王文采
(中國科學(xué)院植物研究所系統(tǒng)與進(jìn)化植物學(xué)國家重點實驗室,北京100093 )
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中國赤車屬新分類
王文采
(中國科學(xué)院植物研究所系統(tǒng)與進(jìn)化植物學(xué)國家重點實驗室,北京100093 )
摘要:( 1)根據(jù)該文中赤車屬與樓梯草屬各種形態(tài)學(xué)的區(qū)別,認(rèn)為赤車屬較為原始,在樓梯草屬之前自冷水花屬衍生而出。( 2)將赤車屬劃分為5組;將C.B.Robinson描述的Elatostemoides屬降為組級群,作為赤車屬中接近原始群全緣赤車組sect.Pileoides的一個組處理,此外還描述了一個與sect.Elatostemoides近緣的1新組,羽脈赤車組sect.Leiolaena,并根據(jù)葉、葉的脈序、雌花序、雌花被片和瘦果等器官的演化趨勢給出顯示赤車屬屬下諸群間親緣關(guān)系的系統(tǒng)發(fā)育樹。( 3)對中國赤車屬的屬下分類群進(jìn)行了修訂,共承認(rèn)32種1亞種7變種,給出檢索表、描述及多幅插圖,并將它們劃分為4組、2亞組和9系,其中除sect.Leiolaena之外,還新描述了2亞組、5系、2種和3變種;在2003年被歸并為同物異名的長柄赤車、曲毛赤車、波緣赤車、小赤車和富寧赤車的種級地位得到恢復(fù)。此外,還寫出了中國赤車屬的分類學(xué)簡史和地理分布。
關(guān)鍵詞:蕁麻科,赤車屬,分類學(xué)修訂,中國
王文采.中國赤車屬新分類[J].廣西植物,2016,36( 1) : 1-29
WANG WT.Nova classification specierum sinensium Pellioniae ( Urticaceae)[J].Guihaia,2016,36( 1) : 1-29
Nova classification specierum sinensium Pellioniae( Urticaceae)
WANG Wen-Tsai
( State Key Laboratory of Systematic and Evolutionary Botany,Institute of Botany,Chinese Academy of Sciences,Beijing 100093,China )
Abstract:( 1) According to the various morphological differences between the two genera,Pellionia and Elatostema,enumerated in the present paper,the genus Pellionis is considered more primitive and to be derived from the genus Pilea before the genus Elatostema.( 2) The genus Pellionia is subdivided into five sections.The genus Elatostemoides described by C.B.Robinson is relegated to sectional rank and treated as a section under Pellionia near its primitive group,sect.Pileoides.Besides,a new section related to sect.Elatostemoides,sect.Leiolaena,is described.On the basis of the evolutionary trends of the leaves,leaf nervation,pistillate inflorescences,pistillate tepals and achenes a putative phylogenetic tree showing the relationships among the infrageneric groups of the genus Pellionia is provided.( 3) The infrageneric taxa of Pellionia in China are revised.Thirty-two species,one subspecies and seven varieties are recognized.They are keyed,described and in most cases illutrated,and are classified into four sections,two subsections,and nine series.Of them,except for sect.Leiolaena just mentioned above,two subsections,five series,two species and three varieties are described as new.The specific statuses of Pellionia tsoongii,P.crispulihirtella,P.subundulata,P.minima and P.funingensis,which were reduced into synonymy in 2003,are reinstated here.Besides,the brief taxonomic history and geographical distribution of the genus Pillionia in China are given. Carex sect. Infossae S. W. Su was recently described from Anhui and Jiangsu provinces of East China,
Key words:Urticaceae,Pellionia,taxonomic revision,China Carex sect. Infossae,scanning electron microscopy,morphological character,new synonym,China
1790年,葡萄牙植物學(xué)家J.Loureiro在其編著的“Flora Cochinchinensis”中根據(jù)分布于我國和中南半島的Polychroa repens Lour.建立了蕁麻科新屬Polychroa Lour.,遺憾的是此新屬自當(dāng)時一直到20世紀(jì)初都遭到國際植物學(xué)界的忽視,最后竟淪為一同物異名( synonym,見下)。1830年,法國植物學(xué)家C.Gaudichaud-Beaupré建立了蕁麻科新屬Pellionia Gaudich.(赤車屬),其命名模式為P.elatostemoides Gaudich.( Hutchinson,1967),此屬名在19世紀(jì)中得到廣泛接受和應(yīng)用,因此被列入1905年《國際植物學(xué)命名法規(guī)》規(guī)定建立的nomina generica conservanda(保留屬名)名錄中,因此比其早40年發(fā)表的Polychroa成為其同物異名。
1861年,英國植物學(xué)家G.Bentham在其編著的“Flora Hongkongensis”中描述了赤車屬2新種: Pellionia scabra Benth.和P.brevifolia Benth.。1868年,另一英國植物學(xué)家H.F.Hance根據(jù)C.F.M.Grijs采自福建的一號標(biāo)本描述了赤車屬又1新種Pellionia grijsii Hance。1869年,瑞士蕁麻科專家H.A.Weddell編寫出當(dāng)時的世界蕁麻科植物志,此志中的冷水花族trib.Procrideae包括Pilea,Pellionia,Lecanthus,Procris和Elatostema等屬。此后,Baillon( 1872),Bentham&Hooker( 1880),Engler ( 1893)和Hutchinson( 1967)等學(xué)者的有關(guān)蕁麻科的著作中均接受了Weddell的trib.Procrideae的內(nèi)容; 在Pellionia屬,Weddell收載了分布于印度、中國、日本和東南亞以及大洋洲一些島嶼的15種。1896年,德國植物學(xué)家H.Hallier不承認(rèn)Pellionia和Procris的屬級地位,將此二屬降級作為Elatostema屬的二亞屬處理。1886-1905年,英國植物學(xué)家F.B.Forbes&W.B.Hemsley主要根據(jù)收藏于英國邱皇家植物園標(biāo)本館的中國標(biāo)本,編著了中國植物名錄; 1899年,C.H.Wright承擔(dān)了其中的蕁麻科部分,在Pellionia屬中,記錄了7種,其中包括根據(jù)湖北和四川標(biāo)本描述的新種Pellionia viridis C.H.Wright,和根據(jù)A.Henry采自云南蒙自的4號標(biāo)本鑒定的P.griffithiana Wedd.。1910年,菲律賓蕁麻科專家C.B.Robinson根據(jù)菲律賓、加里曼丹(原知Borneo)等地區(qū)原置于Pellionia或Elatostema二屬的10余種建立了新屬Elatostemoides C.B.Robinson; 1935年,德國植物學(xué)家H.Schr?ter將此新屬降級作為Elatostema的一個亞屬;本文中,我將其作為Pellionia的一個組處理(見下)。1916年,日本植物學(xué)家B.Hayata描述了中國臺灣赤車屬新種P.a(chǎn)risanensis Hayata; 1925年,又一日本植物學(xué)家Y.Yamamoto描述了中國臺灣赤車屬P.chikushiensis Yamam.和P.keitaoensis Yamam.等新植物。1922年,美國植物學(xué)家E.D.Merrill根據(jù)鐘觀光教授采自廣西東興的一號標(biāo)本描述了新種Polychroa tsoongii Merr.,他了解到Pellionia被列入保留屬名的名錄中后,于1928年將此種名和Polychroa repens Lour.一同轉(zhuǎn)移到Pellionia中。1929年,奧地利植物學(xué)家H.Handel-Mazzetti記錄了中國Pellionia屬6種,他接受Hallier的觀點,將Pellionia歸并于Elatostema,在此6種中,他擬出一新名Elatostema henryanum Hand.-Mazz.以代替E.griffithianum ( Wedd.) Hall.f.( Pellionia griffithiana Wedd.),并根據(jù)他在越南北部與云南河口交界的Laogai采集的一號標(biāo)本和A.Henry在云南采集的二號標(biāo)本描述了后者的一新變種E.henryanum var.oligodontum Hand.-Mazz.; 1936年,H.Schr?ter將此變種升至種級,并擬出新名Elatostema paucidentatum H.Schr?ter; 1963年,錢崇澍教授在編寫《海南植物志》蕁麻科的過程中將此種從Elatostema轉(zhuǎn)移到Pellionia屬中。
1935-1936年,德國植物學(xué)家H.Schr?ter發(fā)表了他編著的Elatostema屬專著的前兩部分。他接受Hallier的觀點,將Pellionia作為Elatostema的一個亞屬,同時將Elatostemoides C.B.Robinson也降級作為Elatostema的另一個亞屬。他的概念的Elatostema屬包括以下4亞屬:
Subgen.1.Pellionia( Gaudich.) Hall.f.退化葉常存在。雄、雌花序均無花序托。雄花序多有長梗,為稀疏的聚傘花序,稀為團(tuán)傘花序。雌花序無?;蛴泄#瑸閳F(tuán)傘花序,稀為疏散的聚傘花序。苞片小,不包住花序。雄花和雌花均為( 4) 5基數(shù)。雌花被片與子房等長,常具長角狀突起。瘦果有瘤狀突起。
Subgen.2.Elatostemoides ( C.B.Robinson) H.Schr?ter退化葉、雄、雌花序的特征同上亞屬。雄花和雌花均5基數(shù)。雌花被片與子房等長,具肋或具突起,但不具長角狀突起。瘦果多光滑。
Subgen.3.Weddellia H.Schr?ter退化葉存在。雄、雌花序均具花序托,雄花序多無梗,雌花序無梗。苞片寬,將花序多少包住。雄花5基數(shù)。雌花3( 4)基數(shù)。雌花被片強(qiáng)烈退化,極小。瘦果具縱肋。
Subgen.4.Euelatostema Baillon退化葉不存在( E.surculosum等少數(shù)種例外)。雄、雌花序均具花序托,最發(fā)育的呈盤狀。苞片寬,多少包住花序。雄花( 4) 5基數(shù)。雌花多為3基數(shù)。雌花被片極退化,或完全消失。瘦果通常具縱肋。
其專著的第一部分( 1935)討論了Elatostema屬植物的葉、花序、花等器官的形態(tài)特征和地理分布,第二部分( 1936)是前3個亞屬的專著。subgen.Pellionia,收載55種(包括我國2新種: Elatostema yunnanense H.Schr?ter和E.incisoserratum H.Schr?ter,及前述的新名E.paucidentatum H.Schr?ter),subgen.Elatostemoides,收載45種,subgen.Weddellia,收載6種。對每一亞屬均給出德文的分種檢索表,每種都給出重要文獻(xiàn)、拉丁文形態(tài)描述、標(biāo)本引證以及與近緣種有關(guān)親緣關(guān)系的討論。對每亞屬未給出正式的亞屬下分類,只將sugen.Elatostemoides根據(jù)雄花序具梗與否等特征將此亞屬的植物劃分為3群: rostratum-Gruppe,vittatum-Gruppe和Weinlandii-Gruppe。此外,subgen.Pellionia,具三出脈的原始種Elatostema latifolium,E.tsonngii和E.heyneanum被放在此亞屬55種中開始的位置,這種情況似乎說明H.Schr?ter注意到了這幾個種較低的演化水平。subgen.Pellionia和subgen.Elatoatemtoides的內(nèi)容加在一起實際上就是Pellionia屬的專著,這為Pellionia屬今后的研究提供了重要的基本資料。
1980年,我編著了我國赤車屬的第一個修訂,將當(dāng)時鑒定的17種劃分為3組、4系,其中包括4新種: Pellionia acutidentata W.T.Wang,P.retrohispida W.T.Wang,P.subundulata和具雌頭狀花序的P.cephaloidea W.T.Wang; 1錯誤鑒定:將2 號P.griffithiana Wedd.的標(biāo)本誤定位P.heteroloba Wedd.。在赤車屬中雌頭狀花序的出現(xiàn)說明Pellionia和Elatostema兩個近緣屬存在平行演化現(xiàn)象。到1995年,我負(fù)責(zé)編寫的中國赤車屬志中的種數(shù)增至24種,被劃分為3組、6系,增加的7種是Pellionia longipedunculata W.T.Wang,P.funingense W.T.Wang,P.crispulihirtella W.T.Wang,P.macrophylla W.T.Wang,P.caulialata S.Y.Liou,P.leiocarpa W.T.Wang和P.brachyceras W.T.Wang。2003年,英文版“Flora of China”的Pellionia屬志由Lin et al承擔(dān)編寫,共收載20種,未進(jìn)行屬下分類,但種的排列次序與《中國植物志》赤車屬的種排列次序相同。這里赤車屬的種數(shù)變少,是作者對7個種名和3個變種名進(jìn)行了歸并的緣故。這些種的歸并,有的正確,有的錯誤[如將具雌頭狀花序的P.cephaloidea W.T.Wang歸并于P.scabra Benth.(王文采,2011)]。近年來,又有赤車屬數(shù)新種發(fā)現(xiàn),由于種數(shù)的增加和錯誤歸并需進(jìn)行改正,我決定編寫本文,即我國赤車屬的第4次修訂。
赤車屬和樓梯草屬在親緣關(guān)系上極為相近,具有相同的體態(tài)和雄花構(gòu)造,大概由于這些相似性,一些植物學(xué)家如H.Hallier( 1896),H.Handel-Matzetti ( 1929),H.Schr?ter( 1935-1936)和I.Friis( 1993)認(rèn)為此二屬實為一屬,應(yīng)將Pellionia歸并于Elatostema。但另一方面,如上節(jié)所述,不少植物學(xué)家則主張承認(rèn)二屬,因此有必要再將二屬的不同特征列出來加以考慮。
2.1赤車屬
雄、雌花序均為聚傘花序,雄聚傘花序多少稀疏分枝,而雌聚散花序分枝短、密集,雌花序具短梗或無梗、極為密集,因此H.Schr?ter稱雌聚散花序為團(tuán)傘花序。2種(特產(chǎn)湖南的Pellionia cephaloidea W.T.Wang和特產(chǎn)印度東北部的P.a(chǎn)mbigua Wedd.),雌花序具花序托(可能由花序密集短分枝發(fā)生愈合而成),在花序托邊緣有總苞,如上所述,這是赤車屬和樓梯草屬二近緣屬的一種平行演化現(xiàn)象。雌花的花被片5枚,稀4或3枚,比子房稍長或近等長,無任何突起或在背面頂端之下有短或長的角狀突起。瘦果具瘤狀突起或光滑,稀具短線紋,或具短線紋和小瘤狀突起,或具縱肋和小瘤狀突起(本文首次發(fā)現(xiàn),見種32頭序赤車P.cephaloidea)(在蕁麻科的其他屬如蕁麻屬Urtica、艾麻屬Laportea、蝎子草屬Girardinia和冷水花屬Pilea,瘦果均具瘤狀突起或光滑)。
2.2樓梯草屬
雌花序均為頭狀花序,具花序托和總苞。絕大多數(shù)種雄花序和雌花序一樣為頭狀花序,只在此屬原始群疏傘樓梯草組sect.Pellioioides的12種(特產(chǎn)我國西南部和南部)為分枝的聚傘花序。雌花的花被片強(qiáng)烈退化,通常不存在,存在時通常為2-3枚,稀4-5枚,極小,比子房短,無任何突起(有2例外,特產(chǎn)云南東南部的五被樓梯草Elatostema quinquetepalum W.T.Wang和長被樓梯草E.longitepalum W.T.Wang的雌花被片呈狹條形,長度超過子房)。瘦果多數(shù)種(包括sect.Pellionioides的12種)具縱肋和具縱肋小瘤狀突起,少數(shù)種具瘤狀突起,具短線紋或光滑。值得注意的是具縱肋及具縱肋小瘤狀突起兩種類型的瘦果原知是樓梯草屬所獨有的,現(xiàn)在頭序赤車P.cephaloidea也發(fā)現(xiàn)了具縱肋和小瘤狀突起的瘦果,這是Pellionia和Elatostema二近緣屬的又一個平行演化現(xiàn)象。
據(jù)上述二屬的區(qū)別特征,可見赤車屬保留了較多的原始特征,如與冷水花屬相似的聚傘花序,雌花被片和瘤狀突起或光滑的瘦果。樓梯草屬則出現(xiàn)了較多的進(jìn)化特征,如頭狀花序,強(qiáng)烈退化的雌花被片,具縱肋及具縱肋小瘤狀突起的瘦果。由此可以推測赤車屬Pellionia和樓梯草屬Elatostema雖然都源于冷水花屬Pilea,在起源時間上樓梯草屬可能在較近時間出現(xiàn),而赤車屬可能早于樓梯草屬自冷水花屬演化而出,并走上了一條與后者不同的演化路線(王文采,1980,2014)。
根據(jù)葉的脈序類型,退化葉的存在與否,雌花序類型,雌花被片和瘦果的形態(tài)特征,可將赤車屬植物劃分為下列5組:
組1.全緣赤車組sect.Pileoides W.T.Wang葉具長柄或稍長柄,具全緣邊緣,具三出脈,葉狹側(cè)的側(cè)生基出脈向上直展達(dá)近葉頂端處。退化葉存在。雌花序聚傘狀。雌花的花被片5枚,在背面無任何突起或頂端之下具短突起,瘦果具瘤狀突起。
4種,我國有2種。
組2.喙尖赤車組sect.Elatostemoides ( C.B.Robinson) W.T.Wang葉具短柄或無柄,具有齒邊緣,具三出脈或半離基三出脈,葉狹側(cè)的側(cè)生基生脈向上直達(dá)近葉頂端處。退化葉通常存在,稀不存在。雌花序聚傘狀。雌花的花被片5枚,在背面無任何突起或頂端之下有短突起。瘦果通常光滑。
