Evolution of genome size across some cultivated Allium species.

Allium L. (Alliaceae), a genus of major economic importance, exhibits a great diversity in various morphological characters and particularly in life form, with bulbs and rhizomes. Allium species show variation in several cytogenetic characters such as basic chromosome number, ploidy level, and genome size. The purpose of the present investigation was to study the evolution of nuclear DNA amount, GC content, and life form. A phylogenetic approach was used on a sample of 30 Allium species, including major vegetable crops and their wild allies, belonging to the 3 major subgenera Allium, Amerallium, and Rhizirideum and 14 sections. A phylogeny was constructed using internal transcribed spacer (ITS) sequences of 43 accessions representing 30 species, and the nuclear DNA amount and the GC content of 24 Allium species were investigated by flow cytometry. For the first time, the nuclear DNA content of Allium cyaneum and Allium vavilovii was measured, and the GC content of 16 species was measured. We addressed the following questions: (i) Is the variation in nuclear DNA amount and GC content linked to the evolutionary history of these edible Allium species and their wild relatives? (ii) How did life form (rhizome or bulb) evolve in edible Allium? Our results revealed significant interspecific variation in the nuclear DNA amount as well as in the GC content. No correlation was found between the GC content and the nuclear DNA amount. The reconstruction of nuclear DNA amount on the phylogeny showed a tendency towards a decrease in genome size within the genus. The reconstruction of life form history showed that rhizomes evolved in the subgenus Rhizirideum from an ancestral bulbous life form and were subsequently lost at least twice independently in this subgenus.     

Morphological and embryological characters of three middle Asian Allium L. species (Alliaceae)

Morphological and embryological characters of the xerophilous species Allium crystallinum Vved., A. filidens Regel, and A. fritschii Khassanov et Yengalycheva (Allium subg. Allium) were compared. The morphology of flowers, the rate of development of male and female embryonic structures and correlations with bud dimensions were studied. The overall similarity of the embryonic processes and structures underlines the close taxonomic relationship of the species investigated as concluded from morphological and anatomical characters. However, several differences in embryological characters between A. filidens and A. fritschii confirm the separation of the latter species at species level.     

Phylogenetic analysis of Allium subg. Melanocrommyum infers cryptic species and demands a new sectional classification

Allium subgenus Melanocrommyum (Alliaceae) from Eurasia comprises about 150 mostly diploid species adapted to arid conditions. The group is taxonomically complicated with different and contradictory taxonomic treatments, and was thought to include a considerable number of hybrid species, as the taxa show an admixture of assumed morphological key characters. We studied the phylogeny of the subgenus, covering all existing taxonomic groups and their entire geographic distribution. We analyzed sequences of the nuclear rDNA internal transcribed spacer region (ITS) for multiple individuals of more than 100 species. Phylogenetic analyses of cloned and directly sequenced PCR products confirmed the monophyly of the subgenus, while most sections were either para- or polyphyletic. The splits of the large sections are supported by differences in the anatomy of flower nectaries. ITS data (i) demand a new treatment at sectional level, (ii) do not support the hypotheses of frequent gene flow among species, (iii) indicate that multiple rapid radiations occurred within different monophyletic groups of the subgenus, and (iv) detected separately evolving lineages within three morphologically clearly defined species (cryptic species). In two cases these lineages were close relatives, while in Allium darwasicum they fall in quite different clades in the phylogenetic tree. Fingerprint markers show that this result is not due to ongoing introgression of rDNA (ITS capture) but that genome-wide differences between both lineages exist. Thus, we report one of the rare cases in plants where morphologically indistinguishable diploid species occurring in mixed populations are non-sister cryptic species.     

Biosystematic studies in Allium L. section Cepa

VAN RAAMSDONK, L. W. D. & DE VRIES, T. 1992. Biosystematic studies in Allium L. section Cepa. Principal component analyses and hybrid analyses, according to Wells, using 31 morphological characteristics were used to study the biosystematic relationships between six Allium species of section Cepa and A. roylei of section Rhizirideum. Seventeen interspecific hybrid populations were included. Based on these analyses and on additional information from geography, chromosome banding pattern, DNA amount and seed epidermis texture, the species of Allium section Cepa can be divided into three groups: 1. A. galanthum and A. pskemense , 2. A. cepa, A. oschaninii and A. vavilovii and 3. A. futulosum and A, allaicum. Presumed evolution of both groups 2 and 3 from the basic group 1 is indicated by the development of new morphological characteristics, a change in genome structure and migration.     

