Substantial Genome Size Variation in Taraxacum stenocephalum (Asteraceae, Lactuceae)

There are only a few exceptions to the rule that polyploidy in Taraxacum is associated with agamospermy. One of them is the sexual, tetraploid species Taraxacum stenocephalum. Incidentally, remarkable variation in karyology was found in this species. The present study aims to confirm this variation by an extensive screen of nuclear DNA content. Individuals from two large populations in the Lesser and Greater Caucasus, Georgia were analyzed using flow cytometry to ascertain intraspecific nuclear DNA content variation. Across the whole data set comprising all 159 individuals, a 1.223-fold difference was detected based on propidium iodide (PI) analyses. To verify this finding, we compared flow-cytometric data obtained using DAPI (4′,6-diamidino-2-phenylindole) and PI staining using a representative subset of individuals. This comparison revealed a 1.194-fold difference in DNA content for DAPI and a 1.219-fold difference for PI. Mean nuclear genome size in absolute terms (2C value ± s.d.) was estimated at 4.38?±?0.21 pg, ranging from 4.01 pg to 4.89 pg, despite the invariable chromosome counts of 2n?=?32. A regression analysis comparing the datasets for DAPI and PI staining found a strong correlation between data obtained by the DAPI and PI dyes (R?=?0.976; P?=?0.0001). Simultaneous high-resolution flow-cytometric analyses also proved the accuracy of our findings. We discuss possible sources of these large differences in DNA content within Taraxacum stenocephalum. Further research is needed to identify the source of this remarkable variation.     

Principal features of the cpDNA evolution in Taraxacum (Asteraceae, Lactuceae): a conflict with taxonomy

Phylogenetic relationships based on the chloroplast genome of Taraxacum were studied. Representative samples of 44 sections or species groups and a number of isolated species were analyzed. On the basis of the sequence variation in psbA–trnH and in trnL–trnF, mutations associated with RFLPs were monitored. Five RFLPs without homoplasy were recognized and used to reconstruct four main cpDNA groups (haplotypes); Group I is ancestral and, contrary to the information in the primary sequences, the RFLPs were not distinct from those of the outgroup species of Agoseris and Prenanthes. This group corresponds to dandelions believed to be ancestral on the basis of morphological data and previous studies of the chloroplast genome. A comparison of parsimony analysis of morphological and chloroplast data showed an overall lack of congruence. The conflict can most probably be accounted for as a consequence of reticulation.     

东北地区蒲公英属瘦果微形态特征及其分类学意义

通过对东北地区10种蒲公英瘦果形态及微形态特征进行观察比较,为蒲公英属（Taraxacum F.H.Wigg.）分类研究提供瘦果形态学证据。利用体式数码光学解剖镜和电子探针显微镜,对蒲公英瘦果进行观察比较,并根据瘦果大小、形状、喙基比例、颜色及微形态表面纹饰特征进行聚类检索。结果表明,（1）瘦果微形态特征为全身具刺,具白色斑点,刺状突起成行且规则排列,但刺大小及密度存在显著差异;（2）检索结果表明,以蒲公英瘦果形态及微形态特征进行检索的结果与《中国植物志》中的分类结果一致。该实验通过对东北地区10种蒲公英瘦果形态及微形态特征进行描述,确定了蒲公英瘦果形态特征差异可作为分类学表征证据,并将喙基长度（B）与瘦果长度（A）（不包括喙基长）之比命名为喙基比（B/A）,该比值可增补为分类要点。研究结果支持《中国植物志》中对蒲公英属的分类结果。     

The genusPicris (Asteraceae,Lactuceae) in Tropical Africa

Three species of the genusPicris L. are native in Tropical Africa:P. abyssinica Sch. Bip. (Ethiopia),P. xylopoda Lack, spec. nova (Nigeria, Ethiopia) andP. humilis DC. (Senegal, Mali). There are indications that the two perennial species,P. abyssinica andP. xylopoda, are related to and have evolved from a primitive Central Asiatic stock in a manner parallel to many African species ofCrepis L.P. humilis, on the other hand, is a small annual plant with a high number of derived characters. The introduced species of European origin growing south of the Sahara are briefly mentioned.     

