对翼蓼(蓼科)属级位置的重新探讨

通过对翼蓼Pteroxygonum giraldii Damm. &; Diels及相关属(蓼属Polygonum、何首乌属Fallopia、虎杖属Reynoutria、荞麦属Fagopyrum和金线草属Antenoron)的形态观察、果实解剖学观察、花被片脉序观察、花粉形态、核型分析, 以及ITS序列的分析确定了翼蓼和荞麦F. esculentum Moench较远的亲缘关系。其中我们发现翼蓼果实基部有三个角状物明显不同于其他属果实的形态特征。翼蓼外果皮明显加厚, 并有零星散布的波状内腔, 而荞麦的外果皮很薄, 细胞不等径, 中果皮极厚。以上证据证明了翼蓼与荞麦属亲缘关系较远。在观察荞麦属和翼蓼的花被片脉络时发现了两种不同的脉序类型, 符合将荞麦属分为两个组的划分。翼蓼花被片脉序为三出状, 支持将翼蓼归为Persicarieae族。对翼蓼及荞麦属植物的花粉进行比较后, 发现荞麦属植物的花粉网孔有明显的内凹穿孔而翼蓼却没有, 结果表明二者亲缘关系较远。通过对nrDNA ITS区域序列分析得出翼蓼及相关属为一个单系类群, 含有两个稳定的分支: 第一个分支由蓼属(萹蓄组sect. Avicularia)、何首乌属、虎杖属的植物组成, 第二个分支由蓼属(刺蓼组sect. Echinocaulon、蓼组sect. Polygonum、分叉蓼组sect. Aconogonon、拳参组sect. Bistorta、翼蓼和荞麦属植物组成。同时第二个分支又分成了两个亚分支, 蓼属(刺蓼组、蓼组、分叉蓼组、拳参组)和翼蓼属Pteroxygonum植物属于第一个亚支而荞麦属植物属于第二个亚支。结果支持翼蓼不属于荞麦属的范畴。实验结果显示翼蓼是个单型属, 属于Persicarieae族。     

Generic limits in Polygonurn and related genera (Polygonaceae) on the basis of floral characters

In order to investigate the taxonomic value of a number of characters in the elucidation of relationships within the collective genus Polygonum L., and segregate genera Atraphaxis L., Calligonum L., Pteropyrum Jaub. & Spach., Oxygonum Burch., Fagopyrum Mill., Harpagocarpus Hutch. & Dandy and Polygonella Michx., 83 species of the tribe Polygoneae emend. Jaretzky were studied. The following floral characters were examined: stamen number, insertion and external morphology; structure and morphology of the floral nectaries; vasculature and surface morphology of the tepals.
Problems in floral morphology in Polygonaceae are discussed, together with relationships within the Family. The tribal division proposed by Haraldson (1978) is supported, but a new and different delimitation of genera is proposed. The present data support a division of Polygonum L. sensu lato and its related taxa into two tribes: Polygoneae (containing Polygonum L. sensu stricto, Fallopia Adans., Oxygonum Burch., Pteropyrum Jaub. & Sparh., Atraphaxis L. and Calligonum L.), and Persicarieae (containing Fagopyrum Mill., Harpagocarpus Hutch. & Dandy, Persicaria Mill. and Koenigia L.). The genus Reynoutria Houtt. is included as a section of Fallopia Adans.; Bislorta Mill., Aconogonon Reichenb. and Tovara Adans. are included within Persicaria Mill.
A number of new combinations are proposed in Persicaria and Fallopia .     

中国荞麦属花粉形态及花被片和果实微形态特征的研究

研究了中国荞麦属Fagopyrum　Mill.的花粉形态、花被片和果实微形态特征。根据沟膜的有无,花粉可以明显地区分为具沟膜者和不具沟膜者2类。透射电镜下显示,荞麦F. esculentum Moench的花粉外壁分化成两个明显的层次,即外壁外层和外壁内层。其外壁外层由覆盖层及柱状层组成,无基层。对花被片腹面的扫描电镜观察表明,根据细胞形状和表面纹饰,这些种类的花被片可分为3类。对果实表面的扫描电镜观察表明,这些种类的果实纹饰也可分为3类。根据上述结果,讨论了荞麦属的种间关系及花粉形态、花被片和果实一些微观性状的演化趋势。荞麦属的花粉形态极为一致,其外壁纹饰均为细网状,萌发孔为三孔沟,这支持将荞麦属从蓼属中分出独立成属的观点。本文还根据花粉形态特征及花被片和果实微形态特征讨论了国产荞麦属的种间关系,探讨了一些特征的演化趋势。     

