Ploidy level selection during germination and early stages of seedling growth in the progeny of allohexaploid facultative apomict,Hieracium rubrum (Asteraceae)

Selection within progeny of a facultative apomict,Hieracium rubrum, was studied using flow cytometry of embryos in seeds (a modified method of Flow Cytometric Seed Screen) and seedlings. Flow cytometric screening of particular progeny classes was based on distinct ploidy categories, reflecting the way of their origin. The results of both estimations of progeny composition significantly differed, which makes direct comparison of proportions detected in seeds or seedling stage impossible. The results suggest that progeny originating from reduced egg cells have higher mortality during germination and the early establishment phase than that from unreduced egg cells. Within the progeny of emasculated plants, the proportion of polyhaploid progeny decreased in favor of apomictically derived plants. Within the progeny of plants pollinated byH. pilosella, the proportion of polyhaploid progeny decreased significantly in favor of apomictically derived plants and 2n + n hybrids. It is argued that at least a proportion of the sexually derived progeny of this facultatively apomictic maternal parent has a lower survival rate than apomictically derived progeny.     

Increased litter decomposition rates of exotic invasive species Hieracium pilosella (Asteraceae) in Southern Patagonia,Argentina

Plant Ecology - Plant invasions can affect key ecosystem processes regulating carbon and nutrient balances, i.e., litter decomposition. Exotic herb Hieracium pilosella (hawkweed, Asteraceae)...     

Diversified impact of mycorrhizal inoculation on mother plants and daughter ramets in the clonally spreading plant Hieracium pilosella L. (Asteraceae)

Plant Ecology - Resource allocation studies of clonal plants whose individuals form networks of interconnected ramets are very challenging. In addition, the presence of mycorrhizal fungi may...     

Predicting the Impact of the Biocontrol Agent Aulacidea subterminalis (Cynipidae) on Growth of Hieracium pilosella (Asteraceae) under Differing Environmental Conditions in New Zealand

A gall wasp, Aulacidea subterminalis, was released into the field in New Zealand in 1999 for biocontrol of the grassland weed Hieracium pilosella. To predict its likely impact in the field, the effect of the gall wasp on plant growth was measured under stress-free conditions in a shade house trial with potted plants. Plants with galls showed a reduction in stolon length of 75%. They had slightly lower total dry matter and reduced root weight. In a glasshouse experiment to measure the impact of water stress, nutrient stress, and plant competition on growth of H. pilosella and performance of the gall wasp, the number and mean diameter of gall clusters were not significantly different between treatment and control plants. However, galled plants produced more, but shorter, stolons in all stress treatments, and stolons that were more branched in nutrient- and water-stressed plants, than ungalled plants. Under the plant competition treatment, total stolon length relative to biomass was greater with, than without, wasps, indicating that stolons were thinner. Galling by A. subterminalis is likely to reduce vegetative reproduction of H. pilosella whether or not the plants are stressed, indicating that the wasp may be a successful biocontrol agent.     

Cytotype distribution and phylogeography of Hieracium intybaceum (Asteraceae)

Using flow cytometry and amplified fragment length polymorphism (AFLP), we explored the cytogeography and phylogeography of Hieracium intybaceum, a silicicolous species distributed in the Alps and spatially isolated in the Vosges Mountains and the Schwarzwald Mountains. We detected two ploidies, diploid and tetraploid, but no triploid or mixed‐ploidy populations. Whereas diploids are sexual and distributed all across the Alps, tetraploids are apomictic and seem to be confined to the western Alps and the Vosges. We detected a low level of genetic variation. Bayesian clustering identified four clusters/genetic groups, which are partly congruent with the ploidal pattern. The first two groups consisting exclusively of diploids dominate the whole distribution range in the Alps and show east–west geographical separation with a diffuse borderline running from eastern Switzerland to the eastern part of North Tyrol. The third genetic group lacks a defined geographical range and includes diploid and tetraploid plants. The last genetic group comprises tetraploid plants in the French Alps and the Vosges. We suppose that diploids colonized the deglaciated areas from source populations most likely located mainly in the southern part of the recent distribution range and occasionally also in the western Alps. Gene flow and further differentiation likely took place. Apomictic tetraploids most likely originated in the western Alps or in the refugium at the south‐western foot of the Alps. Their rather limited geographical range (partly contrasting with the theory of geographical parthenogenesis) can be explained by their rather recent origin. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 487–498.     

A new typification of Hieracium umbellatum (Asteraceae)

The Linnaean name Hieracium umbellatum has generally been understood in a very strict and clear sense. The original material comprises three specimens and a few illustrations, with a lectotype designated here in Burser's Hortus siccus (UPS). The earlier choice published by W. Lack was made from non-original material and was therefore not effective. The application of this name is confined to a variety generally known as H. umbellatum var. commune Fr. nom. illeg. (restricted here to the forms with almost entire, sublinear leaves), most proximate to H. umbellatum var. coronopifolium (Bernh. ex Hornem.) Wimm. et Grab.     

