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The genus Blumea (Asteroideae, Asteraceae) comprises about 100 species, including herbs, shrubs, and small trees. Previous studies have been unable to resolve taxonomic issues and the phylogeny of the genus Blumea due to the low polymorphism of molecular markers. Therefore, suitable polymorphic regions need to be identified. Here, we de novo assembled plastomes of the three Blumea species B. oxyodonta, B. tenella, and B. balsamifera and compared them with 26 other species of Asteroideae after correction of annotations. These species have quadripartite plastomes with similar gene content, genome organization, and inverted repeat contraction and expansion comprising 113 genes, including 80 protein‐coding, 29 transfer RNA, and 4 ribosomal RNA genes. The comparative analysis of codon usage, amino acid frequency, microsatellite repeats, oligonucleotide repeats, and transition and transversion substitutions has revealed high resemblance among the newly assembled species of Blumea. We identified 10 highly polymorphic regions with nucleotide diversity above 0.02, including rps16‐trnQ, ycf1, ndhF‐rpl32, petN‐psbM, and rpl32‐trnL, and they may be suitable for the development of robust, authentic, and cost‐effective markers for barcoding and inference of the phylogeny of the genus Blumea. Among these highly polymorphic regions, five regions also co‐occurred with oligonucleotide repeats and support use of repeats as a proxy for the identification of polymorphic loci. The phylogenetic analysis revealed a close relationship between Blumea and Pluchea within the tribe Inuleae. At tribe level, our phylogeny supports a sister relationship between Astereae and Anthemideae rooted as Gnaphalieae, Calenduleae, and Senecioneae. These results are contradictory to recent studies which reported a sister relationship between “Senecioneae and Anthemideae” and “Astereae and Gnaphalieae” or a sister relationship between Astereae and Gnaphalieae rooted as Calenduleae, Anthemideae, and then Senecioneae using nuclear genome sequences. The conflicting phylogenetic signals observed at the tribal level between plastidt and nuclear genome data require further investigation.

Asteraceae account for 10% of all flowering plant species, and 35%–40% of these are in five closely related tribes that total over 10 000 species. These tribes include Anthemideae, Astereae, Calenduleae, Gnaphalieae, and Senecioneae, which form one of two enormous clades within Subfamily Asteroideae. We took a phylogenomics approach to resolve evolutionary relationships among these five tribes. We sampled the nuclear and plastid genomes via HybSeq target enrichment and genome skimming, and recovered 74 plastid genes and nearly 1000 nuclear loci, known as Conserved Orthologous Sequences. We tested for conflicting support in both data sets and used network analyses to assess patterns of reticulation to explain the early evolutionary history of this lineage, which has experienced whole‐genome duplications and rapid radiations. We found concordance and conflicting support in both data sets and documented four ancient hybridization events. Due to the timing of the early radiation of this five‐tribe lineage, shortly before the Eocene–Oligocene extinction event (34 MYA), early lineages were likely lost, obscuring some details of their early evolutionary history.

We present the hitherto most comprehensive phylogenetic hypothesis for Baccharis. Our results confirm that subtribe Baccharidinae is a monogeneric group and support a broad circumscription of Baccharis as monophyletic with the inclusion of all genera that have been previously segregated from it (Baccharidastrum, Baccharidiopsis, Heterothalamulopsis, Heterothalamus, Lanugothamnus). We provide comprehensive lists of accepted and synonymized infrageneric taxa. Seven main lineages are recognized, which we treat as subgenera. Subgenera Baccharis and Molina roughly match previous circumscriptions, while subgenera Pteronioides and Stephananthus are merged with Molina. In total, we propose that Baccharis comprises 440 species classified into 7 subgenera and 47 sections, while 22 sections are considered synonyms. The phylogeny provided constitutes the basis for further exploration of the evolutionary processes that have allowed the group to attain its wide geographic distribution, morphological variation, variable sexual reproductive strategies, and complex chemical contents.

Haplopappus paucidentatus Phil. (Compositae: Astereae: Machaerantherinae) is described and illustrated in colour and with a black and white text figure. Cultivation and propagation are discussed, most stocks better maintained using cuttings or layering. It is not apparently available commercially, but could be searched for under the ‘catch all’ name of H. glutinosus. Haplopappus Cass. is now recognized as a South American genus, considerably smaller than originally conceived by Hall in his major revision. It consists of 56 species in three subgenera, the largest, subgenus Haplopappus, contains 2 sections, section Haplopappus containing H. paucidentatus and H. glutinosus Cass. within which it has been included in several treatments.

Corethrogyne filaginifolia (Hook. & Arn.) Nutt. (Compositae: Astereae: Machaerantherinae) is illustrated and described. The full synonymy of this monospecific North American genus is provided together with the location of known type material and relevant comments made where applicable. The Conservation Status of the species is considered LC (Least Concern), as previous listings in California are shown to refer to Corethrogyne filaginifolia (Hook. & Arn.) var. incana (Lindl.) D.R. Morgan & R.L. Hartman, which is now considered a synonym of the rather variable Dieteria canescens (Pursh) Nutt. The taxonomic placement of Diplopappus incanus Lindl. is discussed, and a synonymy has been provided for Dieteria canescens subsp. canescens var. incana (Lindl.) D.R. Morgan & R.L. Hartman, of which it is the basionym. The location of the probable holotype of Diplopappus leucophyllus Lindl. ex DC. is discussed, together with the relevance of material mounted on a Kew sheet.