基于核糖体DNA内转录间隔区ITS序列的广义蓼属及近缘属分子系统学研究

通过对广义蓼属及近缘属共32个代表种内转录间隔区ITS序列的分子系统学分析,尝试研究备受争议的广义蓼属及近缘属的物种族、属、组级的划分问题,结果显示,广义蓼属在系统发育树上并不能形成一个单系类群,这些物种共聚为3大支,分别对应春蓼族、蓼族及荞麦族,其中荞麦属与翅果蓼属形成了一支独立于春蓼族及蓼族之外的类群。在春蓼族中,冰岛蓼属与分叉蓼组形成一个单系类群。     

基于叶绿体trnL-F,rbcL序列和核核糖体ITS序列探讨蓼属（蓼科）头状蓼组的系统发育

选用叶绿体基因trnL-F,rbc-L序列和核核糖体ITS序列对蓼属头状蓼组的分类和系统发育进行了分析。以药用大黄为外类群,用最大简约法对3个片段的单独和联合矩阵分别构建系统发育树。结果表明,头状蓼组是一个较自然的类群,与春蓼组、刺蓼组和金线草属的亲缘关系较近;支持头状蓼组作为一组放在春蓼属,细茎蓼和蓝药蓼从头状蓼组转移至冰岛蓼属;不支持将刺蓼组并入头状蓼组;对于小叶蓼的系统学位置有待于进一步研究。     

Pollen morphology of the genera Polygonum s. str. and Polygonella (Polygoneae: Polygonaceae)

The pollen of 30 taxa (27 species, one subspecies and two varieties) in two genera, viz Polygonum s. str. and Polygonella was investigated with LM and SEM, and some selected taxa with TEM. In all genera investigated the pollen is prolate to spheroidal, and the aperture is mostly tricolporate, rarely panto-hexacolporate (especially Polygonum section Polygonum). The exine sculpturing pattern is the most variable feature. Three types of exine can be recognized. Type 1 (Avicularia-Type, sensu Hedberg) - All species of section Polygonum and section Tephis share the smooth tectate exine with spinules, sometimes the surface is more or less rough (Polygonum afromontanum in section Tephis). Type 2 (Pseudomollia-Type, sensu Hong) - Pollen of Polygonum molliaeforme (section Pseudomollia) has the exine, which is verrucose on both poles and nearby the mesocolpium, and mostly psilate around the ectoaperture. Type 3 (Duravia-Type, sensu Hedberg) - Pollen grains of Polygonum section Duravia and Polygonella have the exine which is semitectate-reticulate at the mesocolpium and the poles, and rugulate/reticulate or sometimes foveolate with microspinules around the ectoapertures. The pollen grains in four taxa (viz Polygonum section Pseudomollia, P. section Duravia and genus Polygonella) have a well-marked dimorphism of the ektexine, which is considered to be a synapomorphic condition. The differences of pollen grain between the genus Polygonella and Polygonum section Duravia are almost non existent and clearly interrelated. It is therefore postulated that the similarity in pollen of both taxa is not the result of convergency, but is interpreted as a homology. It is noteworthy that the pollen of Polygonum molliaeforme (section Pseudomollia) appears as intermediate between the Avicularia-type and the Duravia-type, and is well supported the value of separated section for its own. Additionally, in TEM, some exine ultrastructures (e.g. columellae, foot layer, endexine) appear to be valuable characters for comparison between/among taxa. The systematic potentialities of the pollen data of the studied taxa at various systematic levels are also discussed.     

