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.     

Regional gene flow and population structure of the wind-dispersed plant species Hypochaeris radicata (Asteraceae) in an agricultural landscape

Using microsatellites, we investigated population structure and gene flow of the short-lived, wind-dispersed plant species Hypochaeris radicata in a fragmented agricultural landscape where more than 99% of the nutrient-poor grasslands have disappeared over the last century. We sampled populations in the few remaining high density populations in conservation areas, as well as individuals that occurred, with lower densities, in linear landscape elements, at two spatial scales. In a re-inventory of the landscape, after 3 years, both extinctions and colonizations of populations were observed. Contrary to expectations, no differences in genetic diversity between high and low density populations were observed. Both types of populations had relatively high levels of diversity. Overall genetic differentiation (theta) was 0.04 and significantly different from zero (P < 0.01). A significant isolation-by-distance pattern was found when all populations were simultaneously analysed (r = 0.24, P = 0.013). Isolation by distance was (marginally) significant at the small scale (r = 0.32, P = 0.06), whereas nonsignificant at the large spatial scale (r = -0.05, P = 0.66). A maximization-of-explained-variance procedure resulted in a threshold distance of 3.5 km above which populations were effectively genetically isolated. An additional partial exclusion Bayesian-based assignment test showed that overall 32.3% of the individuals were assigned to their population of origin, 48% were assigned to another population in the area and 19.7% were not assigned. Together, these results suggest high levels of gene flow. Seed dispersal contributes to the observed gene flow up to several hundred metres, which is higher than previously modelled using aerodynamic models on seed dispersal of H. radicata. We discuss the consequences of these results for an evaluation of the probability of persistence of this species in the fragmented landscape.     

Lectotypification of the Linnaean name Hypochaeris maculata (Asteraceae)

The typification of the name Hypochaeris maculata L. is discussed. A specimen from the Linnaean Herbarium (LINN) is designated as the lectotype.     

Hypochaeris alliatae group (Asteraceae) in the western Mediterranean Region

A taxonomic, ecological, and phytogeographical revision of the Hypochaeris alliatae group is made. As a result, the name of H. alliatae (vs H. laevigata ) is proposed. The recent taxa H. leontodontoides var. glauca and H. rutea are synonymised with H. alliatae , and H. leontodontoides var. atlantica var. nov ., is described. Using chromosomal counts, morphological and histological characters, H. angustifolia is excluded from the H. alliatae group. A phylogenetic and phytogeographical assay has been made in the context of geologic phenomena from the Miocene and Pliocene in the western Mediterranean Basin.     

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

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

Population genetic structure of the South American species Hypochaeris lutea (Asteraceae)

The genus Hypochaeris has a recent evolutionary history caused by long‐distance dispersal in conjunction with adaptive radiation in the South American continent. Hypochaeris lutea is a perennial herb that grows mostly at altitudes of around 1000 m in cold swamps of the southern regions of Brazil. We investigated the amplified fragment length polymorphism (AFLP) in 270 individuals representing 11 Brazilian populations of H. lutea to elucidate the population genetic structure of this species. The frequencies of polymorphic loci and gene diversity ranged from 83.42% to 91.66% and from 0.26 to 0.34, respectively. Analysis of molecular variance revealed that most of the genetic variability was found within (76.67%) rather than among (23.3%) populations, agreeing with the pattern of genetic distribution within and among populations observed in other allogamous species of Hypochaeris. A Mantel test showed no correlation between genetic and geographic distances when all populations were considered. Simulations performed using a Bayesian approach consistently identified two clusters with different admixture proportions of individuals, as also revealed by a UPGMA dendrogram of populations. The pattern of genetic structure observed in H. lutea is consistent with a process of successive colonization events by long‐distance dispersal resembling the rapid and recent radiation that has been proposed to explain the origin of the South American species of Hypochaeris.     

