Estrella Urtubey

AFLP Phylogeny of South American Species of Hypochaeris (Asteraceae, Lactuceae)

Abstract Hypochaeris is thought to have arrived in South America by dispersal over the Atlantic Ocean from NW Africa during the Pliocene or Pleistocene. We used amplified fragment length polymorphism (AFLP) to unravel specific limits and relationships in the South American group of Hypochaeris (c. 45 species). The Moroccan endemic Hypochaeris angustifolia, which is sister to the entire South American group, was used as outgroup. Our AFLP analysis comprises 415 individuals from 32 South American species and is based on six primer combinations with 670 fragments scored. It provides important information for the delimitation of species and detection of closely related species pairs or groups. Most species are monophyletic and supported with > 90% bootstrap proportion. Hybridization is suggested between Hypochaeris chillensis and H. microcephala in Brazil. The internal nodes (or “backbone”) of the tree are not highly supported, but six major phylogenetic groups (also showing similarity in distribution and growth form) center around Hypochaeris apargioides, H. chondrilloides, H. microcephala, H. pampasica, H. sessiliflora, and H. tenuifolia. These results suggest that rapid migration into different geographical regions played an important role in the initial diversification of Hypochaeris in South America.

Phylogeny of Barnadesioideae (Asteraceae) inferred from DNA sequence data and morphology

Subfamily Barnadesioideae (Asteraceae) consists of nine genera and 91 species endemic to South America. They include annual and perennial herbs, arching shrubs and trees up to 30m tall. Presumed sister to all other Asteraceae, its intergeneric relationships are key to understanding the early evolution of the family. Results of the only molecular study on the subfamily conflict with relationships inferred from morphology. We investigate inter- and intrageneric relationships in Barnadesioideae with novel DNA sequence data and morphological characters using parsimony, likelihood and Bayesian inference. All results verify Barnadesioideae as monophyletic and sister to the rest of the family. A basal split within the subfamily is recognized, with Chuquiraga, Doniophyton and Duseniella in one clade, and Arnaldoa, Barnadesia, Dasyphyllum, Fulcaldea, Huarpea and possibly Schlechtendalia in another. The largest genus, Dasyphyllum, is revealed as biphyletic with the two clades separating along subgeneric and geographic lines. Schlechtendalia, suggested as the earliest diverging lineage of the subfamily by morphological studies and parsimony analyses, is found in a more derived position under model-based inference methods. Competing phylogenetic hypotheses, both previous and present, are evaluated using likelihood-based tests. Evolutionary trends within Barnadesioideae are inferred: hummingbird pollination has developed convergently at least three times. An early vicariance in the subfamilys distribution is revealed. X=9 is supported as the ancestral base chromosome number for both Barnadesioideae and the family as a whole.

Pleistocene refugia and polytopic replacement of diploids by tetraploids in the Patagonian and Subantarctic plant Hypochaeris incana (Asteraceae, Cichorieae)

We report the phylogeographic pattern of the Patagonian and Subantarctic plant Hypochaeris incana endemic to southeastern South America. We applied amplified fragment length polymorphism (AFLP) and chloroplast DNA (cpDNA) analysis to 28 and 32 populations, respectively, throughout its distributional range and assessed ploidy levels using flow cytometry. While cpDNA data suggest repeated or simultaneous parallel colonization of Patagonia and Tierra del Fuego by several haplotypes and/or hybridization, AFLPs reveal three clusters corresponding to geographic regions. The central and northern Patagonian clusters (∼38–51° S), which are closer to the outgroup, contain mainly tetraploid, isolated and highly differentiated populations with low genetic diversity. To the contrary, the southern Patagonian and Fuegian cluster (∼51–55° S) contains mainly diploid populations with high genetic diversity and connected by high levels of gene flow. The data suggest that H. incana originated at the diploid level in central or northern Patagonia, from where it migrated south. All three areas, northern, central and southern, have similar levels of rare and private AFLP bands, suggesting that all three served as refugia for H. incana during glacial times. In southern Patagonia and Tierra del Fuego, the species seems to have expanded its populational system in postglacial times, when the climate became warmer and more humid. In central and northern Patagonia, the populations seem to have become restricted to favourable sites with increasing temperature and decreasing moisture and there was a parallel replacement of diploids by tetraploids in local populations.

