacogetonand other information needed proximal bract erect，s tructure like foliage
for the senior author' seatmentof the genus leaf blades，ca . 1-32 cm long. Spikelets 1t o
位
in an upcoming volume of the Flora of No rth 20 per inflorescence，5 -25 x3 -6 mm. Floral
America (Flora of North America Editorial scales narrowly oblong-ovate，4 -7 mml ong，
Committee 2001). smooth or awn sparsely spinulose，s ome-
In Schoenoplectus three species in eastern times sp selyciliolate distally，f lanks each
訂
No rth America and one in East Asia share usually with 2t o ca. 10 prominent veins (at
December 2001 Journa1 of Japanese Botany Vo .176 No. 6 341
least in proximal part of spikelet)，a pices ing achene; eastern North American
subacute，e ntire or minutely notched，a wn coastal plains …・・ -
(extension of midrib) ca. 0.1-0.5 mm. …...・ ・.Schoenoplectusetuberculatus
H
Perianth bristles stout，f rom ca. 1/2 ofto 3. Leaf blades ca. 1-2 mmw ide; floral
同
twice achene length，r etrorsely to antrorsely scales awned，t he apical midrib exten-
spinulose. Anthers ca. 2-3.5 mmlo ng. Styles sions 0.2-0.5 mm; inflorescence of 1to 4
3-fid or 2-fid. Achenes yellow-to dark spikelets; not forming submerged mead-
brown when ripe，o void to obovoid，p romi- ows; widespread in eastern North
nently beaked， compressed-trigonous or America ・... ・...・ ・.Schoenoplectustorreyi
H H
biconvex，2 -4.5 mm，s mooth. Flowers ab- 3. Leaf blades 1m mo r less wide; floral
sent from basal sheaths. Fresh-water lakes， scales mucronate，t he apical midrib ex-
ponds，s treams，d itches，o ften deeply emer- tensions to 0.1 mm; inflorescence of one
gent or submerged aquatic，i n deep water erect spikelet; forming submerged mead-
flaccid and sometimes entirely vegetative. ows of vegetative plants in deep or flow-
Chromosome numbers reported: 2n = 42， ing water; widespread in No rth America，
70，74，76. mostly eastern...・…
Type species: Schoenoplectus n伊'/)onicus …..S choenoplectus subterminalis
(Makino) Sojak.
The type species is Sch. nipponicus，n ot Koyama (1958，u nder Scirpus) confused
Sch. etuberculatus as stated by Oteng- the species of Sch. sect. Malacogeton with
Yeboah (1974)，b ecause Ohwi (1944)，in his those of Bolboschoenus， which is now
key to species of Scirpus subg. treated as ad istinct genus even when Scirpus
Schoenoplectus， established Scirpus sect. is treated in a broader sense including
Malacogeton，a nd included only Scirpus Schoenoplectus (Kozhevnikov 1988). Oteng-
nipponicus. Koyama (1958) reduced Scirpus Yeboah (1974) regded Sch. sect.
紅
nipponicus to S. etuberculatus subsp. Malacogeton as intermediate between
n伊ponicus，and Oteng-Yeboah (1974) ap- Schoenoplectus and Bolboschoenus but
parently adopted Koyama's treatment of S. closer to the former in having a 'net-like
nipponicus and gave Sch. etuberculatus as ground tissue' and other characters.
the type species of Sch. subg. Malacogeton. Schoenoplectus sect. Malacogeton is here
Wet hink the diagnostic characters of Sch. considered to be distinguishable from the
sect. Malacogeton紅e'not sufficient to回 at other sections as indicated in the key to the
it as as ubgenus，bu t still great enough to rec- sections below (Appendix 1). A close rela-