Cyanide-Resistant Root Respiration and Tap Root Formation in Two Subspecies of Hypochaeris radicata

Root respiration of the tap root forming species Hypochaeris radicata L. was measured during tap root formation. A comparison was made of two subspecies: H. radicata L. ssp. radicata L., a subspecies from relatively rich soils, and H. radicata L. ssp. ericetorum Van Soest, a subspecies from poor acidic soils. Root respiration was high and to a large extent inhibited by hydroxamic acid (SHAM) before the start of the tap root formation, indicating a high activity of an alternative non-phosphorylative electron transport chain. The rate of root respiration was much lower and less sensitive to SHAM when a considerable tap root was present. However, root respiration was also cyanide-resistant when a tap root was present, indicating that the alternative pathway was still present. A decreased rate of root respiration coincided with an increase of the content of storage carbohydrates, mainly in the tap root. The level of reducing sugars was constant throughout the experimental period, and it was concluded that the activity of the alternative oxidative pathway was significant in oxidation of sugars that could not be utilized for purposes like energy production, the formation of intermediates for growth or for storage. Root respiration decreased after the formation of a tap root. This decrease could neither be attributed to a gradual disappearance of the alternative chain, nor to a decreased level of reducing sugars. No differences in respiratory metabolism between the two subspecies have been observed, suggesting that a high activity of the alternative oxidative pathway is not significant in adaptation of the present two subspecies to relatively nutrient-rich or poor soils.     

Karyotype and AFLP data reveal the phylogenetic position of the Brazilian endemic Hypochaeris catharinensis (Asteraceae)

The genus Hypochaeris offers an excellent model for studies of recent adaptive radiation in the South American continent. We used karyotype analysis with chromomycin?A₃ (CMA₃)/4??,6-diamidino-2-phenylindole (DAPI) banding and fluorescence in?situ hybridization (FISH), and amplified fragment length polymorphism (AFLP) fingerprinting to investigate for the first time the Brazilian endemic H.?catharinensis and define its position within the South American group of species. Strong CMA-positive signals were seen at the end of both arms of chromosome?3 and at the end of the long arm of chromosome?4. DAPI bands were only detected in subterminal position on short arm of chromosome?4. FISH with 5S and 35S ribosomal DNA (rDNA) probes revealed a single 5S rDNA locus on short arm of chromosome?2, typical for all other South American Hypochaeris taxa analyzed to date. The 35S rDNA locus was identified at subterminal position on the short arm of chromosome?3, as reported so far for only two of the known species (H.?lutea and H.?patagonica). The AFLP study included 55 individuals, comprising nine species of the South American Hypochaeris plus their putative ancestor H.?angustifolia. Eleven AFLP primer combinations generated a total of 401 fragments, of which 388 (96.7%) were polymorphic. High genetic similarities were observed among taxa, with all South American Hypochaeris species falling into one main cluster [100% bootstrap (BS)]. Hypochaeris catharinensis is closely related to H.?lutea (82% BS), forming a well-separated subcluster within the South American species. Taken together, the karyological and AFLP data contribute to the placement of H.?catharinensis within the phylogenetic framework of South American species of Hypochaeris and allow the definition of a novel and well-resolved phylogenetic group (the Lutea group).     

Isolation and characterization of eight microsatellite loci from the endangered plant species Hypochaeris salzmanniana (Asteraceae)

We report the isolation and characterization of eight microsatellite loci for Hypochaeris salzmanniana, an endangered species endemic to the southwestern coast of Spain and the Atlantic coast of Morocco. A total of 32 alleles were detected across a sample of 45 individuals, with an average number of 4.0 alleles per locus. The average polymorphic information content (PIC) was 0.533 and the observed (H _O) and expected (H _E) heterozygosity values varied from 0.022 to 0.978 and from 0.434 to 0.759, respectively. Five loci exhibited significant deviation from Hardy–Weinberg equilibrium (P ≤ 0.001) and three pairs of loci showed significant linkage disequilibrium (P ≤ 0.01). The eight loci were tested for transferability in three others species (H. arachnoidea, H. glabra, and H. radicata) belonging to the same section of H. salzmanniana. With the exception of locus Hsalz7, all loci successfully amplified in the three species. These preliminary data confirm the usefulness of microsatellite markers for assessing the ecology and genetic structure of H. salzmanniana and to understand the evolution of species within the section Hypochaeris.     