Phylogeny of Chinese Allium (Liliaceae) using PCR-RFLP analysis

Eighteen representative species were selected from all the nine sections of Chinese Allium on the basis of the classification of morphology and cytotaxonomy. The trnK and rpL16 gene fragments of chloroplast DNA were amplified from 18 species by PCR method. The two cpDNA fragments were digested by 26 restriction enzymes, and 303 polymorphic restriction sites were found, of which 163 were informative. The restriction site data were analyzed with PAUP (version 3.1.1) and MEGA (version 1.01) as well as PHYLIP. As a result, the genus Allium could be classified into six subgenera. The recognition of Sect. Anguinum in the Flora of China is reasonable, Sect. Rhizirideum, Sect. Haplostemon and Sect. Cepa are not monophyletic. The infrageneric system of this genus was also discussed.     

Crossing experiments in Allium L. section Cepa

VAN RAAMSDONK, L. W. D., WIETSMA, W. A. & DE VRIES, J. N., 1992. Crossing experiments in Allium L. section Cepa . A full diallel was carried out with six diploid species of Allium section Cepa and A. roylei of section Rhizirideum , High isolation barriers were found between the related species A. cepa and A. oschaninii , between A. oschaninii and A. vavilovii , and between A. galanthum and A. pskemense . On the contrary, the species A. cepa and A. roylei , belonging to different sections, show only slight isolation barriers. The Wallace effect, which is the development of internal isolation barriers as such, is likely to have taken place in the evolution of A. oschaninii and A. vavilovii , and possibly also between A. galanthum and A. pskemense .     

葱属Anguinum亚属4种19居群的核型研究

石蒜科(Amaryllidaceae)葱亚科(Allioideae)葱族(Allieae)葱属(Allium)的Anguinum亚属在中国分布有6种2变种,该亚属植物具有重要的药用价值。该研究在扩大样本量和覆盖范围的基础上,采用常规压片法对葱属Anguinum亚属4种植物19居群的染色体核型进行了研究,首次报道了心叶韭(Allium ovalifolium var.cordifolium)的核型。结果表明:供试类群中茖葱都为四倍体,对叶韭和心叶韭都为二倍体,太白韭具有二倍体和四倍体,染色体基数均为x=8,核型类型都为2A型,每个居群都具有随体染色体,随体染色体多为st染色体,随体均位于短臂末端,并且对叶韭眉县居群和太白韭聂拉木居群存在B染色体。结合前人研究结果讨论了Anguinum亚属植物的染色体特征、随体染色体的多态性和该亚属植物的进化方式,探讨了茖葱(Allium victorialis)和对叶韭(A.listera)的细胞地理学问题。得到如下推论:(1)染色体加倍和结构变异是Anguinum亚属进化的两种重要机制,环境异质性使得Anguinum亚属植物随体染色体具较高的多态性;(2)茖葱通过多倍化和无性繁殖并存来扩大居群;(3)染色体形态变异是对叶韭适应环境变异的重要方式。     

Phylogeny of Chinese Allium (Liliaceae) using PCR-RFLP analysis

Eighteen representative species were selected from all the nine sections of Chinese Allium on the basis of the classification of morphology and cytotaxonomy. The trnK and rpL16 gene fragments of chloroplast DNA were amplified from 18 species by PCR method. The two cpDNA fragments were digested by 26 restriction enzymes, and 303 polymorphic restriction sites were found, of which 163 were informative. The restriction site data were analyzed with PAUP (version 3.1.1) and MEGA (version 1.01) as well as PHYLIP. As a result, the genus Allium could be classified into six subgenera. The recognition of Sect. Anguinum in the Flora of China is reasonable, Sect. Rhizirideum, Sect. Haplostemon and Sect. Cepa are not monophyletic. The infrageneric system of this genus was also discussed.     

Introgression explains incongruence between nuclear and chloroplast DNA-based phylogenies in Allium section Cepa

Phylogenetic relationships between species of Allium section Cepa and A. rqylei (section Rhizirideum) have been inferred from nuclear DNA variation (RAPDs; nDNA dataset) and from morphological, pollen epidermis texture, chromosomal and chemical variation (supranuclear dataset). These sets were complemented with data, taken from the literature, on cpDNA variation and crossability. The trees produced with the supranuclear, nDNA and cpDNA datasets were compared by using the topology of the most parsimonious tree of one dataset as the constraint for the construction of a most parsimonious tree of another dataset. The accuracy of the trees were evaluated by calculating several Consistency and Incongruence Indices. The constrained tree of supranuclear-nDNA datasets showed the highest index values. The tree topologies of the supranuclear and cpDNA datasets were the least similar. The cpDNA tree and crossability dendrograms were identical. The most important difference between the nDNA-supranuclear trees and the cpDNA-crossability trees pertains to the position of Allium roylei , which is much closer to the clade A. cepa/A. vavilovii in the cpDNA tree than in the nDNA tree. This difference is considered to be the result of chloroplast capture from one species to another after an introgression event. A shorter distance between species inferred from a cpDNA tree than from a nDNA or comparable tree might be indicative for the level of crossability.     

Phylogeny of Rubus (rosaceae) based on nuclear ribosomal DNA internal transcribed spacer region sequences

We used nuclear ribosomal DNA internal transcribed spacer region (ITS 1 - 5.8S - ITS 2; ITS) sequences to generate the first phylogeny of Rubus based on a large, molecular data set. We sampled 57 taxa including 20 species of subgenus Rubus (blackberries), one to seven species from each of the remaining 11 subgenera, and the monotypic and closely related Dalibarda. In Rubus, ITS sequences are most informative among subgenera, and variability is low between closely related species. Parsimony analysis indicates that Rubus plus Dalibarda form a strongly supported clade, and D. repens may nest within Rubus. Of the subgenera with more than one species sampled, only subgenus Orobatus appears monophyletic. Three large clades are strongly supported: one contains all sampled species of nine of the 12 subgenera; another includes extreme Southern Hemisphere species of subgenera Comaropsis, Dalibarda, and Lampobatus; and a third clade consists of subgenus Rubus plus R. alpinus of subgenus Lampobatus. Rubus ursinus appears to be a hybrid between a close relative of R. macraei (subgenus Idaeobatus, raspberries) and an unidentified subgenus Rubus species. ITS sequences are generally consistent with biogeography and ploidy, but traditionally important morphological characters, such as stem armature and leaf type, appear to have limited phylogenetic value in Rubus.     

A molecular phylogeny of the wild onions (Allium; Alliaceae) with a focus on the western North American center of diversity

Nuclear ribosomal DNA (ITS and ETS) sequences from 39 native Californian (USA) Allium species and congeners were combined with 154 ITS sequences available on GenBank to develop a global Allium phylogeny with the simultaneous goals of investigating the evolutionary history (monophyly) of Allium in the Californian center of diversity and exploring patterns of adaptation to serpentine soils. Phylogenies constructed with ITS alone or ITS in combination with ETS provided sufficient resolution for investigating evolutionary relationships among species. The ITS region alone was sufficient to resolve the deeper relationships in North American species. Addition of a second marker (ETS) further supports the phylogenetic placements of the North American species and adds resolution within subgenus Amerallium, a clade containing many Californian endemics. Within the global phylogeny, the native North American species were found to be monophyletic, with the exception of Allium tricoccum and Allium schoenoprasum. All native Californian species included in the analysis fell into a monophyletic subgenus Amerallium section Lophioprason, although endemic Californian species were not monophyletic due to the inclusion of species with ranges extending beyond the California Floristic Province. The molecular phylogeny strongly supports previous morphology-based taxonomic groupings. Based on our results, serpentine adaptation appears to have occurred multiple times within section Lophioprason, while the ancestor of the Californian center of diversity may not have been serpentine-adapted.     

Systematic Position of Allium macrostemon Based on nrDNA ITS and cpDNA trnL-F Sequence Data

Allium macrostemon is an important medicinal and edible plant. Its systematic position and taxonmical classification remain controversial to date. To explore this issue, we reconstructed phylogenetic relationships among Amacrostemon and other related taxa using nrDNA ITS and cpDNA trnL F markers. The phylogenetic trees derived from Bayesian inference and maximum parsimony analysis showed that Amacrostemon is monophyletic, and has a close relationship with some species of polyphyletic sections Caerulea and Pallasia instead of sections Codonoprasum and Allium. The including of Amacrostemon within section Allium was not supported by both molecular data and morphological characters of spathe, filaments and ovary. Allium macrostemon should be included in a new section, however further studies using additional samples (especially those from Central Asia) is necessary. In addition, we also provided a discussion on the phylogenetic relationships among four original plants (Amacrostemon, Achinense, Acaeruleum and Aneriniflorum) of Allii Macrostemonis Bulbus and systematic position of partial species of section Pallasia.     

