Systematic consideration of floral microcharacters of the South American genus Chrysolaena (Vernonieae,Asteraceae)

Floral microcharacters of the genus Chrysolaena H. Rob. (Vernonieae, Asteraceae) were analysed in detail for the first time in order to evaluate the taxonomic position of conflictive species in the group. The results were also compared with studies carried out in species of related genera. In addition to distinctive microcharacters previously studied in some species of the genus, other characters such as trichome types of the corolla, style, anthers and cypselae have been analysed for the first time. The presence of glandular apical appendage and cypselae are common characteristics among species Chrysolaena. In addition to these, this study shows that presence of glands on the style and corollas is another consistent characteristic in the genus. However, the absence of basal stylar node would not be a diagnostic character since this varies widely among species. The results indicate that most of the microcharacters of Chrysolaena analysed are quite consistent in the genus, but they are no more consistent than the pollen morphology (type 'C') and chromosome base number (x = 10). Until now, these last two features would be most useful for separating Chrysolaena from the related genera Lessingianthus and Lepidaploa. At species level, the results show that related species can be distinguished by the different combinations of floral microcharacters. The value of microcharacters could be increased if they are combined with other morphological, cytological, and palynological data.     

Chromosome Numbers and Karyotypes of Species of Vernonia sect. Lepidaploa (Asteraceae: Vernonieae)

Vernonia is the largest genus of the tribe Vernonieae (Asteraceae) and comprises more than 1,000 species. In the present study we explore chromosome number and karyotype variation of eight species treated within different subsections of the section Vernonia sect. Lepidaploa. We aimed to explore if these data support the recognition of a single large genus (sensu Baker) or favor its splitting into 22 small genera (sensu Robinson). The species were collected in ??cerrado??, rupicolous and disturbed areas in the states of S?o Paulo and Minas Gerais, Brazil. Chromosome numbers varied from 2n?=?32 to 60. Most chromosomes were small, and the karyotype analysis revealed a predominance of metacentric and some submetacentric chromosomes. The karyotype symmetry in Vernonia was moderate (TF% 32.2 to 45.9), with the most symmetrical karyotype observed in V. rubriramea. The results obtained here did not conclusively support any of the taxonomic proposals for Vernonia due to the absence of distinctive or characteristic karyotype patterns for any of the taxonomic groupings, i.e., sections and subsections (sensu Baker) or new genera (sensu Robinson). Nevertheless, a tenuous relationship was observed between the chromosome numbers reported in the literature, those recorded here, and the taxonomic alterations suggested by Robinson for the genera Lessingianthus, Chrysolaena, and Vernonanthura that were originally part of Vernonia sensu Baker.     

Cytotaxonomy of species of Vernonia, section Lepidaploa, group Axilliflorae (Asteraceae, Vernonieae)

Mitotic chromosomes of eight populations of six species of Vernonia sensu Baker (Asteraceae, Vernonieae, section Lepidaploa , group Axilliflorae ) were studied to assess the validity of maintaining the genus as proposed currently ( sensu Baker) or of splitting it into several smaller genera ( sensu Robinson). The species were collected in areas of 'cerrado' and 'campo rupestre' in southern and central-western Brazil. The chromosome numbers were 2 n  = 20–40 and karyotypic analysis showed a predominance of metacentric chromosomes, with some submetacentrics. B chromosomes were seen in one population of V. geminata . The chromosomes varied in length from 0.9 to 4.6 µm, with a total chromatin length of 29.7–45.0 µm and a symmetry index of 41.2–46.9%. The intrachromosomal asymmetry index varied from 0.13 to 0.29, and the interchromosomal asymmetry index varied from 0.15 to 0.21. The dispersion diagram showed that population 1 of V. geminata had the most asymmetrical karyotype. These results do not provide sufficient evidence for a conclusive cytotaxonomic discussion of Vernonia , mainly because of the impossibility of associating any infrageneric group with specific karyotypic characters, and also because of the small number of species analysed so far.  © 2007 The Linnean Society of London. Botanical Journal of the Linnean Society , 2007, 154 , 99–108.     

Floral microcharacters in Lessingianthus (Vernonieae,Asteraceae) and their taxonomic implications

Micromorphological characters of 113 species of the South American genus Lessingianthus H.Rob. (Vernonieae, Asteraceae) were analyzed to evaluate their reliability as taxonomic markers. The corolla pubescence was studied in detail for the first time in the genus. Glandular and non-glandular trichomes were studied and described. The trichome types allow differentiation among related species, but do not reflect the relationships among species groups. On the other hand, the basal stylar node only occurs in a distinctive group of species within the genus. The apical appendage of the anthers is non-glandular in all species of the genus. Several character states analyzed in Lessingianthus are often widespread in other related genera of Vernonieae. Therefore, Lessingianthus can be circumscribed only by a combination of micro- and macromorphological features.     