Nuclear ribosomal DNA and karyotypes indicate a NW African origin of South American Hypochaeris (Asteraceae, Cichorieae)

Hypochaeris has a disjunct distribution, with more than 15 species in the Mediterranean region, the Canary Islands, Europe, and Asia, and more than 40 species in South America. Previous studies have suggested that the New World taxa have evolved from ancestors similar to the central European H. maculata. Based on internal transcribed spacer (ITS) sequences and fluorescence in situ hybridization (FISH) with 5S and 18S-25S rDNA of the previously overlooked Hypochaeris angustifolia from Moyen Atlas, Morocco, we show that it is sister to the entire South American group. A biogeographic analysis supports the hypothesis of long-distance dispersal from NW Africa across the Atlantic Ocean for the origin of the South American taxa rather than migration from North America, through the Panamian land bridge, followed by subsequent extinction in North America. With the assumption of a molecular clock, the trans-Atlantic dispersal from NW Africa to South America is roughly estimated to have taken place during Pliocene or Pleistocene.     

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.     

Temporal patterns in rates of community change during succession

While ecological dogma holds that rates of community change decrease over the course of succession, this idea has yet to be tested systematically across a wide variety of successional sequences. Here, I review and define several measures of community change rates for species presence-absence data and test for temporal patterns therein using data acquired from 16 studies comprising 62 successional sequences. Community types include plant secondary and primary succession as well as succession of arthropods on defaunated mangrove islands and carcasses. Rates of species gain generally decline through time, whereas rates of species loss display no systematic temporal trends. As a result, percent community turnover generally declines while species richness increases--both in a decelerating manner. Although communities with relatively minor abiotic and dispersal limitations (e.g., plant secondary successional communities) exhibit rapidly declining rates of change, limitations arising from harsh abiotic conditions or spatial isolation of the community appear to substantially alter temporal patterns in rates of successional change.     

Karyotype diversification and evolution in diploid and polyploid South American Hypochaeris (Asteraceae) inferred from rDNA localization and genetic fingerprint data

Background and Aims: Changes in chromosome structure and number play an importantrole in plant evolution. A system well-suited to studying differentmodes of chromosome evolution is the genus Hypochaeris (Asteraceae)with its centre of species' diversity in South America. AllSouth American species uniformly have a chromosome base numberof x = 4 combined with variation in rDNA number and distribution,and a high frequency of polyploidy. The aim of this paper isto assess directions and mechanisms of karyotype evolution inSouth American species by interpreting both newly obtained andprevious data concerning rDNA localization in a phylogeneticcontext. Methods: Eleven Hypochaeris species from 18 populations were studiedusing fluorescence in situ hybridization (FISH) with 35S and5S rDNA probes. A phylogenetic framework was established fromneighbour-net analysis of amplified fragment length polymorphism(AFLP) fingerprint data. Key Results: A single 5S rDNA locus is invariably found on the short armof chromosome 2. Using 35S rDNA loci, based on number (one ortwo) and localization (interstitial on the long arm of chromosome2, but sometimes lacking, and terminal or interstitial on theshort arm of chromosome 3, only very rarely lacking), sevenkaryotype groups can be distinguished; five of these includepolyploids. Karyotype groups with more than one species do notform monophyletic groups. Conclusions: Early evolution of Hypochaeris in South America was characterizedby considerable karyotype differentiation resulting from independentderivations from an ancestral karyotype. There was marked diversificationwith respect to the position and evolution of the 35S rDNA locuson chromosome 3, probably involving inversions and/or transpositions,and on chromosome 2 (rarely 3) concerning inactivation and loss.Among these different karyotype assemblages, the apargioidesgroup and its derivatives constitute by far the majority ofspecies.     