Aster saxicola W. P. Li & Z. Li, a new species of Asteraceae from southeastern Guizhou province, China, is described and illustrated based on morphological, molecular and cytological data. Morphological comparisons showed that A. saxicola is similar to A. oliganthus, but can be distinguished from the latter by its purple abaxial surface of the lower and middle cauline leaves, cordate‐ovate basal leaves with strigose hairs, outer three‐seriate phyllaries with purple or purplish apex, 9–14 ray florets and 10–18 disk florets. Phylogenetic analyses based on nrDNA ITS, ETS and plastid trnL–F DNA sequence data support that A. saxicola is a distinct species and belongs to Aster subgen. Aster sect. Ageratoides. Cytological observation shows that the new species is diploid with a karyotype formula 2n = 2x = 18 = 16m + 2sm. The new species should be considered endangered (EN) based on the IUCN red list criteria because of its restricted geographic range.

Aim Regions with a mediterranean climate are among the most species-rich regions in the world. They are also among the regions with the greatest number of non-native invasive species. However, not all introduced species become invasive. Previously, we found that non-native Asteraceae closely related to native species are more likely to establish and spread in California, USA. We conducted reciprocal studies in another mediterranean-climate region with high species richness, the Italian Peninsula. Mediterranean Basin invaders in the Astereae mostly originated in temperate North America, whereas the invasive Cardueae of California are primarily from the Mediterranean Basin. Location The Mediterranean Basin (Italy). Methods Utilizing the first fossil-calibrated molecular phylogenies of the tribe Astereae, we explored the evolutionary patterns among native and non-native taxa within this group. To elucidate the mechanisms driving the observed patterns, we modelled the ecological niches of Astereae species on a global scale and examined correlations between ecological similarity and evolutionary relatedness among taxa. Results Not only were invasive taxa highly clustered among natives, they were also more closely related to natives than were non-invasive introduced taxa. We found that invasive species had higher degrees of niche overlap with native species than did non-invasive introduced species. Moreover, pairwise niche distance among species was significantly correlated with evolutionary time, indicating that the ecological niche is phylogenetically conserved. Main conclusions The results of our study are consistent with those from our work in California, suggesting that introduced Asteraceae taxa that are closely related to native species are likely to become invasive on both sides of the Atlantic. Furthermore, existing ecological attributes seem to drive these evolutionary patterns. We suggest that potential invaders closely related to natives should be considered as high priorities for management.

The family Asteraceae has a particular inflorescence, the capitulum, consisting of ray florets and disc florets. The ray florets function as petals that attract pollinators. Marked variation in the ray floret morphology is known in a natural population of Aster hispidus var. tubulosus (Asteraceae). We analyzed the variation and found two distinct types in the ray florets, the long tubular ray floret and the ligulate ray floret. In this species, therefore, the variation in floral morphology among capitula, each of which is the basic pollination unit, is caused by the variation in the composition of the two ray floret types among capitula. We evaluated the sources of the observed variation in the floral morphology among capitula within a population using a hierarchical analysis that separated within-individual (i.e. among capitula within each individual) and between-individual components of the variation. We found that the main source of the variation lay at the between-individual level, not at the between-capitulum level nested within individuals. This finding will provide the basic knowledge that enables future study exploring whether the between-individual variation in floral morphology caused by the compositional variation of the ray floret types leads to differential pollination success of individual plants in species of Asteraceae.

Erigeron karvinskianus DC. (Compositae: Astereae: Conyzinae) is described and illustrated, and its cultivation and propagation discussed. The etymology of the species name is explained with a short biography of Baron Karwinsky von Karwin.

The new species names Mairia petiolata, Zyrphelis capensis, Z. glabra, Z. glandulosa, Z. grauii, Z. macrocarpa, Z. nervosa,Z. spathulata, the new subspecies names Mairia hirsuta subsp. robusta, Zyrphelis ecklonis subsp. oligocephala, Z. ecklonissubsp. strictifolia, Z. microcephala subsp. foliosa, Z. microcephala subsp. glandulosa and the new combinations Zyrphelisciliaris, Z. ciliaris subsp. angustifolia, Z. ciliaris subsp. hirsuta, Z. decumbens, Z. fruticosa, Z. gracilis, Zyrphelis sect. Gymnostephium,Z. leiocarpa, Z. levis, Z. montana, Z. monticola, and Z. outeniquae are validly published. These names originallyappeared in the Ph.D. thesis of the second author, Ursula Zinnecker-Wiegand, submitted to the University of Munich, Germany,in 1990. Being printed and distributed to over 50 libraries, this thesis met the formal requirements for effective publicationof the International Code of Botanical Nomenclature then in force, but it is no longer effectively published as a result of aretroactive change in the rules adopted at the International Botanical Congress in Vienna in 2005. As result, the new namesappearing in the thesis were not validly published and they are validated here.

Baccharis paucicostata Joch. Mull. & Giuliano from the Andes of Argentina is here described as a new species and illustrated. Characters for delimitation from similar species and known data on distribution and habitat are provided.

 Premise of the study : The abundance of fossils in Antarctica suggests this continent was a center of diversifi cation and a corridor for migration for many austral plant groups until the late Tertiary and may have played a pivotal role in shaping plant distributions in the southern hemisphere. Although the Antarctic fl ora was largely erased by glaciation during the Pleistocene, at least some Antarctic plant species found refuge on the subantarctic islands.  Methods : We used independent and combined analyses of ITS, ETS, trnK , and trnL DNA sequences to infer phylogenetic relations in Pleurophyllum , a small genus of three species that are endemic to the subantarctic islands of Australia and New Zealand. The inferred phylogeny provided a framework to reconstruct the origin and patterns of diversifi cation in the genus.  Key results : We summarize support for the hypothesis that Pleurophyllum survived episodes of Pleistocene glaciation in the subantarctic islands and that its sisters dispersed northward in response to glacial advance.  Conclusions : The distinctive fl ora of the subantarctic islands includes some of the last remnants of a once-diverse Antarctic fl ora. These plants may still retain distinctive features of their ancestors. Studies of endemic plants such as Pleurophyllum are the key to resolving this puzzle.