45種,我國無此組植物。
組3.赤車組sect.Pellionia葉具短柄或無柄,通常具有齒邊緣,具半離基三出脈(葉狹側(cè)的側(cè)生基出脈向上伸展達(dá)或稍超過葉中部),稀具羽狀脈。退化葉存在或不存在(我國此組的種多無退化葉)。雌花序聚傘狀。雌花的花被片5枚,稀4或3枚,通常其中( 1-) 2-3枚較大,成條狀船形,并在背面頂端之下具長角狀突起。瘦果具瘤狀突起,稀具短線紋(大葉赤車P.macrophylla)。
約62種。我國有22種,根據(jù)雌花被片、退化葉、托葉、脈序和瘦果的特征被劃分為2亞組,5系。
組4.羽脈赤車組sect.Leiolaena W.T.Wang葉具短柄或無柄,具有齒邊緣,具羽狀脈。退化葉通常不存在。雌花序聚傘狀。雌花的花被片5枚,稀4或3枚,在背面通常無任何突起,稀具短突起。瘦果光滑或具小瘤狀突起,稀具短線紋和小瘤狀突起。
8種。我國有7種,據(jù)瘦果類型被劃分為3系。
組5.頭序赤車組sect.Elatostematopsis W.T.Wang葉無柄或有極短柄,具有齒邊緣,具三出脈或半離基三出脈,葉狹側(cè)的側(cè)生基出脈向上伸展稍超過葉中部。退化葉存在或不存在。雌花序頭狀,具盤狀花序托和總苞。雌花的花被片5枚,近等大,無任何突起或背面頂端之下具短突起。瘦果具瘤狀突起或具縱肋和小瘤狀突起。
2種,我國有1種。
從上述組、系的劃分可以觀察到赤車屬有關(guān)形態(tài)特征的演化趨勢: ( 1)葉對生,同一對葉中的一枚正常發(fā)育,另一枚強(qiáng)烈退化成一小葉片→退化葉消失,葉變?yōu)榛ド? ( 2)葉柄:長→短→不存在; ( 3)葉的脈序:三出脈→半離基三出脈→羽狀脈; ( 4)托葉:狹卵形、三角形、披針形→鉆形; ( 5)雌花序:聚傘花序→頭狀花序; ( 6)雌花被片:近等大,背面無任何突起→背面頂端之下有短突起→( 1-) 2-3枚較大,背面頂端之下有長角狀突起; ( 7)瘦果:具瘤狀突起或光滑→具短線紋→具短線紋和小瘤狀突起→具縱肋和小瘤狀突起。
根據(jù)上述有關(guān)形態(tài)特征的演化趨勢,可見在赤車屬的5個組中,全緣赤車組sect.Pileoides具有較多的與冷水花屬相似的原始特征,如葉具較長葉柄,具三出脈,退化葉存在,雌花序聚傘狀,雌花被片無任何突起或背面有短突起,瘦果具瘤狀突起等。因此,可以推測全緣赤車組是赤車屬的原始群,其他組可能均由其演化而出,赤車屬的系統(tǒng)發(fā)育可能如圖1所示。
圖1 赤車屬的系統(tǒng)發(fā)育Fig.1 Phylogeny of Pellioniae
赤車屬約有120種,廣泛分布于亞洲南部和東南部和大洋洲西南部島嶼的熱帶地區(qū)以及亞洲東部亞熱帶地區(qū)。我國約有32種,1亞種,7變種(其中有16特有種,6特有變種),廣泛分布自云南和華南向北達(dá)長江中下游各省,多數(shù)種約30種集中分布于云南東南部和廣西。各省、區(qū)的統(tǒng)計:云南有21種1亞種2變種;廣西有20種1亞種3變種;湖南有8 種;廣東和江西均有6種1變種;福建有6種;四川有5種1變種;貴州有5種;重慶、湖北、安徽和浙江各有4種;臺灣有3種;江蘇和甘肅各有2種。分組敘述如下:
組1.全緣赤車組sect.Pileoides本組有4種。其中Pellionia latifolia( Bl.) Boerl.分布于緬甸東南部、中南半島、馬來半島以及印度尼西亞的蘇門答臘、爪哇等島嶼。全緣赤車P.heyneana分布于印度南部、斯里蘭卡、柬埔寨和我國的云南南部、廣西西北部。長柄赤車P.tsoongii分布于我國(云南南部、廣西和海南)、柬埔寨和越南。第4種P.machaerophylla( Hall.f.) Boerl.特產(chǎn)印度尼亞蘇門答臘島。
組2.喙尖赤車組sect.Elatostemoides本組有45種,分布于東南亞熱帶島嶼和大洋洲西南部島嶼,分布較廣的只有1種,P.rostrata ( Hall.f.) Boerl.,分布于印度尼西亞的明打威群島、蘇門答臘、爪哇、馬魯古群島等島嶼,其他種多為狹域分布,在加里曼丹(過去稱Borneo)有22種,菲律賓有10種,蘇拉威西島(過去稱Celebes)和新幾內(nèi)亞島各有2種,新幾內(nèi)亞島和所羅門島共有2種,蘇門答臘、所羅門島和斐濟(jì)各有1特有種。
組3.赤車組sect.Pellionia本組約有60種,分布自印度北部向東經(jīng)緬甸、我國達(dá)日本,再自中南半島向南達(dá)爪哇島,向東南經(jīng)菲律賓達(dá)大洋洲西南部的新幾內(nèi)亞島和所羅門島。分布較廣的種不多,其中的南亞赤車Pellionia griffithiana自印度東北部向東分布達(dá)我國南部(云南、廣西、廣東)和中南半島、向南達(dá)爪哇。異被赤車P.heteroloba自印度北部和東北部向東北布部達(dá)我國西南部(云南、貴州、湖南、四川、重慶)和中南半島。吐煙花P.repens分布自緬甸向東達(dá)中南半島和我國(云南南部、廣西和海南),向南經(jīng)馬來半島達(dá)蘇門答臘和爪哇。蔓赤車P.scabra自越南北部、我國云南東南部和華南向北達(dá)長江流域中、下游地區(qū),再向東達(dá)朝鮮和日本。赤車P.radicans var.radicans的分布與蔓赤車相似,只是其分布區(qū)西北部沒有達(dá)到甘肅南部;其大葉變種var.grandis分布于越南北部和我國云南東南部和廣西一帶。短葉赤車P.brevifolia和小赤車P.minima的分布與蔓赤車和赤車的分布頗為相似,自廣西、廣東分布達(dá)長江中、下游地區(qū),再向東達(dá)日本,但二者的分布區(qū)均較小,向南沒有分布到越南北部,向東未達(dá)中國臺灣島,此外,在分布區(qū)中的種群分布稀疏,數(shù)目較少。本組的狹域分布種較多:新幾內(nèi)亞有16種,加里曼丹島有9種,中國有9種,菲律賓有4種,所羅門島有3種,馬來半島有2種,越南、日本、斐濟(jì)各有1種。
我國此組有22種5變種。云南有18種2變種(包括5特有種:富寧赤車P.funingensis,尖齒赤車P.a(chǎn)cutidentata,三被赤車P.tritepala,灰被赤車P.calcifera和云南赤車P.yunnanensis) ;廣西有10 種3變種(包括1特有種:長梗赤車P.longipedunculata) ;湖南有7種;海南和江西各有6種1變種;福建有6種;廣東和四川各有5種1變種;貴州有5 種;重慶、湖北和安徽各有4種;臺灣有3種;江蘇南部和甘肅南部各有2種。
組4.羽脈赤車組sect.Leiolaena本組有8種。我國有7種,分布于云南東南部、廣西和廣東北部。其中廣西和廣東北部共有2種(羽脈赤車Pellionia incisoserrata和兩廣赤車P.ronganensis),云南東南部和廣西,以及越南北部共有1種(光果赤車P.leiocarpa),其他4種均特產(chǎn)廣西(龍州赤車P.longzhouensis和柔毛赤車P.mollissima均特產(chǎn)龍州,東蘭赤車P.donglanensis特產(chǎn)東蘭,短角赤車P.brachyceras特產(chǎn)廣西西北部的隆林)。第8種,P.procridioides Gagnep.特產(chǎn)越南北部。
組5.頭序赤車組sect.Elatostematopsis本組有2種,我國有1種,頭序赤車Pellionia cephaloidea,特產(chǎn)湖南衡山。另1種,P.a(chǎn)mbigua Wedd.特產(chǎn)印度東北部( Weddell,1869; Hooker,1888)。
赤車屬的原始群全緣赤車組sect.Pileoides的4種中有3種( P.latifolia,P.heyneana和P.tsoongii)分布于云南高原南部和中南半島中部和北部一帶。另一方面則可以看到樓梯草屬Elatostema的原始群疏傘樓梯草組sect.Pellionioides的多數(shù)種(如短梗樓梯草E.brevipedunculatum,硬毛樓梯草E.hirtellum,三岐樓梯草E.trichotomum,多岐樓梯草E.polystachyoides,鈍齒樓梯草E.obtusidentatum,角萼樓梯草E.subtrichotomum var.corniculatum,長圓樓梯草E.oblongifolium,微齒樓梯草E.microdontum——王文采,2014)分布于云南西部和東南部、廣西西部一帶,與上述赤車屬三原始種的分布區(qū)鄰近或重迭。從上述二原始群植物的分布情況可以推測Pellionia和Elatostema二屬可能起源于云南高原南部一帶,Pellionia沿著三條遷移路線擴(kuò)展其分布區(qū),向西沿喜馬拉雅走廊達(dá)印度、斯里蘭卡,向南經(jīng)馬來半島達(dá)到東南亞及大洋洲西南部熱帶島嶼,向東則到達(dá)日本一帶。(王文采,1992)
赤車屬
PellioniaGaudich.in Freyc.Voy.Bot.494,t.119.1830,nom.gen.cons.; Wedd.in Arch.Mus.Hist.Nat.Paris 9 : 282.1856; et in DC.Prodr.16( 1) : 165.1869; Baill.Hist.Pl.3: 524.1872; Benth.&Hook.f.Gen.Pl.3( 1) : 385.1880; Engler in Engler&Prantl,Nat.Pflanzentam.3 ( 1) : 108.1893; Hutch.Gen.Flow.Pl.2: 184.1967.——Elatostema Forster subgen.Pellionia ( Gaudich.) Hall.f.in Ann.Jard.Bot.Bruitenz.13: 303.1896; H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83( 1) : 2.1935; ( 2) : 1,17.1936.Type: P.elatostemoides Gaudich。
Polychroa Lour.Fl.Cochinch.559.1790.Type: P.repens Lour.
草本或亞灌木。葉二列,兩側(cè)不相等,狹側(cè)向上,寬側(cè)向下,具三出脈、半離基三出脈或羽狀脈,通常具鐘乳體,托葉2;退化葉小,存在或不存在?;ㄐ騿涡裕菩弁昊虍愔?雄花序為聚傘花序,通常具長梗并多少稀疏分枝;雌花序通常為聚傘花序,由于分枝短而密集,多少呈球形,具密集的苞片和雌花,稀為頭狀花序。雄花:花被片( 4-) 5;雄蕊( 4-) 5;退化雄蕊小或不存在。雌花:花被片( 3-4-) 5,通常比子房長,披針形,長圓形或條形,平或船形,無或有角狀突起;退化雄蕊小,與花被片同數(shù);子房橢圓球形,柱頭畫筆頭狀。瘦果具瘤狀突起或光滑,稀具短線紋,具短線紋和小瘤狀突起,或具縱肋和小瘤狀突起。
約有120種,分布于亞洲熱帶和亞熱帶地區(qū)和大洋洲西南部島嶼。我國約有32種(包括16特有種),1亞種,7變種,自云南和華南向北達(dá)長江中、下游地區(qū)廣布。
分種檢索表
1.Pistillate inflorescences cymose,very densely multi-flowered,lacking receptacles and involucres; achenes ellipsoid or ovoid,tuberuculate or smooth,rarely lineolate or lineolate and minutely tuberculate.
2.Leaves long or slightly long petiolate,entire,trinerved,with lateral basal nerve at leaf narrow side extending upwards nearly to leaf apex; reduced leaves present; pistillate flower’s tepals 5,lacking any projections or abaxially below apex shortly corniculate; achenes tuberculate (組1.全緣赤車組sect.Pileoides).
3.Leaf blades 5-15×3-7.4 cm,base obliquely rounded,at leaf broad side inconspicuously auriculate,cystoliths 0.2-0.4 mm long; petioles 0.5-2 cm long; staminate cymes 1.5-3.5 cm long,with peduncles 5.5-12 cm long……………1.全緣赤車P.heyneana
3.Leaf blades larger,12.5-20×5.8-11 cm,base asymmetrical,at leaf broad side with auricles 0.8-2.7 cm long,cystoliths 0.5-0.7 mm long; petioles 4-19 cm long; staminate cymes 3-4 cm broad,with peduncles 3.2-10 cm long……………2.長柄赤車P.tsoongii
4.Leaf blades abaxially on nerves and petioles puberulous……………………………………………2a.模式亞種ssp.tsoongii
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4.Leaf blades abaxially and petioles glabrous……………………………………………………2b.無毛長柄赤車ssp.subglabra
2.Leaves shortly petiolate or sessile,usually with dentate margins,semi-triplinerved or penninerved,with the lateral basal nerve at leaf narrow side extending upwards nearly to or slightly surpassing the leaf middle.
5.Pistillate tepals lacking any projections,or abaxially below apex shortly cornitulate.
6.Leaves semi-striplinerved; reduced leaves present; normal leaves with blades obovate or elliptic,1.8-7×1.2-3.7 cm,apex obtuse,rounded or slightly acute,margin crenate or undulate; stipules triangular,4-8×2-5 mm; staminate cymes 0.6-3 cm broad,with peduncles 2-11 cm long; pistillate flower’s tepals 5; achenes tuberculate (組2.赤車組,吐煙花亞組sect.Pellioniasubsect.Repentes)…………………………………………………………………………………………3.吐煙花P.repens
6.Leaves penninerved(組3.羽脈赤車組sect.Leiolaena ).
7.Achenes not smooth; pistillate tepals lacking any projections; reduced leaves absent.
9.Leaf blades elliptic or long elliptic,4-10×1.6-3.2 cm,denticulate,apex acuminate or long acuminate,with 6-10 pairs of secondary nerves; stipules subulote,2 mm long; staminate cymes 1.6-2.8 cm broad,with peduncles 1.5-2.5 cm long; pistillate flower’s tepals 5……………………………………………………………………………25.羽脈赤車P.incisoserrata
9.Leaf blades long elliptic,oblanceolate or lanceolate,1.5-10×0.5-2.6 cm,denticulate,apex long acuminate or cuspidate; stipules linear or subulate,1-10 mm long; staminate cymes 2 cm broad,with peduncles up to 2.4 cm long; pistillate flower’s tepals 4 or 5………………………………………………………………………………………………26.兩廣赤車P.ronganensis
8.Achenes longitudinally lineolate and minutely tuberculate; stems above and leaf blades on both surfaces densely puberulous; leaf blades larger,oblanceolate,13-17×2-4.8 cm,denticulate,apex long acuminate,secondary nerves ca.5 pairs,with the lowest ones much longer than the upper ones; stipules narrowly lanceolate or linear,9-11.5×2-2.8 mm; pistillate flower’s tepals 5(系4.柔毛赤車系ser.Mollissimae)……………………………………………………………………31.柔毛赤車P.mollissima
7.Achenes smooth.
10.Reduced leaves present,0.6-1.5 cm long; stems and leaves glabrous; normal leave’s blade oblong,0.5-8×1-2 cm,denticulate,apex long acuminate,secondary nerves 4-8 pairs; stipuls subulate,1.8-2.8 mm long; staminate cymes ca.5 mm broad,with peduncles 1-1.5 cm long; pistillate flower’s tepals 5,abaxially below apex with short projections(系2.光果赤車系ser.Leiocarpae)………………………………………………………………………………………………27.光果赤車P.leiocarpa
10.Reduced leaves absent(系3.短角赤車系ser.Brachycerates).
11.Stem branches and leaves abaxially puberulous; leaf blades long elliptic,up to 5 cm long,denticulate,apex acuminate,secondary nerves 5-6 pairs; stipules subulate,2.5-3.5 mm long; staminate cymes 3-8 mm broad,with peduncles 0.7-1.2 cm long; pistillate flower’s tepals 5,abaxially pilose and below apex shortly corniculate…………………………28.短角赤車P.brachyceras
11.Stems and leaves glabrous; leaf blades long elliptic,up to 7-8 cm long; pistillate flower’s tepals 3-4,glabrous,lacking any projections.