中药蓼大青叶及其伪品的nrDNA ITS区序列的测定

中药蓼大青叶的基源为蓼科蓼属植物蓼蓝（Polygonum tinctorium Ait),同属植物的水蓼（辣蓼）,（P.hydropiperL.)的叶常作其伪品,本文采用PCR直接测序法,首次测定了它们的核糖体DNA ITS区序列,结果显示,两者充列长度（ITS1,5,8S,ITS2）均为646bp,序列间共有28个变异位点,本研究为产蓝植物的DNA分子鉴定提供了必要的序列资料。     

Justification for the generic status of Courtoisina (Cyperaceae)

In the past, the two species of Courtoisina were placed either in Cyperus L., Mariscus Vahl, or in Courtoisia Nees (now Courtoisina Sojak). While these species show undoubted external similarities to both Cypm and Muriscus , they differ by their widely winged glumes and the arrangement of the veins of the glumes. They also differ profoundly from Mariscus and most species of Cypm in anatomical characteristics, and it is concluded that these differences warrant separate generic status.     

On a new subsection of Pandanus section Cauliflora (Pandanaceae) with paniculate staminate inflorescence structure,distinctive leaf anatomy and pollen morphology

A new subsection, Pandanus section Cauliflora subsection Paniculiflora, is described to contain P. halleorum, on the grounds of the paniculate structure of its staminate inflorescence, a feature not known elsewhere within the genus Pandanus. Both leaf anatomy and pollen morphology appear to support this taxonomic separation.     

中国蓼属萹蓄组植物果实形态的研究

应用解剖镜和电子扫描显微镜对中国产蓼属萹蓄组Polygonum sect.Polygonum 16种1变种及1变型植物的果实形态进行了研究。发现该组5种一年生植物的果实具有两型性,即有长、短两种果实类型。短果类型为卵形,具三棱,包于宿存花被内或与花被近等长,呈黑褐色。在解剖镜下观察到其表面具小点、无光泽,而在电子扫描显微镜下观察到其表面具有散乱分布或排列成行的瘤状颗粒;长果类型为狭卵形,具三棱,长于宿存花被,黄褐色或淡黄绿色,在解剖镜下观察到其表面平滑、有光泽,而在电子扫描显微镜下观察到其表面上部具有散乱分布的瘤状颗粒,中、下部则具不规则洼点、星花状褶皱或浅波纹状。果实具两型性的植物有萹蓄P.aviculare、乌鲁木齐萹蓄P.urumqiense、展枝萹蓄P.patulum、直立萹蓄P.avicularef.erectum和尖果萹蓄P.rigidum。果实微形态可分为两种类型：1.果实表面无瘤状颗粒或疣状突起、有光泽,此类型又可以分为2种亚类型：（1）果实表面平滑、浅波纹状,如岩萹蓄P.cognatum、松叶萹蓄P.acerosum和帚萹蓄P.argyrocoleum;（2）果实表面具浅洼点或洼点,如铁马鞭P.plebeium和针叶萹蓄P.polycnemoides。2.果实表面有瘤状颗粒或疣状突起,无或具微光泽,此类型又可分为4种亚类型：（1）果实表面具疣状突起,如圆叶萹蓄P.intramongolicum.;（2）果实表面具散乱分布的瘤状颗粒,如普通萹蓄P.humifusum和展枝萹蓄的短果;（3）果实表面具有纵行排列的瘤状颗粒,如萹蓄和乌鲁木齐萹蓄的短果;（4）果实表面上部具散乱分布的瘤状颗粒,中、下部具不规则洼点、星花状褶皱或浅波纹状,如萹蓄、展枝萹蓄、直立萹蓄和尖果萹蓄的长果。因此果实两型性和果实微形态特征对蓼属萹蓄组种、变种的鉴定具有较大的分类学价值。基于果实微形态特征以及其他形态特征,确定了新种晖春萹蓄Polygonum huichunense F.Z.Li,Y.T.Hou＆C.Y.Qu,同时恢复褐鞘萹蓄Polygonum aviculareL.var.fusco-ochreatum（Kom.）A.J.Li为原种,即Polygonum fusco-ochreatum Kom.。     

The genus Koenigia L. emend. Hedberg (Polygonaceae)