Diversity of non-glandular trichomes in subtribe Lychnophorinae (Asteraceae: Vernonieae) and taxonomic implications

Vernonieae is one of the major tribes in Asteraceae (subfamily Cichorioideae) with ca. 1,100 species placed into 129 genera. Currently, 21 subtribes are recognized in Vernonieae and one of them is Lychnophorinae, almost entirely endemic to Brazil, containing 11 genera and ca. 100 species. About 42 % of Lychnophorinae genera are monophyletic, reflecting the poorly understood relationships among the members of the group. Trichomes are one of the most useful anatomical characters to be used in angiosperm taxonomy; they are diverse, exist in many taxa and are not difficult to study. This work intends to illustrate non-glandular leaf trichome diversity in Lychnophorinae and discuss this diversity in the light of the subtribe’s taxonomy. Sampled material included 67 species of 11 genera. Macerations and free hand sections were performed to be analyzed in the light microscope and photographed. A phenogram was generated using a matrix with 67 terminals (species) and 18 characters coded as binary. The subtribe Lychnophorinae displays a great diversity of non-glandular trichomes (5 types and 18 subtypes). The present study reveals the great diversity of non-glandular trichomes in Lychnophorinae. While trichome complement is of little use to distinguish genera, it appears to be a valuable characteristic at a lower taxonomic level to identify closely morphologically related species.     

Pollen morphology of some related genera of Vernonieae (Asteraceae) and its taxonomic significance

Pollen morphology is an important source of information to increase systematic resolution in Asteraceae, especially in Vernonieae. Aiming to investigate if palynological traits give support to Caatinganthus, Strophopappus and Xiphochaeta as separate genera from Stilpnopappus, we used cluster analysis followed by a test of group sharpness. Further, ordination analysis was applied to detect informative pollen traits associated with the revealed groups. The analyses evidenced five groups: (G1) Caatinganthus rubropappus as a single-species group; (G2) species of Stilpnopappus; (G3) Xiphochaeta aquatica as a single-species group; (G4) Strophopappus bicolor, S. glomeratus, S. villosus, S. ferrugineus, S. pohlii and S. speciosus; (G5) Strophopappus bullatus and S. regnelli. The interruption in the middle of the muri in apertural lacunae explains the single-species group Caatinganthus rubropappus. The thickness of sexine, the type of apertures (porate or colporate), and spine dimensions (length, thickness and distance from each other) are the traits explaining differences between species of Stilpnopappus and Strophopappus. Equatorial lacunae give support to Xiphochaeta aquatica as a single-species group, despite the smaller size of pollen grains of this species as compared to the others species analyzed. The differences among pollen morphology give support to Caatinganthus, Stilpnopappus, Strophopappus and Xiphochaeta as genera according to the taxonomic classification currently accepted. The used approach was efficient to reveal individual pollen traits informative to explain the sharp groups, and was an effective alternative to the use of “pollen types”.     

Karyotypes of some species of the genus Lessingianthus (Vernonieae,Asteraceae) and taxonomic implications

We determined the karyotypes of nine species of Lessingianthus, eight of which are here analyzed for the first time. The results include the first chromosome count for L. plantaginoides, which is tetraploid with 2n = 64. All species showed a high proportion of metacentric chromosomes combined with a lower proportion of submetacentric pairs. Only L. coriaceus had a subtelocentric chromosome pair. B chromosomes were observed in L. coriaceus and L. varroniifolius, which were subtelocentric. Differences among the karyotypes of the studied species were small, suggesting that karyotype diversity in the genus evolved by small changes in the structure of chromosomes. Karyotype features appear useful to distinguish Lessingianthus from the closely related genera Chrysolaena and Lepidaploa.     