12.Leaf blades 2.6-7×0.9-2.4 cm,apex long acuminate,secondary nerves 6-8 pairs; stipules subulate,1.5-2.2 mm long; pistillate cymes ca.2.2 mm in diam.,2-3 times branched,ca.7-flowered,with peduncles 0.2 mm long; pistillate flower shortly pedicellate,with 3-4 tepals; achenes ellipsoid,0.9 mm long………………………………29.龍州赤車P.longzhouensis
12.Leaf blades 4-8×1.8-2.6 cm,apex long acuminate,secondary nerves 4-5 at leaf narrow side and 6-7 at broad side; stipules subulate,1.8-3 mm long,pistillate cymes sessile,ca.2.5 mm in diam.,with very short branches,densely multi-flowered; pistillate flower sessile,with 3 tepals; achenes unknown……………………………………30.東蘭赤車P.donglanensis
5.Pistillate flower’s tepals( 3-) 4-5,of them( 1-) 2-3 larger,linear-navicular,abaxially below apex long corniculate(組2.赤車組,赤車亞組sect.Pellionia subsect.Radicantes).
13.Leaves semi-triplinerved; achenes tuberculate.
14.Reduced leaves present; stems prostrate,densely retrorse-hirsute( hairs 1-2.2 mm long) and densely brown-puncticulate; leaf blades obovate,1.2-3.2×0.7-1.7 cm,crenate,apex rounded-obtuse,secondary nerves 1 at leaf narrow side and 2 at broad side; stipules ovate or lanceolate,up to 2-2.5×1 mm; pistillate cymes subglobose,ca.2.5 mm in diam.,densely multi-flowered; pistillate flower’s tepals 5; achenes long ellipsoid,ca.0.8 mm long(系1.婆婆納赤車系ser.Veronicoides)……………………………………………………………………………………………………………4.婆婆納赤車P.veronicoides
14.Reduced leaves absent(系2.赤車系ser.Radicantes).
15.Leaf blades with peltate base,oblong,5-15×1.6-5 cm,crenate or subentire,glabrous,apex long acuminate; stipules subulate,ca.3.5 mm long; stems glabrous; staminate cymes 0.8-2.2 cm long,with pednucles 0.5-1.8 cm long; pistillate flower’s tepals 5………………………………………………………………………………………………………7.綠赤車P.viridis
16.Leaf base symmetrical……………………………………………………………………………7a.模式變種var.viridis
16.Leaf base asymmetrical,obtuse at leaf narrow side,auriculate at broad side…………7b.斜基綠赤車var.basiinaequalis
15.Leaf base not peltate.
17.Stipules lanceolate or triangular,1-3 mm broad; stems with dense hairs.
18.Leaf blades narrowly ovate,10-13×2.8-3.5 cm,crenate,apex long acuminate; stipules broadly lanceolate,11×3 mm; staminate cymes ca.1.4 cm broad,with peduncles 12 cm long……………………………………5.長梗赤車P.longipedunculata
18.Leaf blades elliptic,3.5-7.5×1.1-3.3 cm,denticulate,apex acute or shortly acuminate; stipules triangular,3.2-6.5×1-1.8 mm; staminate cymes 0.8-1.5 cm broad,with peduncles 1-5.5 cm long……………………6.曲毛赤車P.retrohispida
17.Stipules smaller,subulate,0.1-0.3 mm broad.
19.Stems glabrous or very shortly puberulous,with hairs ca.0.1 mm long.
20.Leaf blades 0.9-2×0.4-1.4 cm,broadly elliptic,broadly obovate or suborbicular,apex obtuse or rounded; stipules subulate,1-2 mm long; staminate cymes 0.8-2 cm long,with peduncles 0.5-1.4 cm long; pistillate tepals usualy glabrous…………………………………………………………………………………………………18.小赤車P.minima
20.Leaf blades larger,up to 5-15 cm long,apex acute,acuminate,or caudate.
21.Apical leaves narrowly lanceolate or sublinear,margins undulate…… 12b.狹葉赤車P.subundulata var.a(chǎn)ngustifolia
21.Apical leaves ovate,elliptic,or oblong.
22.Leaf margin undulate or obtusely crenate.
23.Leaf blades elliptic,1-8×0.6-3.5 cm,apex acute,rarely shortly acuminate,with secondary nerves 2 at leaf narrow side and 3 at broad side; stipules subulate,0.8-1×0.1 mm; pistillate flower’s tepals 5,glabrous……………………………………………………………………………………………12a.波緣赤車P.subundulata var.subundulata
23.Leaf blades larger,long elliptic or narrowly obovate,5-15.5×2-6.5 cm,apex cuspidate or acuminate,with secondary nerves 2-3 at leaf narrow side and 4-6 at broad side; stipules larger,subulate,3.5-7 mm long; pistillate flower’s tepals 5,with sparse short hairs…………………………………………19a.滇南赤車P.paucidentata var.paucidentata
22.Leaf margin acutely dentate or denticulate.
24.Leaf blades with cystoliths 0.4-0.7 mm long,conspicuous on adaxial and abaxial surfaces………………………………………………………………………19b.海南赤車P.paucidentata var.hainanica
24.Leaf blades with cystoliths 0.1-0.4 mm long,conspicuous only on adaxial surface.
25.Stems below repent ……………………………………………………………………………13.赤車P.radicans
26.Leaf blades( 1.2-) 2.4-5(-8)×0.9-2(-2.7) cm,ovate or lanceolate,denticulate,apex shortly to long acuminate,secondary nerves 2-3 at leaf narrow side and 3-4 at broad side; stipules subulate,1-4.2×0.2 mm; staminate cymes 1-5(-8) cm long,with peduncles 0.4-3.5(-7) cm long; pistillate flower’s tepals 5… 13a.模式變種var.radicans
26.Leaf blades larger,up to 9-13×3.5-5 cm………………………………………………13b.長莖赤車var.grandis
25.Stems erect,glabrous,rarely with hairs ca.0.1 mm long,usually simple; leaf blades glabrous,obliquely elliptic,ovate or oblong,( 2.5-) 5-10(-14)×( 1.2) 2-4 cm,denticulate,apex acuminate or long acuminate,secondary nerves 1-3 at leaf narrow side and 3-4 at broad side; stipules subulate 1-3 mm long; staminate cymes 1-3.5 cm broad,with peduncles 1-8 cm long………………………………………………………………………14.異被赤車P.heteroloba
19.Stems with hairs( 0.2-) 0.3-1 mm long.
27.Leaf blades 0.5-3.2×0.4-2 cm,elliptic or obovate,denticulate,apex obtuse or rounded; stipules subulate,1.2-2 mm long; staminate cymes 0.8-1.5 cm broad,with peduncles 0.8-4 cm long; pistillate flower’s tepals 5,with spare short hairs……………………………………………………………………………………………17.短葉赤車P.brevifolia
27.Leaf blades larger,up to 4.5-10 cm long,apex acute,acuminate or cuspidate.
28.Leaf apex acuminate,caudate or cuspidate.
29.Subshrubs with ramose stems; leaf blades oblanceolate or narrowly oblong,3.2-8.5×( 0.7) 1.3-3.2 cm,denticulate,apex acuminate or cuspidate; stipules subulate,1.5-3 mm long; staminate cymes up to 4.5 cm long,with peduncles 0.3-3.6 cm long; pistillate flower’s tepals 4-5,glabrous………………………………………………………8.蔓赤車P.scabra
29.Peremial herbs with usually simple stems; pistillate tepals pilose.
30.Stems densely retrorse-strigillose; leaf blades elliptic,3-7.4×1.2-3 cm,acutely denticulate,apex acuminate; stipules subulate,ca.3 mm long; staminate cymes 3 cm long,with peduncles 8 cm long; pistillate flower’s tepals 5………………………………………………………………………………………………9.富寧赤車P.funingensis
30.Stems sparsely spreading-puberulous; leaf blades larger,5-15.5×2-6.5 cm,obtusely crenate; pistillate flower’s tepals 4-5…………………………………………………………………19a.滇南赤車P.paucidentata var.paucidentata
28.Leaf apex acute or shortly acuminate.
31.Stems encrusted by dense minute lumps of whitish lime,and leaves adaxially and abaxially encrusted by a thin layer of lime; staminate cymes singly borne on leafless nodes of lower part of branch,sessile,2.5-4 mm in diam.; staminate flowers tetramerous; leaf blades elliptic,2.4-7×1.4-3.5 cm,crenate,apex acute or shortly acuminate…………………………………………………………………………………………………………16.灰被赤車P.calcifera
31.Stems and leaves without lime; staminate cymes pedunculate; staminate flowers pentamerous.
32.Stems dark-brown-puncticulate and puberulous; pistillate flower’s tepals 3,glabrous,all abaxially below apex usually long corniculate; leaf blades elliptic,or obovate,2.5-5.5×1.5-2.5 cm,dentitulate,apex acute or shortly acuminate; stipules subulate,1-2.5 mm long………………………………………………………15.三被赤車P.tritepala
32.Stems not puncticulate; pistillate flower’s tepals 4-5,of them 2 larger,linear-navicular,abiaxially below apex long
corniculate.
33.Stems densely retrorse-strigillose,with hairs white,straight,0.4-0.8 mm long; leaf blades elliptic,0.5-5.6×0.9-2 cm,acutely dentate,apex acute or shortly acuminate; stipules subulate,1-2 mm long; staminate cymes 0.5-1.2 cm broad,with peduncles 0.8-2.2 cm long; pistillate flowers tepals 5,pilose…………10.尖齒赤車P.a(chǎn)cutidentata
33.Stems densely hirtellous,with hairs fulvous,curved,0.15-0.4 mm long; leaf blades ovate or elliptic,2.8-4.5×2-3.4 cm,crenate,apex acute; stipules subulate,ca.6 mm long; pistillate flower’s tepals 4-5,glabrous or pilose………………………………………………………………………………………11.硬毛赤車P.crispulihirtella
13.Leaves penninerved.
34.Achenes tuberculate.
35.Stems longitudinally winged; reduced leaves prescent; leaf blades broadly ovate,elliptic or obovate,8-18×4.4-8.3 cm,undulate-crenate,apex acuminate,secondary nerves 5-8 pairs; stipules narrowly triangular,6-9×1.5-2 mm; pistillate flower’s tepals 5(系3.翅莖赤車系ser.Caulialatae)……………………………………………………20.翅莖赤車P.caulialata
35.Stems not winged; reduced leaves abscent(系4.南亞赤車系ser.Griffithianae).
36.Stems densely hispid,with hairs 0.5-2 mm long,spreading or retrorse; cystoliths of leaves absent or present,puncticulate; leaf blads long elliptic or oblong-oblanceolate,6-14(-18)×2.4-5(-6) cm,crenate,apex long acuminate,secondary nerves ca.5 pairs; stipules subulate,4 mm long; staminate cymes 0.5-5.5 cm broad,with peduncles 1-8 cm long; pistillate flower’s tepals 5,pilose…………………………………………………………………………………22.華南赤車P.grijsii
36.Stems glabrous,strigillose,sparsely puberulous or villous; leave’s cystoliths either absent or present,bacilliform.
37.Leaves lacking cystoliths; stems glabrous or near apex sparsely puberulous; leaf blades oblanceolate,5.5-12.5×1.8-3.5 cm,dentate,apex cuspidate,secondary nerves 6-7 pairs; stipules subulate,4-6 mm long; staminate cymes 1-2 cm broad,with peduncles 0.7-1.5 cm long; pistillate flower’s tepals 4,glabrous……………23.云南赤車P.yunnanensis
37.Leaves with bacilliform cystoliths.
38.Leaf apex cuspidate,with cusps narrowly linear and entire; pistillate tepals glabrous… 23.云南赤車P.yunnanensis
38.Leaf apex acuminate or long acuminate; pistillate tepals pilose.
39.Leaf base obliquely broadly cuneate………………………………19a.滇南赤車P.paucidentata var.paucidentata
39.Leaf base obtuse at leaf narrow side and auriculate at broad side; leaf blades oblong or narrowly obovate,6-15×2-6 cm,denticulate,apex acuminate or long acuminate; staminate cymes 1-3 cm broad,with peduncles 2-8 cm long; pistillate flower’s tepals 5,pilose…………………………………………………………………21.滇南赤車P.griffithiana
40.Stems densely brownish-strigillose,with hairs 0.1-0.2 mm long; leaf blades abaxially on nerves strigose,secondary nerves 5-6 at leaf narrow side and 6-8 at broad side; stipules narrowly triangular-linear,5.5-12×0.7-1.2 mm………………………………………………………………………………………21a.模式變種var.giriffithiana
40.Stems not densely brownish-strigillose; stipules smaller,subulate,2-4×0.1-0.2mm.
41.Stems glabrous; leaf blades abaxially glabrous,secondary nerves 4 at leaf narrow side and 5-6 at broad side……………………………………………………………………………21b.無毛莖南亞赤車var.glabricaulis
41.Stems hairy.
42.Stems with sparse white long hairs( 0.5-) 1-2 mm long; leaf blades abaxially glabrous,secondary herves 3-4 at leaf narrow side and 4-6 at broad side……………………………………………21c.疏毛南亞赤車var.laxivillosa
42.Stems only near apex with retrorse,white hairs 0.2-0.4 mm long; leaf blades abaxially sparsely puberulous,secondary nerves ca.4 pairs………………………………………………………………21d.少脈南亞赤車var.paucinervis
34.Achenes lineolate; leaf blades obovate-oblong or long elliptic,14-25×5-8.2 cm,crenate,apex acuminate,secondary nerves 9-12 at leaf narrow side and 10-14 at broad side; stipules lanceolate-linear or subulate,6-9×0.5-0.6 mm; staminate cymes 1.5-3 cm broad,with peduncles 1.6-4.6 cm long; pistillate flower’s tepals 5,glabrous(系5.大葉赤車系ser.Macrophyllae)………………………………………………………………………………………………………24.大葉赤車P.macropohylla
1.Pistillate inflorescences capitate,with receptacles and involucres; achenes globose,longitudinally 5-ribbed and minutely tuberculate; stems ramose,branches puberulous; leaf blades long elliptic,up to 2.5×0.9 cm,denticulate,trinerved or semi-triplinerved,apex cuspidate; pistillate flower’s tepals 5,all abaxially below apex shortly corniculate(組4.頭序赤車組sect.Elatostematopsis)……………………………………………………………………………………………………………32.頭序赤車P.cephaloidea
組1.全緣赤車組sect.Pileoides W.T.Wang in Wedd。
Leaves long or slightly long petiolate,with entire margins,trinerved with the lateral basal nerve at leaf narrow side extending upwards nearly to leaf apex.Reduced leaves present.Pistillate inflorescences cymose. Bull.Bot.Lab.N.-E.Forest.Inst.6: 50.1980; et in Fl.Reip.Pop.Sin.23( 2) : 163.1995; Yahara in J.Fac.Sci.Univ.Tokyo.Sect.3,Bot.13( 4) : 484.1984; X.W.Li in F.Yunnan.7: 241.1997.Type: P.heyneanaPistillate flower’s tepals 5,abaxially lacking any projections or below apex shortly corniculate.Achenes tuberculate.
圖2 A-B.全緣赤車 A.葉; B.宿存雌花被和瘦果(據(jù)王啟無77557)。C-E.長柄赤車 C.葉; D.宿存雌花被; E.瘦果(據(jù)C.I.Lei 1028)。F-H.吐煙花 F.葉; G.宿存雌花被; H.瘦果(據(jù)王啟無78046)。I-K.婆婆納赤車 I.二葉; J.宿存雌花被和瘦果; K.瘦果(據(jù)王啟無88690)。L-N.曲毛赤車 L.葉; M.宿存雌花被; N.瘦果(據(jù)邵青,段林東70)。O-Q.綠赤車 O.葉及雌聚傘花序; P.雌花和苞片; Q.雄花(仿王文采,1995)。Fig.2 A-B.Pellionia heyneana A.Leaf; B.Persistent pistillate perianth and achene( from C.W.Wang 77557).C-E.P.tsoongii ssp.tsoongii C.Leaf; D.Persistent pistillate perianth; E.Achene( from C.I.Lei 1028).F-H.P.repens F.Leaf,G.Persistent perianth; H.A-chene( from C.W.Wang 78046).I-K.P.veronicoides I.Two leaves; J.Persistent pistillate perianth and achene; K.Achene( from C.W.Wang 88690).L-N.P.retrohispida L.Leaf; M.Persistent perianth; N.Achene( from Q.Shao&L.D.Duan 70).O-Q.P.viridis var.viridis O.Leaf and pistillate cyme; P.Pistillate flower and bract; Q.Staminate flower( after Wang,1995).