Phylogeography of the invasive weed Hypochaeris radicata (Asteraceae): from Moroccan origin to worldwide introduced populations

In an attempt to delineate the area of origin and migratory expansion of the highly successful invasive weedy species Hypochaeris radicata, we analysed amplified fragment length polymorphisms from samples taken from 44 populations. Population sampling focused on the central and western Mediterranean area, but also included sites from Northern Spain, Western and Central Europe, Southeast Asia and South America. The six primer combinations applied to 213 individuals generated a total of 517 fragments of which 513 (99.2%) were polymorphic. The neighbour‐joining tree presented five clusters and these divisions were supported by the results of Bayesian analyses: plants in the Moroccan, Betic Sierras (Southern Spain), and central Mediterranean clusters are all heterocarpic. The north and central Spanish, southwestern Sierra Morena, and Central European, Asian and South American cluster contain both heterocarpic (southwestern Sierra Morena) and homocarpic populations (all other populations). The Doñana cluster includes two homocarpic populations. Analyses of fragment parameters indicate that the oldest populations of H. radicata are located in Morocco and that the species expanded from this area in the Late Quaternary via at least three migratory routes, the earliest of which seems to have been to the southwestern Iberian Peninsula, with subsequent colonizations to the central Mediterranean area and the Betic Sierras. Homocarpic populations originated in the southwestern Iberian Peninsula and subsequently spread across north and central Spain, Central Europe and worldwide, where they became a highly successful weed.

New hypotheses of phylogenetic relationships in Barnadesioideae (Asteraceae) based on morphology

ISSN 0046-0262. Previous molecular sequence and restriction site data have confirmed subfamily Barnadesioideae as basal within Asteraceae. The subfamily consists of 88 species in nine genera: Arnaldoa, Barnadesia, Chuquiraga, Dasyphyllum, Doniophyton, Duseniella, Fulcaldea, Huarpea, and Schlechtendalia. Two morphological cladistic analyses among genera or subgeneric groups within this subfamily have been completed previously, with contrasting results. Because of the importance of understanding relationships in this group of early Asteraceae, especially for insights on character evolution within the family, comprehensive morphological cladistic analyses were conducted at the specific level. Initial investigations utilising 31 characters with all 88 species led eventually to curtailed studies with 52 species. Five different outgroup hypotheses were employed: Acicarpha laxa, A. spathulata, Calycera leucanthema, C. spinulosa (all Calyceraceae), and Schlechtendalia luzulaefolia (Barnadesioideae; as functional outgroup). The results confirm Schlechtendalia as basal within Barnadesioideae. Monophyly is also confirmed for all genera, sections Chuquiraga and Acanthophyllae of Chuquiraga, and subgenus Archidasyphyllum of Dasyphyllum. All cladograms show two major groups (although statistical support values are low): (1) Chuquiraga, Doniophyton and Duseniella; and (2) Arnaldoa, Dasyphyllum, Fulcaldea, Huarpea and Barnadesia. Although bilabiate (4 + 1) corollas occur in Schlechtendalia, tubular corollas are believed plesiomorphic for the subfamily because they are pervasive in Calyceraceae as well as in all other basal Barnadesioideae. Pollen with no shallow depressions also are hypothesised as plesiomorphic for Barnadesioideae, even though one depression occurs in Schlechtendalia. A southern South American origin of Barnadesioideae (and the entire family) is re-emphasised. The previously hypothesised ancient lineage of Dasyphyllum subg. Archidasyphyllum, consisting of large trees, appears more derived in the new analyses.

AFLP and breeding system studies indicate vicariance origin for scattered populations and enigmatic low fecundity in the Moroccan endemic Hypochaeris angustifolia (Asteraceae), sister taxon to all of the South American Hypochaeris species.

