中国苎麻属大叶苎麻组小赤麻系修订
2016-11-10王文采
王文采
(中国科学院植物研究所,系统与进化植物学国家重点实验室,北京 100093)
中国苎麻属大叶苎麻组小赤麻系修订
王文采
(中国科学院植物研究所,系统与进化植物学国家重点实验室,北京 100093)
1)接受荨麻科苎麻属大叶苎麻亚属小赤车组,但将其降为系级群,作为大叶苎麻组的一个系处理。(2)在小赤麻系中共承认6种,其中包括1新种,给出了6种的检索表,简要形态描述(对新种给出完整描述)和插图,还提供了6种的地理分布和分布地图。(3)讨论了6种的密切亲缘关系,并提出几个推测:小赤麻系中的原始种是细野麻,此系其他种可能均自其演化而出;也可能细野麻和密球苎麻为自同一祖先演化而出的一对姊妹群;也可能异叶苎麻自赤麻演化而出。
荨麻科;苎麻属大叶苎麻组小赤麻系;分类学修订;中国
在1981年我编写出中国苎麻属分类学修订[1],参考了荷兰植物学家C.Blume[4]和日本植物学家Y.Satake[7]关于苎麻属植物分类学的重要著作,在我建立的我国苎麻属分类系统中接受了Blume建立的三个组:Sect.Genuinae Bl.(包括BoehmeriadepauperataWedd.,B.ramifloraJacq.等种。此组植物的团伞花序单生叶腋,不形成穗状、复杂的团伞花序,我认为是苎麻属的原始群,其他组均由其演化而出),Sect.Tilocnide Bl.[包括B.nivea(L.f.) Gaudich.]和Sect.Duretia(Gaudich.) Bl.[包括B.japonica(L.f.)Miq.,B.spicata(Thunb.) Thunb.等多数种],以及Satake建立的一组,Sect. ZollingerianaeSatake(包括B.zollingerianaWedd.)。此外,我将Satake建立的二组,Sect. LongispicaeSatake[包括B.japonica(L.f.) Miq.,B.platanifoliaFranch. & Sav.,B.holoseriseaBl.等种]和Sect. DensifloraeSatake(包括B.densifloraHook. & Arn.等种)降为系级群作为Sect.Duretia之下的二系;同时还将他建立的另外一组Sect. SpicataeSatake[包括B.spicata(Thunb.) Thunb.,B.paraspicataNakai等种]归并于Ser. Longispicae(Satake) W.T.Wang。最近我在鉴定苎麻属植物标本的过程中发现一新种,横断山苎麻(B.hengduanshanicaW.T.Wang),并发现其与细野麻(B.gracilisC.H.Wright),赤麻(B.silvestrii(Pamp.) W.T.Wang),小赤麻(B.spicata(Thunb.) Thunb.),密球苎麻(B.densiglomerataW.T.Wang)和异叶苎麻(B.allophyllaW.T.Wang)在亲缘关系方面甚为相近,这六种拥有以下共同特征:植株被毛较少;茎细,常带红色;叶较小,叶片较薄,薄纸质或膜质,长3~8(~10) cm,宽2~10(~13) cm,通常被少数短毛或近无毛,边缘的牙齿较小,长1~6(~10) mm;穗状花序不分枝,稀在基部有少数短枝,花序轴纤细,直径0.2~0.5 mm;宿存雌花上部被糙伏毛,常变无毛。根据这些特征可与Ser. Longispicae相区别。后者的特征:叶较大,叶片较厚,长度和宽度均达20 cm左右,边缘牙齿也较大,其顶端三角形牙齿长达2(~3) cm,且常为重牙齿;穗状花序常分枝,花序轴稍粗些,直径达1 mm;宿存雌花被顶端密被糙硬毛。根据上述我认识到应该承认Sect. SpicataeSatake,但将其等级下降,作为Sect. Duretia的一个系Ser. Spicatae(Satake) W.T.Wang处理。
六种的主要形态特征和亲缘关系:(1)细野麻(Boehmeriagracilis)为多年生草本;叶片通常宽卵形或圆卵形,3~7(~10)×2~6(~7.5) cm,顶端有1骤尖头,基部圆形或圆截形,边缘每侧具8~16个不等大的牙齿,牙齿呈正三角形,宽三角形或三角形,稀呈狭三角形(如采自重庆市奉节的周洪富,粟和毅110943号标本),1~5(~10)×2~5(~10) mm;穗状花序长2.5~13 cm,花序轴直径0.2~0.4 mm;宿存雌花被具扁平边缘;叶片有时呈卵形,基部宽楔形,如采自重庆市酉阳的谭士贤222、采自湖北宜昌的龙茹090808004和采自陕西平利的采集人不明IV0145等三号标本均具卵形叶而与小赤麻(B.spicata)甚为相似,只是叶缘的牙齿呈正三角形或宽三角形而与小赤麻(叶缘牙齿狭三角形)不同,上列诸号标本均是B.gracilis与B.spicata之间的中间类型的居群的代表(图1:A~B);(2)赤麻(B.silvestrii)为多年生草本;茎顶部少数叶较小,其叶片圆卵形,通常不分裂,顶端具1骤尖头,与B.gracilis的叶极为相似,其下方的叶较大,近五角形,5~8(~13)×4.8~7.5(~13) cm,顶端3浅裂,具3个长骤尖头,基部近截形或宽楔形,边缘每侧有5~12个不等大的牙齿,牙齿呈正三角形或三角形,1~10(~15)×1~9 mm,顶部的牙齿较大,且常为重牙齿;穗状花序长4~11(~20) cm,花序轴直径约0.3 mm;宿存雌花被具扁平边缘。这里我要指出一个情况,就是采自辽宁南部凤凰山的近藤舍五8282号的本种标本只有茎中部的一对叶的叶片顶端具3骤尖头,其他茎下部至顶部的十余个叶的圆卵形叶片顶端均象B.gracilis的叶一样只有1个骤尖头,这个现象有力地说明B.silvestrii是由B.gracilis演化而出的种。在我国苎麻属中存在由具不分裂叶的种演化至具分裂叶的种的现象,如序叶苎麻组(Sect. Phyllostachys W.T.Wang)中的由具不分裂叶的白面苎麻(B.clidemioidesMiq.)演化出具2裂叶的双尖苎麻(B.bicuspisC.J.Chen)和具3裂叶的阴地苎麻(B.umbrosa(Hand.-Mazz.) W.T.Wang)[2]。