4種,我國有2種。
1.全緣赤車圖2: A-B
Pellionia heyneana Wedd.in Arch.Mus.Hist.Nat.Paris 9: 287,332,t.5,1856; et in DC.Prodr.16( 1) : 169.1869 ; Hook f.Fl.Brit.Ind.5: 561.1888; Gagnep.In Lecomte,F(xiàn)l.Gén.Indo-Chin.5: 900.1929; W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 50,1980; et in Fl.Reip.Pop.Sin.23 ( 2) : 164,pl.36: 5-6.1995; X.W.Li in Fl.Yunnan.7: 241,pl.61: 1-6,1997; Q.Lin et al in Fl.China 5: 123.2003; H.N.Qin&Y.Liu,Checklist Vasc.Pl.Guangxi 237.2010.——Procris heyneana Wall.Cat.N.7272.1828,nom.nud.——Elatostema heyneanum ( Wedd.) Hall.f.in Ann.Jard.Bot.Bruitenz.13: 316.1896; H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83( 2) : 22.1936.Holotype: India Kanara,Wallich 7272( K.non vidi).Fig.2: A-B
分布:我國云南南部和東南部、廣西北部;柬埔寨;印度南部;斯里蘭卡。
標(biāo)本登錄*:云南景洪,王啟無77557;馬關(guān),中蘇考察團(tuán)3271,3273;河口,中蘇考察團(tuán)2386。廣西天峨,北京隊891355。
2.長柄赤車圖2: C-E
Pellionia tsoongii ( Merr.) Merr.In Lingnan Sci.J.6 ( 4) : 325.1928; Fl.Hainan.2: 405.1965; Iconogr.Corm.Sin.1: 513,fig.1025.1972; W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 51.1980; in Fl.Reip.Pop.Sin.23( 2) : 164,pl.36: 4.1995,p.p.; et in Fl.Guangxi 2: 844.2005; Y.Z.Ruan in Pl.Guangdong 6: 78.2005——Polychroa tsoongii Merr.in Philip.J.Sci.21.493.1922.——Elatostema tsoongii ( Merr.) H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83( 2) : 21.1936.Isotype:廣西東興,1918-06-22,鐘觀光1870( PE)。Fig.2: C-E
P.cristulata Gagnep.in Bull.Soc.Bot.France 75: 920.1928; et in Lecomte,F(xiàn)l.Gen.Indo-Chin.5: 901.1929 ( fide H.Schr?ter,1936).Holotype: Vietnam Annam,Poilane 10761( P.non vidi).*
P.latifolia auct.non( Bl.) Boerl.: Q.Lin et al in Fl.China 5: 123.2003; H.N.Qin&Y.Liu,Checklist Vasc.Pl.Guangxi 237.2010.
2a.ssp.tsoongii
分布:我國云南西南、廣西西南部、海南;越南;柬埔寨。
標(biāo)本登錄:云南滄源,王啟無73188。廣西扶綏,陳少卿12021;巴馬,李蔭坤01243。海南澄邁,Tsang W.T.17358,Lei C.I.1020,1028。
2b.無毛長柄赤車(亞種)
ssp.subglabra X.W.Li in Fl.Yunnan.7: 242,pl.61: 7-11.1997.Type:云南瀾滄,alt.1 300 m,1936-05,王啟無76619( holotype,KUN; isotype,PE) ;景洪,王啟無79407( paratypes,KUN,PE) ;滄源,李延輝11524,12248( paratypes,KUN) ;耿馬,李延輝2319 ( paratype,KUN)。
P.subpeltata Gagnep.in Bull.Soc.Bot.France 75: 925.1928; et in Lecomte,F(xiàn)l.Gén.Indo-Chine 5: 900.1929.Syntypes: Vitnam Balansa 584&M.Colane 2929( P,non vidi)
分布:我國云南西南及南部、廣西西南部;越南。
標(biāo)本登錄:我國廣西那坡,覃海寧等279,334。越南涼山,1965-01-17,中越考察隊1531。
全緣赤車Pellionia heyneana和P.latifolia( Bl.) Boerl.的葉相似,基部斜圓形或斜淺心形,葉寬側(cè)基部不呈耳形或呈不明顯耳形,而長柄赤車P.tsoongii的葉基部兩側(cè)強(qiáng)烈不對稱,在寬側(cè)呈明顯耳形,耳垂部分長0.8-2.7 cm,這是個穩(wěn)定同時便于利用的形態(tài)區(qū)別特征。H.Schr?ter( 1936)在其Elatostema專著中用此特征在檢索表中區(qū)別P.tsoongii和前二種。此外,Yahara( 1984)在其有關(guān)泰國Pellionia的文章中討論到P.latifolia的近緣種時也指出其與P.tsoongii在葉基部的上述區(qū)別。想不到Lin et al ( 2003)竟將P.tsoongii歸并于P.latifolia,造成一個分類學(xué)混淆,為澄清此一混淆,本文將P.tsoongii的種級地位予以恢復(fù)。
組2.赤車組sect.Pellionia.W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 165.1980; et in Fl.Reip.Pop.Sin.23( 2) : 165.1995; Yahara in J.Fac.Sci.Univ.Tokyo.Sect.3,Bot.13( 4) : 485.1984; X.W.Li in Fl.Yunnan.7: 244.1997.Type: P.elatostemoides Gaudich.[=P.sinuata( Bl.) Boerl.]
Leaves shortly petiolate or sessile,usually with
*本文中記錄的所有標(biāo)本均保存于PE中。dentate margins,semi-triplinerved,with lateral basal nerve at leaf narrow side extending upwards to or slightlysurpassingtheleafmiddle,rarely penninerved.Reduced leaves present or absent( in most species of China absent ).Pistillate inflorescences cymose.Pistillate flower’s tepals 5,rarely 4 or 3,usually ( 1-) 2-3 larger,navicular,abaxially below apex long corniculate,rarely lacking any projections or below apex shortly corniculate ( P.repens).Achenes tubeculate,rarely lineolate( P.macrophylla).
約有62種。我國有22種,根據(jù)雌花被片、退化葉、脈序、托葉和瘦果等器官形態(tài)特征被劃分為2亞組5系。
亞組1.吐煙花亞組subsect.Repentes( W.T.Wang) W.T.Wang,st.et comb.nov.——ser.Repentes W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 51.1980,et in Fl.Reip.Sin.23( 2) : 165.1995; Yahara in J.Fac.Sci.Univ.Tokyo.Sect.3,Bot.13 ( 4) : 485.1984.Type: P.repens ( Lour.) Merr。
Pistillate flower’s tepals 5,slightly unequal in size,lacking any projections or abaxially below apex shortly corniculate.Reduced leaves present.Leaves semi-triplinerved; stipules triangular.Achenes tuberculate.
1種。
3.吐煙花圖2: F-H
Pellionia repens ( Lour.) Merr.in Lingnan Sci.J.6( 4) : 326.1928; F.C.How,F(xiàn)l.Guangzhou.404.1956; Fl.Hainan.2: 406,fig.492.1965; W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 82.1980; Yahara in J.Fac.Sci.Univ.Tokyo,Sect.3,Bot.13( 4) : 484.1984; S.S.Chang in Fl.Guizhou.4: 40,pl.14: 1-2.1989; W.T.Wang in Fl.Reip.Pop.Sin.23( 2) : 165,pl.38: 5.1995; X.W.Li in Fl.Yunnan.7: 244.1997.; Q.Lin et al in Fl.China 5: 124.2003; Y.Z.Ruan in Fl.Guangdong 6: 78,fig.28.2005; F.W.Xing&F.G.Wang,F(xiàn)l.Macao 154.2005; Y.M.Shui&W.H.Chen,Seed Pl.Karst.Reg China 1: 103,fig.256.2006.——Polychroa repens Lour.Fl.Cochinch.559.1790.——Elatostema repens ( Lour.) Hall.f.in Ann.Jard.Bot.Buitenz.13: 316.1986; H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83( 2) : 25.1936; Backer&V.D.Brink,F(xiàn)l.Java 2: 42.1965.No type specimen designated.Fig.2: F-H
P.pulchra Boerl.In Gard.Chron,n.ser.,18( 2) : 104,1873.——Elatostema pulchrum ( Boerl.) Hall.f.1.c.——E.repens( Lour.) Hall.f.var.pulchrum ( Boerl.) H.Schr?ter l.c.26.
P.daveauana Boerl.in Gard.Chron.,n.ser.,14 ( 2) : 262.1880; Hook.f.Fl.Brit.Ind.5: 560.1888; Ridley,F(xiàn)l.Malay.Pen.3: 362.1924; Gagnep.in Lecomte,F(xiàn)l.Gén.Indo-Chin.5: 902.1929.
分布:云南南部和東南部、海南;自緬甸向東達(dá)我國云南、海南和中南半島,向南經(jīng)馬來半島達(dá)印度尼西亞爪哇。
標(biāo)本登錄:云南勐海,王啟無74644,76140;景洪,王啟無77554,78046;勐侖,熱帶植物園,李慶華898;河口,采集人不明2448。海南保亭,侯寬昭72546,73338;尖峰嶺,236隊3800;白沙,金效華5048;崖縣,黃志33063;臨高,曾懷德15761。
亞組2.赤車亞組subsect.Radicantes W.T.Wang,subsect.nov.Type: P.radicans( Sieb.&Zucc.) Wedd。
A subsect.Repentibus( W.T.Wang) W.T.Wang differt floris pistillati tepalis ( 3-4-) 5 valde inaequimagnis( 1-) 2-3 majoribus navicularibus dorso sub apice longe corniculatis.
Pistillate flower’s tepals 5,rarely 4 or 3,strongly unequal in size,( 1-) 2-3 larger,navicular,abaxially below apex long corniculate.Reduced leaves absent,rarely present.Leaves semi-triplinerved,rarely penninerved; stipules subulate,rarely ovate,triangular or lanceolate.Achenes tuberculate,rarely lineolate.
約有59種,我國約有21種。
系1.婆婆納赤車系ser.Veronicoides W.T.Wang,ser.nov.Type: P.veronicoides Gagnep。
Series nova haec est arcte affinis ser.Radicantibus W.T.Wang,a qua foliis reductis praesentibus differt.——Reduced leaves present.Normal leaves semitriplinerved; stipules lanceolate or ovate.Pistillate flower’s tepals 5,of them 2 larger,linear-navicular,abaxially below apex long corniculate.Achenes tuberculate.
1種。
4.婆婆納赤車圖2: I-K
Pellionia veronicoides Gagnep.in Bull.Sec.Bot.France 75: 927.1928; et in Lecomte,F(xiàn)l.Gén.Indo-Chin.5: 896.1929.——Elatostemaveronicoides ( Gagnep.) H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83( 3) : 28,tab.10.1936.Holotype: Vietnam: Tonkin: Cho-bo,1887-11-16,Balansa 2551 (♂,P,non vidi).Fig.2: I-K
Supplementary description to the original one,with the infructescences,persistent pistillate tepals and achenes described for the first time.: Infructescences singly axillary,shortly pedunculate,subglobose,ca.2.5 mm in diam.,densely multi-flowered; bracts numerous,whitish,narrowly triangular-linear or narrowly linear,1-2× 0.1-0.2 mm,margin nearly entire or 1-2-denticulate,glabrous.Persistent pistillate tepals 5,unequal in size,2 larger,linear-navicular,ca.1 mm long,abaxially below apex long corniculate,with horn-like projections ca.1.4 mm long,smaller tepals flat,lanceolate-linear,ca.1×0.1 mm,glabrous,lacking any projections.Achenes brown,long ellipsoid,ca.0.8 mm long,tuberculate.
本種的holotype為雄株標(biāo)本,所以在原始描述中缺雌花序,雌花和瘦果的描述。下引的采自云南西疇的本種標(biāo)本是由著名華裔植物生態(tài)學(xué)家王啟無先生于七十多年前采到,在2003年才由林祁博士定名。此號標(biāo)本為結(jié)果雌株,現(xiàn)在將果序,宿存雌花被片和瘦果描述補(bǔ)充如上。
分布:我國云南東南部;越南北部。
標(biāo)本登錄:云南西疇,Ban-loun,alt.800 m,山谷林中,草本植物平臥地面,1940-04-18,王啟無( C.W.Wang) 88690。
系2.赤車系ser.Radicantes W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 52.1980; et in Fl.Reip.Pop.Sin.23( 2) : 167.1995.Type: P.radicans ( Sieb.&Zucc.) Wedd.
Ser.Virides W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 62.1980; et in Fl.Reip.Pop.Sin.23 ( 2) : 170.1995.Type: P.viridis C.H.Wright.
Reduced leaves absent.Leaves semi-triplinerved; stipules usually subulate,rarely ovate or lanceoltae.Pistillate flower’s tepals 5,rarely 4 or 3,of them( 1-) 2-3 larger,linear,navicular,abaxially below apex long corniculate.Achenes tuberulate.
我國約有20種。
5.長梗赤車圖3: A-B
Pellionia longipedunculata W.T.Wang in Bull.Bot.Res.Harbin 2( 1) : 1,pl.1: 1-2.1982; et in Fl.Reip.Pop.Sin.23( 2) : 167,pl.51: 1-2.1995; Q.Lin et al in Fl.China 5: 124.2003,p.p.excl.syn.P.subundulata var.a(chǎn)ngustifolia W.T.Wang; W.T.Wang in Fl.Guangxi 2: 844,pl.343: 1.2005.Holotype:廣西上思,十萬大山,alt.600 m,1944-04-19,陳少卿4918(♂,IBSC)。Fig.3: A-B
分布:廣西十萬大山特有種。
本種的主要特征為葉片狹卵形,托葉較大,寬披針形,長約11 mm,寬3mm,雄花序梗較長(約12 cm)。Lin et al( 2003)將狹葉赤車P.subundulata var.a(chǎn)ngustifolia W.T.Wang歸并于P.longipedunculata。但狹葉赤車的葉呈披針形或條狀披針形,托葉呈鉆形,長2.5-3.5 mm,寬0.4-0.8 mm。本種的雄花序尚未發(fā)現(xiàn)。從上述情況可見長梗赤車與狹葉赤車之間存在明顯區(qū)別,說明二者不是同一種植物,也說明上述歸并是錯誤的。
6.曲毛赤車圖3: L-N
Pellionia retrohispida W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 54,pl.1: 4-5.1980; et in Bull.Bot.Res.Harbin 3( 3) : 56.1986; Y.Q.He in Fl.Zhejiang 2: 111,fig.2048.1992; W.T.Wang in Fl.Reip.Pop.Sin.23( 2) : 167,pl.50: 11.1995; B.G.Li in Fl.Hunan 2: 288.2000; Q.Lin et al in Fl.China 5: 124.2003; C.X.Yang et al Keys Vasc.Pl.Chongqing 150.2009.Holotype:重慶南川,小河區(qū),alt.1 550 m,1957-06-09,熊濟(jì)華,周子林91324 ( PE).Paratypes:湖南龍山,劉翰林1741( PE)。江西奉新,劉守爐等1292( NAS)。Fig.3: L-N
Elatostema hirticaule W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.7: 54.1980.Holotype:四川都江堰,汪發(fā)瓚20538( PE)。
E.hunanense W.T.Wang 1.c.Holotype:湖南雪峰山,李澤棠1685( PE)。
分布:甘肅南部、四川、重慶、湖北西南、貴州、湖南、江西、浙江、福建。
標(biāo)本登錄:甘肅文縣,白水江隊4377。四川青城山,趙志成87-83。重慶南川,劉正宇4820,13361。湖北鶴峰,李鴻均8089。貴州桐梓,劉正宇70521;正安,劉正宇20522;都勻,溫放070430A;荔波,邵青,段林東59。湖南永順,北京隊01126;芷江,武陵山隊1955;雪峰山,李澤棠1685;武岡,鄧云飛10235;宜章,肖百中3930;衡山,李明紅6。江西上猶,江西隊71-630;修水,譚策銘94304。浙江遂昌,植物資源隊26548。福建建寧,陳家瑞等64。
7.綠赤車圖2: O-Q
Pellionia viridis C.H.Wright in J.Linn.Soc.Bot.26: 481.1899; Iconogr.Corm.Sin.1: 513,fig.1026.1972; Fl.Hupeh.1: 171,fig.222.1976; W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 62.1980; et in Fl.Reip.Pop.Sin.23( 2) : 179,pl.36: 1-3.1995; X.W.Li in Fl.Yunnan.7: 244,pl.62: 1-2.1997; Q.Lin et al in Fl.China 5: 126.2003 ; C.X.Yang et al Keys Vasc.Pl.Chongqing 150.2009——Elatostema viride ( C.H.Wright) Hand.-Mazz.Symb.Sin.7: 143.1929; H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83 ( 2) : 84.1936.Synteypes:湖北宜昌,A.Henry 4098 ( K,non vidi; photo PE).四川Mt.Omei,F(xiàn)aber 439,440( K,non vidi).Fig.2: O-Q
7a.var.viridis
分布:云南東北部、四川、重慶、湖北西部。
標(biāo)本登錄:云南綏江,孫必興0520。四川峨眉山,方文培2594,17487,汪發(fā)瓚23179,杜大華405;雅安,Harry Smith 10037;屏山,俞德浚2979。
7b.斜基綠赤車(變種)
var.basiinaequalis W.T.Wang in Bull.Bot.Lab.N.-E.Foest.Inst.6: 62.1980; et in Fl.Reip.Pop.Sin.23( 2) : 180.1995; Q.Lin et al in Fl.China 5: 126.2003.Holotype:四川筠連,1977-06-30,四川中藥所824( IMC).Paratypes:四川長寧,四川中藥所138 ( IMC) ;敍永,四川中藥所764( IMC)。
分布:四川南部。
8.蔓赤車圖3: C-D
Pellionia scabra Benth.Fl.Hongk.330.1861; Wedd.in DC.Prodr.16( 1) : 166.1869; C.H.Wright in J.Lin.Soc.Bot.26: 481.1899; Gagnep.in Lecomte,F(xiàn)l.Gen.Indo-Chin.5: 896.1929; F.C.How,F(xiàn)l.Guangzhou 404.1956; Fl.Hainan.2: 407.1965; Ohwi,F(xiàn)L.Japan,rev.et.,390.1965; Iconogr.Corm.Sin.1: 514,fig.1027.1972; Fl.Hupeh.1: 172.1976; T.S.Liu&W.D.Huang in Fl.Taiwan 2: 205.1976; W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 52.1980; C.J.Zeng in Fl.Fujian.1: 466.1982; Fl.Jiangsu 2: 82.1982; Z.W.Xuo in Fl.Anhui 2: 123.1986; S.S.Chang et al in Fl.Guizhou.4: 42.1989; Y.Q.He in Fl.Zhejiang 2: 112,fig.2-149.1992; W.T.Wang in Fl.Reip.Pop.Sin.23( 2) : 168,pl.37: 5,44: 3-4.1995; Yang,Shih&Liu in Fl.Taiwan,2nd ed.,2: 231,pl.115.1996; Y.E.Lee,F(xiàn)l.Korea 88,fig.230.1996; B.G.Li in Fl.Hunan 2: 289,fig.2-214.2000; Q.Lin et al in Fl.China 5: 124.2003,p.p.excl.syn.P.cephaloidea W.T.Wang; Y.Z.Ruan in Fl.Guangdong 6: 79.fig.29.2005; W.T.Wang in Fl.Guangxi 2: 844.2005; Z.X.Peng&G.L.Zhang in Fl.Gansu 2: 237,pl.51: 1.2005; Tateishi in Iwatsuki et al Fl.Japan 2a: 100,2006; Q.M.Hu in Fl.Hong Kong 1: 124.2007; C.X.Yang et al Keys Vasc.Pl.Chongqing 150.2009; Z.R.Xu in Fl.Jiangsu 2: 168,fig.2-272.2013.——Elatostema scabrum ( Benth.) Hall.f.in Ann.Jard.Bott.Buitenz.13: 316.1896; H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83 ( 2) : 82,tab.25.1936.——Polychora scabra ( Benth.) Hu in J.Arn.Arb.5: 228.1924.Syntypes香港,Wiford 226,Hance 1095,Harland 326,Wright 447( K.non vidi).