We used Amplified Fragment Length Polymorphism markers (AFLP) and breeding system studies to investigate the population structure and reproductive biology of Hypochaeris angustifolia (Asteraceae: Cichorieae). This species is endemic to altiplanos of the Atlas Mountains (Morocco) where it occurs in scattered populations, and it is the sister species to c. 40 species of this genus in South America. PCoA, NJ, and Bayesian clustering, revealed that the populations are very isolated whilst AFLP parameters show that almost all populations have marked genetic divergence. We contend that these features are more in accord with a vicariance origin for the scattered populations of H. angustifolia, rather than establishment by long-distance dispersal. The breeding system studies revealed that H. angustifolia is a self-incompatible species, with low fecundity in natural and in experimental crosses, probably due to a low frequency of compatible phenotypes within and between the populations.

Gamochaeta beckii (Gnaphalieae, Asteraceae): A New Species from Bolivia

Abstract Gamochaeta beckii, a new species found in La Paz, Bolivia is described and illustrated. Gamochaeta beckii is similar to G. purpurea but it is easily distinguished by stems branched from the base to the inflorescence, oblanceolate leaves, the small capitula arranged in glomerules, forming leafy short and continuous spikes, and short acuminate inner phyllary apices. A key to the species of Gamochaeta occurring in Bolivia is presented.

Karyotype studies on populations of two Hypochaeris species (H. catharinensis and H. lutea), Asteraceae, endemics to southern Brazil.

Hypochaeris is an excellent system for studying different modes of chromosome evolution in plants. We carried out a cytogenetic analysis on populations of 2 Hypochaeris species, comprising 10 populations of H. catharinensis and 5 of H. lutea, to assess possible changes on chromosome organization in this interesting genus. Conventional Feulgen staining and fluorescent banding revealed that the general aspects of chromosome morphology for all populations of both species were similar, evidence of the typical bimodal karyotypes with 2n = 8 chromosomes that characterize the South American Hypochaeris. Comparative analysis of the karyotypes identified minor variations in the absolute size and arm ratio of corresponding chromosome pairs. One population of H. lutea was entirely polyploid adding a novel cytotype to this species. Fluorescent banding revealed strong chromomycin A3 (CMA3)-positive signals on both arms of chromosomes 3 and 4 of H. catharinensis, revealing a new pattern for the distribution of GC-rich heterochromatin in Hypochaeris. A strong CMA-positive signal was observed on the short arm of chromosome 3 in one population of H. lutea, while the other populations validated the CMA3 pattern already described for this species. While the overall karyotype similarities of the 2 species are in compass with all South American Hypochaeris, the presence of unusual large blocks of GC-rich heterochromatin suggests that chromosome rearrangements, related to dispersion of heterochromatin, are taking place in the karyotype of H. catharinensis. The novel polyploid cytotype identified in H. lutea provides support that polyploidization is an active process in the mode of chromosome evolution in Hypochaeris.

ESTUDIOS CROMOSOMICOS EN ESPECIES DE HYPOCHAERIS L. (ASTERACEAE, LACTUCEAE) DE CHILE

The genus Hypochaeris (Asteraceae, Lactuceae) contains twelve species in Europe, three in Asia, and approximately 50 in South America. In this paper, we worked with mitotic chromosomes of three different species of Hypochaeris from Chile: H. acaulis, H. incana and H. scorzonerae. The species are very different in morphology and geographic distribution. The aim of this work was to compare the chromosome morphology of these species. We found that the chromosomes are different in size and morphology in H. acaulis (smaller than 6 µm and only chromosome 3 having a secondary constriction on the short arm). In H. incana and H. scorzonerae the chromosomes are very similar in size, and chromosomes 2 (on the long arm) and 3 (on the short arm) contain secondary constrictions. A tetraploid is reported for H. scorzonerae

CHROMOSOMES REPORT FROM PLANTS ARE GROWING IN CHILE. III

Chromosome numbers of 24 Chilean taxa of vascular plants were studied using root-tip mitotic metaphases. New records for the chilean Flora are: Alstroemeria presliana Herb. subsp. presliana (2n = 16), Chaetanthera moenchioides Less. (2n = 26), Hypochaeris scorzonerae (DC.) F.Muell. (2n = 8), Phycella bicolor (Ruiz et Pav.) Herb. (2n = 16) and Rhodophiala tiltilensis (Traub et Moldenke) Traub (2n = 16).