Histolocalization of chemotaxonomic markers in Brazilian Vernonieae (Asteraceae)

Vernonieae are well represented in South America by subtribe Lychnophorinae, with 104 Brazilian species, some of them rare and endangered. Lychnophorinae are well known for producing metabolites of high pharmaceutical and chemotaxonomic value. Despite the importance of secondary metabolites in Lychnophorinae, there is still little evidence regarding the secretory structures responsible for producing these compounds. Therefore, in this study we investigated 15 species representative of the main lineages of Lychnophorinae, with the following objectives: to identify where the chemotaxonomic markers of Lychnophorinae are secreted and stored and in which developmental stage of the organ these metabolites are found. Samples of stems and leaves were processed according to the usual techniques in plant anatomy. It was found that the main sites of secondary metabolite biosynthesis are glandular trichomes, epidermal cells and parenchyma tissues. Metabolites from glandular trichomes, especially sesquiterpene lactones (STL), are prevalent in the early developmental stages of organs. The metabolite compounds stored in parenchyma tissues are mainly terpenoids, flavonoids and other phenolic compounds; young and expanded leaves are equally rich in metabolites. Thus, the information obtained in this study is essential for conducting chemotaxonomic studies in this group, helping to promote selective collection and conservation of species.     

The foliar micromorphology of Solanum pseudocapsicum

Foliar micromorphology of Solanum pseudocapsicum was investigated by electron microscopical examination. The leaves are characterized by anisocytic stomata which are more abundant on the abaxial surfaces. The leaves have short multicellular glandular trichomes sparsely distributed over the entire leaf surfaces. Crystal deposits were also observed on the surfaces above the stomata. Energy dispersive X-ray spectroscopy-SEM showed that Al, K, Na and Si were the major constituents of the crystals analyzed. The presence of glandular trichomes in this plant could be the source of poisonous compounds that are characteristic of this species.     

Two new species of Vernonia (Asteraceae: Vernonieae) from Panama

Two new species,Vernonia cocleana andV. chiriquiensis, are described from Panama. These endemic species are distinct from those in Central America and most closely resemble taxa in the West Indies and Argentina, respectively. These taxa are described and illustrated; a key is provided to the Panamanian species ofVernonia.     

Two new species of Vernonia (Asteraceae: Vernonieae) from Peru

Two new species,Vernonia sagasteguii andV. paucartambensis, from Peru are described, illustrated, and their relationships discussed.     

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.     

Cytotaxonomic analysis of South American species of Vernonia (Vernonieae: Asteraceae)

Chromosome numbers in 48 populations belonging to 31 species of Vernonia from South America have been determined. First chromosome counts are reported for V. propinqua var. canescens (2 n = 20), V. sceptrum (2 n = 80), V. rufogrisea (2 n = 32), V. heringeri (2 n = 32), V. hovaefolia (2 n = 64), V. hystricosa (2 n = 128), V. obtusata (2 n = 64), V. pseudoincana (2 n = 32), V. rubricaulis var.◊ australis (2 n = 64) and V. rugulosa (2 n = 30). The basic chromosome numbers x = 10, 14, 15, 16 and 17 were confirmed for New World species of Vernonia . Together with previous records, these numbers suggest that the evolution of Vernonieae has occurred through a combination of polyploidy and aneuploidy. Taxonomic implications of certain counts are also discussed.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 401–408.     

Palynological survey of the subtribe Elephantopinae (Asteraceae, Vernonieae)

Despite the taxonomic value of pollen morphology within the tribe Vernonieae, a detailed study of the pollen of the subtribe Elephantopinae is still lacking. The pollen morphology of ten species, representing three of the four genera of the subtribe, Elephantopus, Pseudoelephantopus and Orthopappus, was studied with LM, SEM, and TEM. The pollen of all the species studied was found to be echinolophate, although the differences in aperture features and both sculpture and exine structure allowed recognizing two pollen types. The species of Elephantopus (except for E. elongatus) and Pseudoelephantopus share the regular or more or less regular ridges pattern, the colporate condition but with the ectoapertures little developed, and the acaveate exine structure. From the sculpture and the apertures, the pollen of Caatinganthus harleyi was found to be similar to that of Elephantopus and Pseudoelephantopus. The pollen morphology of the monotypic Orthopappus, which is shared with that of E. elongatus, was further investigated for the first time. We found that it differs from that of the other species in having an irregular pattern of ridges, colporate condition but with a well-developed ectoapertures, and caveate exine. Additional studies of the exine structure and apertures features, coupled with molecular phylogeny, are needed to understand the evolution of pollen characters and re-evaluate the intergeneric relationships within the tribe.     