P.a(chǎn)risanensis Hayata var.caudatifolia Yamamoto,Suppl.Ic.Pl.Formos.1.12.1925.Holoeype:臺灣in Monte Arisan,B.Hayata s.n.( non vidi).
P.scabra subvar.pedunculata Yamamoto,1.c.,17.1925.Syntypes:臺灣S.Ito s.n,B.Hayata s.n.( non vidi).
Elatostema pelliniifolium W.T.Wang in Bull.Bot.Res.Harbin 2( 1) : 12.1982.Holotype:重慶北碚,縉云山,約1940,L.Y.Lin 4138( IBSC).
分布:我國云南東南部、廣西、海南、廣東、福建、臺灣、浙江、江西、湖南、貴州、四川、甘肅南部、重慶、湖北、安徽南部、江蘇南部;越南;朝鮮;日本。
標(biāo)本登錄:云南廣南,稅玉民001487。廣西東興,韋毅剛0670;上思,韋毅剛g06;靖西,韋毅剛g052;那坡,覃海寧等2169;龍州,韋毅剛0642;巴馬,韋毅剛AM6483;百色,華南隊2113;田林,韋毅剛094;天峨,北京隊890399;環(huán)江,北京隊894120;興安,廣西隊53-62;龍勝,花坪隊110673。海南澄邁,Lei C.J.456。廣東深圳,張壽洲,李良千3440;增城,曾懷德20255;平遠(yuǎn),鄧良4294;蕉嶺,鄧良4831;連平,劉媖光00129;翁源,劉心啟25057;連州,B.C.Henry 1095。福建泉州,鐘心煊3087;長汀,采集人不明5130;南平,何國生9650;崇安,236 組2366;順昌,李明生,李振宇4716。臺灣臺北,高瑞卿330;新店,Y.Yamamoto s.n。浙江鄞縣,賀賢育1107;杭州,章紹堯2336。江西遂川,236組173;南豐,聶敏祥2406;玉山,聶敏祥6426;南昌,王文采s.n.;永修,賴書紳2200;廬山,聶敏祥7244。湖南新寧,羅林波465;宜章,莽山,肖百中4119;雪峰山,李澤棠2063;道縣,黃向旭10043;長沙,萬紹賓1。貴州荔波,邵青,段林東27;羅甸,貴州隊402;望謨,貴州隊865;桐梓,蔣英5015;劍河,溫放1053;梵凈山,安明態(tài)s.n.。重慶南川,李國鳳60797;萬州,植物所三峽隊799。安徽休寧,植物資源隊59-2070;祁門,安徽師大生物系89045。
9.富寧赤車圖3: E-F
Pellionia funingensis W.T.Wang in Bull.Bot.Res.Harbin 2 ( 1) : 2.1982; X.W.Li in Fl.Yunnan.7: 248.1997; W.T.Wang in Fl.Reip.Pop.Sin.23( 2) : 169.1995.Holotype:云南富寧,里栱,alt.1 410 m,1965-10-13,文山隊137( PE).
Fig.3 : E-F
P.grijsii auct.non Hance: Q.Lin et al in Fl.China 5:126.2003,p.p.quoad syn.P.funingensis W.T.Wang; H.N.Qin&Y.Liu,Checklist Vasc.Pl.Guangxi 237.2010.p.p.quoad syn.P.funingensis W.T.Wang.
分布:云南富寧特有種。
Lin et al ( 2003 )將本種歸并于P.grijsii Hance.但本種的葉具半離基三出脈,二級脈在葉狹側(cè)2條,在寬側(cè)3條,與P.grijsii葉的脈序(為羽狀脈,二級脈約5對)明顯不同,此外莖密被貼伏、長0.5-0.8 mm的短毛,托葉被短伏毛也不相同。在P.grijsii,莖密被開展或反曲、長0.5-2 mm的硬毛,托葉無毛。
10.尖齒赤車圖3: G-H
Pellionia acutidentata W.T.Wang in Bull.Bot.Lab.N.-E.Forest Inst.6: 53.1980; in Bull.Bot.Res.Harbin 3 ( 3) : 56.1983; et in Fl.Reip.Pop.Sin.23( 2) : 169,pl.38: 6-7.1995; X.W.Li in Fl.Yunnan.7: 248.1997; Q.Lin et al in Fl.China 5: 125.2003.Holotype:云南:西疇,法斗,1939-12-09,王啟無85536( PE).Fig.3: G-H
分布:云南東南部。
標(biāo)本登錄:云南屏邊,稅玉民等10391;西疇,王啟無85514,86815;馬關(guān),古林菁,稅玉民,陳文紅,盛家舒30436,31685;河口,稅玉民等10626;麻栗坡,王啟無86815;綠春,稅玉民,陳文紅13813。
11.硬毛赤車圖3: I-K
Pellionia crispulihirtella W.T.Wang in Bull.Bot.Res.Harbin 3 ( 3) : 57,fig.4.1983; et in Fl.Reip.Pop.Sin.23( 2) : 171,pl.42: 7.1995; X.W.Li in Fl.Yunnan.7: 249.1997.Holotype:我國云南屏邊,Shan-tau-chien,alt.1 500 m,1939-10-09,王啟無82383( PE).Paratype: Vietnam On the way from Laokay to Chapa,alt.710 m,1964-12-24,Sino-Vietnam Exped.856( PE).Fig.3: I-K
P.veronicoides auct.non Gagnep.: Q.Lin et al in Fl.China 5: 125.2003.
分布:我國云南東南部;越南北部。
Lin et al( 2003)將本種歸并于P.veronicoides Gagnep.但二種間存在一些形態(tài)區(qū)別特征:在本種,莖密被彎曲的短硬毛(毛長0.15-0.4 mm),無小斑點,退化葉不存在,托葉鉆形,無脈。在P.veronicoides,莖密被反曲的長硬毛(毛長1-2.2 mm),有密集的褐色小斑點,退化葉存在,托葉披針形或卵形,有2條脈,根據(jù)存在退化葉和具較寬的托葉,Schr?ter&Winkler( 1936)認(rèn)為此種與吐煙花相近緣。根據(jù)上述情況可見本種與P.veronicoides是不同的種,在本文的Pellionia分類系統(tǒng)中此二種分別置于不同的系中(見上)。
12.波緣赤車圖3: L-M
Pellionia subundulata W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 57.1980 ; in Bull.Bot.Res.Harbin 3 ( 3) : 67.1983 ; et in Fl.Reip.Pop.Sin.23 ( 2) : 173.1995 ; X.W.Lin in Fl.Yunnan.7: 245.1997; W.T.Wang in Fl.Guangxi 2: 846.2005.Holotype:廣西上思,南屏,1973-04-17,方鼎35075( GXMI)。Paratypes:廣西橫縣,陳照宙50346( IBSC) ;扶綏,陳少卿12399( PE) ;靖西,張肇騫15069( IBSC)。云南屏邊,王孝等100171( IBSC)。
P.paucidentata auct.non ( H.Schr?ter) Chien: Q.Lin et al in Fl.China 5: 125.2003,p.p.quoad syn.P.subundulata W.T.Wang; H.N.Qin&Y.Liu,Checklist Vasc.Pl.Guangxi 238.2010,p.p.quoad syn.P.subundulata W.T.Wang.
12a.var.subundulata
分布:我國云南東南部、廣西;越南北部。
標(biāo)本登錄:云南西疇,王啟無85407,86080。廣西上思,韋毅剛09019,07414;羅城,韋毅剛07121;欽州,周沛椿80047。
圖3 A-B.長梗赤車 A.葉和雄聚傘花序; B.雄花(仿王文采,1995)。C-D.蔓赤車 C.葉; D.宿存雌花被和瘦果(據(jù)韋毅剛g052)。E-F.富寧赤車 E.葉; F.宿存雌花被和瘦果(據(jù)holotype)。G-H.尖齒赤車 G.葉; H.雌花(據(jù)稅玉民等10297)。I-K.硬毛赤車 I.葉; J.雌花; K.雌蕊(據(jù)paratype)。L-M.波緣赤車 L.葉; M.宿存雌花被和瘦果(據(jù)韋毅剛07414)。N-O.赤車N.葉; O.雌花(據(jù)浙江植物資源隊59-25920)。P-Q.異被赤車 P.葉; Q.雌花被和幼果(據(jù)劉正宇4947)。Fig.3 A-B.Pellionia longipedunculata A.Leaf and staminate cyme; B.Staminate flower( after Wang,1995).C-D.P.scabra C.Leaf; D.Persistent pistillate perianth and achene( from Y.G.Wei g052).E-F.P.funingensis E.Leaf; F.Persistent pistillate perianth and achene( from holotype).G-H.P.a(chǎn)cutidentata G.Leaf; H.Pistillate flower( from Y.M.Shui et al 10297).I-K.P.crispulihirtella L.Leaf; J.Pistillate flower; K.Pistil( from paratype).L-M.P.subundulata var.subundulata L.Leaf; M.Persistent pistillate perianth and achene( from Y.G.Wei 07414).N-O.P.radicans var.radicans N.Leaf; O.Pistillate flower( from Zhejiang Pl.Resour.Exped.59-25920).PQ.P.heteroloba P.Leaf; Q.Pistillate perianth and young fruit( from Z.Y.Liu 4947).
Lin et al( 2003)將本種歸并于P.paucidentata ( H.Schr?ter) Chien.但本種的葉片較小,長1-8 cm,寬0.6-3.5 cm,頂端急尖,稀短漸尖,具半離基三出脈,二級脈在葉片狹側(cè)2條,在寬側(cè)3條,雌花被片無毛,而與后者不同。在P.paucidentata,葉片較大,長5-15 cm,寬2-6 cm,頂端漸近或驟尖,有時具半離基三出脈,二級脈在葉狹側(cè)2-3條,在寬側(cè)4-6條,有時具羽狀脈,雌花被片有少數(shù)短毛。
12b.狹葉赤車(變種)
var.a(chǎn)ngustifolia W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 58.1980; in Bull.Bot.Res.Harbin 3( 3) : 57.1983; in Fl.Reip.Pop.Sin.23 ( 2) : 174.1995; et in Fl.Guangxi 2: 846.2005.Holotype:廣西隆安,1973-03-03,王鑒鈞5487 ( GXMI).Paratypes:廣西扶綏,陳少卿12054 ( IBSC) ;寧明,愛店,蘇宏漢6812( IBSC)。
P.longipedunculata auct.non.W.T.Wang: Q.Lin et al In Fl.China 5: 124,2003,p.p.quoad syn.P.subundulata var.a(chǎn)ngustifolia W.T.Wang; H.N.Qin&Y.Liu,Checklist Vasc.Pl.Guangxi 238.2010,p.p.quoad syn.P.subundulata var.a(chǎn)ngustifolia W.T.Wang.
分布:我國廣西南部;越南北部。
標(biāo)本登錄:廣西上思,十萬大山,韋毅剛0680,07414。
Lin et al( 2003)將本變種歸并于P.longipedunculata W.T.Wang,關(guān)于此問題已在后者的地理分布之后進(jìn)行了討論(見上)。
13.赤車圖3: N-O
Pellionia radicans ( Sieb.&Zucc.) Wedd.in DC.Prodr.16 ( 1) : 167.1869; C.H.Wright in J.Linn.Soc.Bot.26: 481.1899; Fl.Hainan.2: 408.1965; Ohwi,F(xiàn)l.Japan,rev.ed.,390.1965; Iconogr.Corm.Sin.1: 514,fig.1028.1972; Fl.Hupeh.1: 172.1976; T.S.Liu&W.D.Huang in Fl.Taiwan 2: 204.1976; W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 55.1980; C.J.Ceng in Fl.Fujian.1: 465.1982; Fl.Jiangsu 2: 82.1982; W.T.Wang in Fl.Reip.Pop.Sin.23( 2) : 171,pl.37: 6.1995; Yang,Shih&Liu in Fl.Taiwan,2nd ed.,2: 231,pl.114.1996; B.G.Li in Fl.Hunan 2: 288.2000; Q.Lin et al in Fl.China 5: 125.2003,p.p.excl.syn.P.radicans f.grandis Gagnep.; Y.Z.Ruan in Fl.Guangdong 6: 79.2005; W.
T.Wang in Fl.Guangxi 2: 844.2005; Tateishi in Iwatsuki et al Fl.Japan 2a: 99.2006; Q.M.Hu in Fl.Hong Kong.1: 123.2007; C.X.Yang et al Keys Vasc.Pl.Chongqing 150.2009.——Procris radicans Sieb.&Zucc.Fl.Jap.Fam.Nat.218.1846.——Elatostema radicans ( Sieb.&Zucc.) Wedd.in Arch.Mus.Hist.Nat.Paris 9: 322.1856; H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83( 2) : 86.1936.
Holotype: Japan: Siebold s.n.( Ld,non vidi).
Fig.3: N-O
P.cavaleriei Lévl.In Repert.Sp.Nov.Regni Veg.12: 22.1913.——Elatostema cavaleriei ( Lévl.) Hand.-Mazz.Symb.Sin.7: 143.1929.Holotype:貴州Pinfa,1902-03-18,Cavalerie 1295( E,non vidi).
P.chikushiensis Yamamoto,Suppl.Ic.Pl.Formos.1: 14,fig.5.1925.Syntypes:臺灣Mt.Chikushi,T.Soma s.n; inter Heishana et Nimandaira,B.Hayata s.n.( non vidi).