Pollen and chromosome studies in Hieracium sect. Alpestria (Asteraceae)

Potential pollen viability has been investigated in British apomictic Hieracium section Alpestria and related species using Alexander's Stain. New and existing chromosome counts are given. Some species produced no pollen, others varied in the amount they produced, and a few species consistently produced reasonable quantities. The three triploid section Alpestria species produced very little or no pollen, whilst the amount produced by the tetraploids varied widely. Section Alpestria species had significantly lower pollen viability than the non‐section Alpestria species. The generally low pollen production in most species of section Alpestria is consistent with them being of hybrid origin.     

A taxonomic and nomenclatural note on Hieracium caesium (Asteraceae)

The only specimen suitable for typification of Hieracium caesium was discovered in UPS and is here designated as the lectotype. This name appears to have been misapplied and superfluous when originally published at specific rank, but is legitimate according to Art. 52.3. Correct names proposed for three species of the group Caesia (H. caesium = H. basifolium, H. laeticolor and H. plumbeum) are given with infraspecific variants and some more important synonyms. The names H. caesium subsp. laeticolor, H. imitans, H. caesium var. nemorum, H. plumbeum are also lectotypified. Two new combinations H. caesium var. basifolium and H. caesium var. imitans are proposed.     

Genome size in Hieracium subgenus Hieracium (Asteraceae) is strongly correlated with major phylogenetic groups

Background and Aims

Hieracium subgenus Hieracium is one of the taxonomically most intricate groups of vascular plants, due to polyploidy and a diversity of breeeding systems (sexuality vs. apomixis). The aim of the present study was to analyse nuclear genome size in a phylogenetic framework and to assess relationships between genome size and ploidy, breeding system and selected ecogeographic features.

Methods

Holoploid and monoploid genome sizes (C- and Cx-values) of 215 cultivated plants from 89 field populations of 42 so-called ‘basic’ Hieracium species were determined using propidium iodide flow cytometry. Chromosome counts were available for all analysed plants, and all plants were tested experimentally for their mode of reproduction (sexuality vs. apomixis). For constructing molecular phylogenetic trees, the external transcribed spacer region of nuclear ribosomal DNA was used.

Key Results

The mean 2C values differed up to 2·37-fold among different species (from 7·03 pg in diploid to 16·67 in tetraploid accessions). The 1Cx values varied 1·22-fold (between 3·51 and 4·34 pg). Variation in 1Cx values between conspecific (species in a broad sense) accessions ranged from 0·24% to 7·2%. Little variation (not exceeding the approximate measurement inaccurracy threshold of 3·5%) was found in 33 species, whereas variation higher than 3·5% was detected in seven species. Most of the latter may have a polytopic origin. Mean 1Cx values of the three cytotypes (2n, 3n and 4n) differed significantly (average of 3·93 pg in diploids, 3·82 pg in triploids and 3·78 pg in tetraploids) indicating downsizing of genomes in polyploids. The pattern of genome size variation correlated well with two major phylogenetic clades which were composed of species with western or eastern European origin. The monoploid genome size in the ‘western’ species was significantly lower than in the ‘eastern’ ones. Correlation of genome size with latitude, altitude and selected ecological characters (light and temperature) was not significant. A longitudinal component was only apparent for the whole data set, but absent within the major lineages.

Conclusions

Phylogeny was the most important factor explaining the pattern of genome size variation in Hieracium sensu stricto, species of western European origin having significantly lower genome size in comparison with those of eastern European origin. Any correlation with ecogeographic variables, including longitude, was outweighed by the divergence of the genus into two major phylogenetic lineages.Key words: Apomixis, chromosome numbers, Compositae, genome size, hawkweeds, Hieracium subgenus Hieracium, mode of reproduction, nuclear DNA content, phylogeny, polyploidy     

On Hieracium incurrens (Asteraceae) and similar species in central Sweden

Hieracium incurrens Saelan ex Norrl., originally described from Finland, has been reported from Sweden several times and independently by different authors. Still, however, its presence in Sweden has remained dubious and it has become obvious that the different reports are referable to different taxa and the name has thus been, at least partially, misapplied. In the present study, all Swedish material and putative taxa that have been suggested to belong to H. incurrens , as well as some additional closely similar taxa, is critically compared with authentic material of the latter from Finland. It is concluded that H. incurrens do indeed occur in central Sweden, namely in the eastern coastal provinces closest to Finland. However, the relevant material has mostly been referred to H. longilingua Johanss., which is here considered to be a later synonym of H. incurrens. Material previously referred to H. incurrens from other parts of Sweden does, however, represent other taxa. Five accepted species of this affinity are described and discussed in some detail and one species, viz. H. oestmanii T. Tyler is described anew. In addition, the unrelated species H. arnoldii T. Tyler is described anew based on material from the provinces of Hälsingland and Medelpad.     