Karyological features and cytotaxonomy of the tribe Vernonieae (Asteraceae)

Chromosome numbers are reported for 29 populations of 19 Vernonieae taxa collected mainly in the northeastern region of Brazil. Among them, data for five genera (Blanchetia, Rolandra, Pithecoseris, Stilpnopappus and Vanillosmopsis) are here reported for the first time, and the first chromosome counts are presented for 12 species. Chromosome numbers are quite diverse among and sometimes within genera, especially in the controversial and large subtribe Vernoniinae. The numbers varied from 2n = 18 to 2n = ~72. The main karyoevolutionary mechanism seems to be dysploidy, while polyploidy is probably associated with ancient hybridization processes generating most paleotetraploid genera. All studied species presented semi-reticulated interphase nuclei and proximal-early condensing behavior in prophase to prometaphase. In one species (Vernonia condensata with 2n = 40) fluorochrome staining with CMA/DAPI revealed five chromosome pairs bearing subterminal CMA⁺/DAPI^? heterochromatin, probably NOR-associated, revealing the existence of low amounts of satellite DNA. The role of these features in the evolution of the tribe is discussed, revealing some interesting aspects for understanding of the Vernonieae karyoevolution, especially regarding neotropical members.     

A new species of Lessingianthus (Asteraceae: Vernonieae) from Minas Gerais, Brazil

Lessingianthus ibitipocensis, a new species from the Southeastern Brazilian highlands, is described and illustrated.

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Resumen  Se describe e ilustra una especie nueva, Lessingianthus ibitipocensis, del sudeste de Brasil.

     

Comparative anatomy of leaves and stems in some species of the South American genus Chrysolaena (Vernonieae,Asteraceae) and taxonomic implications

Vegetative anatomical features are poorly known in the South American genus Chrysolaena. In this study, leaves and stems of six Chrysolaena species were described and compared morphologically and anatomically using diaphanization, microtome serial sectioning and scanning electron microscopy. The species differed in leaf epidermis, type of stomata, shape of anticlinal walls of epidermal cells, trichome density, and presence or absence in stems of small air spaces in the cortical parenchyma and of druse‐shaped oxalate crystals. Furthermore, glandular trichomes and three types of non‐glandular trichomes with different number of basal cells were identified on leaves and stems. Collectively, these features proved instrumental to discriminate among the six studied species, suggesting that leaves and stems of Chrysolaena can represent a source for taxonomically useful characters. We also discuss anatomical features in relation to the environmental conditions in the species’ habitats.     

Karyotype analysis and DNA content in some species of Chrysolaena (Vernonieae,Asteraceae)

Cytogenetic characterization by karyotyping and determination of DNA content by flow cytometry of five species of Chrysolaena (Vernonieae, Asteraceae) was performed. This is the first study of nuclear DNA content realized in the genus. The 2C-values were compared with the ploidy level and the total karyotype length (TKL) of each species. Mitotic analysis revealed a base chromosome number x = 10 for all entities and different ploidy levels, from diploid (2n = 2x = 20) to octoploid (2n = 8x = 80). All species showed bimodal karyotypes composed of metacentric and submetacentric chromosomes. The average chromosome size (ML) varied from 1.86 μm to 2.70 μm, while the TKL ranged from 18.65 μm to 80.55 μm. The intrachromosomal asymmetry index (A₁) varied from 0.27 to 0.38, while the interchromosomal asymmetry index (A₂) ranged from 0.19 to 0.25. A new cytotype is reported for the first time for C. propinqua. Accessory chromosomes found in C. verbascifolia, C. cognata, C. flexuosa, and C. propinqua are also reported as new.     

A multivariate approach to assess the taxonomic utility of morphometric characters inDoronicum (Asteraceae, Senecioneae)

A multivariate morphometric study based on 22 characters was carried out on samples of the 26 accepted species of the Eurasian and north-African genusDoronicum, to assess whether quantitative, mainly continuous, characters were of use in suggesting, delimiting and structuring natural groups. The study concentrated on three-medium sized to small groups: Mediterranean (D. plantagineum group), European (D. grandiflorum group), and central Asian. Two questions were addressed for each of these groups: species delimitation within them and the inclusion/exclusion of species of uncertain, but putative close affinities. Principal components analysis and discriminant analysis (canonical variates analysis), gave moderately satisfactory results. The addition of doubtfully-related species to presumably natural groups increased cohesiveness of the groups because the between-species differences were minimized when placed in a broader more diverse context. The relative status of the uncertain species, either as outliers or true members, was appropriately depicted by the scatterplots, but more convincingly solved when the species turned out to be outliers. Discrimination between species, which belong to cohesive groups was effective in some cases at least when using discriminant analysis, and the exceptions involved particularly problematic taxa. It was concluded that morphometric characters are a useful complementary source of information in groups where qualitative (in particular, shared) characters are scarce as is the case inDoronicum.