Phylogenetic relationships in the genus Leontopodium (Asteraceae: Gnaphalieae) based on AFLP data

The genus Leontopodium comprises 30–41 species. The centre of diversity is the Sino‐Himalayan region in south‐western China, where about 15 species occur. The two species native to Europe, L. alpinum (known as the common ‘Edelweiss’) and L. nivale, are part of the cultural heritage of the people living there. Despite its importance, very little is known about the systematics of the genus. Because recent molecular studies have shown that species within this genus are closely related and difficult to distinguish with rDNA and cpDNA data, we used AFLPs to obtain a more detailed understanding of the phylogeny of the genus. Our main aims were as follows: (1) to clarify species relationships within the genus; and (2) to reveal information about the biogeography of the genus. We used AFLPs with six primer combinations to investigate 216 individuals in 38 populations of 16 different species. With AFLPs, we were able to recognize 10 different groups, all of which had strong bootstrap support. These results were also congruent with the morphology‐based taxonomy of the genus. Most private and rare fragments were found in the Yunnan region (south‐western China) relative to Europe and Mongolia/central China, suggesting a long‐lasting in situ history of populations in the centre of diversity of the genus. Our results illustrate the utility of AFLPs to resolve phylogenetic relationships between these closely related species. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 364–377.     

Molecular phylogeny of Anaphalis (Asteraceae, Gnaphalieae) with biogeographic implications in the Northern Hemisphere

Anaphalis is the largest Asian genus in the tribe Gnaphalieae (Asteraceae) and has its greatest species diversity in the eastern Himalayas. The nuclear internal and external transcribed spacers were sequenced for Anaphalis species, with an emphasis on the eastern Himalayan taxa to examine the monophyly and construct the phylogenetic relationships of and within the genus. The results suggest that all species of Anaphalis are nested with Helichrysum, showing a close relationship with a Mediterranean–Asian group of Helichrysum. Although the monophyly of Anaphalis is only weakly supported, two clades within the genus are well recognized, each consisting of two subgroups. The inferred phylogenetic relationships within Anaphalis correspond to the shape of leaf base, rather than the morphology of the capitula and phyllaries that are usually used for species delimitation and classification in the genus. All four subgroups of Anaphalis are common and diversified in the eastern Himalayas with multiple dispersals out of this region. The sole North American species of Anaphalis is best hypothesized to be the result of long-distance dispersal or overland migration via Bering land bridge from Asia. Our analyses suggest that the extant distribution of Anaphalis has most likely resulted one radiation into the eastern Himalayas followed by repeated independent dispersals and/or radiations mostly into eastern Asia but also into the western Himalayas, North America, and southeast Asia.     

Chromosome counts and genome size of Leontopodium species (Asteraceae: Gnaphalieae) from south‐western China

Chromosome counts and genome sizes are reported from six species of Leontopodium: five from the centre of diversity in south‐western China, and L. japonicum from cultivation. Previously published chromosome counts for the genus are also compiled. Genome size (1C) in diploids ranges from 0.93 pg (L. dedeckensii) to 1.14 pg (L. cf. stracheyi) and 1.93 pg for tetraploid L. sinense. Leontopodium artemisiifolium had one pair of heteromorphic chromosomes. Leontopodium japonicum accessions showed variation in ploidy levels. Polyploidy, including autopolyploidy, is frequent in the genus. Variation is seen in basic chromosome number, including between species in the centre of diversity of the genus, where x = 12 or 13, but also within species among previously published counts. This variation does not correspond to currently inferred infrageneric groupings, and indicates both the importance of large‐scale chromosome evolution and the need for more in‐depth taxonomic work in a genus that shows little DNA sequence variation. © 2013 The Linnean Society of London     

Trans-oceanic dispersal and evolution of early composites (Asteraceae)