Lectotypification of names of south Swedish Hieracium species (Asteraceae)

Ninety-six south Swedish taxa in Hieracium L. sect. Hieracium and H. sect Vulgata (Griseb.) Willk. & Lange are lectotypified. The authorship of each name is analysed and the identity and synonomy of some dubious taxa are discussed.-Flora Nordica Note no. 24     

A new diploid Hieracium (Asteraceae: Lactuceae) from Bulgaria

Hieracium kittanae Vladimirov, a new diploid (2 n  = 18) species restricted to a few gorges in the Central Rhodope Mountains, south Bulgaria, is described and illustrated. It occurs in crevices of limestone rock in shady to sunny places with high atmospheric humidity. Morphologically, it to some extent resembles taxa from the H. glaucinum and H. schmidtii groups, but is otherwise very distinct, without close relatives in the Rhodope and neighbouring mountains.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003 , 143 , 213−218.     

Genetic structure of the rare and endangered Hieracium wiesbaurianum group (Asteraceae) in Bavaria

The rare and endangered Hieracium wiesbaurianum species group shows a scattered relictual distribution in Bavaria. Recently, a couple of populations were discovered which clearly differ from all other populations. If these must be considered as taxonomically independent units, they would be of crucial conservation interest, because of the sole responsibility that Bavaria has for these worldwide endemics. We therefore analysed the genetic structure of H. wiesbaurianum in a comparative approach. Our analysis comprised 37 populations of 13 taxa of H. wiesbaurianum, H. bifidum and H. laevigatum, including three potentially new taxa. We applied amplified fragment length polymorphism (AFLP) analysis and observed only limited genetic variation within populations and taxa. Nevertheless, each studied individual exhibited a unique genotype. An analysis of molecular variance revealed high levels of genetic variation between taxa, but populations were genetically less different. The clear genetic differentiation between the studied taxa was supported by neighbor‐joining cluster analyses and principal coordinate analyses in which every individual was clearly assigned to its respective taxon. The three potentially new taxa were genetically as well differentiated as the other taxa included in our study. This supports the assumption that they should be treated as taxonomically independent units of high conservation interest. Therefore, the genetic analysis confirmed the morphologically based classification of the studied Hieracium taxa. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 112–123.     

Ploidy level analysis of apomictic Hieracium (Asteraceae) reveal unexpected patterns and variation

The DNA ploidy level of 673 accessions belonging to 238 species of Hieracium have been analyzed by flow cytometry. 222 of the species were of Scandinavian origin as members of the exclusively apomictic H. sectt. Bifida, Hieracium, Oreadea, Tridentata and Vulgata. For the overwhelming majority of the species, the ploidy level (or chromosme number) has never been investigated before. Approximately 50% of the Scandinavian species, previously believed to be exclusively triploid, were found to be tetraploid. In addition, two pentaploid samples, viz. H. intermarginatum Johanss. & Sam. from Sweden and H. cf. plumbeum Blytt & Fr. from Germany, were found. Although two or more accessions from geographically remote sites were analyzed for approximately 50% of the Scandinavian species, only 2 (<2%) taxonomically undisputed species were found to consist of plants with more than one ploidy level. An intriguing pattern was revealed among Scandinavian members of H. sectt. Bifida and Vulgata when ploidy level and morphometric variation was compared, viz. the most typical or extreme representatives of these sections were found to be exclusively triploid whereas tetraploids dominate among species with intermediate morphology and among species morphologically intermediate between these sections and H. sect. Hieracium. This pattern may indicate that the tetraploids, which tend to have mainly northern distributions, have originated as the result of rare sexual hybridizations between triploid members of different sections, plausibly during or after the northward migration of the parental linages after the last glaciation. The results are believed to be highly relevant for understanding the processes of evolution and speciation within the predominantly apomictic genus Hieracium, but it is emphazised that additional data from e.g. molecular markers and pollen viability analysis are needed before any trustworthy conclusions can be made as far as evolutionary processes are concerned.     