ognize it as ad istinct section. tionship between Sch. sect. Malacogeton and
Sch. sect. Schoenoplectus was suggested by
Key to species of sect. Malacogeton van der Veken (1965) based on embryo mor-
1. Inflorescences branched，o f 3 to 25 phology.
spikelets…...・ ・.....・ ・.....・ ・.....・ ・.....・ ・-…2 The leaf architecture of sect. Malacogeton
H H H H H
1. Inflorescences unbranched，o f 1 to 5 appear to be unique in Schoenoplectus. Both
spikelets…...・ ・.....・ ・..…...・ ・..…...・ ・..…..3 Sch. etuberculαtus and Sch. subterminalis
H H H H
2. Styles bifid; achenes biconvex，ca . 2m m may form submerged，f laccid leaf blades
long; perianth bristles two times longer and/or emergent，e rect blades as described
than achene; East Asia... and illustrated by Schuyler (1972). In the
…Schoenoplectus nipponicus emergent blades of Sch. n伊'/)onicus，Sch.
2. Styles trifid; achenes trigonous，2 .5-4.5 etuberculatus，a nd Sch. torreyi，t he cross-
mml ong; perianth brist1es about equal- sections just distal to the ligules are thickly
342 植物研究雑誌第76巻第6号 平成13年12月
V- shaped to trigonous，th en in the central re- ed. 5: 191-317. National Taiwan University，
gions become acutely trigonous or laterally Taipei.
Kozhevnikov A. E. 1988. Cyperaceae. In: Charkevicz
flattened with one edge channeled，t hen to-
S. S. (edよPlantaeVasculares Orientis Extremi
ward the apex become asymmetrically
Sovietici 3: 175-403. Nauka，L eningrad.
flattened-trigonous. In the wider leaf blades
Lye K. A. 1995. Cyperaceae. In: Thulin M. (edよFlora
of Sch. etuberculatus and Sch. torreyi，t he of Somalia 4: 98-147. Royal Botanic Gardens，
apices are blunt and markedly asymmeic， Kew.
住
with the actual apex tumed to one side. Oteng-Yeboah A. A. 1974. Taxonomic studies in Cyp-
eraceae-Cyperoideae. Notes Roy. Bot. Gard.
Schoenoplectus sect. MaIacogeton is an
Edinb. 33: 311-316.
example of flowering plants with ad isjunct
Raynal J. 1976a. Notes Cyperologiques: 25. Le genre
distribution pattern between East Asia and Schoenoplectus 1. Sur quelques especes Sud-
North America，p ticularlyreferred to as Africaines. Adansonia，se r. 2，15 : 537-542.
ぽ
East Asian-eastern North American distribu- 一一一.1976b. Notes Cyperologiques: 26. Le genre
Schoenoplectus 11. L'amphicpieet la sect. Supini.
tion type，s uch as in Panax，S ymplocarpus， 征
Zizania，a nd Hamamelis (Wen 1999). Adansonia，se r. 2，16 : 119ー155.
一一一.1977. Addendum to notes Cyperologiques 26.
Adansonia，se r. 2，1 6: 530.
We thank the curators of Tohoku Schuyler A. E. 1972. Morphological and anatomical
University Herbarium (TUS) and many differences in leaf blades of three North American
No rth American Herbaria for the loan of aquatic bulrushes (Cyperaceae: Scirpus). Bartonia
41: 57-60.
specimens，a nd Mark Strong for helping the
Smith S. G. and Yatskievych.G. 1996. Notes on the
senior author prep'ean early version of an
訂 genus Scirpus sensu lato in Missouri. Rhodora 98:
infrageneric classification of Schoenoplectus. 168-179.
Strong M. T. 1994. Taxonomy of Scirpus，
References Trichophorum，a nd Schoenoplectus (Cyperaceae)
in Virginia. Bartonia 58: 29-68.
Bruhl J. J. 1995. Sedge genera of the world: relation-
Tang T. and Wang F.-T. 1961. Scirpus. In: Chien S.-S.
ships and an ew classification of the Cyperaceae.
and Chun W.-Y. (edsよFlora Reipublicae
Aust. Syst. Bot. 8: 125-305.
Popularis Sinicae 11: 2-34. Science Press，Be ijing.
Flora of North America Editorial Committee 作(eむd吋s..)
van der Veken P. 1965. Contribution al' embryogra-
2001. 日F1J.凹oraぱ0fNorth America north of Mexico. 4+ phie systematique des Cyperaceae-Cyperoideae. 、
vols.，O xford Univ. Press，N ew York and Oxford.