Geographical distribution of wildLactuca species (Asteraceae, Lactuceae)

A broad survey of available world literature showed that at least 98 wildLactuca spp. (Asteraceae) have been described taxonomically. The distribution of the genusLactuca worldwide includes 17 species in Europe, 51 in Asia, 43 in Africa, and 12 in the Americas (mostly the North American subcontinent). Species originating in Asia, Africa, and the Americas form ca. 83% of knownLactuca spp. richness; however, they are very poorly documented from the viewpoint of taxonomic relationships, ecogeography, and variability. The phytogeography ofLactuca spp. regarding their distribution on different continents and in relation to the structure of the lettuce gene pool is discussed. A more detailed analysis of geographical distribution and habitats is given for some species (L. serriola, L. saligna, L. virosa, L. perennis, L. quercina, L. tatarica), which represent the primary, secondary, and tertiary gene pools of cultivated lettuce (L. sativa). Original and synanthropic distributions ofLactuca spp. and their occurrence in natural and secondary habitats are discussed, along with the representation of wildLactuca spp. in world gene-bank collections. Global biodiversity ofLactuca spp. and their representation in germplasm collections are poorly documented. Future studies of taxonomy, phytogeography, ecology, phylogenetic relationships, and genetic diversity are needed for a more complete understanding of this genus and taxonomically related genera.     

Isozymes, and the status of Taraxacum (Asteraceae) agamospecies

HUGHES, L. & RICHARDS, A. J., 1989. Isozymes and the status of Taraxacum (Asteraceae) agamospecies. Genetic identities I and Similarity Indices SI are calculated between 12 samples of Taraxacum, on the basis of 40 isozymes at 15 loci for 10 enzyme systems. Samples included three polyploid agamospermous populations from northern England (group 1), three sexual diploid populations from south-central France (group 2), and six accessions of ‘primitive’ diploid self-compatible sexual taxa from southern Europe and Australia. Samples could be assigned to eight species, classified in seven sections of the genus. Two clusters of high relationship were evident. All the group 1 and group 2 species were very closely related, with pairwise comparisons for I in excess of 0.93. The three group 3 accessions identified as T. bessarabicum showed pairwise comparisons for SI in excess of 0.71. Comparisons for SI between the other group 3 species, and between all the group 3 species and the group 1 and 2 species were all very low, not exceeding 0.45. It is concluded that dissimilarity between samples as assessed by isozymes is probably related to the time of evolutionary divergence of those samples. Although allopolyploid, and morphologically very diverse, the group 1 agamospecies may have very recently diverged asexually from a common stock. The group 2 diploids may have resulted from rediploidization and regained sexuality from the same originally agamospermous stock. In areas of Europe in which such ‘modern’ sexuality is common, it is probable that all ‘modern’ Taraxaca, including at least five sections of the genus, should be included within a single taxon. In contrast, ‘primitive’ self-compatible sexual species in group 3 appear to have diverged from each other several million years ago, and with the exception of the disjunct accessions of T. bessarabicum, are genetically highly distinct. Such species should be maintained in the taxonomies of all areas. It is probable that an agamospecies classification of ‘modern’Taraxacum will continue to convey much useful information in areas, such as northern Europe, in which sexuality is absent.     

Hypervariable DNA Fingerprint Loci in Microseris pygmaea (Asteraceae,Lactuceae)

Oligo-GATA containing probes reveal DNA fingerprinting patterns in DNA of Microseris pygmaea after restriction enzyme digestion. There are only a few major restriction fragments. These are longer than 2 kb, and they differ completely among plants from different populations, among most members of a single local inbreeding population, and even among some parent and offspring plants. In the F2 of an interpopulation hybrid, major fingerprint bands could be assigned to 2 unlinked loci by 3: 1 and 1: 2: 1 segregations respectively, one further band segregated roughly 1: 1, one band appeared irregularly. The hypervariability of the GATA-fingerprint loci contrasts with a low variability of other genetic markers in M. pygmaea and with a more complex but less variable GATA-fingerprint pattern in related species.     

Sesquiterpenoids as chemosystematic markers in the subtribe Hypochaeridinae (Lactuceae,Asteraceae)