再如大叶苎麻系(Ser. Longispicae)中的由具不分裂叶的大叶苎麻(B.japonica(L.f.) Miq.)演化出具3裂叶的悬铃叶苎麻(B.tricuspis(Hance) Makino)[2]。而在本系Ser. Spicatae则出现了由具不裂叶的B.gracilis演化出具3裂叶的B.silvestrii和具2裂叶的B.hengduanshanica的现象。上述情况显示了苎麻属中在物种形成方面的平行演化现象(图1:C~D);(3)横断山苎麻(B.hengduanshanica)为多年生草本或小灌木;叶片卵形或椭圆形,3~10×1.5~5.5 cm,顶端不等2浅裂,具2不等大的骤尖头,基部圆形或宽楔形,边缘每侧具6~13个牙齿,牙齿呈宽三角形或三角形,1~5×2~6 mm;雄穗状花序长2~5.2 cm,花序轴直径约0.2 mm。本种与B.gracilis近缘,区别主要在于叶片顶端2裂,可能由后者演化而出(图2:A~C);(4)小赤麻B.spicata为小灌木;茎具分枝;叶对生,叶片卵状菱形或宽菱形,2.4~7.5×1.5~5 cm,顶端长骤尖,基部宽楔形,边缘每侧具(3~)4~9(~16)个牙齿,牙齿呈狭三角形或三角形,1~6×0.8~6 mm;雄穗状花序长2.5 cm,雌穗状花序长4~10 cm,花序轴直径0.2~0.4 mm;宿存雌花被具扁平边缘。上面介绍了在重庆酉阳、奉节,湖北宜昌和陕西平利等地出现了B.gracilis和B.spicata之间的中间类型,由此推测本种可能自B.gracilis演化而出(图1:E~F);(5)密球苎麻(B.densiglomerata)为多年生草本;叶对生,叶片圆卵形,5~9.4×5.2~8 cm,顶端短骤尖或渐尖,基部浅心形,稀圆形,边缘每侧具7~17个牙齿(牙齿呈正三角形或宽三角形,1~5×1~6 mm),下面疏或密被糙伏毛,有时无毛;穗状花序长2.5~5.5 cm,花序轴直径约0.5 mm,其上的团伞花序极为密集,彼此紧密邻接,这是本种的一个重要特征,在小赤麻系的其他种,花序轴上的团伞花序互相间均多少分离;宿存雌花被无扁平边缘,或有狭扁平边缘,或顶端部分扁平。本种根据叶片基部通常浅心形和穗状花序轴上紧密排列的团伞花序二特征可与B.gracilis区别,但这二种形态很相似,常难于区别。我曾将本种标本误定为B.gracilis,因此认为二种的亲缘关系很紧密,推测本种是由B.gracilis演化而出,向南分布到华南地区的种,也可能本种与B.gracilis是由同一祖先演化而出的一对姊妹群(图2:D~I);(6)异叶苎麻(B.allophylla)为多年生草本;茎顶端叶互生,叶片椭圆形,4~5.7×1.4~1.7 cm,不分裂,顶端短骤尖,基部宽楔形;其他下部叶对生,较大,叶片近五角形,6~7×4~5.8 cm,顶端不等3微裂,不等3短骤尖,基部宽楔形,边缘每侧具8~20个牙齿,牙齿呈宽三角形或三角形,1~2.5×1.5~4 mm;穗状花序长3~8 cm,花序轴直径0.3~0.5 mm。本种茎顶端互生叶形状似B.gracilis的叶,下部的对生叶具3骤尖头,则与B.silvestrii相似,由此推测本种可能由后二者之一演化而出(图2:J~M)。
图1 A~B.细野麻:A.开花茎上部;B.瘦果和宿存花被 C~D.赤麻:C.开花茎上部;D.瘦果和宿存花被 E~F.小赤麻:E.叶;F.瘦果和宿存花被(自王文采,1995)Fig.1 A-B.Boehmeria gracilis: A. Upper part flowering stem; B. Achene with persistant perianth C-D.B.silvestrii: C. upper part of flowering stem; D. Achene with persistent perianth E-F. B.spicata: E. Leaf; F.Chene with persistent perianth(from Wang 1995)
图2 A~C.横断山苎麻:A.开花雄茎上部;B.雄团伞花序;C.雄花蕾(据holotype) D~I.密球苎麻:D.叶和穗状花序:E. 2雄团伞花序;F. 3雌团伞花序;G.雌花(据李洪钧3231);H.瘦果和宿存花被(据曹子余,王忠祷292);I.另1瘦果和宿存花被(据李学根202260) J~M.异叶苎麻:J.开花茎上部;K.雄团伞花序;L.雌团伞花序;M.雌花(据holotype)Fig.2 A-C. Boehmeria hengduanshanica:A. Upper part of flowering staminate stem; B. Staminate glomerule; C. Staminate flower bud(from holotype) D-I. B.densiglomerata: D. Leaf and axillary spike; E. Two staminate glomerules; F. Three pistillate glomerules; G. Pistillate flower(from H.J.Li 3231); H. Achene with persistent perianth(from Z.Y.Can & Z.T.Wang 292); I. Another achene with persistent perianth(from X.G.Li 202260) J-M. B.allophylla: J. Upper part of flowering stem; K. Staminate glomenule; L. Pistillate glomerule; M. Pistillate flower(from holotype)