Aqueous and organic extracts of 25 selected species from four tribes of Hungarian Asteraceae were screened in vitro for antiproliferative activity against HeLa (cervix epithelial adenocarcinoma), A431 (skin epidermoid carcinoma) and MCF7 (breast epithelial adenocarcinoma) cells, using the MTT assay. Twenty five of the 228 tested extracts from different parts of the species of Astereae (6), Inuleae (3), Heliantheae (5) and Anthemideae (11) demonstrated a substantial antiproliferative effect (at least 50% inhibition of cell proliferation) at 10 microg/mL against one or more of the cell lines. Complete dose-response curves were generated and IC(50) values were calculated for these active extracts, and their direct cytotoxic effects were determined. In summary, 11 of the tested 25 plants were found to be active and 4 of them (Anthemis ruthenica, Erigeron canadensis, Erigeron annuus and Inula ensifolia) had not been studied previously for either active compounds or anticancer properties.

The embryological features of microsporangiate wall formation, microsporogenesis and male gametogenesis in Aster subulatus, Kalimeris indica, Heteropappus arenarius and Erigeron annuus are described for the first time. Although external morphology might suggest that these four species differ from each another, we found no distinctly different embryological features. They all share similar characteristics, such as a tetrasporangiate anther, coexisting amoeboid and glandular tapeta, simultaneous cytokinesis (except for 12% occurrence of successive cytokinesis in E. annuus), 3-celled mature pollen grains and endothecium of anther wall that does not develop fibrous thickenings. Based on the embryological data, we suggest that Kalimeris and Heteropappus may be included within Aster, and that Erigeron is closely related to Aster sensu lato.

The systematic position of the monoecious rheophyte, Sheareria (Asteraceae), a genus endemic to China, has been long disputed and never rigorously examined. In this study, two molecular datasets (trnL-F, ITS), Plus morphological and cytological characters, were used to assess the systematic position of Sheareria. The broader phylogenetic analyses based on the trnL-F data placed Sheareria in the tribe Astereae with strong support. This placement was also supported by the lanceolate style, "Asteroid type" ligule epidermis, yellow anthers, green achenes, and basic chromosome number of x = 9. Analysis of the tribe Astereae based oil the ITS dataset showed that Sheareria and three East Asian genera (Kalimeris, Aster, Rhynchospermum) formed a strongly Supported clade characterized by flat, 2-ribbed, glandular disc achenes. Detailed morphological comparisons indicate that Sheareria is more similar to Kalimeris than to Rhynchospermum. Within this clade, Sheareria is distinguished from other genera by its monoecious breeding system (vs. gynomonoecious). Based on fossil and geological evidence, the development of monoecy in Sheareria is discussed. Within Sheareria, only one species, S. nana, is recognized; S. leshanensis, a newly described species, is placed in synonymy.

Aim The aims of this study were (1) to investigate whether the two growth forms of Darwiniothamnus Harling (Asteraceae) originated from the colonization of a single ancestor, (2) to identify the closest relative(s) of Darwiniothamnus, and (3) to review molecular phylogenies from other plant groups to infer the origin of Galapagos endemics. Location Darwiniothamnus is endemic to the Galapagos Islands. Methods All putative relatives of Darwiniothamnus plus 38 additional species were included. Nucleotide sequences of the internal transcribed spacers of the nuclear ribosomal DNA were used for Bayesian and parsimony analyses. Results Darwiniothamnus is polyphyletic. Two species (D. lancifolius (Hook. f.) Harling and D. tenuifolius (Hook. f.) Harling) are woody shrubs that usually grow to 1–2 m in height; they belong to a clade composed of species otherwise restricted to the Caribbean. These two species are sister to Erigeron bellidiastroides Griseb., a herbaceous species endemic to Cuba. The third species (D. alternifolius Lawesson & Adsersen) is a perennial herbaceous plant, woody at the base and reaching only up to 50 cm in height. It is sister to two Chilean (Coquimbo–Valparaiso region) species that also have a perennial herbaceous habit: E. fasciculatus Colla and E. luxurians (Skottsb.) Solbrig. They are placed in an assemblage restricted to South America. The review of previous molecular phylogenetic studies revealed that two of the endemic genera and endemic species of three non-endemic genera have their closest relatives in South America. Endemic species belonging to three non-endemic genera have sister species in North America or the West Indies. One endemic genus and endemic species in three non-endemic genera have sister taxa with a widespread continental distribution, or their molecular phylogenies yielded equivocal results. Main conclusions The flora of Galapagos has affinities with both North America (including the Antilles) and South America. Darwiniothamnus exhibits both patterns: two species of this genus are sister to a taxon endemic to Cuba, supporting a connection between the Cocos plate and the West Indies; the third species, D. alternifolius, provides a link with the Coquimbo–Valparaiso region, suggesting a biogeographical connection between the Nazca plate and southern South America.

Nardophyllum cabrerae (Asteraceae: Astereae), a new species from western Argentina is described. The new species differs from its closest relative, Nardophyllum lanatum (Meyen) Cabrera, in its non-revolute margins, white corollas, and pappus of unequal scabrid bristles. Two names, Nardophyllum obtusifolium Hook. et Arn. and N. bracteolatum Hauman, were shown not to represent distinct species and accordingly placed in the synonymy of Nardophyllum bryoides (Lam.) Cabrera and Nardophyllum chiliotrichioides (J. Remy in Gay) A. Gray, respectively.