13a.var.radicans
分布:我國云南、廣西、海南、廣東、福建、臺灣、浙江、江西、湖南、貴州、四川、湖北、重慶、安徽、江蘇南部;越南;日本。
標(biāo)本登錄:云南河口,稅玉民001401;元陽,綠春隊1722;昆明,E.E.Maire s.n.廣西十萬大山,秦仁昌5302;武鳴,韋毅剛07674;那坡,覃海寧等2053;百色,華南分隊2122;西林,紅水河隊89-1444;融水,陳少卿17003;臨桂,花坪隊110185;環(huán)江,北京隊891093;天峨,北京隊896350;興安,廣西隊53-62;龍勝,劉標(biāo)0637。海南保亭,吊羅山,吊羅山隊3131。廣東鼎湖山,W.Y.Chun 6310;深圳,張壽洲,李良千0387;英德,梁向日61382;乳源,黃志5812;樂昌,左景烈20291;連縣,譚沛祥59867。福建南平,何國生15743;連城,王大順1044;崇安,王名金3095。臺灣臺北,陳家瑞97428;宜蘭,C.M.Wang 05482;新竹,黃雅怡等814;花蓮,陳添財?shù)?2842。浙江平陽,鐘觀光152;諸暨,陳詩98;景寧,章紹堯5160;遂昌,植物資源隊59-25920。江西龍南,236組936;遂川,236組648;武功山,江西隊54-314;萍鄉(xiāng),江西隊54-2818;鉛山,陳躍東等1129;黎川,楊祥學(xué)650010;武寧,陳策銘97021;廬山,胡先骕2208。湖南宜章,肖百中3931;新寧,羅林波345;芷江,武陵山隊2364;武岡,鄧云飛10428;邵陽,段林東2556;衡山,李明紅8;長沙,關(guān)克儉74618;永順,北京隊958。貴州荔波,邵青,段林東40;雷山,簡焯坡等50153;都勻,溫放070430;黃平,韋毅剛,溫放1070;松桃,武陵山隊485;梵凈山,黔北隊972;習(xí)水,畢節(jié)隊1593;赤水,安明態(tài)32;道真,劉正宇20516;桐梓,劉正宇20517;遵義,川黔隊1471。四川雷波,管中天7819;峨邊,趙子孝64;峨眉山,關(guān)克儉等1777;汶川,阿壩藥檢所719;都江堰,傅德志等87-1407。重慶縉云山,黃志明1097;涪陵,劉正宇180979;金佛山,韋毅剛1133;萬州,植物所三峽隊0876;巫山,植物所三峽隊1366。湖北鶴峰,李洪鈞685;宜昌,董佩萱20;鄂州,金仁龍324。安徽黃山,植物所華東工作站6494;祁門,鄧樊彬5088。
13b.長莖赤車(變種)
var.grandis ( Gagnep.) W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 56.1980; X.W.Li in Fl.Yunnan.7: 247.1997; W.T.Wang in Fl.Guangxi 2: 846.2005.——P.radicans f.grandis Gagnep.in Lecomte,F(xiàn)l.Gén.Indo-Chin.5: 899.1929; Fl.Hainan.2: 408.1965; C.J.Ceng in Fl.Fujian.1: 465.1982; W.T.Wang in Fl.Reip.Pop.Sin.23( 2) : 173.1995; Y.Z.Ruan in Fl.Guangdong 6: 79.2005.——Elatostema radicans var.grandis( Gagnep.) H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83 ( 2) : 89.1936.Syntypes: Vietnam Tonkin: prov.de Ninh-yen,Eberhardt s.n.; prov.Bac-kan et Thai-nguyen,Eberhardt s.n.( P,non vidi).
分布:我國云南東南部、廣西、海南、廣東;越南北部。
標(biāo)本登錄:廣西上思,SCNG3392。
14.異被赤車圖3: P-Q
Pellionia heteroloba Wedd.in Arch.Mus.Hist.Nat.Paris 9: 283,t.9,fig.15.1856 ; et in DC.Prodr.16( 1) : 166.1869 ; Hook.f.Fl.Brit.Ind.5: 559.1888 ; Gagnep.in Lecomte,F(xiàn)l.Gén.Indo-Chin.5: 899.1929; W.T.Wang in Fl.Reip.Pop.Sin.23( 2) : 176.1995,p.p.excl.syn.P.griffithianum Wedd.et Elatostema henryanum Hand.-Mazz.; X.W.Li in F.Yunnan.7: 245.1997,p.p.excl.pl.62: 3-6.1997; Q.Lin et al in Fl.China 5: 126.2003,p.p.excl.syn.P.griffithiana Wedd.,P.menglianense Y.Y.Qian et Elatostema henryanum Hand.-Mazz.Holotype: Sikkim,J.D.Hooker&T.Thomson s.n.( K.non vidi, photo PE).Fig.3: P-Q
P.heteroloba var.minor W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 60.1980; S.S.Chang et al in Fl.Guizhou.4: 40.1989.Holotype:貴州:習(xí)水,alt.650 m,1959-09-24,畢節(jié)隊1593( PE)。
分布:我國云南、貴州、湖南西南、重慶、四川南部;越南;印度北部和東北部。
標(biāo)本登錄:云南騰沖,Gaoligong Shan Biodiv.Survey 30146;景東,哀牢山,楊國平393;金平,周哲坤等215;廣南,王啟無87305。貴州冊亨,曹子余1032;望謨,韋毅剛,溫放1032;荔波,邵青,段林東60;正安,劉正宇15594,20381;道真,劉正宇16043;懷仁,向邦華s.n.;赤水,韋毅剛,溫放1142。湖南武岡,云山,張彩飛1709。重慶江津,劉正宇183073;南川,金佛山,李國鳳64319,劉正宇4947;豐都,三峽隊2665。
最近,我看到Pellionia heteroloba Wedd.和P.griffithiana Wedd.的模式照片,并參閱了這二種的原始描述,了解到前者的莖無毛,葉較小,也無毛,具半離基三出脈,而后者的莖、葉都有毛,葉較大,具羽狀脈,二者是兩個不同的種。因此,H.Schr?ter ( 1936)將P.heteroloba Wedd.和P.grijsii Hance都?xì)w并于P.griffithiana是制造了一個植物分類學(xué)混亂,對此,本文進(jìn)行了改正。
15.三被赤車圖4: A-D
Pellionia tritepala W.T.Wang,sp.nov.Holotype:云南屏邊,Shuiweicheng,alt.2 000 m,at water side near woods,2000-03-27,稅玉民等( Y.M.Shui et al) 12136(♀,PE)。Fig.4: A-D
Species nova haec est affinis P.radicanti( Sieb.&Zucc.) Wedd.,a qua foliorum laminis apice breviter acuminatis vel acutis subtus atrobrunneo-puncticulatis,flore pistillato tepalis tribus omnibus dorso sub apice longe raro breviter corniculatis praedito differt.In P.radicanti,foliorum laminae apice acuminatae vel longeacuminataesubtushaudatrobrunneopuncticulatae,et flos pistillatus tepalis quinque vulgo tribus majoribus dorso sub apice longe corniculatis ceteris minoribus haud corniculatis praeditus est.
Perennial herb.Stems 1-2,20-28 cm tall,ca.1 mm in diam.,minutely puberulous( hairs 0.1-0.25 mm long),dark-brown-puncticulate,simple or below shortly branched.Leaves alternate,shortly petiolate; blades pa-pery,obliquely elliptic or obliquely obovate,2.5-4.5(-5.5)×1.5-2.5 cm,apex shortly acuminate or acute,base at leaf narrow side obtuse and at broad side auriculate,margin crenate or denticulate; surfaces adaxially sparsely strigillose,abaxially subglabrous,densely dark-brownpuncticulate; nervation semi-triplinerved,with basal nerves and ca.2 pairs of secondary nerves abaxially slightly prominent; cystoliths sparse,inconspicuous,thinly bacilliform,0.2-0.4 mm long; petioles 1-2 mm long,glabrous; stipules subulate or narrowly linear,1-2.5×0.1-0.3 mm,glabrous.Staminate cymes and staminate flowers unknow.Pistillate cymes singly axillary,5-7 mm in diam.,shortly 3-4-branched,densely multiflowered; peduncle 0.5-4 mm long,glabrous; bracts numerous,white,linear,1-1.5×0.1-0.2 mm,glabrous.Pistillate flower sessile: Tepals 3,unequal in size,white,linear-navacular,0.3-0.8×0.25 mm,glabrous,abaxially below apex long rarely shortly corniculate,with horn-like projections( 0.2-) 0.5-1.5 mm long.Pistil ca.0.4 mm long; ovary ellipsoid,0.3-0.4 mm long,glabrous; stigma penicillate,ca.0.15 mm long.
在赤車系ser.Radicantes,多數(shù)種的雌花具5枚花被片,少數(shù)種(如蔓赤車P.scabra,滇南赤車P.paucidentata)的雌花具4或5枚花被片。本種的雌花具3枚花被片,在本系中是唯一的情況,因此,根據(jù)這一特征即可與本系的其他種相區(qū)別。
16.灰被赤車圖4: E-G
Pellionia calcifera W.T.Wang,sp.nov.Holotype:云南麻栗坡,豆豉店( Malipo,Douchidian),alt.950 m,山坡( on slope),1998-10-30,王印政,彭華,鄧云飛,稅玉民,朱大海( Y.Z.Wang,H.Peng,Y.F.Deng,Y.M.Shui&D.H.Zhu) 5155( PE)。
Fig.4: E-G
Species nova haec caulibus calci glebulis minutis densissimis albidis incrustatis,foliorum laminis utrinque calcis strato tenui albido incrustatis,cymis staminatis sessilibus,floribus staminatis tetrameris insignis est,et cumcharacteribus his a congeneribus omnibus sinensibus ceterisdistat.Inejuscongeneribus sinensibus,caules et foliorum laminae calce carentes,cymae staminatae pedunculatae,et flores staminati pentameri rarissime tetrameri[in P.paucidentata ( H.Schr?ter) Chien]sunt.
Perennial herbs.Stems ca.25 cm tall,branched,on branch surface encrusted by numerous very dense minute lumps of whitish lime,and on upper part of branch densely puberulous.Leaves alternate,ca.5 borne on apical part of branch,shortly petiolate or sessile; blades papery,obliquely elliptic,2.4-7×1.4-3.5 cm,apex shortly acuminate or acute,base at leaf narrow side cuneate and at broad side broadly cuneate,margin above the middle crenate; surfaces both adaxially and abaxially encrusted by a thin grey-whitish layer of lime,glabrous; nervation semi-triplinerved or trinerved,with basal nerves and ca.3 pairs of secondary nerves abaxially prominent; cystoliths dense,inconspicuous,thickly bacillioform,0.1-0.25 mm long; petioles 0-1.5 mm long; stipules subulate,ca.2×0.2 mm,glabrous.Staminate cymes singly borne on leafless nodes of lower part of branch,sessile,2.5-4 mm in dian.,densely 10-20-flowered; bracts brown,triangular or linear,0.6-0.8×0.3-0.5 mm,ciliolate.Staminate flower: Pedicel up to 1 mm long,glabrous.Tepals 4,free,broadly elliptic,ca.1.2× 0.7 mm,glabrous.Stamens 4,glabrous; filaments subterete,ca.0.6 mm long; anthers oblong,ca.0.6 mm long; pistilllode wanting.Pistillate cymes and pistillate flowers unknown.
本種的特征:莖被極為密集的白色石灰小塊包著,葉片的腹面和背面均被一薄層灰白色石灰包著,雄聚傘花序無花序梗,雄花四基數(shù)。根據(jù)上述特征,本種可與我國赤車屬的其他種區(qū)別開。在我國赤車屬的其他種,莖和葉均無石灰,雄聚傘花序均有花序梗,雄花均為五基數(shù),稀四基數(shù)[滇南赤車P.paucidentata( H.Schr?ter) Chien.]。
17.短葉赤車圖4: H-I
Pellionia brevifolia Benth.Fl.Hongk.330.1861; Wedd in DC.Prodr.16( 1) : 167.1869; C.H.Wright in J.Linn.Soc.Bot.26: 481.1899; Fl.Hainan.2: 409.1965; W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 58.1980; C.J.Ceng in Fl.Fujian.1: 466.1982; W.T.Wang.in Fl.Reip.Pop.Sin.23( 2) : 174,pl.38: 4.1995; B.G.Li in Fl.Hunan 2: 289.2000; Q.Lin.in Fl.China 5: 125.2003,p.p.excl.syn.P.minima Makino; Y.Z.Ruan in Fl.Guangdong 6: 79.2005; W.T.Wang in Fl.Guangxi 2: 846.2005; Tateishi in Iwatsuki et al Fl.Japan 2a: 78.2006; Q.M.Hu in Fl.Hong Kong1: 123.2007.——Elatostema brevifolium ( Benth.) Hall.f.in Ann.Jard.Bot.Bruitenz.8: 316.1896; H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83 ( 2) : 89.1936.Holotype:香港,Mt.Gough and Victoria,Wilford 227( K,non vidi; photo,PE).Fig.4: H-I
圖4 A-D.三被赤車 A.開花雌莖頂部; B.雌聚傘花序; C,D.雌花(據(jù)holotype)。E-G.灰被赤車 E.葉; F.雄聚傘花序; G.另一雄聚傘花序(據(jù)holotype)。H-I.短葉赤車 H.葉; I.宿存雌花被和瘦果(據(jù)廈門大學(xué)隊929)。J-K.小赤車 J.葉; K.宿存雌花被和瘦果(據(jù)浙江植物資源隊59-25760)。L-M.滇南赤車 L.葉; M.雌花(仿王文采,1995)。N-P.南亞赤車 N.葉和雌聚傘花序; O.雌花; P.瘦果(據(jù)蔡希陶52440)。Fig.4 A-D.Pellionia tritepala A.Apical part of flowering pistillate stem; B.Pistillate cyme,C,D.Pistillate flowers( from holotype).EG.P.calcifera E.Leaf; F.Staminate cyme; G.Another staminate cyme( from holotype).H-I.P.brevifolia H.Leaf; I.Persistent pistillate perianth and achene( from Xiamen Univ.Exped.929).J-K.P.minima J.Leaf; K.Persistent pistillate perianth and achene( from Zhejiang Pl.Resour.Exped.59-25760).L-M.P.paucidentata var.paucidentata L.Leaf; M.Pistillate flower( after Wang,1995).N-P.P.griffithana var.griffithiana N.Leaf and pistillate cyme; O.Pistillate flower; P.Achene( from H.T.Tsai 52440).
分布:我國廣西、海南、廣東、福建、江西、湖南、安徽南部;越南北部;日本。
標(biāo)本登錄:廣西融水,元寶山隊Y1381;環(huán)江,北京隊895144;興安,韋毅剛0912。福建南靖,廈門大學(xué)隊931。江西武功山,江西隊54-442;廬山,楊穗生262。湖南新寧,紫云山,劉林翰15106;桑植,八大公山,李洪鈞4371。安徽潛山,沈保安398。
18.小赤車圖4: J-K
Pellionia minima Makino in Bot.Mag.Tokyo 23: 85.1909; Ohwi,F(xiàn)l.Japan,rev.ed.,390.1965; P.S.Hsu in Obs.Pl.Hwangshan.109.1965; W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 158.1980; Z.W.Xiuo in Fl.Anhui 2: 122,fig.288.1986; Y.Q.He in Fl.Zhejiang 2: 111,fig.2-147.1992; W.T.Wang in Fl.Reip.Pop.Sin.23( 2) : 175.1995; Y.Z.Ruan in Fl.Guangdong 6: 178.2005; W.T.Wang in Fl.Guangxi 2: 846.2005; Tateishi in Iwatsuki et al Fl.Japan 2a: 98.2006.——Elatostema radicans var.minima ( Kakino) H.Schr?ter in Repert.Sp.Nov Regni Veg.Beih.83 ( 2) : 88.1936.——Pellionia radicans var.minima( Makino) Hatusima in Sci.Rep.Yokosuka City Mus.n.13: 37.1967.Described from Japan,no type specimen designated.Fig.4: J-K
P.brevifolium auct.non Benth.: Q.Lin et al in Fl.China 5: 125.2003,p.p.quoad syn.P.minima Makino.
分布:我國廣西、廣東、湖南、江西、福建、浙江、安徽;日本。
標(biāo)本登錄:廣西龍勝,韋毅剛s.n.廣東:從化,曾懷德20445;新豐,鄧良8211;豐順,李學(xué)根201424。湖南瀏陽,嚴(yán)岳鴻5814。江西奉新,劉守爐1192;玉山,廬山植物園6125;婺源,李啟和62。福建延平,鐘心煊3580;寧化,黃鋤荒s.n.;南平,何國生9645;建寧,陳家瑞等79。浙江遂昌,植物資源隊59-25760;昌化,賀賢玉22628。安徽黃山,植物所華東工作站6111;祁門,錢士心s.n。
Lin et al( 2003)將Pellionia minima Makino歸并于P.brevifolia Benth.但本種的莖被長0.1 mm的小毛,葉呈寬橢圓形,寬倒卵形或近圓形,雌花被片無毛;而在P.brevifolia,莖被長0.3-1 mm的較長毛,葉呈橢圓形或狹倒卵形,雌花被片疏被短毛。這些區(qū)別說明二者不是同一分類群,本文仍將P.minima作為種級分類群處理。
19.滇南赤車圖4: L-M
Pellionia paucidentata ( H.Schr?ter) Chien in Acta Phytotax.Sin.8: 354.1963; W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 60.1980; et in Fl.Reip.Pop.Sin.23( 2) : 175,pl.37: 34,pl.39.1995; X.W.Li in Fl.Yunnan.7: 251,pl.62.1997; Q.Lin et al in Fl.China 5: 125.2003,p.p.excl.syn.P.subundulata W.T.Wang et P.paucidentata var.hainanica Chien&S.H.Wu ; Y.M.Shui&W.H.Chen,Seed Pl.Karst Reg.China 1: 103,fig.255.2006; H.N.Qin&Y.Li,Checklist Vasc.Pl.Guangxi 288.2010,p.p.excl.syn.P.subundulata W.T.Wang.——Elatostema henryanum var.oligodotum Hand.-Mazz.Symb.Sin.7: 144.1929.——E.paucidentatum H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83 ( 2 ) : 80.1936.Syntypes: Vietnam Tonkin: Phomoi near Laogai,alt.150 m,1914-02-02,Handel-Mazzetti 10 ( W.non vidi).China.Yunnan: Mengzi,A.Henry 11261( K,non vidi) ; Simao,A.Henry 12678( K,non vidi).
