小檗科的花粉演化

以APG III定义的基部真双子叶分支（Basal Eudicots）中毛茛目（Ranunculales）小檗科（Berberidaceae）为研究对象,选取4个DNA片段（rbcL、matK、trnLF和26S rDNA）,利用最大似然法构建分子系统树,结合已报道的花粉形态数据,分析了该科16个属的花粉形态。选择花粉分散单位、极性、形状、大小、萌发孔数目、萌发孔位置、外萌发孔形状、覆盖层上元素、覆盖层纹饰和外壁厚度共10个关键性状,采用简约法推断了该科花粉的祖征、共衍征和演化式样。研究表明：单粒、等极、近球形、中等大小是小檗科花粉的祖征。无极、多萌发孔和周面孔是小檗亚科（Berberidoideae）的共衍征,支持其为一个单系。三萌发孔分别为鬼臼亚科（Podophylloideae）、南天竹亚科（Nandinoideae）各自的共衍征;覆盖层上元素不存在是小檗亚科和南天竹亚科的共衍征,将它们与鬼臼亚科区分开来,同时也支持了小檗亚科和南天竹亚科之间的姐妹关系。此外,对一些属花粉形态的演化意义进行了讨论,提出一些特殊的花粉性状可以用来定义某些属,如Bongardia和兰山草属（Ranzania）。     

从叶绿体DNA trnL-F序列论双参属的归属问题

双参属Triplostegia Wall．ex DC．由分布于东南亚地区的2个种组成,为多年生草本植物。它的归属一直存在争议,有时置于川续断科Dipsacaceae或败酱科Valerianaceae,有时单立一科,即双参科Triplostegiaceae。本研究对广义川续断目Dipsacales s．l．的21种植物(分别来自于败酱科、川续断科、双参属、刺参属Morina、广义忍冬科Caprifoliaceae s. l．、五福花科 Adoxaceae)和外类群人参Panax schin-seng Nees．的叶绿体 DNA trnL-F区进行了测序,并建立系统发育树状图。结果显示,败酱科、川续断科、双参属、刺参属和广义忍冬科的4个属(双盾木属Dipelta、虫胃实属Kolkwitzia、六道木属Abelia和北极花属Linnaea)形成 了一个单系群并得到了很强的支持(100％ bootstrap);双参属与川续断科有更近的关系,建议作为一个亚科置于川续断科;广义忍冬科为一多系类群;而刺参属与其他广义川续断目类群之间的关系尚不能确定。     

中国鸡矢藤属花粉形态

用扫描电子显微镜对中国鸡矢藤属(Paederia L.)6种1变种植物的花粉进行观察。结果表明:鸡矢藤属植物的花粉均为单粒,辐射对称,小型或中型,极面观3-裂圆形至钝三角形,赤道面观呈长圆球形或近长球形,具3个萌发沟,无内萌发孔。外壁纹饰网状、细网状或穿孔状,孔边缘具小刺状突起或无。臭鸡矢藤(P.foetida)和白毛鸡矢藤(P.pertomentosa)具有花粉二型现象,其中白毛鸡矢藤(P.pertomentosa)是首次报道。花粉二型现象与花柱二型现象可能没有直接的关联性,与前人的观点一致。推测鸡矢藤属外壁纹饰的可能演化趋势为:穿孔、网状、细网状→粗网状;网眼内无棒状突起→网眼内有棒状突起。鸡矢藤属花粉的外壁纹饰变化较大,且无内萌发孔,是茜草科花粉过渡类型的特征。     

湖北省泽泻科、水鳖科、眼子菜科及茨藻科植物花粉形态研究

本文利用光镜及扫描电镜对湖北省泽泻科、水鳖科、眼子菜科及茨藻科11属29种1变种1变型植物(另加采于湛江的软骨草)的花粉形态进行了研究,发现泽泻科植物花粉具多个圆形萌发孔,外壁表面为小刺状纹饰;茨藻科植物花粉具远极单槽,表面为绉波状纹饰;眼子菜科及本文研究的水鳖科植物花粉均无萌发孔,分别具网状和小刺状饰纹饰。1.茨藻科植物花粉最原始,泽泻科花粉较进化,眼子菜科花粉较水鳖科花粉进化;2.泽泻属与泽苔草属花粉较慈姑属花粉原始;3.鞘叶眼子菜亚屈花粉较眼子菜亚属的花粉处于更高演化阶段;4.多孔茨藻花粉在该科中最原始。本文工作尚对易变形水生植物花粉形态研究方法进行了尝试。     

中国紫草科破布木属花粉形态和外壁超微结构

为了深入探讨紫草科（Boraginaceae）的分类问题,用光学显微镜和扫描电子显微镜观察了该科破布木属（CordiaL.) 10种植物的花粉形态和外壁超微结构。发现该属花粉具三孔、三孔沟、三拟孔沟和三合沟4种萌发孔类型。外壁表面具微刺状纹饰、刺状纹饰、网状纹饰和不规则的条纹网状纹状。破布木属的花粉特征表明,该属花粉在紫草科中既是独特的分类群,又是比较原始的属种。     

国产爵床科芦莉花族植物的花粉形态

报道了国产爵床科Acanthaceae芦莉花族Ruellieae芦莉花亚族Ruelliinae 2属7种、假杜鹃亚族Barlerinae 1属3种和马蓝亚族Strobilanthinae 16属34种植物扫描电镜下的花粉形态.芦莉花亚族的地皮消属Pararuellia和喜花草属Eranthemum的花粉均为圆球形,具3孔或3孔沟,外壁为不同的网状结构; 假杜鹃亚族的假杜鹃属Barleria的花粉为长球形,具3孔沟,外壁亦为网状结构;马蓝亚族植物(包含广义的马蓝属Strobilanthes s.l.)花粉形态多样,结构复杂.依据花粉萌发孔和外壁纹饰特征,可将马蓝亚族16属植物和上述两亚族3属植物的花粉形态归纳成3大类型: 1. 具3孔类型.其中又有(1)外壁具网状纹饰者,见于地皮消属; (2)外壁具芽胞状纹饰者,见于黄猄草属Championella; (3)外壁具刺状(棒状)纹饰者,见于南一笼鸡属Paragutzlaffia、叉花草属Diflugossa和假蓝属Pteroptychia.2. 具3孔沟及具3孔沟与假沟类型(肋条带型).其中又有(1)具3孔沟和网状纹饰者,见于喜花草属和假杜鹃属; (2)具刺状(棒状)纹饰者,见于南一笼鸡属、叉花草属和假蓝属; (3)具3孔沟与假沟,外壁纹饰具节隔、肋条带状或网状,网眼纵向排列成行,网眼内有细网纹者,见于耳叶马蓝属Perilepta、马蓝属Pteracanthus(大部分)、金足草属Goldfussia、紫云菜属Strobilanthes(部分)和合页草属Sympagis; (4)具3孔沟与假沟类型,肋条带状,但不具节隔,外壁纹饰网状,网眼不成行或不明显纵向排列,网内无细网纹者,见于尖蕊花属Aechmanthera、板蓝属Baphicacanthus、马蓝属(部分)和糯米香属Semnostachya; (5)具双脊及细网状纹饰者,见于环毛紫云菜Strobilanthes cycla.3. 具(4-)5孔沟及假沟类型(肋条带型),外壁具网状或拟网状纹饰,见于腺背蓝属Adenacanthus.另外兰嵌马蓝属Parachampionella、山一笼鸡属Gutzlaffia和肖笼鸡属Tarphochlamys的花粉有无萌发孔尚不清楚,有待进一步研究.综上所述,芦莉花族植物的花粉形态具有较高的多样性,是重要的分类性状.利用花粉形态特征能较好地区分高级分类群如亚科、族以及亚族,有时也有助于阐明类群之间的相互关系,甚至也能用于区分属、种和阐明其关系.     

甘薯属10种植物花粉形态扫描电镜观察

对用于育种研究的甘薯属(Ipomoea)10种植物花粉形态进行扫描电镜观察研究,以补充该属植物花粉形态学信息. 结果表明,甘薯属植物花粉粒的形状均为近球形,体积较大;花粉萌发孔多为散孔,少数萌发孔不明显;表面纹饰有3种:刺状类型、网状带刺类型和网状类型.根据花粉粒大小、刺的长短、萌发孔特征和表面纹饰类型等可以把这10种植物区分开来,同时发现I.wrightii与已有报道存在较大差异.     

水雍科植物的花粉形态研究

应用光学显微镜、扫描电镜和透射电镜对世界水雍科6种植物的花粉形态进行了观察。水雍科植物花粉为舟形或船形,具远极单沟萌发孔类型,外壁纹饰通常为浅网状至网状,稀为小刺状纹饰,外壁外层由覆盖层、柱状层和基层组成,覆盖层厚。水雍科植物花粉外壁纹饰表现了从小刺状向网状的过渡。该科花粉为远极单沟,覆盖层厚,具小刺等特征反映了其与水鳖科和泽泻目花蔺科的密切联系,而该科花粉外壁纹饰多为网状则与茨藻目植物接近。这一结果支持将水雍科从茨藻目中分出作为一个独立的目处理的观点。此外,水雍科植物的花粉大小、纹饰类型、网眼大小与深浅等方面的差异对种级分类有一定意义。     

泽泻科的花粉形态研究

本文对泽泻科11属27种代表植物的花粉进行了光学显微镜、扫描电镜和透射电镜观察。在系统描述了该科及各属植物花粉形态的基础上,将泽泻科植物的花粉划分为3种类型,即少果泽苔草型、慈菇型和泽泻型。根据花粉形态特征的比较,并依据泽泻科植物祖先类群的花粉具有船形、具单沟萌发孔、花粉外壁具明显的刺状纹饰、覆盖层完整无通道等特征,作者认为泽泻科植物花粉形态的如下演化趋势是明显的：由船形演化为卵球形、球形和多面体球形;由单沟萌发孔经过一无孔的中间类型演化为散孔类型;孔膜由光滑演化为具颗粒和小刺;萌发孔不内陷进化到内陷;花粉粒外壁的刺状纹饰逐渐过渡为颗粒状纹饰或者消失,以及覆盖层由无通道到具细通道和通道。     

红树(Rhizophora a piculata Bl.)的花粉形态与多态现象

通过光学显微镜(LM)和扫描电子显微镜(SEM)观察表明,红树(Rhizophora apiculata)花粉粒赤道面的形状为球形-近球形,极面观为圆三角形,偶见圆四方形,3-4孔沟,具有连续的环赤道内孔,花粉外壁的典型纹饰为细网状-皱纹状(microreticulate-rugulate)。作者首次报道红树花粉的多态现象,其花粉外壁纹饰和萌发孔数量存在显著的变异,SEM观察到花粉外壁纹饰的变异主要是孔状(perforate)、皱纹状(rugulate)和穴状(foveolate)等类型,LM观察发现4个萌发孔的花粉变异类型。花粉形态的观察与描述为化石花粉的鉴别提供了不可或缺的对比依据。研究红树的花粉形态和发现多态现象有助于了解红树科红树属的花粉外壁演化。花粉的多态现象表明单个花粉形态特征并不能完全代表种的特征。花粉的分类也应该充分考虑花粉性状的间断和连续性,以期正确认识花粉性状在种群内的变异和变异式样,达到客观认识和正确划分植物种下等级的目的。花粉的多态现象为化石花粉的种类鉴定增加了新的参考信息,作者也讨论了花粉多态现象在植物系统演化和古生态学等研究中的可能价值与意义。     

Phylogenetic relationships of the disputed genus Triplostegia based on trnL-F sequences

The phylogenetie relationships of Triplostegia Wall. ex DC., comprising two species of perennial herbs from southeastern Asia, have long been in dispute. This genus was placed in either Dipsacaceae or Valerianaceae or in a family of its own, Triplostegiaceae. In this paper, the chloroplast DNA (cpDNA) trn L-F regions of 21 species in the Dipsacales s. l. (including Valerianaceae, Dipsacaceae, Triplostegia, Morina, Caprifoliaceae s. l. and Adoxaceae) and an outgroup Panax schin-seng Nees. were amplified and sequenced. The phylogenetic relationships among these 22 species were constructed based on trn L-F sequences. The results demonstrated that Valerianaceae, Dipsacaceae, Triplostegia, Morina and four genera from the Caprifoliaceae s. l. form a monophyletic group with a strong support (100% bootstrap). Triplostegia, a sister group to Dipsacaceae, is close enough to be placed in the Dipsacaceae as a subfamily. The traditional Caprifoliaceae s.l. are polyphyletic, and relationships of Morina among the groups within Dipsacales s. l. are uncertain. Key words Triplostegia; Caprifoliaceae s. l.; Morina; Dipsacales s. l.; trnL-F sequences; Sys-tematic position     

Pollen Morphology of Caprifoliaceae from China and Its Taxonomic Signi-ficance

The pollen morphology of 54 samples representing 12 genera and 31 species was investigated with the aid of scanning electron microscope. Observed were pollen grains of Sambucus, Viburnum, Lonicera, Leycesteria, Heptacodium, Linnaea, Abelia, Dipelta, Kolkwitzia, Symphoricarpos, Triosteum, Weigela. Based on the shape, size, position and number of aperture, exine sculpture, three types are recognized: 1. Pollen grains subprolate, less frequently prolate, rather small, 3-colporate, exine reticulate, as in Sambucus, Viburnum. 2. Mostly spheroidal, subolate, bigger than the former, also 3-colporate, exine spinulose as in Lonicera, Leycesteria, Heptacodium, Triosteum, Linnaea, Abelia, Dipelta, Kolkwitzia, Symphoriocarpos, Weigela. 3. Spheriodal, more or less flattend, exine scabrous as in Abelia section Zabelia and Lonicera section Isoxylosteum. 1. The systematic position of Caprifoliaceae: It has been generally treated as a member of the order Rubiales together with Rubiaceae, Valeriaceae and Dipsacaceae on floral characters. In respect to serological character, it has a close relationship with Cornaceae, and was placed in Araliales. The above stated 2nd and 3rd types of pollen grains are similar to those of Patrinia (Valerianaceae), Scabiosa (Adoxaceae), Cornus (Cornaceae), and the pollen grains of the 1st type are similar to those of Styraceae, Genetianaceae and Araliaceae. Taking the information so far available into consideration, the authors agree to the Cronquists treatment retaining Caprifoliaceae in the order Dipsacales together with Adoxaceae, Valerianaceae and Dipsacaceae. 2. The division of tribes: Formerly Sambuceae included the genera Sambucus and Viburnum. Fritsch (1891) segregated Viburnum from Sambuceae and suggested a new tribe Viburneae including Triosteum. There is distinct difference in palynological features between these two genera. The exine sculpture of Viburnum is reticulate, but that of Triosteum is spinulose. It is reasonable to separate another new tribe, Triosteae, from Viburneae. 3. The pollen morphorlogy of several Chinese endemic genera, such as Heptacodium, Dipelta, Kolkwitzia resembles that of Lonicera, Leycesteria, Linnaea, Symphoricarpos, Abelia, Triosteum. This evidence supports the foregoing treatment including them in Caprifoliaceae. 4. Two different exine sculptures are shown in sections of the genera Abelia and Lonicera. In Abelia the exine of the section Euabelia is spinulose, but that of the section Zabelia is scabrous. Likewise, in Lonicera, the exine of the section Isoxylosteum is scabrous, while that of other sections such as Nintooa, Isika, Lonicera, subgenus Caprifolium, is spinulose. It shows that pollen morphology is one of diagnostic characters for section division.     

Dating the Dipsacales: comparing models, genes, and evolutionary implications

Dipsacales is an asterid angiosperm clade of ca. 1100 species, with most of its lineages occupying temperate regions of the Northern Hemisphere. A recent phylogenetic analysis based on 7593 nucleotides of chloroplast DNA recovered a well-resolved and strongly supported phylogenetic hypothesis, which we use here to estimate divergence times within the group. A molecular clock is strongly rejected, regardless of data partition. We used recently proposed methods that relax the assumption of rate constancy among lineages (local clocks, nonparametric rate smoothing, penalized likelihood, and Bayesian relaxed clock) to estimate the ages of major lineages. Age estimates for Dipsacales varied widely among markers and codon positions, and depended on the fossils used for calibration and method of analysis. Some methods yielded dates for the Dipsacales diversification that appear to be too old (prior to the presumed 125 my [million years] age of eudicots), and others suggested ages that are too young based on well-documented Dipsacales fossils. Concordant penalized likelihood and Bayesian studies imply that Dipsacales originated in the Cretaceous, as did its two major lineages, Adoxaceae and Caprifoliaceae. However, diversification of crown Adoxaceae and Caprifoliaceae mainly occurred in the Tertiary, with the origin of major lineages within these clades mainly occurring during the Eocene. Another round of diversification appears to have occurred in the Miocene. Several radiations, such as Valerianaceae in South America and Dipsacaceae around the Mediterranean, are even more recent. This study demonstrates the wide range of divergence times that can be obtained using different methods and data sets, and cautions against reliance on age estimates based on only a single gene or methodology. Despite this variance, significant conclusions can be made about the timing of Dipsacales evolution.     

Phylogeny of the Dipsacales s.l. based on chloroplast trnL-F and ndhF sequences

Sequences of the chloroplast trnL-F region and 3(') end ndhF gene were used to elucidate phylogenetic relationships and the delimitation of families within Dipsacales s.l. Parsimony analyses of individual and combined data were conducted using maximum parsimony method. The most parsimonious tree based on combined trnL-F and 3(') end ndhF data set recognizes seven major clades of Dipsacales s.l. with the following relationships: Apiales (Adoxaceae ((Diervillaceae, Caprifoliaceae s.str.) (Linnaeaceae (Morinaceae (Dipsacaceae, Valerianaceae))))). Both Sambucus and Viburnum have close relationships with Adoxaceae, supporting their inclusion in this family. Caprifoliaceae s.l. (excluding Sambucus and Viburnum) is polyphyletic, and comprises three clades or families, i.e., Linnaeaceae (Abelia, Dipelta, Kolkwitzia, and Linnaea), Diervillaceae (Weigela and Diervilla) and Caprifoliaceae s.str. (Heptacodium, Leycesteria, Lonicera, Symphoricarpos, and Triosteum). This study focuses on the systematic position of Heptacodium, Triplostegia, and Morinaceae; and suggests that Heptacodium is closely related to the other Caprifoliaceae s.str.; Triplostegia is a sister to Dipsacaceae; Morinaceae, which has an affinity with Dipsacaceae, is possibly a sister group with Dipsacaceae-Valerianaceae clade. Our results are highly congruent with those of and.     

Revisiting the phylogeny of Dipsacales: New insights from phylogenomic analyses of complete plastomic sequences

Phylogenetic relationships in Dipsacales have long been a major challenge. Although considerable progress has been made during the past two decades, questions remain; the uncertain systematic positions of Heptacodium, Triplostegia, and Zabelia, in particular, impede our understanding of Dipsacales evolution. Here we use 75 complete plastomic sequences to reconstruct the phylogeny of Dipsacales, of which 28 were newly generated. Two primary clades were recovered that form the phylogenetic backbone of Dipsacales. Seven of the primary clades correspond to the recognized families Adoxaceae, Caprifoliaceae s. str., Diervillaceae, Dipsacaceae, Linnaeaceae, Morinaceae, and Valerianaceae, and one corresponds to Zabelia, which was found to be the closest relative of Morinaceae in all analyses. Additionally, our results, with greatly increased confidence in most branches, show that Heptacodium and Triplostegia are members of Caprifoliaceae s. str. and Dipsacaceae, respectively. The results of our study indicate that the complete plastomic sequences provide a fully‐resolved and well‐supported representation of the phylogenetic relationships within Dipsacales.     