近年来,Ser. Spicatae中所有已发表的种的种级地位遭遇到变动。2003年,《Flora of China》vol.5[5]中第二版的中国苎麻属志将B.gracilis归并于B.spicata。2013年,英国荨麻科专家C.M.Wilmot-Dear和丹麦荨麻科专家I.Friis合作发表了旧世界苎麻属的分类学修订[9],最近我阅读了这篇130多页、图文并茂的大文章,了解到旧世界苎麻属植物的情况,感到此文是荨麻科植物分类学的一个重要文献,但此文对Ser. Spicatae诸种的分类学处理令我甚感诧异:(1)将异叶苎麻B.allophylla与B.grandifoliaWedd.、B.longispicaSteud.、B.tricuspis(Hance) Makino、B.macrophylla(Thunb.) Sieb. & Zucc.等多数学名一起放进B.japonica(L.f.) Miq. var.japonica的同物异名的长名录中。如上所述,B.allophylla与B.gracilis和B.silvestrii近缘,在叶、花序等器官方面与B.japonica存在多数区别,因此,二者不可能是同一分类群。(2)将B.silvestrii恢复为变种等级,作为B.japonica的一个变种B.japonica(L.f.)Miq. var.silvestrii(Pamp.) Friis & Wilmot-Dear。如上所述,B.silvestrii由B.gracilis演化而来,而与B.japonica也存在营养和生殖器官方面的多数区别特征,但却被作为B.japonica的变种,由此,我感到在种的概念(species conception)上,我与这二位专家存在不小区别。(3)将B.spicata(Thunb.) Thunb. var.teneraBl.(1857,此变种名多年来被不少日本植物学家归并于B.spicata)转移至B.japonica之下作为后者的一个变种:B.japonica(L.f.) Miq. var.tenera(Bl.) Friis & Wilmot-Dear,同时还将B.spicata和B.gracilis放入此变种的同物异名的名录中。如上所述,B.gracilis和B.spicata在亲缘关系上极为相近,存在中间类型,但二者还是两个分类群,并不是同一分类群。此外,这两个分类群和近缘的B.silvestrii和B.allophylla一样,与B.japonica存在不少形态区别,由此说明这些分类群与B.japonica不是同一种植物。(4)将B.densiglomerata降级作为B.virgata(G.Forster) Guill.的一个变种:B.virgatassp.macrophylla(Hornem.) Friis & Wilmot-Dear var.densiglomerata(W.T.Wang) Friis & Wilmot-Dear。我不了解B.virgata这种植物,从二位专家的著作中了解到这个种包括2亚种,14变种,广布于亚洲东南部和南部热带地区,其中2变种分布于非洲热带地区,还有1变种分布于澳大利亚东部,只有1变种var.densiglomerata自我国华南北部一带向北达长江中游地区。我对B.virgata缺乏了解,但对B.densiglomerata有一定了解,如上所述,此种与B.gracilis在亲缘关系上甚为相近,有时难于区别,我曾推测此种可能在Ser. Spicatae的分布中心云贵高原北缘一带由B.gracilis分化而出,或与后者为一对姊妹群一同起源于上述Ser. Spicatae的分布中心,然后向东南分布达云南东南部和华南一带,根据这种认识,我对将B.densiglomerata作为旧世界热带分布的B.virgata的一个变种深感诧异。
地理分布。小赤麻系(Ser. Spicatae)是苎麻属分布于东亚植物区系[8]中的一个具6种的小群,6种中有3种广泛分布于中国、朝鲜和日本,其他3种为我国特有种。本系的原始种细野麻B.gracilis是本系分布最广的种,西自四川横断山区东缘,南自华南北部(南岭山区)向北、向东分别达到了辽宁南部和吉林东南部,以及朝鲜和日本四岛(九州、四国、本州和北海道)[6]。第二个分布最广的种是赤麻(B.silvestrii),西自四川横断山区东缘,南自湖南向北经过华中、华北,朝东北方向到达黑龙江东南部,向东和细野麻一样也到达朝鲜和日本四岛[6]。小赤麻(B.spicata)是本系第三个分布广的种,西自四川东部,南自贵州东南、江西北部和浙江向北比较星散地分布到山西南部和山东东北部,向东分布到朝鲜和日本三岛(九州、四国和本州)[6],并在日本分化出一小叶变种var.microphyllaNakai ex Satake[6~7]。日本诸岛是在第三纪中新世(Miocene)与大陆分离的,可见上述三种可能在中新世或其前已在我国大陆出现了。本系第四个分布广的种是我国特有种密球苎麻(B.densiglomerata),在上面讨论本种的种级地位问题时曾谈到其分布情况,此种可能自云贵高原北部起源后向东南分布达华南一带以及云南东南部,这里形成Ser. Spicatae的南界。最后是特产我国的两个狭域分布种:横断山苎麻(B.hengduanshanica)可能是细野麻(B.gracilis)向西演化出的种,出现在横断山区的云南维西和西藏察隅,这里形成Ser. Spicatae的西界。另一种,异叶苎麻(B.allophylla)特产广西西北部凌云县,此种可能起源自赤麻(B.silvestrii),因此可能是由长江中游地区向南分布过来的。从6种上述分布情况看,在云贵高原北缘一带分布有细野麻(B.gracilis),赤麻(B.silvestrii),小赤麻(B.spicata)和密球苎麻(B.densiglomerata),从采集到的大量标本看到有关四种的居群在这一带大量密集分布,云贵高原北缘无疑是小赤麻系(Ser. Spicatae)的分布中心。
分类学处理
苎麻属Boehmeria Jacq.大叶苎麻组小赤麻系
Sect.Duretia(Gaudich.) Bl. Ser.Spicatae(Satake) W.T.Wang, st. et comb. nov. ——BoehmeriaJacq. subgen.Duretia(Gaudich.) Satake sect.SpicataeSatake in J. Fac. Sci. Univ. Tokyo, sect. 3, Bot. 4: 480. 1936; Ohba in Iwatsuki et al. Fl. Japan 2a: 102. 2006. Lectotype: Boehmeria spicata(Thunb.) Thunb.