基于trnK基因的葱属植物分子系统研究

在形态和细胞分类研究的基础上选择产于中国的9组47种葱属植物(含外类群5种),运用PCR方法扩增叶绿体trnK基因,选择26种限制性内切酶对PCR扩增片段进行了RFLP分析.结果表明:trnK基因的PCR产物在各分类群间几乎不存在长度变异,约为2 520 bp,PCR扩增片段酶切后,共得到247个变异位点,其中信息位点201个.运用PAU P 4.0 B 10.0和M EGA 3.1软件进行分析,构建葱属系统发育的D o llo和W agner最简约(M P)树及邻接(N J)树.分析表明:(1)宽叶组类群组成比较自然的单系群,洋葱组和葱组也各自形成独立分枝,表明这3个组的划分是比较自然的.多籽组和合被组在本次分析中形成1个单系群,表明这2个组具有较密切的亲缘关系.而粗根组、根茎组和单生组的划分是不自然的,需进一步研究后作适当的调整.粗根组的类群在trnK基因的RFLP分析中,得到很好的分辨,可按其染色体基数分为3个大的类群.(2)中国葱属植物可以划分为6个亚属的新等级,在各亚属下可以再分组.(3)本文还对葱属的种间亲缘、进化关系等问题进行了讨论.     

黄耆属簇毛黄耆亚属系统学研究

簇毛黄耆亚属以其柱头上具簇毛等形态特征与黄耆属其它类群相区别,约含49种（变种）。主要分布在青藏高原及其邻近地区,绝大多数种类分布于横断山区。本文基于形态特征和分支系统学方法分析亚属内种系发生关系。结果表明,49种（变种）可分为9组。其中双小苞组Sect.Bibracteolati含种类最多,是一个并系类群。借助于分支图,分析了6个较重要的鉴定特征的演化,习性,茎的着生方式,小苞片,果实形态,柱头具簇毛,果实膨胀状况等。最后作出了这个亚属的一个分类纲要。     

Sequence analysis of a chloroplast intergenic spacer for phylogenetic estimates in Allium section Cepa and a PCR-based polymorphism detecting mixtures of male-fertile and male-sterile cytoplasmic onion

Restriction-enzyme analysis of the chloroplast (cp) DNA yielded maternal phylogenies supporting a close phylogenetic relationship among normal (N) male-fertile and male-sterile (S) cytoplasmic bulb onion (Allium cepa), Allium altaicum, Allium fistulosum, Allium galanthum, Allium roylei, and Allium vavilovii. The S cytoplasm of onion is most likely an alien cytoplasm introduced in antiquity into onion populations. We previously showed that size differences in an intergenic spacer in the cp DNA distinguish N and S cytoplasms of onion. We cloned and sequenced this intergenic spacer from the N and S cytoplasms of onion, A. altaicum, A. fistulosum, A. galanthum, Allium pskemense, Allium oschaninii, A. roylei, and Allium ampeloprasm (outgroup) to identify the nature of previously described RFLPs and to develop a PCR-based marker revealing N-cytoplasmic contamination of S-cytoplasmic hybrid seed lots. Phylogenies based on restriction-enzyme analysis of the entire cp DNA were similar, but not identical, to those based on sequence divergence in this intergenic region. Received: 29 November 1999 / Accepted: 28 April 2000     

A Review of Current Theories and Methods in Cladistics and a Cladistic Study of Twelve Lindera Species in Eastern China

Cladistics has become a widely used method for phylogenetic reconstruction．Because of rapid improvement Of cladistic theories and methodologies,and application of new data,especially,molecular data,it is becoming realistic to reconstruct phylogenies of organisms,and to establish natural classifications based on these phylogenies．This paper reviews some current cladistic theories and methods in a practical way,such as choosing characters,defining character states,polarizing characters,analyzing data matrices, calculating consensus cladograms,choosing among multiple equally most parsimonious cladograms,estimating reliability of cladograms,and applying cladograms to classification, character evolution,and biogeography． Based on 36 morphological characters．a parsimony analysis of 12 species representing six sections in subgenus Lindera and an outgroup species from subgenus lteodaphne of the genus Lindera(Lauraceae)was conducted．The results suggest a close relationship between section Lindera and section Sphaerocarpae,which is different from the previous phylogenetic hypothesis within the genus．In the strict consensus cladogram,two species,L．megaphylla and L.chienii,from section Cupuliformes are in the most primitive and the most advanced clades respectively,indicating that the section is polyphyletic．The cladogram also suggests that section Lindera be a polyphyletic group．     

A cladistic study of Arillastrum, Angophora and Eucalyptus (Myrtaceae)

