基于多基因序列和形态性状的牡丹组种间关系

牡丹被认为是中国的国花,具有很高的医学、观赏和经济价值.野生牡丹被认为是栽培牡丹的野生祖先,因此弄清牡丹组的种间亲缘关系具有重要的理论和实践意义.由于受到信息量的限制,根据单基凼数据或形态数据往往无法对牡丹组的种间关系得到明确的结果.本研究用12份样品代表野生牡丹组(Paeonia section Moutan DC.,Paeoniaceae)8个种,利用包括核基因(Adh1A、Adh2和GPAT)和叶绿体基因(trnS-trnG和rps16-trnQ)的DNA序列以及形态性状的多套数据来探讨野牛牡丹的种间关系.合并分析得到具高支持率的牡丹组物种间的系统发育关系.结果表明,芍药属牡丹组8个野生种分为两个亚组,即肉质花盘亚组subseet.Delavayanae和革质花盘亚组subsect.Vaginatae.肉质花盘亚组包括滇牡丹P delavayi和大花黄牡丹P.ludlowii;革质花盘亚组包括其余6个种.革质花盘亚组中,四川牡丹P.decomposita ssp.decomposita和紫斑牡丹P. rockii ssp. rockii关系密切;卵叶牡丹P.qiui和矮牡丹P. jishanenMs关系密切;银屏牡丹P. suffruticosa ssp.yinpingmudan与风丹P. ostii关系 密切,并且后两个分支为姊妹群.     

芍药属牡丹组基于形态学证据的系统发育关系分析

对芍药属牡丹组Paeonia L.sect.Moutan DC.（全部野生种）40个居群进行了基于形态学证据的系统学分析,试图建立组内种间的系统发育关系。利用PAUP （4.0）计算机程序分别构建了建立在25个形态学性状基础上的所有研究类群的距离树（UPGMA、NJ）和最大简约树（MP）。所得树的拓扑结构基本一致,差异只发生在距离树和简约树之间,在由形态和细胞学关系都很近的5个种（牡丹P.suffruticosa、矮牡丹P.jishanensis、卵叶牡丹P.qiui、紫斑牡丹P.rockii和凤丹P.o     

芍药属牡丹组分类补注

把沈保安2001年发表的6个新组合和新等级处理为异名,这些异名是Paeonia linyanshanii (S. G. Haw &; L. A. Lauener) B. A. Shen (不合法名),P. linyanshanii ssp. taibaishanica (D. Y. Hong) B. A. Shen (不合法名),P. ostii T. Hong &; J. X. Zhang ssp. lishizhenenii (lishizhenii) (B. A. Shen) B. A. Shen (=P. ostii), P. delavayi Franch. ssp. angustiloba (Rehder &; E. H. Wilson) B. A. Shen 和ssp. lutea (Delavay ex Franch.) B. A. Shen (=P. delavayi)以及P. delavayi ssp. ludlowii (Stern &; Taylor) B. A. Shen (=P. ludlowii (Stern &; Taylor) D. Y. Hong)。     

Systematic studies on Paeonia sect. Moutan DC. based on RAPD analysis

Plants in Paeonia sect. Moutan DC., whose wild types are endemic to China, are deciduous subshrubs. Taxonomic treatments of most species in this section have long been in dispute. To address this question, both intraspecific and interspecific relationships of the species in this section were analyzed using RAPD markers. The dendrogram constructed by UPGMA showed that the accessions of the same species were always grouped together earlier than those of different species. The intraspecific similarity coefficients ranged from 0.60 to 0.90, grouping precisely those species of the same subsection together. Hence, the seven species under question can be well distinguished from each other. The similarity coefficient between P. delavayi and P. ludlowii was 0.60, and they were clustered in a clade. The similarity coefficients between P. jishanensis and the three species P. rockii, P. ostii, P. qiui, and between P. jishanensis and P. decomposita were both 0.48. These five species were clustered in another clade. These two clades corresponded well to Subsect. Delavayanae and Subsect. Vaginatae. Our results support the taxonomic treatment of Sect. Moutan re- cently proposed by Hong (1998,1997).     