Studies of herbivorous insects have played a major role in understanding how ecological divergence can facilitate genetic differentiation. In contrast to the majority of herbivorous insects, grasshoppers as a group are largely polyphagous. Due to this relative lack of intimate grasshopper-plant associations, grasshopper-plant systems have not played a large part in the study of host-associated genetic differentiation. The oligophagous grasshopper, Hesperotettix viridis (Thomas), is endemic to North America and feeds on composites (Asteraceae) within the tribe Astereae. Previous work has shown both preference and performance differences between H. viridis individuals feeding on either Solidago mollis or Gutierrezia sarothrae. Using 222 AFLP markers, we examined the genetic relationships among 38 H. viridis individuals feeding on these plants both in sympatry and allopatry. Neighbour-joining analysis resulted in two distinct host-associated clades with 71% bootstrap support for host-associated monophyly. Analyses of molecular variation (amova) revealed significant genetic structuring with host plant accounting for 20% of the total genetic variance while locality accounted for 0%. Significant genetic differentiation was detected between S. mollis-feeders and G. sarothrae-feeders even when the two were present at the same locality. These results are consistent with observed differences in preference and performance between H. viridis grasshoppers feeding on either G. sarothrae or S. mollis and indicate that H. viridis is comprised of at least two genetically distinct host plant-associated lineages.

Galatella anatolica Hamzaoglu & Budak sp. nov. (Asteraceae), collected from Osmaniye (Turkey) is here described as a new species. It is similar to G. angustissima (Tausch) Novopokr. in general habit. Both have stems with few branches and 1-veined middle leaves, but are distinguished by involucral features, series of phyllaries, and lengths of disc florets, achenes and pappus.

As currently delimited, Solidagininae are a large (approximately 190 species) subtribe of tribe Astereae. Recent molecular and morphological studies have prompted a new definition of the subtribe, but the lack of absolutemorphological synapomorphies raises the possibility that this assemblage may not be monophyletic. Cladistic and likelihood-based analyses were conducted on a nuclear rDNA ITS sequence dataset derived from 23 of the 24 genera included in recent Solidagininae circumscriptions. Cladistic analyses identified two clades entirely composed of proposed Solidagininae genera. The data were not able to support deeper relationships, and these two clades might or might not form one monophyletic lineage. Topology testing indicated compatibility between the taxonomic definition of Solidagininae and molecular data.

Guynesomia (Asteraceae: Astereae), a new genus restricted to the western slopes of the Andes mountains in central Chile, is described. Guynesomia is a shrub characterised by raceme-like or panicle-like capitulescences, disciform capitula, alveolate receptacles, pistillate zygomorphic marginal florets with three or four short lobes, and a pappus of two series of subequal scabrid bristles. Diplostephium is the putative closest relative of Guynesomia. The new combination Guynesomia scoparia is based on Nardophyllum scoparium.

Previous molecular investigations of the phylogenetic relationship for Xylothamia employed exemplar taxa and indicated its affiliation is either with the Gutierrezia or the Ericameria alliances. The present 3' ETS and ITS nrDNA gene phylogenies yielded the unexpected result that Xylothamia is polyphyletic. Species within its circumscription are placed into two well-supported evolutionary lineages with other genera of Astereae. Of nine species of Xylothamia, four species plus Gundlachia constitute one clade. The other five species are placed in a sister clade with several other genera including Amphiachyris, Bigelowia, Euthamia, Gutierrezia, and others. Except for X. johnstonii and X. palmeri, these five species of Xylothamia have little affinity for one another or for other genera in the same clade. Morphological similarities in foliar features between certain species in both lineages are possibly from a convergence resulting in adaptation to arid habitats. Such similarities were also the basis, in part, for their taxonomic treatment within Xylothamia or their earlier affiliation with Ericameria, which has similar features. The uncharacteristic leaf form in Xylothamia purpursii has a pattern of adaptation considerably different from its allies. Whether the Xylothamia-containing clades are related to Solidaginineae or represent an independently derived lineage remains equivocal because this outcome rested with the choice of data and optimality criteria.

Morphological and ITS (internal transcribed spacer) sequence data for 40 species of the Austral-Pacific genera Camptacra, Kippistia, Minuria, Peripleura, Tetramolopium, and Vittadinia as well as one semiherbaceous species of Olearia were subjected to cladistic analysis, separately and together. Minuria, Peripleura, and Tetramolopium are paraphyletic as currently defined. Tetramolopium vagans from Australia appears to represent an undescribed genus. Both Kippistia suadefolia and Peripleura diffusa show close affinity to Minuria species, and Minuria macrorhiza appears to contain two distinct but closely related species. Vittadinia and the remaining species of Tetramolopium and Peripleura form a strong affinity group. The distribution of indels and the combined analysis each provide evidence that the Hawaiian and Cook Island species of Tetramolopium are descended from New Guinea species. The combined analysis also suggests that the Cook Island species T. mitiaroense is sister to the Hawaiian clade. Olearia arguta groups strongly with Camptacra and shows no close affinity with either of the arborescent species of Olearia used to root these analyses. Marked homoplasy among morphological characters indicates why generic delimitation in the group has been problematic.