How did Asteraceae (the daisy family) expand from its area of origin and become so widespread? This question has challenged generations of evolutionary botanists. Molecular phylogenetic and biogeographic analyses indicate a South American origin of Asteraceae, a view supported by the recent discovery of the earliest fossils of the family in Middle Eocene (ca. 50 Ma) deposits in southern South America. The early-branching lineages in the phylogenetic tree of Asteraceae are South American and African, suggesting that the earliest successful colonization of areas outside South America may have involved long-distance dispersal to Africa. However, one particularly challenging unanswered question is how early members of Asteraceae reached Africa at a time when the Atlantic Ocean constituted a barrier between the two continents. Morphological, phylogenetic, geographic, paleogeographic, and paleontologic data have been combined to propose scenarios on possible geographical and dispersal routes and vectors of dispersion of early-branching lineages of Asteraceae from South America to Africa. Of the different scenarios proposed here, two concern alternative geographical routes: (1) via the Rio Grande Rise-Walvis Ridge axis in the South Atlantic; or (2) via Antarctica, possibly including the Subantarctic islands. Three scenarios consider different dispersal routes: (1) stepping-stones; (2) single-step; and (3) sweepstakes. Finally, three vectors of dispersion are considered: (1) birds; (2) wind; and (3) floating islands. Evaluation of these scenarios suggests that early-branching lineages of Asteraceae probably dispersed from South America to Africa along an island chain formed by the Rio Grande Rise and the Walvis Ridge, transported by birds, possibly combined with rafting and/or sweepstakes. Morphological changes typically associated with evolution on islands characterize many African carduoid descendants, providing indirect evidence for step-wise dispersal along the island chain.     

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.     

Polyploidy and new chromosome counts in Helichrysum (Asteraceae,Gnaphalieae)

Mitotic chromosome numbers are reported for 31 populations representing 28 taxa of Helichrysum. Twelve are new and eight others provide confirmation of a unique previous reference. A new chromosome number, 2n = 42, is reported for H. odoratissimum. Polyploidy is confirmed as the most significant evolutionary trend in chromosome number within the genus. Chromosome data agree with trends observed in phylogenetic studies: a South African and diploid origin of the genus, followed by a radiation and diversification in southern Africa and several migrations towards the north of the African continent, the Mediterranean basin and Asia. Expansion and diversification of the genus have been accompanied by several genome duplications which have led to the acquisition of the tetraploid, hexaploid and octoploid levels, all in several independent events. Both autopolyploidy and allopolyploidy are suggested as probable speciation agents within the genus. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 511–521.     

Karyological evolution in Rhaponticum Vaill. (Asteraceae, Cardueae) and related genera

Chromosome counts are reported in 29 populations representing 20 species and one subspecies of the genera Callicephalus (one sp.), Klasea (seven spp.), Myopordon (two spp.), Oligochaeta (one sp.), and Rhaponticum (nine spp., one ssp.). Eleven reports are new, the others supplement limited previous data. New and published cytological data are summarized and evaluated critically in light of current taxonomic treatments and evolutionary hypotheses. Basic chromosome numbers are a valuable source of taxonomic information and relate well to the phylogeny. They are quite conserved in Klasea (only x  = 15), the sister group of the remaining genera, and in the early branching genera of the Rhaponticum group ( x  = 14 for Centaurothamnus and Callicephalus ). However, a more complex pattern of genome evolution is present among the more derived clades, Oligochaeta plus Myopordon ( x  = 12, 14) and the genus Rhaponticum ( x  = 12, 13). The occurrence of x  = 12 in Oligochaeta divaricata and Rhaponticum carthamoides seems to be the result of a recent event of chromosome fusion, as suggested by the fragility of particularly long pairs of chromosomes. The use of chromomycin staining, including an enzymatic digestion step that facilitates spreading and permits a reduction of potential fragmentation, only partially resolves this problem.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 193–201.     

Jalcophila boliviensis,a new species of South American Asteraceae (Gnaphalieae)

The new species, Jalcophila boliviensis (Asteraceae: Gnaphalieae) is described and illustrated. A discussion of the systematic position of Jalcophila Dillon & Sagást. and a key to the three species of the genus are presented.     