芍药属牡丹组的系统学研究——基于RAPD分析

芍药属牡丹组(Paeonia sect．Moutan DC)是落叶亚灌木,其野生类群为我国特有。长期以来不同 学者根据形态性状对这个组中种的分类处理不断修正,不断有新种描述。我们采用RAPD标记分析了 牡丹组种内与种间遗传关系。从10个RAPD引物获得121个多态位点。用UPGMA方法构建的树系 图表明每个种的所有个体都各自聚为一支,种内的相似性系数为0.60～0.90,因此现有的7个种能很好 地区分开来。P．delavayi与P．ludlowii相似性系数为0.60,聚为一支;P．jishanensis与P．rockii、P. ostii、P．qiui以及P．decomposita之间的相似性系数为0.48,聚为一大支。这两支与肉质花盘亚组和革 质花盘亚组相对应。这些结果与洪德元根据形态性状对该组所做的分类处理基本相符。我们认为RAPD技术用于牡丹基因组分析是灵敏而行之有效的工具。     

芍药属牡丹组的分类历史和分类处理

芍药属Paeonia木本类群，即牡丹组sect．Moutan DC.，为中国特有；广泛栽培的牡丹也起源于 中国。遗憾的是我国学者对牡丹类群的分类一直未予重视，野生类群的标本也很少。90年代以来对该 类群的分类突然热起来，中外学者连续发表了多个新种、新亚种和新等级，但仍然没有一个全面的修订。 在美国地理学会基金资助下，我们几乎踏遇了该类群分布区的各个角落，拜访了所有模式产地(仅一个 例外)，对64个居群进行了考察和取样。在性状分析的基础上检查了各个分类群的问题，考证了全部模 式，最后对该类群作出分类修订。本文是对分类历史的回顾，并表述了我们的分类处理：牡丹组共有8个种，其中3个种各包含两个亚种，另有两个杂种。     

Taxonomical history and revision of Paeonia sect. Moutan (Paeoniaceae)

The taxonomical history of the woody group(peonies) in the genus Paeonia is reviewed in the present paper. The group is endemic to China, but Paeonia suffruticosa is commonly cultivated throughout the north temperate region and has long been known as “the King of Flowers” in China. However, the taxonomy of the group had been neglected before the 1990' s. Since 1990, a number of new species and subspecies have been published. With the support from the National Geographic Society, the senior author of the present paper and his coworkers have made expeditions to all parts of the distribution area of the group, visited all the type localities and sampled a total of 64 populations. Based on the character analysis and examination of the type specimens or photos, each taxon is reviewed with its name checked nomenclaturally, and finally the taxonomy of the whole group is revised. As a result, eight species, three of which each contains two subspecies, are recognized. They are Paeonia suffruticosa Andrews subsp. suffruticosa and subsp. yinpingmudan D. Y. Hong, K. Y. Pan et Z. W. Xie, P. jishanensis T. Hong et W. Z. Zhao, P. qiui Y. L. Pei et D. Y. Hong, P. ostii T. Hong et J. X. Zhang, P. rockii (S. G. Haw et L. A. Lauener )T. Hong et J. J. Li subsp. rockii and subsp. taibaishanica D. Y. Hong, P. decomposita Hand.-Mazz. Subsp. decomposita and subsp. rotundiloba D. Y. Hong, P. delavayi Franeh. and P. ludlowii (Stern et Taylor)D. Y. Hong. P. baokangensis Z. L. Dai et T. Hong and P. yananensis T. Hong et M. R. Li are considered as hybrids between P. rockii and P. qiui and between P. rockii and P. jishanensis respectively. In addition, the reduction of a number of names recently published to synonyms is explained.     

芍药属牡丹组分类新注

自我们近年发表一系列牡丹组分类文章以来, 国内外基本上赞同我们8个种的分类系统,但对一些问题仍有不同见解。本文进一步申述牡丹Paeonia suffruticosa Andrews 是一个独立的种而不是人工杂种, 以及银屏牡丹P. suffruticosa ssp. yinpingmudan是牡丹P. suffruticosa的野生类型而不是逸生的P. ostii的理由。上述论点也得到了分子树的支持。P. jishanensis T. Hong &; W. Z. Zhao是一个合法名称, 而P. spontanea (Rehder) T. Hong &; W. Z. Zhao则确实是一个多余名。太白山紫斑牡丹的学名应是P. rockii ssp. atava (Brühl) D. Y. Hong &; K. Y. Pan, 因此P. moutan Sims ssp. atava Brühl不应是可疑的分类群。本文还对Halda的6个组合和两个杂交种名作了处理。结果, 本文包括了5个新异名和一个新组合。     