Series haec est arcte affinis Ser.Longispicis(Satake) W. T. Wang, quae foliis majoribus crassioribus utrinque pilis pluribus obtectis margine valde grosse dentatis, dentibus apicalibus triangularibus usque ad 2(-3) cm longis saepe duplicatis, spicarum rhachidibus crassioribus usque ad 1 mm in diam., perianthiis fructiferis apice dense hispidis differt.
Perennial herbs, rarely small shrubs. Stems often reddish in colour. Leaves opposite, rarely alternate; blades usually thinly papery or membranous, 3-8(-10)×2-10(-13) cm, usually covered with few short hairs or subglabrous, with marginal teeth 1-6(-10) mm long. Spikes usually simple, rarely at base with a few short branches; rachis slender, 0.2-0.5 mm in diam. Persistent pistillate perianth above strigose, often glabrate.
Six species, of them three species wide-spread in China, Korea and Japan, and the other three ones endemic to China.
1.细野麻 图1:A-B
BoehmeriagracilisC. H. Wright in J. Linn. Soc. Bot. 26: 485. 1899; Hand.-Mazz. Symb. Sin. 7: 151. 1929; Fl. Econ. Pl. China 1: 98, fig. 62. 1961; Iconogr. Corm. Sin. 1: 520, fig. 1039. 1972; Fl. Tsinling. 1(2): 115, fig. 96. 1974; S. X. Fu, Fl. Hupeh. 1: 180. 1976; B. Z. Ding et al. Fl. Honan. 1: 301. 1981; W. T. Wang in Acta Bot. Yunnan. 3(4): 409. 1981; Fl. Jiangsu 2: 86, fig. 854. 1982; Fl. Dalian Distr. 1: 137, fig. 143. 1982; Z. T. Yin in Fl. Beijing, rev. ed., 1: 126. 1984; et in Fl. Hebei. 1: 290. 1986; Z. W. Xue in Fl. Anhui 2: 128, fig. 396. 1987; S. Z. Liu in Fl. Liaoning 1: 307, pl. 121. 1988; X. S. Zhang in Fl. Guizhou. 4: 70. 1989; Fl. Shandong 1: 993, fig. 655. 1992; X. Y. Yu et al. in Fl. Shanxi 1: 352, pl. 216. 1992; Y. Q. He in Fl. Zhejiang 2: 119, fig. 2-160. 1992; W. T. Wang in Fl. Reip. Pop. Sin. 23(2): 346, pl. 78: 5-6. 1995; B. G. Li in Fl. Hunan 2: 304, fig. 2-227. 2000; S. J. Fan et al. Fl. Laoshan 1: 162, fig. 148. 2003; F. L. Zhang in Fl. Tianjin 97, fig. 94. 2004; Z. X. Peng & G. L. Zhang in Fl. Gansu 2: 245. 2005; W. T. Wang in Fl. Guangxi 2: 870. 2005; Ohba in Iwatsuki et al. Fl. Japan 2a: 103. 2006; X. Z. Zi & D. C. Zhang, Fl. Dabieshan 329, fig. 459. 2006; C. X. Yang et al. Keys Vasc. Pl. Chongqing 156. 2009; Z. H. Chen in Fl. Tianmushan 1: 284, fig. 1-299. 2009; Y. Zhou, Pl. Resour. Changbai Mount. China 420, pl. 204: 5. 2010; Q. L. Gan, Fl. Zhuxi., Suppl. 176. 2011; F. Z. Li et al. Fl. Taishan 1: 314, pl. 352. 2012. Syntypes: 湖北:巴东,A. Henry 4692(K, non vidi),4728(K, non vidi; photo, PE);Changlo, A. Henry 6258(K, non vidi).
B.paraspicataNakai, Rep. Veg. Mt. Apoi 19. 1930; Satake in J. Fac. Sci. Univ. Tokyo, sect. 3, Bot. 4: 483, fig. 9-10. 1936; Kitagawa, Lineam. Fl. Manch. 171. 1939; B. S. Xu in Obs. Fl. Hwangshan. 108. 1965. Holotype: Japan: Hokkaido, Isihari Prov., 1931-08, G. Koidzumi s. n.(TI, non vidi).