Ploidy level and origin of the European invasive weed Senecio inaequidens (Asteraceae)

Native to South-Africa, species of the Senecio inaequidens complex are presently invasive in Europe, Australia and South-America. Previously, different ploidy levels have been found in these different areas, with only tetraploid individuals reported in Europe, and only diploids in South-Africa and Australia. In the present study chromosome counts and flow cytometry were used to survey DNA ploidy levels in a large sample of 66 native and 21 European invasive populations. One Mexican individual was also added to the study. We found only tetraploid individuals occurring in Europe, whereas both ploidy levels, diploid and tetraploid, were found in South-Africa. Moreover, based on genome size, we suggest that two largely allopatric varieties of diploids exist in South-Africa. The Mexican individual was diploid. We suggest that European tetraploid individuals come from South-Africa and hypothesize that a hybridization event between the two DNA types of diploids occurred in the Lesotho area. The taxonomic difficulties surrounding species of theS. inaequidens complex are briefly discussed.     

Microbial community succession and lignocellulose degradation during agricultural waste composting

The changes of microbial community during agricultural waste composting were successfully studied by quinone profiles. Mesophilic bacteria indicated by MK-7 and mesophilic fungi containing Q-9 as major quinone were predominant and seemed to be important during the initial stage of composting. Actinobacteria indicated by a series of partially saturated and long-chain menaquinones were preponderant during the thermophilic period. While Actinobacteria, fungi and some bacteria, especially those microbes containing MK-7(H4) found in Gram-positive bacteria with a low G+C content or Actinobacteria were found cooperate during the latter maturating period. Since lignocellulsoe is abundant in the agricultural wastes and its degradation is essential for the operation of composting, it’s important to establish the correlation between the quinone profiles changes and lignocellulose degradation. The microbes containing Q-9 or Q-10(H2) as major quinone were found to be the most important hemicellulose and cellulose degrading microorganisms during composting. While the microorganisms containing Q-9(H2) as major quinone and many thermophilic Actinobacteria were believed to be responsible for lignin degradation during agricultural waste composting.     

Quadrat variance analysis and pattern development during primary succession

Abstract. The interpretation of Hill's ‘Two Term Local Quadrat Variance’ analysis to detect the scale of spatial pattern in vegetation is improved by an equation that relates the block size of a variance peak to the scale of the pattern that gave rise to it. (Contrary to previous belief, the two are not the same, especially for large block sizes.) Deviations of pattern from a perfectly regular alternation of equally sized gaps and patches of uniform density cause changes in the variance. To aid in the interpretation of these changes, two indices of pattern regularity are proposed, one based on density and one based onpresence/absence, in orderte distinguish the effects of irregularities of patch density from irregularities of patch size and position. These methods are applied to a study of primary succession on glacial moraines near Mt. Robson, British Columbia, Canada, in order to evaluate certain hypotheses about the development of pattern. Other researchers have proposed that during succession, the pattern at first intensifies at the scales initially observed, then as succession proceeds some scales of pattern are lost due to coalescence of patches and eventually the intensity of those that remain decreases as the patterns become more and more irregular. The vegetation on the Mt. Robson moraines confirms this sequence of changes in vegetation pattern, only to the extent that patterns intensify initially in the chronosequence; the number of scales of pattern in the vegetation remains about the same throughout and there is no evidence that the patterns become more irregular. The variance-block size graphs derived from presence / absence data matched those from density data well, indicating that the simpler data, in this case, are almost as informative about pattern as the more detailed data.     

Ectomycorrhizal networks and seedling establishment during early primary succession

Ectomycorrhizal (ECM) fungal mycelia are the main organs for nutrient uptake in many woody plants, and often connect seedlings to mature trees. While it is known that resources are shared among connected plants via common mycorrhizal networks (CMNs), the net effects of CMNs on seedling performance in the field are almost unknown. CMNs of individual ECM fungal species were produced in an early succession volcanic desert by transplanting current-year seedlings of Salix reinii with ECM mother trees that had been inoculated with one of 11 dominant ECM fungal species. Most seedlings were connected to individual CMNs without being infected by other ECM fungi. Although control seedlings showed poor growth under severe nutrient competition with larger nonmycorrhizal mother trees, nutrient acquisition and growth of seedlings connected to CMNs were improved with most fungal species. The positive effects of CMNs on seedling performance were significantly different among ECM fungal species; for example, the maximum difference in seedling nitrogen acquisition was 1 : 5.9. The net effects of individual CMNs in the field and interspecific variation among ECM fungal species are shown.