P.longgangensis W.T.Wang in Bull.Bot.Res.Harbin 8( 4) : 77.1988.Holotype:廣西龍州,弄崗,1985-11-09,李振宇,傅德志,覃海寧27( PE)。
19a.var.paucidentata
分布:我國云南中南部和東南部、廣西西南部;越南北部。
標(biāo)本登錄:云南南澗,無量山,劉恩德3534;勐臘,王啟無80685;勐侖,熱帶植物園,李慶華462;孟鑫,王啟無80050;金平,武素功等4343;河口,稅玉民等00716;馬關(guān),稅玉民等30577,31722;西疇,王啟無85235;個舊,稅玉民等14424。廣西龍州,韋毅剛07292;防城,鐘觀光2665;百色,韋毅剛百色4號。
19b.海南赤車(變種)
var.hainanica Chien&S.H.Wu in Acta Phytotax.Sin.8 ( 4) : 354,1963; Fl.Hainan.2: 467.1965; W.T.Wang in Fl.Reip.Pop.Sin.23 ( 2) : 176.1995; Y.Z.Ruan in Fl.Guangdong 6: 80.2005.Holotype:無準(zhǔn)確地址,Bot.Inst.Agr.Trop.Austr.-Sin.0011( IBSC)。
與滇南赤車的區(qū)別:葉較小,長達(dá)12 cm,寬達(dá)4.2 cm,鐘乳體在兩面均明顯。
分布:海南特有變種。
系3.翅莖赤車系ser.Caulialatae W.T.Wang in Acta Phytotax.Sin.30 ( 3) : 196.1992; et in Fl.Reip.Pop.Sin.23( 2) : 180.1995.Type: P.caulialata S.Y.Liou.
Reduced leaves present.Stems longitudinally winged.Normal leaves penninerved; stipules narrowly triangular or narrowly lanceolate.Pistillate flower’s tepals 5,of them 2 larger,abaxially below apex long corniculate.Achenes tuberculate.
1種。
20.翅莖赤車圖5: A-D
Pellionia caulialata S.Y.Liou in Guihaia 3( 4) : 317,cun fig.1983; W.T.Wang in Fl.Reip.Pop.Sin.23( 2) : 180.1995; Q.Lin et al in Fl.China 5: 126.2003.Isotype:廣西桂平,太平山,alt.400 m,1982-12-14,劉壽養(yǎng)等D00322( PE).Fig.5: A-D
分布:云南東南部、廣西西部和東南部;越南北部。
標(biāo)本登錄:云南馬關(guān),古林菁,稅玉民,陳文紅,盛家舒30803。廣西龍州,韋毅剛14;大新,韋毅剛6673;靖西,韋毅剛AM6356;凌云,華南分隊1378;東蘭,稅玉民,陳文紅B2004-167;天峨,北京隊897064。Vietnam: Cho Don Distr.,Bac Kan Province,2004-10-27,L.Q.Li et al 0447.
系4.南亞赤車系ser.Griffithianae W.T.Wang,ser.nov.Type: P.griffithiana Wedd.
Series nova haec est affinis ser.Caulialatis W.T.Wang,a qua foliis reductis absentibus,stipulis subulatis vel anguste linearibus,caulibus haud alatis differt.
Reduced leaves absent.Stems not winged.Leaves penninerved; stipules subulate or narrowly linear.Pistillate flower’s tepals 5,rarely 4,of them 2-3 larger,abaxially below apex long corniculate.Achenes tuberculate.
我國有3種。
21.南亞赤車圖5: G-I
Pellionia griffithiana Wedd.in DC.Prodr.16 ( 1) : 165.1869; Hook.f.Fl.Brit.Ind.5: 559.1888; C.H.Wright in J.Linn.Soc.Bot.26: 481.1899; Gagnep.in Lecomte,F(xiàn)l.Gén.Indo-Chin.5: 895.1929.——Elatostema griffithianum( Wedd.) Hall.f.in Ann.Jard.Bot.Bruitenz.13: 316.1896; H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83( 2) : 76.1936,p.p.excl,syn.Pellionia heteroloba Wedd.et P.grijsii Hance; Backer&v.d.Brink,F(xiàn)l.Java 42.1965.——Elatostema henryanum Hand.-Mazz.Symb.Sin.7: 144.1929.Holotype:印度India E Bengal,Palkaya,W.Griffith 4544( K,non vidi,photo PE)。Fig.5: G-I
P.a(chǎn)risanensis Hayata,Ic.Pl.Formos.6: 53.1916.Holotype:臺灣Mt.Arisan,0914-03,B.Hayata s.n.( non vidi)。
P.keitaoensis Yamamoto,Suppl.Ic.Pl.Formos.1: 15,cum tab.1925.Holotype:臺灣Keitao et Goshorin,1916-04,B.Hayata s.n.( non vidi).
P.menglianensis Y.Y.Qian in Acta Phytotax.Sin.37( 5) : 523,fig.1.1999.Isotype:云南孟連,alt.2 000 m,0992-04-29,錢義詠2563( PE)。
P.heteroloba auct.non Wedd.: W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 59.1980,p.m.p.quoad H.T.Tsai 51824 et C.W.Wang 78418 ; et in Fl.Reip.Pop.Sin.23( 2) : 176.1995,p.p.quoad syn.Pellionia griffithiana Wedd.; X.W.Li in Fl.Yunnan.7: 245.1997,p.p.quoad pl.62: 3-6.; Q.Lin et al in Fl.China 5: 126.2003,p.p.quoad syn.Pellionia griffithiana Wedd.et P.menglianensis Y.Y.Qian.
21a.var.griffithiana
分布:我國云南、廣西、廣東、臺灣;越南;老撾;印度東北部、印度尼西亞(爪哇)。
標(biāo)本登錄:云南貢山,獨龍江隊4532;景東,哀牢山,周仕順1471;勐海,中蘇考察團(tuán)5119;景洪,王啟無78415;蒙自,王啟無83433,A.Henry 11405;個舊,稅玉民等14388;屏邊,蔡希陶52440,61828,王啟無82942;硯山,王啟無84795;文山,蔡希陶51824;馬關(guān),蔡希陶51912,馮國楣13668,稅玉民等30781,31380,31656;綠春,陶德定313,857,稅玉民等70434;麻栗坡,王啟無86866,馮國楣13968;河口,稅玉民等10641;金平,武素功等3164,4448。廣西那坡,覃海寧等2175;靖西,韋毅剛g054;百色,百色隊1477;凌云,華南隊1573;田林,韋毅剛0736;天峨,北京隊890335。廣東深圳,田心山,韋毅剛,溫放1120。
21b.無毛莖南亞赤車(變種)
var.glabricaulis W.T.Wang,var.nov.Holo type:云南( Yunnan) :綠春( Luchun),alt.800 m,2000-10-23,稅玉民,陳文紅等( Y.M.Shui,W.H.Chen et al) 14123( PE)。
P.heteroloba auct.non Wedd.: W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 59.1980,p.p.quoad L.Deng 8077.
A var.griffithiana differt caulibus glabris vel subglabris,foliorum laminis subtus glabris,nervis secundariis paucioribus 4 latere angustiore 5-6 latere latiore,stipulis minoribus subulatis 3-4 mm longis ca.0.1 mm latis.In var.griffithiana,caules densissime minute appresso-brunneolo-puberuli,pilis 0.1-0.2 mm longis ; foliorum laminae subtus ad nervos puberulae,nervis secundariis 5-6 latere angustiore et 6-8 latere latiore; et stipulae majores anguste trangulari-lineares,5.5-12 mm longae,0.7-1.2 mm latae sunt.
分布:云南、廣西、廣東。
標(biāo)本登錄:云南騰沖,alt.2 400 m,1955-07-10,吳征鎰等( C.Y.Wu et al) 867;景東,無量山( Jingdong,Wuliang Shan),alt.2 800 m,1956-11-12,邱炳云( B.Y.Qiou) 53370;南澗,無量山( Nanjian,Wuliang Shan),alt.2 232 m,2012-05-22,劉恩德( E.D.Liu) 3534 ;綠春( Luchun),alt.1 600 m,2006-03-10,稅玉民,陳文紅等70298;金平( Jinping),alt.2 600 m,1996-09-30,武素功等( S.K.Wu et al) 3693。廣西那坡,德孚保護(hù)區(qū)( Napo,Defu nature reserve),alt.1 400 m,1998-11-03,覃海寧等( H.N.Qin et al) 2096;百色,大王嶺( Baise,Dawangling),2009-03-21,韋毅剛,溫放( Y.G.Wei&F.Wen) 1186。廣東新豐( Xinfeng),alt.650 m,1958-10-14,鄧良( L.Deng) 8077.
本變種的莖無毛或近無毛,葉片下面無毛,二級脈較少,在葉狹側(cè)4條,在寬側(cè)5-6條,托葉較小,呈鉆形,長3-4 mm,寬約0.1 mm而與南亞赤車不同。在南亞赤車,莖被極密而貼伏的淡褐色極短毛(毛長0.1-0.2 mm),葉片下面在脈上被短毛,二級脈在葉狹側(cè)5-6條,在寬側(cè)6-8條,托葉較大,狹三角狀條形,長5.5-12 mm,寬0.7-1.2 mm。
21c.疏毛南亞赤車(變種)
var.laxivillosa W.T.Wang,var.nov.Holotype:云南( Yunnan)福貢( Fugong),alt.2 000 m,林中( in forest),1934-10-21,蔡希陶( H.T.Tsai) 58874( PE).
P.heteroloba auct.non Wedd.: W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 59.1980,p.p.quoad H.T.Tsai 58874 et C.W.Wang 72294.
A var.griffithana differt caulibus pilis sparsis albis tenuibus( 0.5-) 1-2 mm longis obtectis,foliorum laminis subtus glabris,nervis secundariis paucioribus 3-4 latere angustiore et 4-6 latere latiore,stipulis minoribus subulatis 2-4 mm longis 0.1-0.2 mm latis.
分布:云南特有變種。
標(biāo)本登錄:云南福貢( Fugong),alt.2 300 m,1934-09-17,蔡希陶( H.T.Tsai) 54276 ; Lu-se,alt.1750 m,1934-03-02,蔡希陶( H.T.Tsai) 56382;鎮(zhèn)康( Zhenkang) alt.2 200-2 700 m,1936-03,王啟無( C.W.Wang) 72294,72421,72553;普文,瘴氣河( Puwne,Zhangqi River),alt.1 060 m,密林中石上( on rocks in dense forest),1957-04-01,中蘇考察團(tuán)( Sino-Russian Exped.) 3008。
本變種的莖疏被白色細(xì)長柔毛[毛長( 0.5-) 1.0 -2 mm],葉片下面無毛,二級脈較少,在葉狹側(cè)3-4條,在葉寬側(cè)4-6條,托葉較小,呈鉆形,長2-4 mm,寬0.1-0.2 mm,而與南亞赤車相區(qū)別。
21d.少脈南亞赤車(變種)
var.paucinervis W.T.Wang,var.nov.Holotype:江西上猶,陵水樹木園( Shangyou,Lingshui arboretum),alt.300 m,竹林中( in bamboo thicket),2007-11-01,贛南采集隊( S.Jiangxi Exped.) 928( PE).
A var.griffithiana differt caulibus tantum prope apicem retrose albo-strigosis ( pilis 0.2-0.4 mm longis),folio nervis secundariis 4-jugatis praedito,stipulis minoribus subulatis 3-4 mm longis 0.1-0.2 mm latis.
分布:江西上猶特有變種。
本變種的莖只在近頂部被白色反曲糙伏毛(毛長0.2-0.4 mm),葉有4對二級脈,托葉較小,呈鉆形,長3-4 mm,寬0.1-0.2 mm,而與南亞赤車相區(qū)別。
22.華南赤車圖5: E-F
圖5 A-D.翅莖赤車 A.葉和雌聚傘花序; B,C.雌花被片; D.瘦果(仿劉壽養(yǎng),1983)。E-F.華南赤車 E.葉; F.宿存雌花被和瘦果(據(jù)溫放070506B)。G-I.云南赤車 G.葉; H.宿存雌花被和瘦果; I.瘦果(稅玉民等11636)。J-M.大葉赤車 J.葉; K.雌花; L.瘦果; M.雄花(仿王文采,1995)。N-P.羽脈赤車 N.葉和雌聚傘花序; O.宿存雌花被和瘦果; P.瘦果(據(jù)isotype)。Q-S.兩廣赤車Q.葉和雌聚傘花序; R.宿存花被和瘦果; S.瘦果(仿Wei&Wang,2010)。Fig.4 A-D.Pellionia caulialata A.Leaf and pistillate cyhme; B,C.Pistillate tepals; D.Achene( after S.Y.Liu,1983).E-F.P.grijsii E.Leaf; F.Persistent pistillate perianth and achene( from F.Wen 070506B).G-I.P.yunnanensis G.Leaf; H.Persistent pistillate perianth and achene( from Y.M.Shui et al 11636).J-M.P.macrophylla J.Leaf; K.Pistillate flower,L.Achene; M.Staminate flower( after Wang,1995).N-P.P.incisoserrata N.Leaf and pistillate cyme; O.Persistent pistillate perianth and achene; P.Achene( from isotype).Q-S.P.ronganensis Q.Leaf and pistillate cyme; R.Persistent pistillate perianth and achene; S.Achene( after Wei&Wang,2010).
Pellionia grijsii Hance in J.Bot.6: 49.1868 et 12: 262.1874; C.H.Wright in J.Linn.Soc.Bot.26: 481.1899; W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 61.1980; C.J.CenginFl.Fujian.1:467.1982; W.T.Wang in Fl.Reip.Pop.Sin.23( 2) : 178,pl.37: 1-2.1995; X.W.Lin in Fl.Yunnan.7: 252.1997; B.G.Li in Fl.Hunan 2: 289.2000; Q.Lin et al in Fl.China 5: 126.2003,p.p.excl.syn.P.funingensis W.T.Wang; Y.Z.Ruan in Fl.Guangdong 6: 81.2005; W.T.Wang in Fl.Guangxi 2: 846.2005; Y.M.Shui&W.H.Chen,Seed Pl.Karst Reg.China 1: 102,fig.255.2006; Q.M.Hu in Fl.Hong Kong 1: 123,fig.90.2007.Holotype:福建1861,C.F.M.Grijs 6704( BM,non vidi; photo,PE)。Fig.5: E-F
Elatostema griffithianumauct.non ( Wedd.) Hall.f.: H.Schr?ter in Repert.Sp.Nov Regni Veg.Beih.83( 2) : 76.1936,p.p.quoad syn.Pellionia grijsii Hance.
Pellionia griffithiana auct.non Wedd.: Fl.Hainan.2: 406.1965.