芍药属牡丹组（Paeonia sect. Moutan）种间关系的分子证据：GPAT基因的PCR-RFLP和序列分析

为了探讨芍药属牡丹组Paeonia sect. Moutan的种间关系,对采自15个野生居群,代表牡丹组全部8个野生种的15份材料的GPAT 基因片段（外显子5和6之间2 kb的内含子）进行了PCR-RFLP分析,并对代表牡丹组全部8个野生种的9份材料进行了测序。 根据12个限制性内切酶的PCR-RFLP数据,使用Network 3.0计算机程序的RM (reduced-median)法建立了牡丹组种间亲缘关系网络树。同时根据8个种9份材料的GPAT基因片段序列,利用PAUP*4.0计算机程序建立了牡丹组GPAT基因的最大简约（MP）树和邻接(NJ)树。结果获得了具有很高自展值支持、分辨良好的牡丹组种间关系（GPAT基因）树。最重要的是,该基因树所显示的牡丹组种间关系与根据形态学证据提出的牡丹组的种间关系基本吻合,并得到其他研究证据的支持。根据这一结果,对牡丹组的种间关系进行了详细的讨论。     

Chloroplast DNA phylogeny, reticulate evolution, and biogeography of Paeonia (Paeoniaceae)

The coding region of the mat K gene and two intergenic spacers, psb A-trn H and trn L(UAA)-trn F(GAA), of cpDNA were sequenced to study phylogenetic relationships of 32 Paeonia species. In the psb A-trn H intergenic spacer, short sequences bordered by long inverted repeats have undergone inversions that are often homoplasious mutations. Insertions/deletions found in the two intergenic spacers, mostly resulting from slipped-strand mispairing, provided relatively reliable phylogenetic information. The mat K coding region, evolving more rapidly than the trnL-trn F spacer and more slowly than the psb A-trn H spacer, produced the best resolved phylogenetic tree. The mat K phylogeny was compared with the phylogeny obtained from sequences of internal transcribed spacers (ITS) of nuclear ribosomal DNA. A refined hypothesis of species phylogeny of section Paeonia was proposed by considering the discordance between the nuclear and cpDNA phylogenies to be results of hybrid speciation followed by inheritance of cpDNA of one parent and fixation of ITS sequences of another parent. The Eurasian and western North American disjunct distribution of the genus may have resulted from interrruption of the continuous distribution of ancestral populations of extant peony species across the Bering land bridge during the Miocene. Pleistocene glaciation may have played an important role in triggering extensive reticulate evolution within section Paeonia and shifting distributional ranges of both parental and hybrid species.     

薯蓣属根状茎组基于3个cpDNA序列的分子系统研究

Seventeen species, one subspecies and one variety of Dioscorea sect. Stenophora Uline were investigated for their phylogenetic relationships based on a sequence analysis of chloroplast matK and rbcL genes and trnL-F intergenic spacer by maximum parsimony and maximum likelihood methods. The results showed that (a) sect. Stenophora was a strongly supported monophyletic group; (b) D. rockii, D. membranacea, D. banzhuana, and D. simulans formed a moderately supported monophyletic group, and D. prazeri was weakly supported to be sister to this group; (c) D. althaeoides and D. nipponica ssp. nipponica formed a moderately supported clade, and D. nipponica ssp. rosthornii was not a member of this clade; (d) D. zingiberensis and D. sinoparviflora showed a moderate to strong sister relationship; and (e) D. collettii var. hypoglauca and D. collettii var. collettii were sister to each other, but with only weak support.     

Reexamination of series relationships of South American wild potatoes (Solanaceae: Solanum sect. Petota): evidence from chloroplast DNA restriction site variation

Chloroplast DNA (cpDNA) restriction enzyme site analysis was used to test hypotheses of series and superseries affiliations of 76 taxa, representing 11 of the 13 South American series (material unavailable for two series) of wild potatoes (Solanum sect. Petota) recognized in the latest classification by Hawkes. The cladistic results, combined with those from earlier cpDNA studies of 30 taxa of the Mexican and Central American species (representing eight series; ser. Conicibaccata and ser. Tuberosa have representatives in Mexico and in South America), support four main clades for 17 of the 19 series examined in sect. Petota: (1) the Mexican and Central American diploid species, exclusive of S. bulbocastanum, S. cardiophyllum, and S. verrucosum, (2) S. bulbocastanum and S. cardiophyllum (ser. Bulbocastana, ser. Pinnatisecta), (3) South American diploid species constituting all of ser. Piurana, but also members of ser. Conicibaccata, ser. Megistacroloba, ser. Tuberosa, and ser. Yungasensia, (4) all Mexican and Central American polyploid species (ser. Longipedicellata, ser. Demissa), S. verrucosum (diploid Mexican species in ser. Tuberosa), and South American diploid and polyploid members of ser. Acaulia, ser. Circaeifolia, ser. Commersoniana, ser. Conicibaccata, ser. Cuneoalata, ser. Lignicaulia, ser. Maglia, ser. Megistacroloba, ser. Tuberosa, and ser. Yungasensia. Each of these clades contains morphologically and reproductively very diverse species, and there are no evident morphological features that unite members within a clade to therefore distinguish them. These results strongly suggest a need for a reevaluation of the series and superseries classifications of sect. Petota.     