Sesquiterpene lactones are well established as chemosystematic markers in the Asteraceae family. From the Lactuceae tribe of the Asteraceae family a large number of sesquiterpene lactones – mainly of the guaiane type – have been isolated. One of the 11 subtribes of the Lactuceae recognized by Bremer is the subtribe Hypochaeridinae, which encompasses 10 genera with approximately 170 species. The present communication summarizes the sesquiterpene derivatives reported from these 10 genera, points out, which constituents are characteristic for particular groups, and discusses the occurrence of these secondary metabolites in a chemosystematic context. To this end, each of the reported sesquiterpenoids reported for the Hypochaeridinae is classified into one of three main compound classes (MCCs; eudesmane-, germacrane-, guaiane-derivatives) and into one of a number of compound classes (CCs) within these principal groups. The distribution of sesquiterpenoids belonging to these particular classes of sesquiterpenoids generally follows the currently accepted generic limits. However, the genus Helminthoteca, which is included into Picris by many authors, possesses an array of compounds implying a closer relationship to the genus Hypochaeris than to the remainder of the genus Picris. Furthermore, Leontodon subgenus Oporinia shows closer similarity in secondary metabolite patterns to the genus Picris than to Leontodon subgenus Leontodon. On the other hand Leontodon subgenus Leontodon has more chemical characters in common with the genus Hedypnois than with Leontodon subgenus Oporinia. These findings are in-line with recent results of molecular analyses, which imply that current generic limits within the Hypochaeridinae might not reflect the phylogeny of the subtribe.     

Karyotype evolution in South American species of Hypochaeris (Asteraceae, Lactuceae)

The genus Hypochaeris (Asteraceae, Lactuceae) contains ten species in Europe, three in Asia, and approximately 50 in South America. Previous cytotaxonomic studies have shown two groups of taxa: (1) European species with different basic chromosome numbers and differentiated karyotypes, and (2) South American species with x=4 and uniform asymmetric and bimodal karyotypes. Karyotypic data are synthesized for South American species of Hypochaeris with new information for six Chilean species: H. acaulis, H. apargioides, H. palustris, H. spathulata, H. tenuifolia and H. thrincioides. Four main groups can be distinguished based on presence and localization of secondary constrictions – SCs (bearing Nucleolar Organizer Regions – NORs) on chromosomes 2 and 3, and 18S–25S and 5S rDNA loci number, localization, and activity. We propose karyotypic evolution of South American Hypochaeris (x=4) from H. maculata-like (x=5) European ancestors. The original South American karyotype would have possessed two SCs, one on the long arm of chromosome 2, and the other on the short arm of chromosome 3 (in terminal position). Further evolution would have involved inversion within the short arm of chromosome 3 and inactivation/loss of the SC on chromosome 2.     

Anatomy of ovary and ovule in dandelions (Taraxacum, Asteraceae)

The genus Taraxacum Wigg. (Asteraceae) forms a polyploid complex within which there are strong links between the ploidy level and the mode of reproduction. Diploids are obligate sexual, whereas polyploids are usually apomictic. The paper reports on a comparative study of the ovary and especially the ovule anatomy in the diploid dandelion T. linearisquameum and the triploid T. gentile. Observations with light and electron microscopy revealed no essential differences in the anatomy of both the ovary and ovule in the examined species. Dandelion ovules are anatropous, unitegmic and tenuinucellate. In both sexual and apomictic species, a zonal differentiation of the integument is characteristic of the ovule. In the integumentary layers situated next to the endothelium, the cell walls are extremely thick and PAS positive. Data obtained from TEM indicate that these special walls have an open spongy structure and their cytoplasm shows evidence of gradual degeneration. Increased deposition of wall material in the integumentary cells surrounding the endothelium takes place especially around the chalazal pole of the embryo sac as well as around the central cell. In contrast, the integumentary cells surrounding the micropylar region have thin walls and exhibit a high metabolic activity. The role of the thick-walled integumentary layers in the dandelion ovule is discussed. We also consider whether this may be a feature of taxonomic importance.     

Phylogeographic patterns in Hypochaeris section Hypochaeris (Asteraceae, Lactuceae) of the western Mediterranean

Aim  To analyse phylogeographic patterns in the four species of Hypochaeris sect. Hypochaeris , evaluating possible areas of origin and the microevolutionary processes that have shaped their morphology, genetics and distribution.
Location  Western Mediterranean area.
Methods  We applied amplified fragment length polymorphism (AFLP) markers to a total of 494 individuals belonging to 82 populations of Hypochaeris arachnoidea , H. glabra , H. radicata and H. salzmanniana to determine population structure.
Results  Populations with the largest proportion of private and rare AFLP fragments were found in Morocco. This region was consequently inferred to be the ancestral area for H. arachnoidea , H. glabra , H. radicata and H. salzmanniana . The Guadalquivir River (southern Spain) was inferred to be an effective dispersal barrier for H. glabra and H. radicata. The Strait of Gibraltar was inferred to be a somewhat weaker barrier than the Guadalquivir River for H. radicata and a much weaker barrier for H. glabra . The main barrier for H. salzmanniana coincides with the extension of the Rif Mountains to the Atlantic coast in Morocco, and the Strait of Gibraltar is a much weaker barrier for this species. Hypochaeris arachnoidea appears to have originated in the Atlas Mountains.
Main conclusions  The highest levels of genetic variation in La Mamora forest ( H. glabra and H. salzmanniana ) or the adjacent central Middle Atlas ( H. arachnoidea and H. radicata ) in Morocco suggest that these areas were a centre of origin of Hypochaeris sect. Hypochaeris . All three potential barriers – the Guadalquivir River, the Strait of Gibraltar, and the Rif Mountains – have been important in shaping genetic diversity in species of section Hypochaeris .     