Pollen morphological studies revealed the occurrence of the characteristic spinulose pollen type of Koenigia not only in the three species earlier recognized in the genus (A" islandka, K. nepalensis and K. pilosd) but also in three additional species earlier treated under Polygonum , viz. K delicatula (Meisn.) Hara, K. forrestii (Diels) Mesicek & Soják, and A" nummularifolia (Meisn.) Mesicek & Soják. Further studies of flower morphology, fruit and petiole anatomy, basic chromosome number, etc., revealed additional similarities between those species, which led to a taxonomic revision of the genus Koenigia. This genus seems to be most closely related to Persicaria Mill, sections Cephalophilon (Meisn.) Gross and Echinocaulon (Meisn.) Gross, with Koenigia delicatula as a connecting link. There arc also interesting similarities with the genus Aconogonon (Meisn.) Rchb. Koenigia exemplifies the derivation from montane ancestors of a high mountain-dwelling genus displaying adaptive radiation to fit diverse alpine niches. Five species out of six are confined to high mountain areas in southeastern Asia, primarily in the Himalayas, whereas the sixth has spread to Arctic and alpine areas in the northern hemisphere and even penetrated to southern South America. The latter species shows progressive reduction in size in combination with adaptation to a very short summer under severe climatic conditions.     

Unexpected high divergence in nrDNA ITS and extensive parallelism in floral morphology of Pedicularis (Orobanchaceae)

Internal transcribed spacer (ITS) region sequences of nuclear ribosomal DNA (nrDNA) were used in the phylogenetic reconstruction of Pedicularis, a genus with strong adaptive radiation. Forty-two species representing 12 greges of the genus were all, except P. resupinata, sampled from the Hengduan Mountain region, China. A high level of ITS sequence variation was found among the species distributed in such a small geographical area, which had been rarely reported in other groups. The great ITS divergence in Pedicularis could be explained by a relatively ancient origin and diversification of the genus followed by migration of different floristic components into the Hengduan Mountains, or accelerated rates of molecular evolution in parasitic lineages of Orobanchaceae. In the present ITS phylogeny, almost all the main clades are not consistent with the high hierarchical taxa within Pedicularis, which implies that significant parallel evolution occurred in floral morphology of the genus, and that undue attention has been paid to corolla characters in the intragenus classifications.We thank the two anonymous reviewers for valuable comments and suggestions; Professor Guo You-Hao, Drs. Tan Dun-Yan, Luo Yi-Bo, Bao Ying, Luo Yan, Mr. Mao Jian-Feng and Mr. Qin Wei for their great help in the field collection; Drs. Song Bao-Hua, Wei Xiao-Xin and Miss Sun Ying-Xue for their help in the laboratory work. This study was supported by State Key Basic Research and Development Plan (Grant No. G2000046804) and the National Natural Science Foundation of China (Grant No. 30121003).     

A systematic study of North American Angelica species (Apiaceae) based on nrDNA ITS and cpDNA sequences and fruit morphology

Angelica is a taxonomically complex genus widespread throughout the North Temperate Zone. Previous phylogenetic studies of the genus have focused primarily on its East Asian species. The relationships among its North American members, the monophyly of these species, and the value of fruit morphology in circumscribing its taxa have yet to be examined. This study represents the most comprehensive sampling of Angelica to date (100 species) and includes all 26 species in North America. Relationships are inferred using Bayesian inference, maximum likelihood, and maximum parsimony analyses of ITS sequences and, for multiple accessions of each North American species, cpDNA ndhF-rpl32, rpl32-trnL, and psbM-psbD sequences. The fruit morphological characters examined were those considered phylogenetically important in East Asian Angelica. The results revealed that the North American species fell into three major clades: North American Angelica clade, Archangelica clade, and the Eurasian Angelica clade. Angelica dawsonii has affinities with Lomatium brandegeei. Fourteen species within the North American Angelica clade were strongly supported as monophyletic. Two paraphyletic species resulted in new combinations in A. lineariloba and A. venenosa. Conflict between the ITS-derived and cpDNA-derived phylogenies and the lack of resolution in portions of the trees may be due to chloroplast capture and rapid species radiation. Fruit morphology supported some interspecific relationships based on molecular data, and relationships revealed by ITS and cpDNA data were roughly in accordance with fruit classification type and geographic distribution region, respectively. A diagnostic key based on fruit morphology is provided for the identification of the North American Angelica taxa.     