川赤芍的化学成分研究

从川赤芍Paeonia anomala subsp·veitchii根皮的70%丙酮提取物中,分离鉴定了22个化合物,其中包括一个新的24,30位降常春藤皂苷三萜衍生物,命名为paeonenolide H(1)。化合物2,4,9,10为首次从该植物中分离得到。     

Cytogeography and taxonomy of the Paeonia obovata polyploid complex (Paeoniaceae)

 The taxonomy of the Paeonia obovata polyploid complex in E-Asia has been controversial with treatments ranging from one species and two varieties to five species and numerous infraspecific taxa. Extensive observations in the field, analysis of morphological characters based on 32 populations sampled, and the examination of a large amount of herbarium specimens show that all the characters used by previous authors for distinguishing the species are variable between or even within populations: Leaf position, petal colour, stamen number (from 21 to 110 in the red-flowered and from 58 to 239 in the white-flowered form), filament and anther colour, style length as well as follicle number and position. Therefore, all these characters are of no value for species delimitation. However, an extensive chromosome survey and the analysis of morphological characters show that the ploidy level is basically correlated with the geographical distribution and with the indumentum on the lower leaf surface. Thus, the recognition of one species with two subspecies is proposed: Paeonia obovata subsp. obovata, mostly diploid, widely distributed in the north, east and south of the distribution range and glabrous to sparsely pubescent or hirsute on the lower leaf surface, and subsp. willmottiae, tetraploid, limited to the west and with leaves densely hirsute or pubescent below. Received October 30, 1998 Accepted June 21, 2000     

基于ITS和matK基因对牡丹组序列分析及其亲缘关系的研究

以芍药属牡丹组全部9个野生种、5个紫斑牡丹栽培品种及3个中原牡丹品种为试材,进行核糖体内转录间隔区（ITS）和叶绿体成熟酶K （matK）基因片段测序分析,探讨ITS和matK序列为牡丹组所有野生种种间关系提供分子证据。从GeneBank中选取了1个牡丹及3个外类群芍药、川赤芍和草芍药的ITS及matK序列。对试验样品进行DNA提取、PCR扩增并双向测序得到44条序列,人工校正后将所得44条序列进行比对;计算碱基组成频率、变异位点、简约信息位点数、转换/颠换比率、种内及种间遗传距离,以邻接法进行系统发育分析。结果表明,牡丹组所有个体ITS序列长度在750~800 bp,含有86个多态位点,74个简约信息位点,转换/颠换比率（R）为1.2;而matK序列含有20个简约信息位点,转换/颠换比率（R）为1.7。ITS序列分析将牡丹组野生种分为两大枝,稷山牡丹、紫斑牡丹、卵叶牡丹和杨山牡丹聚为一枝,狭叶牡丹、滇牡丹、黄牡丹和大花黄牡丹聚为另一枝,这两枝分别与革质花盘亚组和肉质花盘亚组相对应,而四川牡丹位于革质花盘亚组最底端,支持前人研究将四川牡丹归为革质花盘亚组。matK序列分析的牡丹组野生种间遗传距离结果不理想,未能清晰的表明野生种之间的亲缘关系。由此说明,ITS序列更适合牡丹组野生种间亲缘关系的研究分析。     

中国薹草属(莎草科)研究随记(Ⅳ).隐匿薹草组的分类修订(英文)

隐匿薹草组(Carex sect.Infossae)为最近建立的薹草属(Carex)的组,当时报导分布于我国安徽和江苏,有2种和1变种,即隐匿薹草(C.infossa)、显穗薹草(变种)(C.infossa var.extensa)和矮秆薹草(C.minuticulmis)。根据描述,另有15种和2变种应归属本组,这些分类群具有较为一致的特征:果囊椭圆球状卵球形或卵球形,通常疏被短毛或近无毛,先端渐狭成中等长的喙,喙口具2小齿;瘦果卵球形,先端钝或微凹。通过文献考证、标本查阅、野外采集,并结合果囊和瘦果的扫描电镜观察,对隐匿薹草组所涉及的种(含种下类群)进行了分类修订。该组仅有2种:百里薹草(C.blinii)和尖叶薹草(C.oxyphylla)。将Carex elmeri、C.granifera、矮秆薹草(C.minuticulmis)、上海薹草(C.shanghaiensis C.blinii subsp.shanghaiensis)、C.sublateralis、C.taihokuensis和锐果薹草(C.tatsutakensis)并入百里薹草;将沟囊薹草(C.canaliculata)、樟木薹草(C.changmuensis)、C.distantiflora、隐匿薹草(C.infossa)、显穗薹草(C.infossa var.extensa)、C.lateralis、C.loheri、无芒长嘴薹草(C.longerostrata var.exaristata)、城湾薹草(C.longerostrata var.hoi)、C.lyi、和平菱果薹草(C.macrandrolepis)和C.sharyotensis并入尖叶薹草。     