B.tricuspis(Hance) Makino var.unicuspisMakino, Ill. Fl. Japan, rev. ed., 641. 1948, nom. subnud.; Ohwi, Fl. Japan 441. 1956 et rev. ed., 391. 1965; Y. C. Zhu in Herb. Pl. N.-E. China 2: 5, fig. 3. 1959; Satake in Satake et al. Wild Fls. Japan 2: 9, pl. 7: 2. 1982; Y. N. Lee, Fl. Korea 91. 1996.
B.spicataauct. non(Thunb.) Thunb.: Chen, Friis & Wilmot-Dear in Fl. China 5: 173. 2003, p. p. quoad syn.B.gracilisC. H. Wright; B. Liu, Field Guide Wild Pl. China: Shandong 292, fig. 3. 2009; J. C. Zhao et al. Fl. Xiaowutaishan 1: 176, fig. 126. 2011; Z. L. Xu in Fl. Jiangsu 2: 172, fig. 2-279. 2013, p. p.
B.japonica(L. f.) Miq. var. tenera(Bl.) Friis & Wilmot-Dear in Blumea 58(2): 193. 2013, p. p. quoad syn. B. gracilis C. H. Wright.
Perennial herbs. Leaves opposite; blades broadly ovate or orbicular-ovate, rarely ovate, 3-7(-10)×2-6(-7.5) cm, apex cuspidate, base rounded, rounded-truncate or broadly cuneate, margins on each side 8-16-dentate, with teeth deltoid or triangular, 1-5(-10)×2-5(-10) mm. Spikes 2.5-13 cm long; rachis 0.2-0.4 mm in diam.; glomerules along rachis among themselves more or less separate. Persistent perianth with complanate margins.
地理分布:四川、贵州、广西北部、广东北部、福建、浙江、江西、湖南、湖北、重庆、甘肃南部、陕西南部、山西南部、河南、安徽、江苏东北部、山东、河北、辽宁南部和东部、吉林东南部;朝鲜、日本。
标本登录(均存PE):四川:宝兴,关克俭,王文采2597。贵州:纳雍,毕节队423;雷山,简焯坡等51079;凯里,黔南队1966;石阡,黔南队2608;道真,刘正宇20507;梵净山,黔北队2672。广西:融水,陈少卿12861;罗城,秦仁昌5479。广东:乳源,粤73队603。福建:南靖,叶国栋1623;漳平,林镕4491。浙江:天目山,贺贤育25397;天台山,陈诗3920。江西:上犹,江西队71-522;井冈山,熊杰3104;铅山,聂敏祥4424;靖安,张吉华97004;修水,赖书绅2922;庐山,关克俭74242。湖南:新宁,罗林波317;衡南,王小溪117;桑植,刘林翰9246;石门,壶瓶山队1185。湖北:咸丰,王映明6548;鹤峰,李洪钧6749;利川,戴伦膺1602;宜昌,龙茹090808004,090819018;巴东,鄂植考队24490;神农架,鄂植考队25249;罗田,何其果U1。重庆:南川,金佛山,李国凤63545;酉阳,谭士贤170,222;武隆,刘正宇180330;石柱,三峡队1919;开县,巴山队2526;奉节,周洪富,粟和毅109229;巫溪,三峡队290;城口,戴天伦101149。甘肃:文县,王新402;康县,张志英16897。陕西:岚皋,采集人不明I0364;紫阳,李培元4656;平利,采集人不明IV0145;汉中,汪劲武376;太白山,秦岭队1410;终南山,王作宾2072;户县,郭本兆674;蓝田,苏贵兴278。山西:晋城,包士英1763。河南:新县,植物资源队D0917;商县,植物资源队D0582;栾川,河南队2028;西峡,河南队1446;林州,经济植物队1189。安徽:休宁,刘晓龙279;绩溪,黄成林1129;黄山,钱崇澍1214;潜山,沈保安149;岳西,刘晓龙476;舒城,谢中瑶97232;金寨,刘晓龙438。山东:崂山,金维坚s.n.。河北:内丘,刘鑫源1283;正定,采集人不明T36;易县,杨朝广3227;兴隆,俞德浚366;迁西,迁西队467。北京:卧佛寺,卧佛寺队476;百花山,王启无60783;十三陵,刘继孟643。辽宁:丹东,王薇1181;凤凰山,J. Sato 5508。
《天目山植物志》苎麻属作者陈征海根据细野麻(B.gracilis)和小赤麻(B.spicata)在叶形、叶片顶端尾状尖头,叶缘牙齿数目等方面的区别特征明确指出细野麻“明显不同于小赤麻”,而不同意将B.gracilis归并于B.spicata[3],我对他的种的概念完全赞同。
2.赤麻 图1:C-D
Boehmeriasilvestrii(Pamp.) W. T. Wang in Acta Phytotax. Sin. 20(2): 204. 1982; Z. T. Yin in Fl. Beijing, rev. ed. 1: 127, fig. 160. 1984; et in Fl. Hebei. 1: 291, fig. 257. 1986; Z. X. Peng & G. L. Zhang in Acta Bot. Bor.-Occ. Sin. 5(2): 130. 1985; S. Z. Liu in Fl. Liaoning 1: 308, pl. 122. 1988; Fl. Shandong 1: 993, fig. 054. 1992; J. Xing & Z. Lu in Fl. Heilongjiang 4: 180, pl. 46. 1992; W. T. Wang in Fl. Reip. Pop. Sin 23(2): 348, pl. 78: 1-2. 1995; B. G. Li in Fl. Hunan 2: 307, fig. 2-228. 2000; Chen, Friis & Wilmot-Dear in Fl. China 5: 172. 2003; S. J. Fan et al. Fl. Laoshan 1: 161, fig. 147. 2003; F. D. Zhang in Fl. Tianjin 98, fig. 95. 2004; Z. X. Peng & G. L. Zhang in Fl. Gansu 2: 246. 2005; C. X. Yang et al. Keys Vasc. Pl. Chongqing 155. 2009; Q. L. Gan, Fl. ZHuxi., Suppl. 178, fig. 207. 2011. ——B.platanifoliaFranch. & Sav. var.silvestriiPamp. in Nouv. Giorn. Bot. Ital., n. s., 22: 278. 1915. ——B.japonica(L. f.) Miq. var.silvestrii(Pamp.) Friis & Wilmot-Dear in Blumea 58(2): 191, fig. 34: g. 2013. Syntypes: 湖北西北部:P. C. Silvestri 4070(FI, non vidi), 4070a(FI, non vidi; photo, PE).