The Astereae is the largest tribe of Asteraceae in North America. Morphological diversity suggests that the North American assemblage is polyphyletic as 12 endemic genera, as well as lineages of the genus Erigeron and Conyza (Conyzinae), have been hypothesized to represent at least five separate invasions of North America from Africa, Australia, Eurasia, and South America. This hypothesis was tested with a phylogenetic analysis of nucleotide sequence data from the internal transcribed spacers (ITS) of nuclear ribosomal DNA. Sequences for 62 taxa represent seven outgroup taxa and all major Northern and Southern Hemisphere groups of Astereae, including broad taxonomic and geographic sampling of Conyzinae and Aster s.l. (sensu lato). Parsimony analyses indicate that all North American Astereae are members of a strongly supported clade, and that a diverse group of predominantly woody taxa from Africa, Australia, and South America, are basal Astereae. Furthermore, Aster s.l. is deeply polyphyletic as Eurasian taxa, including Aster s.s. (sensu stricto), appear more closely related to Southern Hemisphere taxa than to North American Aster segregates. There is only low to moderate agreement between proposed higher level Astereae relationships based on ITS and those based either on morphology or chloroplast restriction site data.

A new subspecies of the widespread and polymorphic B. salicifolia from the Andes of central and southern Peru is described and illustrated. The morphological delimitation of the new taxon is discussed.

Intermediate specimens between Grindelia chiloensis and Haplopappus pectinatus occur in Patagonia (Argentina). They were analysed using the methods proposed by Wells (distance diagram) and Wilson (character count procedure). With both methods the hypothesis of hybridity is supported. A close proximity of H. pectinatus to the genus Grindelia is also suggested by a high percentage of stainable pollen grains in the putative hybrids, chromosome morphology, and achene anatomy. Based on such evidence, the new combination G. anethifolia (H. pectinatus Phil., non G. pectinata Baker) is proposed.

Anderberg, A. A. & Karis, P. O. 1995. Psednotrichia, a genus of the tribe Senecioneae hitherto misplaced in the Astereae (Asteraceae). — Nord. J. Bot. 15: 375–379. Copenhagen. ISSN 0107–055X. The tribal position for the little known genus Psednorrichia, is established. This monotypic genus, hitherto placed in the tribe Astereae, is a congener of Xyridopsis of the tribe Senecioneae (Asteraceae). The two species of Xyridopsis are here transferred to Psednotrichia and the new combinations P. xyridopsis and P. newtonii are made. A brief discussion of the morphology of the genus, and its systematic position within the tribe Senecioneae is provided.

Premise of the study: Simple sequence repeat (SSR) markers were developed for Aster savatieri (Asteraceae) and the serpentine variety A. savatieri var. pygmaeus to re-evaluate their taxonomic status. Methods and Results: Using RNA-Seq data, 22 expressed sequence tag (EST)-SSR markers were developed. Polymorphisms were assessed in A. savatieri and in A. savatieri var. pygmaeus. The average number of alleles ranged from four to 15, and expected heterozygosity ranged from 0.417 to 0.870. Transferability was examined in six representative species of Japanese Aster and in Solidago virgaurea subsp. asiatica var. asiatica, a member of the tribe Astereae (Asteraceae); most of the loci were transferable to these examined species. Conclusions: These markers will be useful for genetic studies of variation in A. savatieri and other Aster species that occur in Japan.

The genus Kalimeris with a diagnostic character of short or inconspicuous pappus consists of two sections, Asteromoea and Cordifolium. As a result of 6PGD isozyme analysis, sect, Asteromoea, including 2 × and poly-ploid taxa from 5 × to 8 ×, show similar cytosolic isozyme multiplicity and share a monomorphic locus. The data suggest that gene duplication of polyploid members was derived from a common ancestor. K. miqueliana, belonging to sect. Cordifolium. also possessed a gene duplication in 6PGD, though significant differences were detected in electrophoretic mobility between the sections. The occurrence of gene duplication in East Asian diploid Astereae leaves intact the validity of the allopolyploid-origin hypothesis of n= 9, which was rejected by Gottlieb (1981a) in American Astereae.

Summary Robinson, H.: Notes on the tribes Eremothamneae, Gundelieae, and Moquinieae, with comparisons of their pollen. - Taxon 43: 33-44. 1994. - ISSN 0040-0262. The new tribe Moquinieae is established, and scanning electron micrographs of pollen are presented of Hoplophyllum of the Eremothamneae, Gundelia of the Gundelieae, and Pseudostifftia of the Moquinieae. For most of a hundred years the number of recognized tribes in the Asteraceae remained rather constant as defined by Bentham (in Bentham & Hooker, 1873). Bentham accepted only thirteen tribes: Vernonieae, Eupatorieae, Astereae, Inulae, Heliantheae, Helenieae, Anthemideae, Senecioneae, Calenduleae, Arctotideae, Cynareae, Mutisieae, and Cichorieae. Of these, only the Helenieae were described as new by Bentham. Some additional tribes recognized by Cassini (Solbrig, 1963) were placed in the synonymy of the Senecioneae, Cynareae, and Mutisieae by Bentham. In spite of numerous errors of delimitation, the Bentham concepts were maintained by Hoffmann (1890-1894) and in the review of the Asteraceae by Cronquist (1955), and they were the working basis for the symposium on the family edited by Heywood & al. (1977). Twentieth Century corrections on the Bentham concepts include the transfer of Ursinia Gaertn. from the Arctoteae to the Anthemideae (Beauverd, 1915). The latter position is accepted today in spite of a difference in the pollen on which a new tribe Ursinieae (Robinson & Brettell, 1973b) was based. The value of the Helenieae was considered dubious by many authors (Cronquist, 1955), but Stuessy (1977) was the first since Bentham to place Helenium L. in the Heliantheae. Robinson (1981) subsequently treated the Helenieae and many genera from the Senecioneae as members of the Heliantheae. Descriptions of new tribes after Bentham & Hooker (1873) started with Rydberg's Adenocauleae (Rydberg, 1917), which are tentatively accepted here; Iveae (Rydberg, 1917), now placed in the Heliantheae, subtribe Ambrosiinae; Liabeae (Rydberg, 1927), redescribed and accepted by Robinson & Brettell (1973a, 1974) and Robinson & Marticorena (1986); Neurolaeneae (Rydberg, 1927), now placed in the Heliantheae; and Ursineae (Robinson & Brettell, 1973b, now placed in the Anthemideae. The Eremothamneae, described by Robinson & Brettell (1973c), have been recently reviewed and expanded (Karis, 1992; Robinson, 1992). The Gnaphalieae, named by Bentham (1867), but included in the Inuleae in Bentham & Hooker (1873), were renamed by Rydberg (1917), and are now accepted by Anderberg (1991a). Strother (1977) resurrected the Tageteae Cass. In the same symposium, Turner & Powell (1977) named the tribe Coreopsideae, emphasizing cytochrome C differences between Guizotia Cass., traditionally placed in the subtribe Coreopsidinae, and Helianthus L.