Floral development and evolution of capitulum structure in Anacyclus (Anthemideae,Asteraceae)

Background and Aims

Most of the diversity in the pseudanthia of Asteraceae is based on the differential symmetry and sexuality of its flowers. In Anacyclus, where there are (1) homogamous capitula, with bisexual, mainly actinomorphic and pentamerous flowers; and (2) heterogamous capitula, with peripheral zygomorphic, trimerous and long-/short-rayed female flowers, the floral ontogeny was investigated to infer their origin.

Methods

Floral morphology and ontogeny were studied using scanning electron microscope and light microscope techniques

Key Results

Disc flowers, subtended by paleae, initiate acropetally. Perianth and androecium initiation is unidirectional/simultaneous. Late zygomorphy occurs by enlargement of the adaxial perianth lobes. In contrast, ray flowers, subtended by involucral bracts, initiate after the proximal disc buds, breaking the inflorescence acropetal pattern. Early zygomorphy is manifested through the fusion of the lateral and abaxial perianth lobes and the arrest of the adaxials. We report atypical phenotypes with peripheral ‘trumpet’ flowers from natural populations. The peripheral ‘trumpet’ buds initiate after disc flowers, but maintain an actinomorphic perianth. All phenotypes are compared and interpreted in the context of alternative scenarios for the origin of the capitulum and the perianth identity.

Conclusions

Homogamous inflorescences display a uniform floral morphology and development, whereas the peripheral buds in heterogamous capitula display remarkable plasticity. Disc and ray flowers follow different floral developmental pathways. Peripheral zygomorphic flowers initiate after the proximal actinomorphic disc flowers, behaving as lateral independent units of the pseudanthial disc from inception. The perianth and the androecium are the most variable whorls across the different types of flowers, but their changes are not correlated. Lack of homology between hypanthial appendages and a calyx, and the perianth double-sided structure are discussed for Anacyclus together with potential causes of its ray flower plasticity.     

Cypsela micromorphology and anatomy in Cirsium sect. Epitrachys (Asteraceae,Carduoideae) and its taxonomic implications

Cirsium Miller, one of the largest genera of the Asteraceae (Carduoideae), contains more than 250 taxa. Turkey is one of its diversity centers and according to recent studies this genus is represented by 80 taxa. In the present study, cypsela morphologies, micromorphologies and anatomical structures of 17 C. sect. Epitrachys taxa distributed in Turkey are characterized in detail by using stereomicroscope, light microscope (LM) and scanning electron microscope (SEM). Cypselas are found to be obovate to narrowly oblong, or rarely oblanceolate. Based on SEM characters, six cypselas surface ornamentation types are identified, pericarp anatomical structures are described, as well as the structure of the testa. Crystal formations were observed in three taxa and secretory materials in the mesocarp were evaluated. Data obtained from this study are compared with previously published results and it is concluded that fruit micromorphological and anatomical characters are useful for the delimitation of taxa in Cirsium and may be useful for supraspecific classification.     

Pollen studies in the genus Echinops L. and Xeranthemum group (Asteraceae)

The phylogeny of the genus Echinops and the Xeranthemum complex were the subject of a previous molecular study of the tribe Cardueae. An electronic microscope survey of pollen morphology and ultrastructure is presented for all the genera belonging to these groups, allowing four pollen types to be identified. The exine pattern indicates that there are two pollen types in the genus Echinops (including Acantholepis ), although no differences have been found between sections in Echinops . First, the annual species of Echinops and Acantholepis orientalis show a pollen type that differs from that of perennial species of the former genus and, second, the genera Chardinia and Xeranthemum exhibit a pollen type distinct from that Amphoricarpos and Siebera . The internal structure of the pollen wall is the same for all the genera. Only slight differences were observed in the tectum, but in all cases these were of the anthemoid type. Statistically significant correlations between pollen size and the life cycle were found.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 549–557.     

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.     

