中国野生牡丹研究(三)芍药属牡丹组新分类群

芍药属牡丹组新分类群,红斑牡丹Paeonia ridleyi Z.L.Daiet T.Hong sp.nov.;保康牡丹Paeonia baokangensis Z.L.Dai et T.Hong sp.nov.     

中国野生牡丹研究（一）芍药属牡丹组新分类群

牡丹为我国特产珍贵花树和药用树种,已有1500余年栽培历史,建国以来,各地栽培品种已达500余个。 有关牡丹分类的主要研究成果多为西方科学家根据18—19世纪从我国引种到英、美、法等国的栽培牡丹和腊叶标本加以描述和定名。 作者近几年来在安徽、河南、湖南、山西、陕西、甘肃、四川、云南等地对我国野生牡丹进行了较广泛的调查和研究。 本文发表3个新种和1个新等级,这对研究我国栽培牡丹的起源和栽培品种的自然分类,发掘、保护、利用我国珍稀野生牡丹基因资源,培育新品种,扩大牡丹栽培地区等方面提供了科学理论依据。     

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。     

中国野生牡丹研究 二：芍药属牡丹组新分类群

本文发表一个新亚种和一个新等级。新亚种林氏牡丹和紫斑牡丹同属一个种系,花瓣基部均具紫斑,花丝,花盘及柱头均为白色;但林氏牡丹的小叶多为披针形或窄卵形,全缘,而紫斑牡丹的小叶多为卵形或卵圆形,常１－３深裂。     

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。     

芍药属牡丹组革质花盘亚组的系统演化探讨

芍药属牡丹组革质花盘亚组全部原产中国,它们对中国、日本、美国和欧洲的栽培牡丹品种影响甚大。作者在对本亚组9种牡丹(含3个亚种)形态学比较分析的基础上,结合花粉、染色体、蛋白质和DNA水平上的研究证据,提出了本亚组的系统演化的可能途径为:(1)卵叶牡丹→银屏牡丹→矮牡丹→圆裂四川牡丹→四川牡丹,(2)卵叶牡丹→凤丹→紫斑牡丹→太白山紫斑牡丹。另外作者对肉质花盘亚组的系统演化地位也进行了讨论。     

秦岭芍药属植物及其地理分布修订

在野外调查的基础上,结合查阅标本和文献比对,对秦岭地区芍药属(Paeonia)植物及其地理分布进行修订。结果表明,秦岭地区分布有芍药属8个野生种及1个栽培种,可分为牡丹组(Section Mouton DC.)和芍药组(Sect. Paeonia)。地理分布格局研究表明：芍药组内4个种：草芍药(P. obovata)、美丽芍药(P. mairei)、芍药(P. lactiflora)、川赤芍(P. anomala subsp. veitchii)和牡丹组中的牡丹(P. suffruticosa Andrews)、紫斑牡丹(P. rockii)为秦岭广布种,卵叶牡丹(P. qiui)和杨山牡丹(P. ostii)为秦岭中东部分布种,矮牡丹(P.jishanensis)为秦岭东部分布种。根据修订,给出了秦岭地区芍药属植物分种检索表。     

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。     

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.     

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.     

Phylogenetic analysis of Paeonia sect. Moutan (Paeoniaceae) based on multiple DNA fragments and morphological data

Tree peony, being crowned the title “King of Flowers” in China, is of great medicinal, ornamental, and economic values. In the present study, the phylogeny of the wild tree peony species (section Moutan, Paeonia, Paeoniaceae), represented by twelve accessions collected from all eight species in the section, was investigated based on the DNA sequence in five DNA fragments from both nuclear (Adh1A, Adh2 and GPAT) and chloroplast (trnS-trnG and rps16-trnQ) genomes, as well as morphological characters. Both maximum parsimony (MP) and Bayesian inference of phylogeny (BI) trees were reconstructed based on the combined data of the DNA sequences and morphological data, respectively. The MP and BI trees have the similar topology, and the sect. Moutan clearly branched into two clades. One clade consists of two species, P. delavayi and P. ludlowii, corresponding to the subsect. Delavayanae, and another clade is composed of other six species. Within the second clade, the six species can be divided into three subclades consisting of P. rockii and P. decomposita, P. jishanensis and P. qiui, P. suffruticosa and P. ostii, respectively. Among the three subclades, P. jishanensis/P. qiui is most closely related to P. suffruticosa/P. ostii. These results provide up to date the clearest picture of the phylogeny of wild tree peony species in the sect. Moutan.     

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).     

Paeonia anomala 的核型研究

报道了Paeonia anomala L. 的核型, 澄清了新疆阿尔泰地区分布的该物种的染色体数目。分布于该地区的Paeonia anomala L. 的核型组成: 2n = 2x = 10 = 6m+ 2sm+ 2st。该类群核型与该属其它类群一致———2A 型。在综合比较分析该属染色体参数以及核型不均一性数, 包括最长􊄯最短染色体比(L1/Ln ) 和染色体不对称系数( CKOA) 的基础上, 我们发现该属三个组在核型上没有明显分化, 仅在木本类群(Sect . Moutan DC .) 和草本类群(Sect. Onaepia Lindley 和Sect. Paeonia) 之间存在微小差异。此外, 作为二倍体类群, 新疆阿尔泰地区分布的Paeonia anomala L. 很可能是二倍体杂种, 这将为研究被子植物的父母本同倍化杂交式物种起源提供一个很好的研究材料。