The anatomy of internal secretory spaces in immature and mature aerial stems, rhizomes, and roots of common goldenrod, Solidago canadensis (Asteraceae, tribe Astereae), was studied from resin-embedded samples cut 1–2 μm thick. Oil-filled cavities of different lengths, each lined by a uniseriate epithelium, form in young stems and rhizomes. Epithelial cells enlarge and elongate, and some continue to divide mitotically. The cavities thereby expand and accommodate themselves to the growing stem and rhizome. Septa are sometimes forced open by this enlargement, which merges adjacent cavities. Oil reservoirs in roots are probably also cavities, but their lack of densely cytoplasmic epithelial cells prevented us from locating the end walls. Cavities in stem, root, and the cortex of rhizomes occur only as one median file located just outside of each phloem strand; in the pith of rhizomes, however, several files of cavities of larger diameter seem randomly distributed. This study extends our earlier findings that the oil reservoirs in leaves of this species are cavities; we have therefore verified that cavities, not ducts, permeate the goldenrod plant.

Phylogenetic relationships and composition of Ericameria were investigated by parsimony, maximum likelihood and Bayesian analyses of ITS and 3'ETS sequences for 50 taxa including all known species in Ericameria and several related taxa. The independent analyses of the data sets led to highly congruent, though not identical, results. Analyses of the combined sequence data generally improved resolution and clade support. Ericameria, Pentachaeta, Rigiopappus and Tracyina are resolved in a monophyletic group distinct from Chrysothamnus, Xylothamia, and representatives of the Solidago lineage. The expanded circumscription of Ericameria to include sections Asiris, Macronema and Stenotopsis, and species transferred from Chrysothamnus including E. albida, is supported. Within Ericameria the alignment of species into sections is not in total agreement with previously proposed relationships. Also, the molecular data do not support the treatment of section Stenotopsis as distinct from section Ericameria.

A B S T RA C T Relative amounts of nuclear DNA were determined in root tip cells of seven species of Astereae: Aster hydrophilus Greene, A. oblongifolius Nutt., A. riparius H.B.K., Machaeranthera boltoniae (Greene) Turner and Horne, M. brevilingulata (Sch-Bip.) Turner and Horne, M. parviflora Gray, and M. tenuis (S. Wats.) Turner and Horne. The results show that A. hydrophilus and M. brevilingulata, with a chromosome number of n = 9, have less nuclear DNA than other closely related species which are either n = 4 or n = 5. Cytological analyses of meiosis in the intergeneric hybrid M. parviflora X A. hydrophilus showed cells with two or more small chromosomes of the latter species pairing with single large chromosomes of the former. Pachytene cells of the hybrids M. parviflora X A. hydrophilus, M. parviflora X A. riparius, and M. boltoniae X M. tenuis showed some unpaired chromosome segments. The significance of these results to chromosome evolution in the tribe Astereae is discussed.

Variation in the frequency of individuals of Xanthisma texanum with B chromosomes among populations at the local level and variation in the frequency and distribution through time of individuals with B chromosomes within populations was investigated in a small area of southwestern Oklahoma and adjacent Texas between 1974 and 1987. The chromosome numbers of 1,224 individuals were determined from meiotic material; 15.8% of these had B chromosomes, less than 1% had three or four B chromosomes; none had more than four B chromosomes. The frequency of plants with B chromosomes among all sites sampled ranged from 0 to 33% and from 0 to 25% in sites separated by only a hundred meters; large differences in sample size occurred. Even at the subsite level large differences in frequencies in plants with B chromosomes were found. Within single sites, B chromosome frequency was not constant through time: the pattern of change differed among subsites within populations. In some cases the frequency went to zero. No correlations between environmental parameters and B chromosome frequency were

A B S T R A C T Reports of 209 original chromosome counts are made for the tribe Astereae of Compositae, including first counts for two genera and 46 species or subspecies. With over 80 % of the species counted, chromosome numbers are now available for all North American sections of Haplopappus. Two major groupings are apparent: one, with basic numbers of x = 4, 5, or 6, is basically herbaceous; the other, with x = 9, consists of shrubs or subshrubs. Aneuploidy is known only in the "herbaceous" group of Haplopappus, and polyploidy is more extensive there than in the woodier group of sections.