Molecular Phylogeny and Species Speciation of Adoxaceae. s. s

It remains unclear about the speciation and phylogeny of Adoxaceae s. s ., a small family with 3 genera and 4 species . In this paper , ITS ( nuclear DNA internal transcribed spacer) regions of Adoxa orientalis and Sambucus adnata were firstly sequenced . Phylogenetic trees were constructed for all species of Adoxaceae (four species ) , Sambucus, Viburnum and four genera of Caprifoliaceae . The divergences among four species of this family were further calculated based on the calibration of the fossil records of the Caprifoliaceae and the general evolutionary rate of herbs for ITS . The phylogenetic analyses did not support the previous assumptions on the phylogeny and species divergence of Adoxaceae s. s . based on the morphological evidence : Tetradoxa is not the firstly diverged and it clustered with two species of Adoxa as a monophylogenetic group , paralleling to the other lineage comprising of monotypic Sinadoxa . The allopatric speciation at the diploid level might have contributed to the differentiation among Sinadoxa corydalifolia, Tetradoxa omeiensis and Adoxa moschatellina and the polyploidy to the origin of A. orientalis . The crude timing based on ITS sequence differentiation suggested a recent divergence among all four species probably between the late Miocene and the Tertiary and this speciation process might be closely correlated with habitat fragmentation and change due to the extensive uplifts of the Qinghai-Tibetan Plateau and climatic oscillation during the glacial and interglacial ages occurred at this stage .     

A search for the phylogenetic position of the seven-son flower (Heptacodium, Dipsacales): Combining molecular and morphological evidence

A first report on the problematic phylogenetic position ofHeptacodium (2 spp.; China) using molecular data from chloroplast DNA is presented. Amplification of ORF2280 homolog region was executed in a number of representative taxa in order to determine ifHeptacodium shows similar structural rearrangements as other Dipsacales. DNA sequences ofndhF were generated to clarify the phylogenetic position ofHeptacodium among Caprifoliaceae (s.l.). Six outgroup taxa and fifteen representatives of Dipsacales were sampled and more than 2100 basepairs ofndhF sequence were used in a cladistic analysis. Parsimony analysis produced two shortest trees and showedHeptacodium as sister to all members of Caprifoliaceae (s.str.), although weakly supported. Additionally, trees were constructed withndhF data supplemented with availablerbcL sequences and a morphological data set. Results of all analyses support an unresolved basal position forHeptacodium among Caprifoliaceae (s.l.), which in part explains the difficulty experienced previously in classifying the genus.     

狭义五福花科的分子系统学和物种分化

狭义五福花科(Adoxaceae.s.s)仅含3属4种,但该科的物种分化、系统发育和分类一直存在争议。本文通过测定东方五福花和血满草的ITS(核糖体DNA内转录间隔区)序列,构建了包括狭义五福花科(4种)、广义忍冬科接骨木属、荚属以及其余4属植物在内的系统发育树。研究结果不支持狭义五福花科内根据形态学证据做出的系统发育假设四福花不是该科中最早分化出来的种;该物种与五福花属的两个物种形成一个单系群,与另一分支华福花属相对应。该科中3个物种,四福花、五福花和华福花之间的分化主要是在二倍体水平上的异域分化,而东方五福花则是通过多倍化形成的。粗略的时间估算表明这些物种之间的分化较晚,可能在第三纪末至第四纪早中期,与青池高原近期强烈隆升以及冰期气候反复变化形成的环境变迁密切相关。     

A phylogenetic analysis of 100+ genera and 50+ families of euasterids based on morphological and molecular data with notes on possible higher level morphological synapomorphies

 A data matrix of 143 morphological and chemical characters for 142 genera of euasterids according to the APG system was compiled and complemented with rbcL and ndhF sequences for most of the genera. The data were subjected to parsimony analysis and support was assessed by bootstrapping. Strict consensus trees from analyses of morphology alone and morphology + rbcL + ndhF are presented. The morphological data recover several groups supported by molecular data but at the level of orders and above relationships are only superficially in agreement with molecular studies. The analyses provide support for monophyly of Gentianales, Aquifoliales, Apiales, Asterales, and Dipsacales. All data indicate that Adoxaceae are closely related to Dipsacales and hence they should be included in that order. The trees were used to assess some possible morphological synapomorphies for euasterids I and II and for the orders of the APG system. Euasterids I are generally characterised by opposite leaves, entire leaf margins, hypogynous flowers, “early sympetaly” with a ring-shaped corolla primordium, fusion of stamen filaments with the corolla tube, and capsular fruits. Euasterids II often have alternate leaves, serrate-dentate leaf margins, epigynous flowers, “late sympetaly” with distinct petal primordia, free stamen filaments, and indehiscent fruits. It is unclear which of these characters represent synapomorphies and symplesiomorphies for the two groups, respectively, and there are numerous expections to be interpreted as reversals and parallelisms. Received August 28, 2000 Accepted August 7, 2001