Bull. Jard. Bot. Etat Bruxelles 35: 285-354.
Goetghebeur P. 1998. Cyperaceae. In: Kubitzki K.
Wen J. 1999. Evolution of eastem Asian and eastem
(edよThefamilies and genera of vascular plants 4:
North American di司unctdistributions in flowering
141-190. Springer，B erlin.
plants. Ann. Rev. Ecol. Syst. 30: 421-455.
一一一.and Simpson D. A. 1991. Critical notes on
Wilson K. L. 1981. A synopsis of the genus Scirpus
Actinoscirpus， Bolboschoenus，I solepis，P hyllo-
sens. la .t(Cyperaceae) in Australia. Telopea 2:
scirpus and Amphiscirpus (Cyperaceae). Kew Bull.
153-172.
46: 169-178.
一一.1989. (931) Proposal to conserve 468 Scirpus L.
Hayasaka E. and Ohashi H. 2000. New combinations
(Cyperaceae) with S. sylvaticus L. as type. Taxon
in Japanese Schoenoplectus (Cyperaceae). J. Jpn.
38: 316-320.
Bot. 75: 223-225.
Hooper S. S. 1985. Cyperaceae. In: Townsend C. C.
Appendix 1. Key to the Sections of Schoeno・
and Guest E. (edsよFloraof Iraq 8: 331-406，
plectus
Ministry of Agriculture & Agrarian Reform，
1. Achenes rugulose，b lackish when ripe; floral scale
Republic of Iraq.
apices entire; plants densely tufted，r hizomes very
Koyama T. 1958. Taxonomic study of the genus
short，o r in Sch. lineolatus spreading by long rhi-
Scirpus L. J. Fac. Sci. Univ. Tokyo，se ct. 3，7: 271-
zomes...・・.....・ ・-…・・ ・・-… ・・-… ・・-…....・ ・...・・-・.2
366. H H H H H H H H H H
1. Achenes smooth，ye llow-to dark brown when ripe;
一一一，Kuoh C.-S. and Leong W.-C. 2000. Cyperaceae.
In: Huang T.ーC.& al. (eds.)，Fl ora of Taiwan，2 nd eflnotriarle socra len eaarpliyc ess o cliena rlsye cet. mM紅agliancaotgeettoo n;bi fpilda，nto sr
December 2001 Joumal of Japanese Botany Vo .l76 No. 6 343
widely spreading by rneans of rhizornes ....・ ・-……3 (Torr.) Sojak; Sch. torreyi (Olney) Palla
H
2. One (rarely 2) node and leaf usually present on It is possible that Sch. ehrenbergii (Bock.) S吋ak
culrn above basal leaf sheaths; plants often with (see fig. 7i n Tang and Wang 1961) of China and S. E.
flowers in bas}l leaf sheaths; perianth bristles ab- Russia rnight be included in sect. Malacogeton，b ut we
sent frorn spikelet flowers，s ornetirnes present in exclude it here pending further inforrnation.
basal flowers.…...・ ・.....・ ・-…....・ ・.....・ ・.sect.Supini
H H H H
2. All nodes crowded at base of culrn; plants never Schoenoplectus sect. Actaeogeton (Rchb.) J.Raynal
forrning flowers in basalleaf sheaths; perianth bris- Sch. かscorubens (T.Koyarna) T.Koyarna; Sch.
tles present or absent....・ ・・・..…sect.Actaeogeton hondoensis (Ohwi) Sojak; Sch. hotarui (Ohwi) Holub;
H H H
3. Floral scale flanks each with 2t o ca. 10 prorninent Sch. juncoides (Roxb.) Palla; Sch. komarovii (Roshev.)
veins，at least in proxirnal ptof spikelet，a pices Sojak; Sch. lineolatus (Franch. & Sav.) T.Koyarna;
紅
entire or obscurely ernarginate; rhizornes weak， Sch. mucronatus (L.) Palla [type species]; Sch.