Patterns and consequences of self-pollen deposition on stigmas in heterostylous Persicaria japonica (Polygonaceae)

In order to test the "anti-interference" hypothesis for adaptive significance of reciprocal herkogamy, patterns of illegitimate pollination and effects of self-pollen on the legitimate pollination and fertility were investigated in a naturally pollinated experimental population of distylous Persicaria japonica. Pollen deposition was compared among the emasculation treatments, i.e., emasculation of a single flower from individual inflorescences, emasculation of all the flowers of individual inflorescences, and no emasculation control. In both morphs, considerable illegitimate pollination was found to occur in all the treatments, and there was no significant difference in incompatible pollen load among the treatments. Therefore, it is suggested that herkogamy of P. japonica can sufficiently reduce both intraflower and intra-inflorescence self-pollinations, but not interinflorescence geitonogamous and-or interclonal illegitimate pollination. Measurements of pollen load after the repetitive pollen addition showed that space on the stigma surface may not limit the legitimate pollination under natural pollination conditions. Seed sets after legitimate pollination following prior self-pollination did not differ from the controls without self-pollination. Therefore, there is little possibility that incompatible pollen load interferes with either pollination or fertilization by compatible pollen, suggesting that "anti-interference" is unimportant for adaptive significance of reciprocal herkogamy at least in P. japonica.     

A phylogenetic study ofPolygonum sect.Tovara (Polygonaceae) based on ITS sequences of nuclear ribosomal DNA

Polygonum sect.Tovara comprises three morphologically very similar species;P. virginianum,P. filiforme, andP. neofiliforme. Sequences of internal transcribed spacers (ITSs) of nuclear ribosomal DNA of these were determined to examine phylogenetic relationships and the levels of differentiation among them. The size of ITS 1 was 241 bp inP. filiforme andP. neofiliforme, and 242 bp inP. virginianum. The size of ITS 2 was 243 bp, and that of the 5.8S rRNA coding region was 163 bp. The ITS sequences clearly separate North AmericanP. virginianum from the eastern Asian species. Nucleotide divergence between them ranges from 3.3% to 3.8% for ITS 1 and from 9.3% to 10.7% for ITS 2. The molecular data also revealed that two eastern Asian species are closely related but should be treated as distinct species.     

Molecular authentication of Anthemis deserti Boiss. (Asteraceae) based on ITS2 region of nrDNA gene sequence

The dried plant material of medicinally important Anthemis deserti Boiss. (family: Asteraceae) especially when it remains in the powdered form often look similar to Anthemis melampodina Del.; and therefore, difficult to distinguish, finally lead to chances of adulteration. The adulteration in medicinal plants effects on the efficacy of the drugs. The molecular authentication of herbal plant materials such as based on the internal transcribed spacer 2 (ITS2) sequences of nuclear ribosomal DNA (nrDNA) is considered as more reliable method compared to other the biochemical or histological methods. The present study aims to molecular authentication ofA. deserti based on molecular phylogenetic analyses of ITS2 gene sequence of nrDNA region. The ITS2 region of nrDNA of A. deserti were sequenced, and the molecular phylogenetic analyses were performed together with the GenBank sequences. The Maximum Parsimony tree revealed the close relationships of A. deserti with A. melampodina; however, the Neighbor-Joining and Maximum Likelihood tree clearly revealed that A. deserti is distinct from A. melampodina, which is also supported by the differences in nucleotides at five diffident positions (i.e. 22, 28, 87, 175 and 198) in the DNA sequence alignment.     

中国蓼族(蓼科)植物区系的研究

根据蓼族植物的生境及分布状况,在全国范围内进行多年的野外考察和标本采集,同时参考前人的研究成果对中国蓼族植物区系进行深入研究。结果表明,中国蓼族植物有12属、170余种及变种,包括发现的多个新种(其中3新种已正式发表,1新种已被植物分类学报接受);同时揭示了其区系特点:(1)分布广泛,在全国各省区从低海拔到高海拔的区域内均有分布;(2)区系具温带性质,以北温带成分为主;(3)全国范围内,四川拥有该族在中国分布的全部12属,种及变种数占全国的51.7%,特有种数占全国的44.4%;云南分布11属占全国的91·7%,种及变种数占全国的62.2%,特有种数占全国的55.6%。这表明西南地区的云南、四川是中国蓼族植物的现代分布中心和多样化中心;(4)中国有该族植物12属,170余种及变种,分别占世界属种数的85.7%和48.0%～58.2%。可见中国(尤其是四川、云南)是世界蓼族植物的现代分布中心和多样化中心。