B.tricuspisauct. non(Hance) Makino: Y. C. Zhu in Herb. Pl. N.-E. China 2: 5. 1959; Ohwi, Fl. Japan, rev. ed. 391. 1965; Iconogr. Corm. Sin. 1: 520, fig. 1040. 1972; Fl. Tsinling. 1(2): 115, fig. 97. 1974; S. X. Fu, Fl. Hupeh. 1: 182. 1976; B. Z. Ding et al. Fl. Henan. 1: 299. 1981; W. T. Wang in Acta Bot. Yunnan. 3(4): 410. 1981; Fl. Dalian Distr. 1: 137, fig. 144. 1982; Satake in Satake et al. Wild Fls. Japan 2: 9, pl. 7:1. 1982; Y. N. Lee, Fl. Korea 91. 1996; Ohba in Iwatsuki et al. Fl. Japan 2a: 103. 2006
Perennial herbs. Leaves opposite; blades of apical leaves often undivided, those of other lower leaves 3-lobed and 3-cuspidate, broadly ovate or subpentagonal, 5-8(-13)×4.8-7.5(-13) cm, base subtruncate or broadly cuneate, margins on each side 5-12-dentate, with teeth deltoid or triangular, apical ones often double, 1-10(-15)×1-9 mm. Spikes 4-11(-20) cm long; rachis ca. 0.3 mm in diam.; glomerules along rachis among themselves separate. Persistent pistillate perianth with complanate margins.
地理分布:四川、重庆、湖南、湖北、甘肃南部、陕西南部、河南西部、山东东部、河北、辽宁东部、吉林东南部和黑龙江东南部;朝鲜、日本。
标本登录(均存PE):四川:洪溪,管中天6942;宝兴,C.S.Chang, H. Kim等S11120;茂县,汪发缵22043。重庆:城口,戴天伦101582,102442。湖北:宜昌,赵常明等70;兴山,赵常明等1757;神农架队11380。甘肃:文县,陈伟烈等8812;康县,张志英16558;天水,张珍万195。陕西:石泉,西北大学队238;宁陕,孔宪武3031;佛坪,傅坤俊4616;太白山,王作宾1600;商洛,王作宾16058;蓝田,王作宾15986;渭南,王作宾15794;华山,郝景盛4114。河南:卢氏,刘继孟4571;西峡,河南队1341。山东:青岛,周太炎1010。河北:内丘,刘鑫源745;涿鹿,杨朝广160;昌黎,夏纬瑛1929。北京:西山,王忠涛,马欣堂124;百花山,王文采294;延庆,陈家瑞83015。辽宁:凤凰山,J. Sato 5506,近藤舍吾8282。
3.横断山苎麻 图2:A-C
BoehmeriahengduanshanicaW. T. Wang, sp. nov. Type: 西藏(Xizang):察隅县,上察隅,格容(Chayu Xian, Shangchayu, Gerong),alt. 2200 m,山谷常绿阔叶林下(under evergreen broad-leaved forest in valley),多年生草本(perennial herbs),1973-07-01,自然资源综合考察委员会调查队(Expedition from Commission for integrated survey of natural resources)401(♂,holotypeand isotype, PE)。云南(Yunnan):维西县,白济汍,两岩瓦(Weixi Xian, Baijiwan, Liangyanwa),alt. 1 880 m;常绿阔叶林,水边(by streams under evergreen broad-leaved forest),灌木高1 m,1981-07-10,植物所横断山队(Hengduan Shan Exped. Inst. Bot.)1257(♂, PE).
Species nova haec est arcte affinisB.gracilisC. H. Wright, a qua foliis omnibus apice inaequaliter 2-lobatis et 2-cuspidatis differt.