Summary The style of the Asteraceae varies in the direction of its branching. In examples studied in the subfamily Asteroideae the disk flowers or flowers of homogamous heads have style branches diverging radially to the center of the head. In that subfamily, the style branches in the ray flowers are usually tangentially directed in the Heliantheae, Inuleae and Senecioneae, and are radially directed in the Astereae and Anthemideae. In examples studied in the subfamily Cichorioideae, the Lactuceae and Vernonieae have tangential style branches in all flowers, Carduus of the Cardueae has styles all radial, they vary in the Liabeae, and they have a Helianthean pattern in Haplocarpha of the Arctoteae. It seems significant that tribes with basically different style orientations in the ray and disk flowers of the same head are usually tribes where many members have sterile gynoecia in either the rays or the disk. The style is functionally and taxonomically the most important structure of the Asteraceous flower; it has been extensively studied in relation to its stigmatic surfaces, its apical appendages, its basal pubescence and enlargements, and its internal ducts (Fig. 1). Nevertheless, various aspects of the style remain in need of study, one of which is the alignment of the style branches in relation to the orientation of the flower in the head. This aspect has been briefly reviewed in the Asteraceae by the present author to determine the variation in style orientation and to discover whatever order there might be in such variation. The most significant references to style orientation in the literature involve the analysis of style traces in the achene walls of various Heliantheae by Carlquist (1957) and Robinson (1981). In these papers, the styles of the disk flowers are shown to have branches diverging along the radii of the flowering head, and the style traces were shown to be continuous with veins on the outer and inner edges or surfaces of the achenes. These vascular traces were cited as evicence that the achenes of Helianthus having veins along their edges were morphologically compressed, whereas the achenes of Fitchia and the Coreopsidinae with the veins along the middle of the outer and inner faces were obcompressed. Because of the emphasis of these papers on one tribe of the Asteraceae, and the emphasis on genera such as Helianthus and Fitchia having only one type of style-bearing flower, the impression might have arisen that the styles of the Asteraceae are always oriented in the same radial direction. However, in some other tribes, and even in ray flowers of some Heliantheae in such subtribes as the Melampodiinae and Ecliptinae, the styles are oriented tangentially. Not only do the styles rotate to different positions in different Asteraceae, but they rotate in different flowers in the same head in some Asteraceae. The difficulties in determining the style orientation in Asteraceous flowers are sufficient to explain the previous lack of any survey of the character. The position of the style branches as seen in the live flowering heads is often misleading because of the twisting in the shaft. Dissection and microscopic examination are often needed. In radially symmetrical flowers

Chromosome counts are reported for 167 taxa representing 160 species and 76 genera of Compositae. First reports for 27 species and 2 infraspecific taxa are included. New chromosome numbers are reported for 9 additional taxa. The genus Carminatia DC. is merged with Brickellia Ell. resulting in two new combinations: Brickellia tenuiflora (DC.) Keil & Pinkava and Brickellia recondita (McVaugh) Keil & Pinkava. CHROMOSOME COUNTS serve as a reservoir of data for biologists concerned with systematic and evolutionary investigations. Unlike the morphological information content of specimens, which can be measured and remeasured by succeeding generations of botanists, chromosomal information is ephemeral and is not directly retrievable from a dried specimen. Consequently, documentation of chromosome numbers and meiotic irregularities serves as a unique source of information for both current and future investigators. In this publication, we are presenting 270 chromosome counts representing 76 genera, 160 species, and 7 additional infraspecific taxa from the Compositae (Table 1). Counts are published here for the first time for 27 species and for 2 infraspecific taxa and previously unpublished chromosome numbers are reported for 9 additional taxa. Both Table 1 and the accompanying discussion are arranged according to a modification of the tribal system of Hoffmann (1894). Discussion is provided only where it is necessary to add to or clarify material presented in the table. METHODS-Immature capitula were killed and fixed in the field in modified Carnoy's fixative (4 [or 6] chloroform: 3 ethanol: 1 glacial acetic acid; v:v). In a modification of the methods of Beeks (1955), floret buds were stained with ironacetocarmine and were squashed in a drop of Hoyer's solution. All chromosome counts were determined from meiotic microsporocytes and were documented with camera lucida drawings. OBSERVATIONS AND DISCUSSION-EUPATORIEAE-Barroetea is very closely related to Brickellia (Robinson, 1917). Both genera have the 1 Received for publication 22 January 1975. We acknowledge the cooperation of the curators of the following herbaria for making specimens available for our studies: ASU, GA, GH, MICH. We thank those collectors who contributed plant materials in support of our research. same base number (x = 9) and are very similar in habit. Barroetea apparently differs from Brickellia only in having setiferous teeth on the leaves and flattened rather than terete achenes. Further investigations may result in the merger of the two genera. Our first report of n = 9 for the plant long known as Carminatia tenuiflora DC. prompted a re-evaluation of the genus Carminatia. Carminatia is morphologically very similar to Brickellia (x = 9) in many respects but differs in having 5angled rather than 10-angled achenes and plumose rather than merely scabrous pappus bristles (Robinson, 1917). Flyr (1968) stated that the number of angles on the achene is not a consistent character for distinguishing Brickellia from its close relatives, and Shinners and McVaugh (1968), in arguing for the merger of Kuhnia and Brickellia, indicated that a pappus of plumose bristles is not a satisfact-ory generic character either. The presence or absence of hairs on the style bases, a character used by King and Robinson (1972) to separate Brickellia from Carminatia in Nueva Galicia, is not consistent since several species of Brickellia (e.g., B. scoparia (DC.) A. Gray, B. spinosa A. Gray) have glabrous styles bases. We have concluded that there is no justification for maintaining Carminatia as a genus distinct from Brickellia. We therefore propose the following new combinations: Brickellia tenuiflora (DC.) Keil & Pinkava, comb. nov. Carminatia tenuiflora DC. Prodr. 7: 267. 1838. Brickellia recondita (McVaugh) Keil & Pinkava, comb. nov. Carminatia recondita McVaugh, Contr. Univ. Mich. Herb. 9(4): 384. 1972. ASTEREAE-Aster is cytologically one of the most complex genera of the Astereae. Chromosomal evolution has apparently involved both aneuploidy and polyploidy, with polyploid series arising from several base numbers. Our new re