Molecular differentiation between coexisting species of Taraxacum sect. Erythrosperma (Asteraceae) from populations in south-east and west Germany

The present study focuses on the genetic differentiation between six coexisting species of Taraxacum sect. Erythrosperma . Twenty-six individuals of T. rubicundum , T. brachyglossum , T. tortilobum , T. lacistophyllum , T. parnassicum and T. scanicum were sampled at 14 locations in south-east (Bavaria) and west Germany (Eifel). Using random amplified polymorphic DNA (RAPD) analysis 139 fragments were amplified. Nineteen of these were found to be species specific. The level of polymorphic bands within the different Taraxacum species varied between 0% and 15.8%. An analysis of molecular variance (AMOVA) revealed strong differentiation between the investigated species: 14.3% of the molecular variance were found within the species, while 85.7% of the genetic variability was observed between the species. However, a geographical differentiation between the different regions could not be observed. All analysed species were clearly separated from each other in a cluster analysis using the neighbour joining technique. The observed strong genetic differentiation between the Taraxacum species clearly supports the agamospecies concept of Taraxacum sect. Erythrosperma , which is at present mainly based on morphological and cytological criteria  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 109–117.     

Species relationships in Lactuca s.l. (Lactuceae, Asteraceae) inferred from AFLP fingerprints

An AFLP data set comprising 95 accessions from 20 species of Lactuca s.l. (sensu lato) and related genera was generated using the primer combinations E35/M48 and E35/M49. In phenetic analyses of a data subset, clustering with UPGMA based on Jaccard's similarity coefficient resulted in the highest cophenetic correlation, and the results were comparable to those of a principal coordinates analysis. In analyses of the total data set, phenetic and cladistic analyses showed similar tree topologies for the well-supported parts of the trees. The validity of cladistic analysis of AFLP data is discussed. The results do not support a distinction among the serriola-like species L. sativa, L. serriola, L. dregeana, and L. altaica, which is in line with previous results. Therefore, we postulate that these species are conspecific. The serriola-like species L. aculeata occupies a clearly separate position, making it an ideal outgroup for studies of the closest relatives of L. sativa. The subsect. Lactuca as a group is well supported by our data, but the positions of L. saligna and L. virosa relative to the serriola-like species remain unclear. The close relationship between the sect. Mulgedium species L. tatarica and L. sibirica is corroborated by the present AFLP results and by additional crossability data.     

Evolution of microsporangium numbers in Microseris (Asteraceae: Lactuceae)

The genus Microseris contains species with disporangiate stamens and species with tetrasporangiate stamens. We determined the number of microsporangia per stamen in serial sections of heads from all 13 species of Microseris, its close relative Uropappus lindleyi, and the two allopolyploid species of Slebbinsoseris. Four Microseris species, three diploid and one tetraploid, have two microsporangia per stamen; all other species investigated have four. The most parsimonious assumption is that the disporangiate condition is derived and arose once in the evolution of Microseris. The inheritance of the number of microsporangia per stamen in crosses between M. bigelovii (disporangiate) and M. douglasii (tetrasporangiate) was determined. Segregation of microsporangium number per stamen in F2s derived from these crosses is quantitative rather than Mendelian. The average number of microsporangia per stamen in the F2 plants ranges from 2.0 to 4.0. There is a predominance of tetrasporangiate stamens in the F1 and in most F2 plants. The observed pattern of inheritance suggests a major gene with dominance and quantitative modifiers.     