Density and the commitment of apical meristems to clonal growth and reproduction in Hieracium pilosella

Summary We examined responses to population density in the commitment of apical meristems to reproduction and clonal growth in a rosette-forming, stoloniferous herb (Hieracium pilosella). Despite close physiological coupling between the evocation of the terminal inflorescence bud and the development of one or more axillary buds into stolons, the allocation of meristems was extremely plastic.Genets at the higher sowing densities showed density-dependent mortality consistent with self-thinning along a-3/2 trajectory. The probability of inflorescence evocation and associated stolon development was negatively dependent on surviving density. The proportinal distribution of primary stolons amongst genets became strikingly more unequal (expressed as the Gini coefficient) with increasing density. Clonal growth was resolved into the number of primary stolons per stoloniferous genet and the extent of stolon branching (i.e. number of apices per primary stolon); both showed strongly negative density-dependence. Reproduction, expressed as the mean number of flowering capitula per stoloniferous genet, declined 15-fold with increasing density; although theoretically expected to be unity, greater values resulted from capitulum production by attached secondary rosettes and lower values reflected the increasing abortion rate of inflorescence buds with increasing density.Both the total number of apices produced per unit area and the corresponding number of reproductive apices were maximal at intermediate surviving densities (700–1,000 m^-2). The balance between reproductive and clonal growth may be expressed as the probability of an apical meristem producing a capitulum, that also peaked sharply at intermediate density. This finding does not conform with linear models that predict a shift from vegetative growth to sexual reproduction with increasing population density.     

Chromosome numbers in Hieracium and Pilosella species (Asteraceae) from the Iberian Peninsula and the Balearic Islands

Mitotic chromosome numbers are reported from 16 Hieracium and two Pilosella species from the Iberian Peninsula and the Balearic Islands that are little known cytogenetically. The chromosome numbers of Hieracium aragonense Scheele (2 n  = 27, 36), H. compositum Lapeyr. (2 n  = 27), H. murcandidum G. Mateo (2 n  = 27), H. spathulatum Scheele (2 n  = 27), H. segurae Mateo (2 n  = 27), H. teruelanum Mateo (2 n  = 27), H. valentinum Pau (2 n  = 27), Pilosella pseudovahlii (De Retz) Mateo (2 n  = 18), and P. tardans (Peter) Sójak (2 n  = 36) were determined for the first time. New cytotypes were detected in H. cordifolium Lapeyr. (2 n  = 27) and H. loscosianum Scheele (2 n  = 36). The karyotype of the studied species was similar in overall morphology and comprised metacentric and submetacentric chromosomes; secondary constrictions (up to two) were detected in the chromosome complements of some species.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 311–320.     

Three hawkweeds (Hieracium,Asteraceae) from the Netherlands typified and raised to species rank

After a short sketch of Dutch hieraciology, three taxa belonging to Hieracium sectt. Vulgata and Tridentata that were described from the Netherlands at the rank of subspecies or variety by Zahn are typified and raised to species rank. Hieracium meppelense (Zahn) Haveman comb. et stat nov. is found in the province of Drenthe in the northeastern part of the country, H. limburgense (Zahn) Haveman comb. et stat nov. in south‐Limburg in the southernmost part and H. macrodontophyllum (Van Soest et Zahn) Haveman comb. et stat nov. in the surroundings of Nijmegen and Arnhem, and further north in the central sand area. Photographs of the types as well as maps with the hitherto known distribution of these three species are included.     

Reproductive strategy as population density varies in Daphnia magna (Cladocera)

• 1 In this study an experiment was carried out to test whether Daphnia magna (Cladocera) changed its reproductive strategy in response to varying population density but constant food supply.
• 2 As population density increased there was a reduction in the age at which females reproduced. Early reproduction was offset by smaller body length and clutch size (number of eggs per female) relative to later reproducing females grown at lower population densities.
• 3 This pattern was interpreted as a strategy carried out by females to reduce the time at which they release their offspring. One potential advantage of early reproduction is to reduce the risk of severe food limitation for neonates born during a period of rapid population increase.

     

Sexual and apomictic reproduction in Hieracium subgenus pilosella are closely interrelated developmental pathways

Seed formation in flowering plants requires meiosis of the megaspore mother cell (MMC) inside the ovule, selection of a megaspore that undergoes mitosis to form an embryo sac, and double fertilization to initiate embryo and endosperm formation. During apomixis, or asexual seed formation, in Hieracium ovules, a somatic aposporous initial (AI) cell divides to form a structurally variable aposporous embryo sac and embryo. This entire process, including endosperm development, is fertilization independent. Introduction of reproductive tissue marker genes into sexual and apomictic Hieracium showed that AI cells do not express a MMC marker. Spatial and temporal gene expression patterns of other introduced genes were conserved commencing with the first nuclear division of the AI cell in apomicts and the mitotic initiation of embryo sac formation in sexual plants. Conservation in expression patterns also occurred during embryo and endosperm development, indicating that sexuality and apomixis are interrelated pathways that share regulatory components. The induction of a modified sexual reproduction program in AI cells may enable the manifestation of apomixis in HIERACIUM: