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。     

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.     

Preliminary phylogeny of Valerianaceae (Dipsacales) inferred from nuclear and chloroplast DNA sequence data

Valerianaceae is a relatively small (ca. 350 species), but morphologically diverse angiosperm clade. Sequence data from the entire ndhF gene, the trnL-F intergenic spacer region, the trnL intron, the matK region, the rbcL-atpB intergenic spacer region and internal transcribed spacer (ITS) region of nuclear ribosomal DNA were collected for 21 taxa within Dipsacaceae and Valerianaceae (1 and 20, respectively). These data were included in several phylogenetic analyses with previously published sequences from Dipsacales. Results from these analyses (maximum parsimony, maximum likelihood, and Bayesian analysis) are in strong agreement with many of the conclusions from previous studies, most importantly: (1) Valerianaceae is sister to Dipsacaceae; (2) Triplostegia is more closely related to species of Dipsacaceae than to Valerianaceae; and (3) Valeriana appears not to be monophyletic, with Valeriana celtica falling outside the remainder of the species of Valeriana sampled here (with very strong support). With the exception of V. celtica, these data support two major clades within Valeriana; one that is exclusively New World and another that is distributed in both the Old and New World. Although the species of Valerianaceae and its sister group Dipsacaceae plus Triplostegia, are widely distributed in the Northern Hemisphere, and the data imply that Valerianaceae diversified initially in Asia (the Himalayan Patrinia and Nardostachys falling at the base of the clade), the center of modern species diversity for the group is in the Andes of South America with as many as 175 species restricted to that region. Although the exclusively South American taxa form a clade in the chloroplast and combined ITS and chloroplast analyses, support values tend to be low. Future studies will need to include additional data, in the form of both characters and taxa, before any strong conclusions about the character evolution, diversification, and biogeography of the South American valerians can be made.     

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.     

蹄盖蕨科的系统发育: 叶绿体DNA trnL-F区序列证据

蹄盖蕨科Athyriaceae是蕨类植物中一个复杂的大科,由于属间关系不甚清楚,该科分类系统还有一些问题,比如新蹄盖蕨、拟鳞毛蕨属、假冷蕨属、肠蕨属、短肠蕨属和菜蕨属的系统位置常有争议。根据蹄盖蕨科34种植物和3种外类群植物的叶绿体DNA trnL-F区序列建立了系统发育树,结果显示:1.trnL_F区序列分析的结果与rbcL基因序列分析的结果几乎一致。2.新蹄盖蕨属Neoathyrium Ching&Z.R.Wang不应成立,该属应与角蕨属Cornopteris Nakai合并。3.假冷蕨属Pseudo     

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.     

Petal venation in the Asterales and related orders

Petal venation in families related to the Asteraceae was studied by means of light microscopy. The group of study was delimited on the base of previously published molecular data and includes the Alseuosmiaceae, Apiaceae, Aquifoliaceae, Araliaceae, Argophyllaceae, Bruniaceae, Brunoniaceae, Calyceraceae, Campanulaceae sensu lato , Caprifoliaceae, Donatiaceae, Dipsacaceae, Escalloniaceae, Goodeniaceae, Griseliniaceae, Menyanthaceae, Pentaphragmataceae, Pittosporaceae, Sambucaceae, Sphenocleaceae, Stylidiaceae, Valerianaceae, and Viburnaceae. The Calyceraceae, Goodeniaceae, and Menyanthaceae are very similar to the Asteraceae in their petal venation. One feature common to these four families but not found in any other group is the prominent marginal veins entering the petal independently of the midvein, but meeting it at the apex of the petal. Other morphological data as well as gene sequence data suggest that these four families form a monophyletic group. The other investigated families show a wide array of venation types, but they are all very different from the Asteraceae. Typical for die Campanulaceae sensu lato is a densely reticulate pattern. The Dipsacaceae, Valerianaceae and some Caprifoliaceae are characterized by well-developed transpetal veins.     

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.     

Molecular phylogeny reveals the non-monophyly of tribe Yinshanieae(Brassicaceae) and description of a new tribe,Hillielleae

The taxonomic treatment within the unigeneric tribe Yinshanieae(Brassicaceae) is controversial, owing to differences in generic delimitation applied to its species. In this study, sequences from nuclear ITS and chloroplast trn L-F regions were used to test the monophyly of Yinshanieae, while two nuclear markers(ITS, ETS) and four chloroplast markers(trnL-F, trn H-psbA, rps16, rpL32-trnL) were used to elucidate the phylogenetic relationships within the tribe. Using maximum parsimony, maximum likelihood, and Bayesian inference methods, we reconstructed the phylogeny of Brassicaceae and Yinshanieae. The results show that Yinshanieae is not a monophyletic group, with the taxa splitting into two distantly related clades: one clade contains four taxa and falls in Lineage I, whereas the other includes all species previously placed in Hilliella and is embedded in the Expanded Lineage II. The tribe Yinshanieae is redefined, and a new tribe, Hillielleae, is proposed based on combined evidence from molecular phylogeny, morphology, and cytology.     

川续断目的花粉演化

以APGⅢ定义的川续断目(Dipsacales)忍冬科(Caprifoliaceae)和五福花科(Adoxaceae)为研究对象,基于已有的川续断目分子系统树和花粉形态数据,分析了忍冬科25属和五福花科5属的花粉形态多样性,采用简约法(Fitch Parsimony)推测花粉祖征和演化式样,寻找共衍征和分类性状.选择花粉萌发孔数目、萌发孔类型、花粉形状、大小和外壁纹饰五个关键性状推断了其演化式样.研究表明:三孔沟、近球形、较小花粉、网状纹饰是川续断目花粉祖征.长球型、中等和较大花粉,以及刺状纹饰是忍冬科的共衍征,支持忍冬科和科下分支为单系群.忍冬科花粉刺状纹饰和五福花科花粉网状纹饰明显将两个科区分开来.通过追溯性状演化分析,支持将七子花属(Heptacodium)置于忍冬族(Caprifolieae),以及Zabelia置于刺续断科的观点.     

Polyphyly of Limoniastrum (Plumbaginaceae): evidence from DNA sequences of plastid rbcL, trnL intron and trnL-F intergene spacer

Phylogenetic relationships of Limoniastrum and other genera of subfamily Staticoideae (Plumb-aginaceae) were studied using parsimony analysis of the plastid gene rbc L, the intron of trn L and the intergene spacer of trnL-trn F. Our analysis showed that Limoniastrum was polyphyletic. Limoniastrum ifniense , in both rbc L and combined data analyses, is sister to Armeria and Psylliostachys , whereas in the trn L-F (intron and spacer combined) analysis it is sister to a clade composed of Acantholimon, Dictyolimon and the remaining species of Limoniastrum . In all analyses, the five remaining species of Limoniastrum (excluding Limoniatrum ifniense ) formed a clade with two groups of species: L. monopetalum+L. guyonianum and those sometimes considered as the segregate genus Bubania ( L.feei, L. weygandiorum and L. rechingeri ). Levels of sequence divergence among these three groups of Limoniatrum were greater than for other well supported genera in the family and, in combination with morphological differences and paucity of synapomorphies, led us to conclude that separate generic status for each of the three clades is warranted.     

A molecular phylogenetic analysis of the bloodroot and kangaroo paw family, Haemodoraceae: taxonomic, biogeographic and conservation implications

A phylogenetic analysis of plastid DNA sequences from the trn L-F region corroborates the hypothesis that Haemodoraceae, a small monocotyledonous family centred in southwestern Australia, are monophyletic with relationships to Philydraceae, Pontederiaceae and Comme–linaceae. It also supports the long-standing recognition of two subfamilies. In Conostylidoideae Tribonanthes falls in an isolated position, thus supporting its segregation as a recently recognized monogeneric tribe. Tribal status for Phlebocarya is not supported as this taxon is unexpectedly placed in Conostylideae as sister to Conostylis–Blancoa. Macropidia falls as sister to Anigozanthos . The DNA tree permits continued recognition of Macropidia and Blancoa as distinct genera, contrary to a recent morphological cladistic analysis. Haemodoroideae fall into two clades: ( Dilatris(Lachnanthes+Haemodorum )) and ( Xiphidium(Schiekia+Wachendofia )). It is unlikely that Haemodoraceae are of Gondwanan origin, and the phylogenetic pattern indicates a largely relictual distribution with a recent radiation in Western Australia.     

A revision and phylogenetic analysis of Stapeliopsis (Apocynaceae)

A survey of morphological characters is carried out for Stapeliopsis . The information obtained from this is combined with molecular data from the plastid trn L-F DNA region and ITS1 of the nuclear encoded 18S−26S rRNA cistron, to obtain a hypothesis of the evolutionary relationships among the species. It is shown that Stapeliopsis is monophyletic in a combined molecular and morphological analysis. Stapeliopsis is sister to a clade containing Huernia , Orbea and Tromotriche . The species of Stapeliopsis group into two clades. One contains S. khamiesbergensis , S. neronis and S. urniflora , and this is highly supported. The remaining species fall into an unsupported clade in which S. exasperata is sister to the others. The genera Hermanschwartzia Plowes and Neopectinaria Plowes are rejected. It is shown that a synapomorphy for Stapeliopsis is the laterally flattened inner corona-lobes, which touch the anthers only at their bases. Eight species of Stapeliopsis are recognized, with no subgeneric divisions.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 125–155.     

单种属弥勒苣苔属系统位置研究：基于分子和细胞学数据

弥勒苣苔属是苦苣苔科的单种属,仅分布于中国西南部。为探讨弥勒苣苔在苦苣苔亚科中的系统位置,我们选择了苦苣苔亚科116个类群,外类群为苦苣苔亚科以外的7个物种。用最大简约法（MP）和贝叶斯分析（BI）,对以上类群的核基因ITS以及两个叶绿体基因trnL-F、atpB-rbcL数据进行了独立和联合分析。在三个片段联合分析的结果中,弥勒苣苔与马铃苣苔属、后蕊苣苔属、金盏苣苔属、直瓣苣苔属以及川鄂粗筒苣苔构成一个强烈支持的分枝。MP树中,此分枝为并系,而在BI分析中,弥勒苣苔与川鄂粗筒苣苔、直瓣苣苔属互为姐妹类群。同时,第一次报道了弥勒苣苔的染色体数目（2n=34）。根据前人报道,马铃苣苔属、后蕊苣苔属、粗筒苣苔属和直瓣苣苔属的染色体数目同为2n=34,这进一步支持我们的分子系统发育分析。     

Correlates of diversification in the plant clade Dipsacales: geographic movement and evolutionary innovations

We explore patterns of diversification in the plant clades Adoxaceae and Valerianaceae (within Dipsacales), evaluating correlations between biogeographic change (i.e., movements into new areas), morphological change (e.g., the origin of putative key innovations associated with vegetative and reproductive characters), and shifts in rates of diversification. Our findings indicate that rates of diversification in these plants tend to be less tightly correlated with the evolution of morphological innovations but instead exhibit a pronounced correlation with movement into new geographic areas, particularly the dispersal of lineages into new mountainous regions. The interdependence among apparent novelties (arising from their nested phylogenetic distribution) and the correlation between morphological and biogeographic change suggests a complex history of diversification in Dipsacales. Overall, these findings highlight the importance of incorporating biogeographic history in studies of diversification rates and in the study of geographic gradients in species richness. Furthermore, these results argue against a simple deterministic relationship between dispersal and diversification: like other factors that may influence the probability of speciation and/or extinction, the impact of dispersal on diversification rates depends on being in the right place at the right time.     

Phylogeny and biogeography of Valerianaceae (Dipsacales) with special reference to the South American valerians

Species of Valerianaceae are a common component of the alpine flora throughout the Northern Hemisphere as well as the Andes of South America. Sequence data from three chloroplast markers (psbA-trnH intron, trnK-matK intron, and the trnL-F region) along with the internal transcribed spacer region (ITS) of nuclear ribosomal DNA were used to infer relationships within Valerianaceae. Both genomes, as well as a combined data set, provide support for the major clades within the group and do not support a monophyletic Valeriana. In addition, these data indicate that Plectritis is nested within South American Valeriana, as opposed to being sister to Centhranthus as previously hypothesized. Valerianaceae appear to have originated in Asia, probably in the Himalayas, and subsequently to have dispersed several times to Europe and to the New World. Our results imply that Valerianaceae colonized South America on multiple occasions from the north. In one of these cases there appears to have been a substantial and rapid radiation, primarily in the high elevation paramo habitat. A variety of methods were used to estimate divergence times to determine when Valerianaceae might have colonized South America. Regardless of the method and fossil constraints applied, our estimates suggest that Valerianaceae colonized South America prior to the formation of the Isthmus of Panama.