Genetic components of heterocarpy inMicroseris hybrid B87 (Asteraceae,Lactuceae)

Microseris B87 is derived from a single hybrid specimen betweenM. pygmaea with few, weakly hairy peripheral achenes and aM. bigelovii with many, strongly hairy peripheral achenes. Offspring through the F4 and F5 generations obtained by spontaneous selfing were analyzed for the segregation of quantitative and qualitative characters relating to achene dimorphism. The phenotypic effects of two previously identified unlinked genes determining the relative number of outer achenes are characterized in partially and completely homozygous sublines. We show that two morphological markers genetically linked to one of these genes are themselves regulated by the system inducing heterocarpy. Not more than two more unlinked genes are involved in the genetic basis of the heterocarpic response. The interaction of these genes in determining the heterocarpy phenotypes is discussed in the framework of a model postulating genes for a morphogen gradient across the capitulum and genes responding to this gradient.     

The intriguing complexity of parthenogenesis inheritance in Pilosella rubra (Asteraceae, Lactuceae)

Neither the genetic basis nor the inheritance of apomixis is fully understood in plants. The present study is focused on the inheritance of parthenogenesis, one of the basic elements of apomixis, in Pilosella (Asteraceae). A complex pattern of inheritance was recorded in the segregating F(1) progeny recovered from reciprocal crosses between the facultatively apomictic hexaploid P. rubra and the sexual tetraploid P. officinarum. Although both female and male reduced gametes of P. rubra transmitted parthenogenesis at the same rate in the reciprocal crosses, the resulting segregating F(1) progeny inherited parthenogenesis at different rates. The actual transmission rates of parthenogenesis were significantly correlated with the mode of origin of the respective F(1) progeny class. The inheritance of parthenogenesis was significantly reduced in F(1) n?+?n hybrid progeny from the cross where parthenogenesis was transmitted by female gametes. In F(1) n?+?0 polyhaploid progeny from the same cross, however, the transmission rate of parthenogenesis was high; all fertile polyhaploids were parthenogenetic. It appeared that reduced female gametes transmitting parthenogenesis preferentially developed parthenogenetically and only rarely were fertilized in P. rubra. The fact that the determinant for parthenogenesis acts gametophytically in Pilosella and the precocious embryogenesis in parthenogenesis-transmitting megagametophytes was suggested as the most probable explanations for this observation. Furthermore, we observed the different expression of complete apomixis in the non-segregating F(1) 2n?+?n hybrids as compared to their apomictic maternal parent P. rubra. We suggest that this difference is a result of unspecified interactions between the parental genomes.     

Revision of the genus Clibadium (Asteraceae, Heliantheae)

Clibadium L. (Asteraceae, Heliantheae) is a genus of 29 species distributed throughout latin America, from Mexico to Peru, and in the West Indies, with high numbers of species in Costa Rica, Colombia, and Ecuador.Clibadium includes shrubs and small trees; usually with loosely aggregated capitula; herbaceous phyllaries arranged in 1–5 series; receptacles usually paleaceous throughout; corollas of pistillate florets 2–4-lobed; corollas of the staminate florets 4–5-lobed; purple to black anthers; and chromosome numbers alln=16. Two sections of species previously recognized are here considered as subgenera (subg.Paleata and subg.Clibadium) containing two and four sections, respectively.Clibadium subg.Paleata contains five species distributed in sects.Eggersia (3 spp.) andTrixidium (2 spp.), and subg.Clibadium has 24 species distributed among sects.Clibadium (6 spp.),Glomerata (9 spp.),Grandifolia (5 spp.), andOswalda (4 spp.).     

RAPD mapping of three QTLs determining trichome formation in Microseris hybrid H27 (Asteraceae:Lactuceae)

Segregation for 289 random amplified polymorphic DNA markers (RAPDs) has been determined in 106 F₂ plants of an interspecific hybrid (H27) between Microseris douglasii (strain B14) and M. bigelovii (C94). Multicelluar trichomes (type D, specific for Microseris) occur on the leaf teeth of early vegetative rosettes of the B14 parent and on the leaf blades of later rosettes in both parents. Trichomes on the leaf blades appear earlier and eventually more densely in B14. Segregation for trichome appearance is quantitative and strongly transgressive in the F₂ hybrid. Cosegregation between RAPDs and trichome phenotypes combined with linkage data have revealed a main gene (quantitative trait locus A, QTL-A) with a pleiotropic effect on all trichome characters and two unlinked additive modifiers (QTL-B, QTL-C). Alleles of both modifiers reduce the main gene effect in each parent. Their recombination explains the occurrence of plants with transgressive phenotypes in the hybrid offspring. Additional QTLs affecting trichomes are at and below the level of statistical significance.