A B S T R A C T A survey of over 30 species of Aster and 61 species in other genera of Astereae indicates that the somewhat flattened style branches of perfect disk florets characteristically unfold and spread in a pronate, rather than a supinate manner. While not exhibited in all genera (Baccharis and Conyza are among the exceptions), this phenomenon appears to be restricted to and, therefore, diagnostic for the tribe. The slender style branches of pistillate ray florets display the supinate expansion which is regularly observed in other tribes of Compositae. THE CAPITULUM in genera of Astereae can be distinguished from that of other tribes in the Compositae by a combination of the following characteristics. Typically the flowers are of two kinds, both fertile. The ligulate zygomorphic or filiformtubular florets forming the outer series lack stamens; the inner disk florets are actinomorphic and perfect or, if sterile, are at least structurally hermaphroditic. Receptacles lack chaff but may be alveolate or somewhat paleaceous. Styles have no hairy or thickened ring below the branches. The two style branches show well-defined stigmatic lines; those of the disk florets typically are somewhat flattened and are provided with deltoid or lanceolate, hairy, non-stigmatic appendages of varying length; those of the ligulate florets are more slender, not flattened, and have marginal papillate stigmatic lines continuous over the whole length of the bifurcation. Anthers are truncate to sagittate at the base, not tailed. The pappus varies with different genera. In some it consists of capillary bristles, in others it is awned or somewhat paleaceous. Sometimes it is absent. The fact that style branches of the ligulate florets are morphologically distinct from those of the disk florets has been discussed in the literature (Cassini, 1826, 1834; Cronquist, 1955) but generally receives no mention in floristic treatments. Only the styles of hermaphroditic florets, with their sterile appendages, provide significant characteristics useful for diagnostic keys. Style branches of the pistillate ray florets vary for different species only in size, the degree of tapering, and exsertion from the corolla tube. This structural difference of styles is determined by function. The perfect disk florets in Astereae are proterandrous. Pollen is released before the stigmatic lines on the inside of the closely appressed style branches become exposed and receptive. At anthesis the style elongates rapidly and pushes

In leaves of goldenrod, Solidago canadensis (Asteraceae, tribe Astereae), numerous internal oil reservoirs with a uniseriate epithelium occur as a single file above or below veins or as isolated cavities in the mesophyll. Reservoirs are abaxial to major veins (vein orders 1–3), either above, below, or superimposed in intermediate 4th order veins, but strictly adaxial to 5th and 6th order minor veins. Reservoirs are initiated as discrete cavities, but those below 1st and 2nd order veins are in a single crowded file, each separated only by epithelial cells. Elongation of these cavities, accompanied by stretching and separation of septa, gives a false impression at maturity of an indefinitely long duct instead of a series of tubular cavities. Reservoirs of vein orders 3–6 are mostly more widely separated and less subject to elongation, thus they are shorter and remain discrete at maturity. The overall foliar pattern is one of successively shorter reservoirs, a sequence that is in concert with the successively narrower and progressively less elongated vein orders. The shift from abaxial to adaxial reservoirs in minor veins may be related to different phloem functions: sugar transport in major veins and photosynthate assimilation in minor veins.

Summary The tribal affinities and inter-relationships of the Californian-Chilean Blennosperma and the western American Crocidium (Compositae) have been the subject of controversy since the mid-nineteenth century. Most recent workers have placed Blennosperma in the Helenieae and Crocidium in the Senecioneae. Investigations of the flavonoids of these genera indicate that at the generic level both contain quercetin-3-o-galactoside, (probably isomeric) quercetin-3-o-glucoside, a quercetin-3-o-polyglucoside, and luteolin-7-o-glucoside. The first two compounds were present in all samples tested, but the other two glycosides were not. These striking biochemical similarities and the weight of morphological evidence suggest that the two genera are closely related to each other. Both should probably be placed in the Senecioneae. Blennosperma is a genus of 3 annual species, two of which occur in California and the third of which is restricted to Chile (Ornduff, 1963, 1964). Most taxonomists considering the affinities of this small genus have placed it in Helenieae (e.g. Bentham, 1876; Hoffman, 1889; Rydberg, 1914; Keck, 1959; Ferris, 1960), although other authors have allied it with Heliantheae (Gray, 1884), Anthemideae (Lessing, 1832; Remy, 1849), and Senecioneae (DeCandolle, 1838; Skvarla and Turner, 1966). Crocidium, a monotypic genus, ranges from southern British Columbia southward to central California. It has generally been placed in Senecioneae, although Bentham (1876) indicated that the relationships of the genus were obscure. In general habit Crocidium resembles Blennosperma and, on occasion, the two have been confused with each other by collectors. Nevertheless, rather few authors have suggested that the two genera are related. Gray (I884) was equivocal about the tribal affinities of Blennosperma and suggested that "perhaps it should stand next to Crocidium", although he placed Crocidium in Senecioneae. Both Bentham and Hoffman remarked on the helenioid aspect of Crocidium, and Bentham commented on the similarities of its styles to certain Astereae. Recently, Ornduff et al. (1963) suggested that Crocidium "surely belongs" elsewhere than Senecioneae. Skvarla and Turner (1966) alluded to similarities between these genera in various floral characteristics such as vascularization patterns of phyllaries and receptacles, and their "similar" ray flowers, styles, and achenes. On the basis of a study of pollen wall ultrastructure, these authors concluded that "Blennosperma and Crocidium possess Senecioid-type pollen and . . . are perhaps best treated together as members of the tribe Senecioneae". It should