Karyomorphology of Turkish species in Centaurea sections Centaurea and Phalolepis (Asteraceae) and implications for taxonomy

In this paper, 31 mostly endemic and locally distributed Turkish Centaurea taxa belonging to Centaurea and Phalolepis sections were examined in terms of their karyomorphology. The basic chromosome number for all of the studied species was concurringly determined as 9(x = 9) for both sections, excluding C. hierapolitana. We also determined tetraploid and hexaploid species in spite of many species having diploid chromosome numbers. Twelve chromosome counts are reported for the first time and most of the karyotyping analyses are described for the first time via the KAMERAM program. The karyotypes had a predominance of metacentric (m) chromosomes. However, in the karyotyping of six taxa, submetacentric (sm) chromosomes were dominant. Five quantitative asymmetric indices were used to evaluate karyological features of the species. A meaningful dendrogram was carried out to assess the karyotype-symmetry conditions and describe the karyotyping relationships between different taxa.     

Genome size in Echinops L. and related genera (Asteraceae, Cardueae): karyological, ecological and phylogenetic implications

Genome size was assessed by flow cytometry in 33 species belonging to seven genera of the tribe Cardueae (Asteraceae), which can be grouped in three taxonomic complexes. 2C nuclear DNA content ranged from 1.49 to 16.98 pg, which is more than elevenfold variation. Genome size correlated well with some karyological traits. Nuclear DNA amount variations also have systematic and evolutionary implications and/or are linked to adaptations to ecological conditions.     

Elucidative studies on the generic concept of Senecio (Asteraceae)

VINCENT, P. L. D. & GETLIFFE, F. M., 1992. Elucidative studies on the generic concept of Senecio (Asteraceae) . This paper presents the results of studies of the generic concept of Senecio sensu stricto. The sample of taxa studied consisted of 93 Natal senecios (including seven varieties and two forms), five Cape heterochromous senecios and nine non-southern African senecios, including the type of the genus, S. vulgaris L. Also included in the study were six species from taxonomically closely related genera in the tribe Senecioneae and one Senecio of uncertain taxonomic position. The phenotype of these taxa was investigated with respect to a large number (122) of morphological and micromorphological characters. Six characters were selected as being taxonomically important with respect to elucidating the generic concept of Senecio sensu stricto. The generic concept of Senecio has been provisionally re-circumscribed and the generic status of each of the senecios and non-senecios studied has been tested according to this concept of Senecio sensu stricto sensu Vincent. In the light of this concept of Senecio , the following species are recommended for exclusion from Senecio sensu stricto sensu Vincent: S. cissampelinus, S. transvaalensis, S. syringifolius and S. hockii. The following species are considered to be peripheral to Senecio sensu stricto sensu Vincent: S. tanacetopsis, S. seminiveus, S. medley-woodii, S. tamoides, S. helminthioides, S. barbertonicus, S. brevilorus, S. viminalis, S. radicans and S.fulgens. Before any taxonomic changes are made to the current composition of Senecio , the concept of Senecio sensu stricto sensu Vincent, is being tested on a worldwide sample of the genus.     

Classification, diversity, and distribution of Chilean Asteraceae: implications for biogeography and conservation

This paper provides a synopsis of the Chilean Asteraceae genera according to the most recent classification. Asteraceae is the richest family within the native Chilean flora, with a total of 121 genera and c . 863 species, currently classified in 18 tribes. The genera are distributed along the whole latitudinal gradient in Chile, with a centre of richness at 33°–34° S. Almost one-third of the genera show small to medium-small ranges of distribution, while two-thirds have medium-large to large latitudinal ranges of distribution. Of the 115 mainland genera, 46% have their main distribution in the central Mediterranean zone between 27°–37° S. Also of the mainland genera, 53% occupy both coastal and Andean environments, while 33% can be considered as strictly Andean and 20% as strictly coastal genera. The biogeographical analysis of relationships allows the distinction of several floristic elements and generalized tracks: the most marked floristic element is the Neotropical, followed by the antitropical and the endemic element. The biogeographical analysis provides important insights into the origin and evolution of the Chilean Asteraceae flora. The presence of many localized and endemic taxa has direct conservation implications.