亚洲西部和欧洲东南部芍药属的一个亚种

根据存于欧洲一些大标本馆的大量标本的观察,分布于希腊爱琴海东部、土耳其南部、塞浦路斯、黎巴嫩、叙利亚、伊拉克北部的Paeonia mascula与欧洲中部至巴尔干半岛的均不相同,可确认为一个独立的亚种。绝大多数个体总有一些小叶全裂,因此小叶及裂片数为(9)12～18(23);小叶宽椭圆形至卵圆形,通常两面无毛或背面疏生柔毛。     

基于形态性状的葱属粗根组植物系统发育关系

以环地中海地区和北美葱属Allium的Amerallium亚属种类为外类群,基于36个形态学性状对葱属粗根组sect. Bromatorrhiza进行系统发育分析。简约分析获得的分支拓扑结构与分子系统学的研究结果相吻合,粗根组与环地中海地区的Amerallium亚属种类形成姐妹群关系,表明它们具有密切的亲缘关系。粗根组明显分为两支,粗根韭A. fasciculatum (x = 10,2胚珠)与染色体基数为11的种类(单胚珠)形成一支,另一支由染色体基数为7的大花韭A. macranthum与多星韭A. wallichii (2胚珠)组成,从而否定了粗根亚组subsect. Bromatorrhiza(粗根韭+大花韭+多星韭)的成立。综合已有研究结果,本研究认为粗根组内存在4个相对独立的演化支系,即多星韭及其变种、大花韭、粗根韭,以及染色体基数为11的种类(5种1变种),尽管前人建议的单胚珠亚组subsect. Uniovulatum是合理的,但没有必要在物种数量相对较少的粗根组下进一步细分。     

Phylogenetic analysis of the genus Hexachlamys (Myrtaceae) based on plastid and nuclear DNA sequences and their taxonomic implications

Myrtaceae are one of the most species‐rich families of flowering plants in the Neotropics. They include several complex genera and species; Hexachlamys is one of the complex genera. It has not been recognized as a distinct genus and has been included in Eugenia, based on morphological grounds. Therefore, molecular systematic studies may be useful to understand and to help to solve these relationships. Here, we performed a molecular phylogenetic analysis using plastid and nuclear data in order to check the inclusion of Hexachlamys in Eugenia. Plastid (accD, rpoB, rpoC1, trnH‐psbA) and nuclear (ITS2) sequence data were analysed using Bayesian and maximum parsimony methods. The trees constructed using ITS2 and trnH‐psbA were the best able to resolve the relationships between species and genera, revealing the non‐monophyly of Hexachlamys. The molecular phylogenetic analyses were in agreement with previous morphological revisions that have included Hexachlamys in Eugenia. These results reinforce the importance of uniting knowledge and strategies to understand better issues of delimitation of genera and species in groups of plants with taxonomic problems. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 532–543.     

A revision of Dryopteris sect. Diclisodon (Dryopteridaceae) based on morphological and molecular evidence with description of a new species

Dryopteris sect. Diclisodon is a small section of ferns with about 12 species mainly distributed in East Asia. Here, we carried out morphological and phylogenetic analyses of this section. A new species from southwest China, D. gaoligongensis, is described and illustrated. Dryopteris gaoligongensis resembles D. indonesiana and D. sparsa, but differs by having a creeping rhizome and large 4-pinnate fronds. We also show that D. glabrior Ching & Z.Y. Liu is a distinct species; however, because it is a later homonym of D. glabrior Copel., it should be renamed D. renchangiana. We conclude that a species previously known as D. nitidula, also an illegitimate homonym, should be recognized with a new name, D. sinonepalensis. We resolve the phylogenetic position of D. yoroii as sister to other sampled species of D. sect. Diclisodon. Our phylogenetic analyses confirm the distinctiveness of D. gaoligongensis, D. renchangiana, and D. sinonepalensis. A key to species of D. sect. Diclisodon in China is provided.