soft when rnature，t ubers often terrninating rhi- ohwianus (T.Koyarna) Holub; Sch. orthorhizomatus
zornes; often subrnerged aquatic; North Arnerica (Arai & Miyarn.) Hayasaka & H.Ohashi; Sch.
and East Asia...・・・・..…...・ ・..….sect.Malacogeton purshianus (Fem.) M.T.Strong; Sch. smithii (A.Gray)
H H H H
3. Floral scale flanks without veins (except often on S吋ak;Sch. subbisetosus (T.Koyarna) Sojak; Sch.
basal scales)，a pices clearly ernarginate to deeply triangulatus (Roxb.) Sojak; Sch. wallichii (Nees)
bifid; rhizornes strong，h ard when rnature，t ubers T.Koyarna
absent; ernergent aquatic or on wet soils (Sch.
lacustris sornetirnes subrnerged aquatic); world- Schoenoplectus sect. Supini (Cherrn.) J.Raynal
wide..・・.....・ ・...・ ・....・ ・...・ ・...sect. Schoenoplectus Sch. articulatus (L.) Palla; Sch. erectus (Poir.) Palla ex
H H H H H
J.Raynal; Sch. hallii (A.Gray) S.G.Srnith; Sch.
Appendix 11. List of Sections and Included αsximontanus (Fem.) J.Raynal; Sch. supinus (L.) Palla
Species from North America and East Asia [type species]
Schoenoplectus sect. Schoenoplectus
Sch. acutus (Muhl. ex Bigelow) A.Love & D.Love; The infrageneric classification of Schoenoplectus
Sch. americanus (Pers.) Volkart; Sch. californicus given herein is based rnainly on East Asian and North
(C.A.Mey) Sojak; Sch. deltarum (Schuyler) Sojak; Sch. Arnerican species. A rnore cornplete infrageneric sys-
heterochaetus (Chase) Sojak; Sch. lacustris (L.) Palla tern based on world-wide species is needed. It is possi-
[type species]; Sch. litoralis (Schrad.) Palla; Sch. ble that rnore than four sections rnight be recognized
pungens (Vahl) Palla; Sch. tabemaemontani (C.C. within the genus，b ecause sorne African species，e. g.，
Grnel.) Palla; Sch. triqueter (L.) Palla Sch. muriculatus (Kuk.) J.Browning and Sch.
paludicola (Kunth) Palla ex J.Raynal cannot definitely
Schoenoplectus sect. Malacogeton (Ohwi) S.G.Srnith be assigned to either of the four sections. In仕asectional，
& Hayasaka taxa rnight also be recognized after ar nore detailed
Sch. etuberculatus (Steud.) Sojak; Sch. nipponicus study of species，e specially in sect. Schoenoplectus.
(M北ino)S吋ak[type species]; Sch. subterminalis
スミス S.G人早坂英介 .フトイ属シズイ節(カ
b
ヤツリグサ科)の学名の新組み合わせ，節の定義
と種の区別 点を明らかにした.また フトイ属に 4節 sect.
フトイ属シズイ節の学名の新組み合わせ Schoenoplectusフトイ節;sect. Malacogetonシズイ
Schoenoplectus (Rchb.) Palla sect. Malacogeton 節， sect. Actaeogeton (Rchb.) J.Raynalカンガレイ
(Ohwi) S.G.Srnith & Hayasakaを提唱し，節の形態 節， sect. Supini (Cherrn.) 1. Raynalホソガタホタル
的な特徴を記載した.シズイ節には北米に 3種 イ節を認めてこれらの節への検索表を示し，それ
Sch. etuberculatus (Steud.) S吋ak，Sch. subterminalis ぞれの節に含まれる北米と東アジアの種を一覧し
(Torr.) Sojak，Sc h. torreyi (Olney) Palla，および東ア た.
ジアに 1種Sch.nipponicus (Makino) S吋akシズイ (aウイスコンシン大学生物学部，
がある.これらの種への検索表を示し，種の区別 b東北大学大学院理学研究科付属植物園)