分布:云南南部和東南部、廣西、海南、廣東、福建、江西西部、湖南、貴州東南部。
標(biāo)本登錄:云南思茅,周仕順972;西疇,王啟無35991。廣西那坡,覃海寧等556;大新,李振宇等86;扶綏,陳少卿12064;武鳴,韋毅剛AM6395;橫縣,陳照宙503460;金秀,覃海寧68;融水,北京隊891722,溫放070506B;百色,華南分隊2142;臨桂,花坪隊H0451;環(huán)江,北京隊895009。海南保亭,吊羅山隊3135;陵水,海南東隊920;安定,黃志35348;白沙,海南東隊632。廣東鼎湖山,鐘觀光1007;清遠(yuǎn),黃志161447;增城,曾懷德20178;新豐,鄧良8081;翁源,韋毅剛,溫放1017;連平,衛(wèi)兆芬120063。福建南靖,廈大采集隊937。江西井岡山,熊杰3193。湖南桂東,湖南隊A183;道縣,黃向旭10025;永興,王福德s.n。貴州荔波,邵青,段林東62。
23.云南赤車圖5: G-I
Pellionia yunnanensis( H.Schr?ter) W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 61.1980; et in Fl.Reip.Pop.Sin.23( 2) : 179.1995; X.W.Li in Fl.Yunnan.7: 249,pl.63: 1-3.1997; Q.Lin et al in Fl.China 5: 126.2003.——Elatostema yunnanense H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83 ( 2) : 79,tab.29.1936.Type:云南蒙自之南,alt.1 700 m,A.Henry 9163D( holotype,B; isotypes,E,K,non vidi)。Fig.5: G-I
Supplenmentary description to the original one, with the infructescences,persistent pistillate tepals and achenes described for the first time.: Infructescences singly axillary,depressed-globose,4-10 mm in diam.,densely multi-flowered; peduncles 1-4 mm long,glabrous; bracts numerous,narrowly triangular-linear or narrowly linear,1-2×0.1-0.15 mm,glabrous.Pistillate flower’s tepals 4,persistent,whitish,unequal in size,glabrous,3-4 larger,linear-navicular,0.8-1 mm long,abaxially below apex long corniculate,with horn-like projections 0.8-1 mm long,other 1-2 tepals smaller,flat,narrowly lanceolate-linear,0.7-1.1 mm long.Achenes brown,long ellipsoid,ca.1×0.4 mm,minutely tuberculate.( from Y.M.Shui et al 11636)
本種的模式標(biāo)本A.Henry 9163D為雄株,因此原始描述中缺雌花序,雌花和瘦果的描述,最近看到采自云南,河口,大圍山的稅玉民等11636號標(biāo)本為雌株,遂對其果序、宿存雌花被片和瘦果做出如上補(bǔ)充描述。
分布:云南東南部特有種。
標(biāo)本登錄:云南綠春,黃連山,武素功等2040(♂) ;河口,萬丈崖,稅玉民等10907(♂) ;河口,大圍山,稅玉民11636(♀) ;馬關(guān),古林菁,稅玉民等31070(♂) ;屏邊,蔡希陶61140(♂),馮國楣5108(♂) ;麻栗坡,王啟無86850(♂),王印政等4840(♂)。
系5.大葉赤車系ser.Macrophyllae W.T.Wang,ser.nov.Type: P.macrophylla W.T.Wang.
Series nova haec est affinis ser.Griffithianis W.T.Wang,a qua acheniis longitudinaliter dense lineolatis differt.
Reduced leaves absent.Stems not winged.Leaves penninerved; stipules subulate or narrowly linear.Pistillate flower’s tepals 5,of them 2-3 larger,abaxially below apex long cornitulate.Achenes longitudinally densely lineolate.
1種。
24.大葉赤車圖5: J-M
Pellionia macrophylla W.T.Wang in Acta Bot.Yunnan.5( 3) : 271,fig.1.1983; et in Fl.Reip.Pop.Sin.23( 2) : 181,pl.40.1995; X.W.Li in Fl.Yunnan.7: 249,pl.63: 6-7.1997; Q.Lin et al in Fl.China 5: 127.2003.Type:云南綠春,alt.2 000 m,1973-10-15,陶德定845( holotype,KUN; isotype,PE),857( paratype,KUN),1973-09-09,141,195( paratypes,KUN).Fig.5: J-M
分布:云南東南部、廣西西部。
標(biāo)本登錄:云南金平,武素功等4301;河口,稅玉民778,1031;馬關(guān),古林菁,稅玉民,陳文紅,盛家舒30763;硯山,王啟無84988。廣西那坡,覃海寧等2270;百色,百色隊1508。
組3.羽脈赤車組sect.Leiolaena W.T.Wang,sect.nov.Type: P.incisoserrata ( H.Schr?ter) W.T.Wang。
Sectio nova haec est affinis sect.Elatostemoidibus ( C.B.Robinson) W.T.Wang**,a qua foliis penninervibus,foliis reductis absentibus,raro praesentibus differt.
Leaves shortly petiolate or sessile,with dentate margins,penninerved.Reduced leaves absent,rarely present.Pistillate inflorescences cymose.Pistillate flower’s tepals 5,rarely 4 or 3,abaxially usually lacking any projections,rarely below apex shortly corniculate.A-chenes smooth or tuberculate,rarely lineolate and minutely tuberculate.
我國有7種,根據(jù)瘦果和退化葉等特征被劃分為4系。
系1.羽脈赤車系ser.Incisoserratae W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 63.1980; et in FL.Reip.Pop.Sin.23 ( 2) : 181.1995,p.p.excl.P.macrophylla W.T.Wang; Yahara in J.Fac.Sci.Univ.Tokyo,Sect.3,Bot.13( 4) : 386.1984.Type: P.incisoserrata( H.Schr?ter) W.T.Wang。
Reduced leaves abaent.Pistillate flower’s tepals 5,rarely 4,all lacking any projections.Achenes minutely tuberculate.
2種。
25.羽脈赤車圖5: N-P
Pellionia incisoserrata( H.Schr?ter) W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 63.1980; in Fl.Reip.Pop.Sin.23( 2) : 183,pl.41: 1-5.1995; et in Fl.Guangxi 2: 847.2005.——Elatostema incisoserratum H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83 ( 2) : 90,tab.27.1936.Type:廣東乳源,1933-05-10,高錫朋S.P.Ko 52629( holotype,W,non vidi,photo,PE; isotype,IBSC)。Fig.5: N-P
分布:廣西、廣東北部。
標(biāo)本登錄:廣西巴馬,甲篆,韋毅剛AM6465;融水,溫放070505。廣東翁源,韋毅剛,溫放1015。
26.兩廣赤車圖5: Q-S
Pellionia ronganensis W.T.Wang&Y.G.Wei in Nordic.J.Bot.28: 54,fig.2010.Type:廣西融安,浮石鎮(zhèn),Shangliuliao village,alt.200 m,2009-04-28,韋毅剛09185( holotype,PE; isotype,IBK)。Fig.5: Q-S
P.tenuicuspis W.T.Wang,Y.G.Wei&F.Wen in Ann.Bot.Fennici 49: 190,fig.2.2012,syn.nov.
Type:廣東乳源,Luoyang Town,Tianjingshan Mount,alt.408 m,2010-03-08,韋毅剛,溫放1019( holotype,IBK,non vidi; isotype,PE) ;同地,alt.410 m,2011-03-10,韋毅剛,溫放1103( paratype,IBK,non vidi)。
分布:廣西北部、廣東北部。
系2.光果赤車系ser.Leiocarpae W.T.Wang in Guihaia 2( 3) : 115.1982; et in Fl.Reip.Pop.Sin.23 ( 2) : 183.1995,p.p.excl.P.brachyceras W.T.Wang.Type: P.leiocarpa W.T.Wang。
Reduced leaves present.Pistillate flower’s tepals 5,abaxially lacking any projections or below apex with short projections.Achenes smooth.
1種。
27.光果赤車圖6: A-C
Pellionia leiocarpa W.T.Wang in Guihaia 2( 3) : 115,fig.1-4.1982; in Fl.Reip.Pop.Sin.23( 2) : 183,pl.42: 1-4.1995; in Fl.Guangxi 2: 847,fig.344: 1-4.2005; et in Bull.Bot.Res.*
Harbin 26( 1) : 15.2006; Q.Lin et al in Fl.China 5:127.2003; Y.M.Shui&W.H.Chen,Seed Pl.Karst Reg.China 1: 102,fig.253.2006.Holotype:廣西那坡,百都,弄花,alt.1 100 m,1981-08-21,方鼎,覃德海,王振剛25142( GXMI)。Fig.6: A-C * Pellionia Gaudich.sect.Elatostemoides ( C.B.Robinson ) W.T.Wang,st.et comb.nov.——Gen.Elatostemoides C.B.Robinson in Philip.J.Sci.Bot.5( 6) : 497,1910.——Elatostema Forster subgen.Elatostemoides( C.B.Robinson) H.Schr?ter in Repert.Sp.Nov.Regni Veg.Beih.83( 1) : 2.1935; ( 2) : 2,8,92.1936.Lectotype: P.rostrata ( Hall.f.) Boerl.,here designated.
Leaves shortly petiolate or sessile,usually with dentate margins,trinerved or semi-triplinerved,with lateral basal nerve at leaf narrow side extending upwards nearly to leaf apex.Redueed leaves usually present,rarely absent.Pistillate inflorescnes cymose.Pistillate flower’s tepals 5,abaxially lacking any projections or below apex shortly corniculate.Achenes usually smooth.
Forty five species,wide-spread in tropical islands of SE Asia and SW Oceania.
圖6 A-C.光果赤車 A.莖頂部; B.宿存雌花被和瘦果; C.瘦果(仿王文采,1995)。D-E.短角赤車 D.開花莖頂部; E.雌花(仿王文采,1995)。F-H.龍州赤車 F.具二葉的分枝; G.果序; H.瘦果(仿王文采,2010)。I-K.東蘭赤車 I.葉; J.雌聚傘花面,下面觀; K.雌花(仿王文采,2010)。L-N.柔毛齒車 L.葉; M.果序; N.瘦果(仿王文采,2014)。O-R.頭序赤車 O.三葉和三雌頭狀花序; P.雌頭狀花序,下面觀; Q.雌花(仿王文采,2010) ; R.瘦果(據(jù)李明紅,曠月秋841)。Fig.6 A-C.Pellionia leiocarpa A.Apical part of stem; B.Persistent pistillate perianth and achene; C.Achene( after Wang,1995).DE.P.brachyceras D.Apical part of flowering pistillate stem; E.Pistillate flower( after Wang,1995).F-H.P.longzhouensis F.A branch with two leaves; G.Infructescence; H.Achene( after Wang,2010).I-K.P.donglanensis I.Leaf; J.Pistillate cyme,seen from beneath; K.Pistillate flower( after Wang,2010).L-N.P.mollissima L.Leaf; M.Infructescence; N.Achene( after Wang,2014).O-R.P.cephaloideaO.Three leaves and three pistillate capitula; P.Pistillate capitulum,seen from beneath; Q.Pistillate fower( after Wang,2010).R.Achene( from M.H.Li&Y.Q.Kuang 841).
分布:云南東南部、廣西西部;越南北部。
標(biāo)本登錄:云南河口,稅玉民,陳文紅等20612;麻栗坡,稅玉民,陳文紅等20376,吳增源等10395;馬關(guān),古林菁,稅玉民等30804。廣西那坡,中植聯(lián)廣西隊4125;靖西,韋毅剛,溫放1171;凌云,韋毅剛101;樂業(yè),韋毅剛g102。Vietnam Tam Dao保護(hù)區(qū),韋毅剛v-36。
系3.短角赤車系ser.Brachycerates W.T.Wang,ser.nov.Type: P.brachyceras W.T.Wang.Series nova haec est affinis Ser.Leiocarpis W.T.Wang,a qua foliis reductis nullis differt。
Reduced leaves absent.Pistillate flower’s tepals 3 -5,abaxially lacking any projections,rarely with short projections.Achenes smooth.
3種。
28.短角赤車圖6: D-E
Pellionia brachyceras W.T.Wang in Bull.Bot.Res.Harbin 3 ( 3) : 185,fig.2-3.1983; in Fl.Reip.Pop.Sin.23( 2) : 185,pl.42: 5-6.1995; et in Fl.Guangxi 2: 846.2005; Q.Lin et al in Fl.China 5: 127.2003.Holotype:廣西隆林,大吼豹自然保護(hù)區(qū),alt.1 420 m,1984-11-10,梁健英,覃民府K0442 ( IBK)。Fig.6: D-E
分布:廣西西北部特有種。
標(biāo)本登錄:廣西隆林,大吼豹自然保護(hù)區(qū),韋毅剛,溫放g081。
29.龍州赤車圖6: F-H
Pellionia longzhouensis W.T.Wang in Guihaia 30( 1) : 3,fig.2: A-C,1020.Holotype:廣西龍州,高山村,板閉,alt.330 m,2005-05-11,稅玉民,陳文紅,張美德B2005-043( PE)。Fig.6: F-H
分布:廣西龍州特有種。
30.東蘭赤車圖6: I-K
Pellionia donglanensis W.T.Wang in Guihaia 30 ( 1) : 1,fig.1.2010.Holotype:廣西東蘭,波豪鄉(xiāng),alt.300 m,石灰?guī)r山山洞口,2004-02-18,稅玉民,陳文紅B2004-434( PE)。Fig.6: I-K
分布:廣西東蘭特有種。
系4.柔毛赤車系ser.Mollissimae W.T.Wang,ser.nov.Type: P.mollissima W.T.Wang。
Series nova haec acheniis longitudinaliter dense lineolatis et minute tuberulatis a seriebus tribus ceteris sectionis Leiolaenae distat.
Reduced leaves absent.Pistillate flower’s tepals 5,abaxially lacking any projections.Achenes longitudinally densely lineolate and minutely tuberculate.
1種。
31.柔毛赤車圖6: L-N
Pellionia mollissima W.T.Wang in Guihaia 34 ( 1) : 1,fig.1.2014.Holotype:廣西龍州,金龍,alt.250 m,石灰?guī)r山林中,2003-09-15,稅玉民,陳文紅B2003-038( PE)。Fig.6: L-N
分布:廣西龍州特有種。
組4.頭序赤車組sect.Elatostematopsis W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6: 64.1980; in Fl.Reip.Pop.Sin.23( 2) : 185.1995; et in Guihaia 31( 1) : 2.2011.Type: P.cephaloidea W.T.Wang。
Leaves sessile or very shortly petiolate,with dentate margins,trinerved vel semi-triplinerved,with lateral basal nerve at leaf narrow side extending upwards slightly surpassing the leaf middle.
Reduced leaves present or absent.Pistillate inflorescences capitate,with discoidreceptacles and involucres.Pistillate flower’s tepals 5,subaequal in size,longer than ovary,lanceolate or narrowly oblong,lacking any projections or abaxially below apex shortly corniculate.Achenes ellipsoid or globose,tuberculate or longitudinally ribbed and minutely tuberculate.
2種,我國有1種。
32.頭序赤車圖6: O-R
Pellionia cephaloidea W.T.Wang in Bull.Bot.Lab.N.-E.Forest.Inst.6 64.1980; et in Fl.Reip.Pop.Sin.23( 2) : 185,pl.38: 1-3.1995; B.G.Li in Fl.Hunan 2: 289.2000; W.T.Wang in Guihaia 31 ( 2) : 2,fig.1.2011.Holotype:湖南衡山,1925-06-08,湖南博物館s.n.( PE)。
P.scabraauct.non Benth.: Q.Lin,L.D.Duan&M.H.Li in Acta Phytotax.Sin.40( 5) : 178.2002,p.p.quoad syn.P.cephaloidea W.T.Wang; Q.Lin et al in Fl.China 5: 124.2003,p.p.quoad syn.P.cephaloidea W.T.Wang.
Supplementary description to the original one,with the achenes described for the first time.Achenes yellowish-white,globose,ca.0.8 mm in diam.,longitudinally 5-ribbed and minutely tuberculate.( from M.H.Li&Y.Q.Kuang 841)
分布:湖南衡山特有種。
標(biāo)本登錄:湖南衡山,玉板橋,alt.350 m,1996-03-14,夏江林59;同地,半山亭,alt.600 m,2003-05-03,李明紅,曠月秋841。
致謝蒙傅德志教授,楊永博士和楊志榮博士提供一些種的模式標(biāo)本照片和文獻(xiàn),孫英寶先生為本文繪圖,作者謹(jǐn)表示衷心感謝。
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CLC number: Q949.71Document code: AArticle ID: 1000-3142(2016)01-0030-14
Notes on Carex (Cyperaceae) from China (IV). the identity and revision of sect. Infossae
CEN Jia-Meng1, JIN Shui-Hu2, JIN Xiao-Feng1*
( 1. College of Life & Environmental Sciences,Hangzhou Normal University,Hangzhou 310036,China; 2. School of Foresrty and Bio-technology,Zhejiang Agricultural and Forestry University,Lin’an 311300,China )
containing two species with one variety: C. infossa var. infossa,C. infossa var. extensa and C. minuticulmis. According to the sectional description,fifteen additional species and two varieties should be placed in this section. Based upon literature survey,specimen examination,field work,and SEM observation of perigynia and achenes,a concise table of different treatments between Florae Reipublicae Popularis Sinicae and Flora of China was provided,as well as the morphological characters distinguishing these taxa and micromorphology of perigynia and achenes for the selected taxa were compared. Among the diagnostic characters,arrangement of spikes and perigynium indumentum are inseparable from each other,and two species in sect. Infossae are consequently recognized in the present paper. Carex elmeri,C. granifera,C. minuticulmis,C. shanghaiensis,C. blinii subsp. shanghaiensis,C. sublateralis,C. taihokuensis and C. tatsutakensis were synonymized to C. blinii. Carex canaliculata,C. changmuensis,C. distantiflora,C. infossa,C. infossa var. extensa,C. lateralis,C. loheri,C. longerostrata var. exaristata,C. longerostrata var. hoi,C. lyi,C. macrandrolepis,
C. sharyotensis were reduced as synonyms to C. oxyphylla.
作者簡介:王文采( 1926-),男,研究員,中國科學(xué)院院士,植物分類學(xué)家,長期從事有花植物分類學(xué)研究。
收稿日期:2015-09-17修回日期: 2015-10-28
DOI:10.11931/guihaia.gxzw201509011
中圖分類號:Q949
文獻(xiàn)標(biāo)識碼:A
文章編號:1000-3142( 2016) 01-0001-29