Perennial herbs or small shrubs. Stems 40-100 cm tall, on internodes very sparsely and on nodes densely strigose. Leaves opposite, those of the same pair slightly unequal in size, the larger one with longer petiole and the smaller one with shorter petiole; blades thinly papery, ovate or elliptic, 3-10×1.5-5.5 cm, apex obliquely and unequally 2-lobed with each lobe at apex cuspidate, base rounded or broadly cuneate, margins on each side 6-13-dentate, with teeth broadly triangular or triangular, 1-5×2-6 mm; surfaces adaxially finely strigose, abaxially on nerves strigose; nervation trinerved, with the midrib having 2-3 pairs of secondary nerves; cystoliths dense, minutely puncticulate; petioles 0.5-5 cm log, very sparsely strigose; stipules brown, narrowly lanceolate, 4.5-6×0.5-0.8 mm, adaxially glabrous, abaxially on midrib strigose. Staminate spikes singly axillary, 2-5.2 cm long, simple or below with a short branch, with 12-30 glomerules; rachis 0.5 mm in diam., strigose; bracts similar to stipules, but smaller, 1-1.8 mm long; glomerules 1-2 mm in diam., 2-5-flowered. Staminate flower bud subglobose, ca. 1 mm in diam., appressed-hispid, tetramerous. Pistillate spikes with pistillate flowers and achenes unknown.
4.小赤麻 图1:E-F
Boehmeriaspicata(Thunb.) Thunb. In Trans. Linn. Soc. 2: 330. 1794; Bl. Mus. Bot. Lugd.-Bat. 2: 220. 1856; C. H. Wright in J. Linn. Soc. Bot. 26: 488. 1899; Chien in Contr. Biol. Lab. Sci. Soc. China 9: 270. 1934; Satake in J. Fac. Sci. Univ. Tokyo. sect. 3, Bot. 4: 482. 1936; Ohwi, Fl. Japan, rev. ed. 391. 1965; S. X. Fu, Fl. Hupeh. 1: 181. 1976; W. T. Wang in Acta Bot. Yunnan. 3(4): 411. 1981; Satake in Satake et al. Wild Fls. Japan 2: 9, pl. 7: 3. 1982; Fl. Jiangsu 2: 86, fig. 383. 1982; W. T. Wang in Iconogr. Corm. Sin., Suppl. 1: 203, fig. 8463. 1982; Yahara in J. Jap. Bot. 59: 129, fig. 1. 1984; Y. Q. He in Fl. Zhejiang 2: 119, fig. 2-159. 1992; W. T. Wang in Fl. Reip. Pop. Sin. 23(2): 350, pl. 78: 3-4. 1995; Y. N. Lee, Fl. Korea 91, fig. 237. 1996; Chen, Friis & Wilmot-Dear in Fl. China 5: 173. 2003, p. p. excl. syn.B.gracilisC. H. Wright,B.paraspicataNakai ex Satake etB.tricuspisvar.unicuspisMakino; Ohba in Iwatsuki et al. Fl. Japan 2a: 103. 2006; Z. H. Chen in Fl. Tianmushan 1: 283, fig. 1-298. 2009; Q. L. Gan, Fl. Zhuxi., Suppl. 177, fig. 206. 2011; Z. R. Xu in Fl. Jiangsu 2: 172. 2013, p. p. excl. fig. 2-279 et syn.B.gracilisC. H. Wright. ——UrticaspicataThunb. in Murray, Syst. Veg., ed. 14, 850. 1784. Lectotype: Japan: C. P. Thunberg 22115(UPS-fide Yahara, 1984).
B.spicata(Thunb.) Thunb. var.akariBl. l. c. Type: Japan: Unknown collector s. n.(holotype, L; isotype, K. ——fide Wilmot-Dear & Friis, 2013).
B.spicata(Thunb.) Thunb. var.teneraBl. l. c. ——B.japonica(L. f.) Miq. var.tenera(Bl.) Friis & Wilmot-Dear in Blumea 58(2): 193, fig. 35. 2013, p. p. excl. syn.B.gracilisC. H. Wright,B.paraspicataNakai ex Satake etB.tricuspisvar.unicuspisMakino. Holotype: Japan: Herb. Lugd.-Bat. 908186(L-fide Yahara, 1984).
Small shrubs. Stems with branches. Leaves opposite; blades ovate-rhombic, 2.4-7.5×1.5-5 cm, apex long cuspidate, base broadly cuneate, margins on each side(3-)4-9(-16)-dentate, with teeth narrowly triangular or triangular, 1-6×0.8-6 mm. Staminate spikes ca. 2.5 cm long, pistillate spikes 4-10 cm long; rachis 0.2-0.4 mm in diam.; glomerules along rachis among themselves more or less separate. Persistent pistillate perianth with complanate margins.