A cladistic study of Anllastrum, Angophora and Eucalyptus (Myrtaceae). Transformed cladistic; character compatibility; branch and bound, and Farris-Wagner methods gave similar solutions in a cladistic study of Arillastrum, Angophora and Eucalyptus. These analyses, based on morphological characters, indicate that Eucalyptus is a monophyletic group and that its sister taxon is Angophora.
Within Eucalyptus , subgenera Blakella and Corymbia are sister taxa to all other groups; subgenera Monocalyptus, Idiogenes and Gaubaea form a monophyletic group with subgenus Monocalyptus sister to subgenera Idiogenes and Gaubaea ; subgenera Symphyomyrtus and Telocalyptus together also form a monophyletic group and, with Eucalyptus similis (subgenus Eudesmia group 4), are sister to the Monocalyptus group. Eucalyptus subgenus Telocalyptus (4 species), Eucalyptus subgenus Idiogenes (1 species) and Eucalyptus subgenus Gaubaea (2 species) should not be recognized as subgenera and some individual species need further examination. Eucalyptus subgenus Eudesmia is a paraphyletic group.
Some characters are identified as parallelisms, e.g. axillary inflorescences, sepaline operculum, bristle glands, and clustered anthers. A more congruent interpretation of the single operculum of Eucalyptus subgenus Monocalyptus as at least partly petaline rather than solely sepaline in origin is suggested.
The area relationships for the taxa are concordant with those derived from geological and climatological information. New Caledonia is sister area to Australia, and within Australia southwestern Australia is sister area to south-eastern and north-eastern Australia.     

分支系统学当前的理论和方法概述及华东地区山胡椒属十二个种的分支系统学研究

本文概述了当前分支系统学研究中涉及的主要理论和方法,包括性状的选取、性状状态和极性的确定、数据矩阵的分析计算、结果分支图的处理、分支图可靠性的评价及分支图的应用。本文同时以华东地区樟科山胡椒属Linderal2个种的分支系统学研究为例,讨论了用形态性状进行分支系统学研究中可能遇到的问题,也揭示了一些分支系统学与传统的系统学在应用性状推导进化关系上的不同点。对这12个种的分支系统学研究得出了一些不同于传统系统学方法所推测的山胡椒属内的系统发育关系,如分支系统学研究显示山胡椒组和球果组很近缘。在严格一致性分支图上,杯托组的黑壳楠和江浙山胡椒分别位于最原始和最进化的分支,表明这个组是复系类群。分支图也显示山胡椒组可能是复系类群。     

用matK序列分析探讨木兰属植物的系统发育关系

用木兰科Magnoliaceae 57种植物的matK基因序列构建了该科的系统发育分支图。结果表明: (1)木兰属Magnolia L.是一个因为性状的趋同演化而建立的多系类群; (2)木兰亚属subgen. Magnolia和玉兰亚属subgen. Yulania (Spach) Reichenb.亲缘关系较远, 支持将后者从该属中分出建立玉兰属Yulania Spach, 木兰亚属作为木兰属保留; (3)木兰亚属的sect. Splendentes Dandy ex Vazquez组与皱种组sect. Rytidospermum Spach的两个美洲种M. macrophylla Michaux和M. dealbata Zucc.亲缘关系较近, 荷花玉兰组sect. Theorhodon Spach与常绿组sect. Gwillimia DC.的亲缘关系较近; (4)盖裂木属Talauma Juss.可以成立, 而其分布于亚洲的Blumiana Blume组可归入木兰属; (5)拟单性木兰属Parakmeria Hu &; Cheng、华盖木属Manglietiastrum Law以及单性木兰属Kmeria (Pierre) Dandy形成一个单系群, 与玉兰亚属和含笑属Michelia L.的亲缘关系较近。花的着生位置不足以作为木兰科的分族依据, 含笑族Michelieae和木兰族Magnolieae的特征及其界定应做修改。将玉兰亚属从木兰属分出后, 木兰属与含笑属无性状交叉,成为两个区别明显的属。     

Taxonomic data supporting differences between Allium elmaliense and Allium cyrilli

The first specimens of Allium elmaliense were collected in Elmal? (Antalya/Turkey) in 2001 and were described as new species in 2004. However, Allium elmaliense was claimed as synonym of Allium cyrilli in 2006. This study was undertaken to reveal the differences between these taxa utilizing morphological, palynological, and chromosome characters and genomic differences based on the DNA analyses along with the ecological preference studies conducted during 2006 and 2011. The results clearly indicated differences between these two taxa in terms of morphological characters, pollen, seed surfaces and niche preferences. Chromosome morphology and Td-DAMD-PCR fingerprinting studies revealed that Allium elmaliense Deniz & Sümbül is a distinct species and not a synonym of A. cyrilli Ten.