地理分布:四川、重庆、贵州东南部、湖北、江西北部、浙江、江苏北部、河南西部、陕西南部、山西南部、山东东部;朝鲜、日本。
标本登录(均存PE):四川:万源,李培元6115。重庆:石柱,三峡队1920;开县,巴山队2318;奉节,三峡队2874;巫山,周洪富,粟和毅109728;巫溪,三峡队191。贵州:雷山,简焯坡等50769。湖北:恩始,傅国勋1431;神农架,鄂植考队25249。江西:庐山,王名金788。浙江:丽水,章绍尧6078;天目山,刘慎谔146。江苏:江阴,采集人不明20359;云台山,刘昉勋10792。河南:内乡,河南队837;西峡,河南队969;嵩县,河南队905。陕西:宁陕,孔宪武303;太白山,Zhu, Chen, Xu & Wang 1615。山西:垣曲,H. Smith 6373。山东:栖霞,采集人不明179;牟平,刘慎谔,刘继孟1199,1383。
5.密球苎麻 图2:D-I
BoehmeriadensiglomerataW. T. Wang in Acta Bot. Yunnan. 3(4): 408, pl. 1; 1-2. 1981; Fl. Fujian. 1: 474. 1982; X. S. Zhang in Fl. Guizhou. 4: 72. 1989; W. T. Wang in Fl. Reip. Pop. Sin. 23(2): 346, pl. 77:1-2. 1995; B. G. Li in Fl. Hunan 2: 307, fig. 2-230. 2000; Chen, Friis & Wilmot-Dear in Fl. China 5: 172. 2003; Y. Z. Ruan in Fl. Guangdong 6: 102, fig. 44. 2005; W. Q. Chen in Fl. Luochang 1: 604. 2005; W. T. Wang in Fl. Guangxi 2: 870. 2005; C. X. Yang et al. Keys Vasc. Pl. Chongqing 156. 2009. ——B.virgata(G. Forster) Guill. ssp.macrophylla(Hornem.) Friis & Wilmot-Dear var.densiglomerata(W. T. Wang) Friis & Wilmot-Dear in Blumea 58(2): 173, fig. 29. 2013. Holotype: 广西: 大瑶山, 1928-06-24, 辛树帜844(IBSC). Paratypes: 广西:融水,陈少卿14808(IBK,IBSC);东兰,陈立卿92186(IBK)。广东:连南,谭沛祥54321;英德,黄成163450;翁源,刘心祈2102;乐昌,陈少卿2551(上列标本均存IBSC)。福建:南靖,曾沧江417(AU)。江西:龙南,刘心祈4619(IBSC)。贵州:兴仁,贵州队7941(PE)。四川:峨眉山,贺贤育5584(IBSC);长宁,四川中药所515(IMC);珙县,四川中药所275(IMC);合江,四川中药所260(IMC)。
Perennial herbs. Leaves opposite; blades orbicular-ovate, 5-9.4×5.2-8 cm, abaxially sparsely or densely strigose, sometimes glabrous, apex shortly cuspidate or acuminate, base subcordate, rarely rounded, margins on each side 7-17-dentate, with teeth deltoid or broadly triangular, 1-5×1-6 mm. Spikes 2.5-5.5 cm long; rachis ca. 0.5 mm in diam.; glomerules along rachis among themselves tightly contiguous. Persistent pistillate perianth without complanate margins or with narrow complanate margins, or apically complanate.
地理分布:四川、重庆、贵州、云南东南部、广西北部和西南部、广东北部和东北部、福建南部、江西西南部、湖南和湖北西南部。
标本登录(均存PE):四川:雷波,赵清威117431。重庆:江津,刘正宇183054;金佛山,刘正宇10235;酉阳,谭士贤255。贵州:赤水,曹子余,王忠涛292;梵净山,武陵山队295。云南:屏边,大围山,中苏云南考察队3624。广西:兴安,广西队53-947;那坡,方鼎,覃德海79663。广东:蕉岭,李学根202260。福建:漳平,林镕4491。江西:崇义,赖占钧等赣-14。湖南:新宁,罗林波132;绥宁,席建明,万成志81320;永顺,北京队459。湖北:宣恩,李洪钧3231;建始,王从荣1010。
6.异叶苎麻 图2:J-M
BoehmeriaallophyllaW. T. Wang in Acta Bot. Yunnan. 3(4): 112, pl. 1: 7. 1981; et in Fl. Reip. Pop. Sin. 23(2): 350. 1995; Chen, Friis & Wilmot-Dear in Fl. China 5: 173. 2003; W. T. Wang in Fl. Guangxi 2: 870. 2005. Holotype: 广西:凌云,小东兰,1928-08-17,陈立卿02643(IBK).
B.japonicaauct. non(L. f.) Miq.: Wilmot-Dear & Friis in Blumea 58(2): 189. 2013, p. p. quoad syn.B.allophyllaW. T. Wang tantum.
Perennial herbs. Apical leaves alternate, their blades undivided, elliptic, 4-5.7×1.4-1.7 cm, apex shortly cuspidate, base broadly cuneate; other lower leaves opposite, their blades subpentagonal, 6-7×4-5.8 cm, apex unequally 3-lobulate and shortly 3-cuspidate, base broadly cuneate, margins on each side 8-20-dentate, with teeth broadly triangular or triangular, 1-2.5×1.5-4 mm. Spikes 3-8 cm long; rachis 0.3-0.5 mm in diam.; glomerules on rachis among themselves more or less separate. Achenes unknown.
地理分布:广西凌云特有种。
致谢深切感谢钟小红女士打印本文稿件。
1.王文采.中国苧麻属校订[J].云南植物研究,1981,3(3):307-328.
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ArevisionofBoehmeriasect.Duretiaser.Spicatae(Urticaceae)ofChina
WANG Wen-Tsai
(State Key Laboratory of Systematic and Evolutionary Botany,Institute of Botany,Chinese Academy of Scienses,Beijing 100093)
(1)Boehmeriasubgen.Duretiasect.Spicataeis recognized, but relegated to serial rank, and treated as one of the series under Sect.Duretia. (2)In Ser.Spicataeare recognized 6 species, including 1 new species, which are keyed, except the new species briefly described, and illustrated. Besides, the geographical distributions and a distribution map of them are provided. (3)The close relationships among the 6 species are discussed, and several speculations are thought out: In Ser.Spicatae,Boehmeriagracilisis the primitive species, and all other species of this series might originate from it; orB.gracilisandB.densiglomeratamight be a pair of sister groups derived from a common ancestry; orB.allophyllamight be derived fromB.silvestrii.
Urticaceae;Boehmeriasect.Duretiaser.Spicatae;taxonomic revision;China
王文采(1926—),男,研究员,中国科学院院士,从事有花植物分类学研究。
2016-04-18
Q949.737.5
A
10.7525/j.issn.1673-5102.2016.06.002