Differences in Esterase Distribution Detected by Electrophoresis as Evidence of Continuing Interspecific Hybridization in the Genus Senecio

The distribution of esterases of different mobility, locatedby acrylamide gel electrophoresis, was compared for differentpopulations of plants of the genus Senecio in Central Scotland.Evidence was obtained of continued interspecific hybridization,with gene flow from Senecio squalidus L. to Senecio vulgarisL.     

Effects of Intercropping on Growth and Reproductive Capacity of Late-emerging Senecio vulgaris L., with Special Reference to Competition for Light

Due to increased emphasis on long-term management of weed populationsin cropping systems with a reduced reliance on herbicides, theproduction of seeds by weeds that emerge after the criticalperiod for weed control is increasingly important. It was hypothesizedthat increased soil cover and light interception by a crop canopywould shorten the critical period for weed control and reducegrowth and fecundity of late-emerging weeds. This hypothesiswas tested in a series of field and glasshouse experiments inwhich competition for light was manipulated. Senecio vulgaris,an important weed in vegetable production systems, was chosenas the target plant, and canopies of pure and mixed stands ofleek and celery were used to provide shade. The time courseof light interception differed among the crop canopies. Increasingcompetition for light caused morphological changes to S. vulgaris,including a vertical shift in leaf area distribution. Increasedshading reduced biomass, capitula:shoot ratio and seed productionof S. vulgaris. However, the viability of seeds produced bythe shaded weed plants was not affected. Results indicate thatintercropping can increase light interception in a weakly competitivecrop such as leek and can contribute to weed suppression ina long-term strategy for weed management. Copyright 2001 Annalsof Botany Company Competition for light, late-emerging weeds, critical period, Apium graveolens L., celery, Allium porrum L., leek, Senecio vulgaris L., common groundsel, seed production, weed management, intercropping     

Origins of the new allopolyploid species Senecio cambrensis (asteraceae) and its relationship to the canary islands endemic Senecio teneriffae

The distribution of a 330 bp cpDNA insertion was reexamined in British material of Senecio cambrensis (2n = 6x = 60), its two putative parental taxa, S. squalidus (2n = 2x = 20) and S. vulgaris (2n = 4x = 40), and the closely related Canary Islands' endemic S. teneriffae (2n = 6x = 60). This formed part of a test of the hypothesis that the Welsh form of S. cambrensis is derived from introduced S. teneriffae rather than having originated in Wales via allopolyploidy as previously supposed. It was established that the 330 bp insertion was carried by all plants of Welsh S. cambrensis and also S. teneriffae, but was absent from the cpDNA of Edinburgh S. cambrensis and all plants of British S. squalidus and S. vulgaris var. vulgaris surveyed. However, two of 19 individuals tested of S. vulgaris var. hibernicus also possessed the cpDNA insertion, indicating that it is present in British material of S. vulgaris, although at low frequency. The close similarity between S. teneriffae and S. cambrensis, especially the Welsh form of S. cambrensis, was confirmed by the results of a restriction analysis of rDNA, and also morphometric and crossing studies. However, isozyme analysis showed that S. teneriffae is monomorphic for βEST-3 and ACO-1 phenotypes that are not present in Welsh and Edinburgh S. cambrensis, nor in S. squalidus and S. vulgaris material surveyed. It is concluded that S. teneriffae and S. cambrensis are two closely related allohexaploid taxa that have very similar, but different origins. It is postulated that whereas S. cambrensis (in Wales and Edinburgh) is the allohexaploid of S. vulgaris and S. squalidus, S. teneriffae is possibly the allohexaploid of S. vulgaris and S. glaucus, the latter being a diploid species closely related to S. squalidus.     

Hybridization of Senecio squalidus and S. viscosus and Introgression of Genes from Diploid into Tetraploid Senecio Species

Natural triploid hybrids (Senecio x londinensis Lousley) betweenS. squalidus L. (2n = 20) and 5. viscosus L. (2n = 40) are fairlyfrequently found in Britain. Under glasshouse conditions bothnatural and artificial hybrids displayed very low levels ofseed fertility and gave rise to morphologically diverse F₂ plantsat about the triploid or pentaploid chromosome levels. By theF₄ generation, progeny of a F₂ pentaploid plant had somaticchromosome numbers near to the tetraploid level and considerablyincreased pollen and seed fertilities. Such fertile tetraploidsegregants of S. x londinensis permit the introgression of S.squalidus genes into S. viscosus, and may indicate the courseof introgression into other tetraploid species of Senecio. Senecio, hybridization, introgression     

Recent hybrid origin and invasion of the British Isles by a self-incompatible species, Oxford ragwort (Senecio squalidus L., Asteraceae)

Senecio squalidus is a diploid hybrid species which originated in the British Isles following the introduction of material collected from a hybrid zone on Mount Etna, Sicily, approximately 300 years ago. Introduced hybrid material was cultivated in the Oxford Botanic Garden and gave rise to the stabilized diploid hybrid species, which later spread throughout much of the UK and into some parts of Ireland. Unusually for an invasive species, S. squalidus has a strong system of sporophytic self-incompatibility (SSI) that may have become modified as a result of its recent hybrid origin and spread. First, S. squalidus contains relatively few S alleles (between 2 and 6 S alleles within individual UK populations) compared to other species with SSI (estimates average ~17 S alleles per population). This most probably reflects the population bottleneck experienced by introduced hybrid material. Second, dominance relationships among S. squalidus S alleles are more extensive than those reported in other species with SSI. Third, although pseudo-self-compatibility occurs sporadically in S. squalidus, it is not widespread, indicating that SSI is maintained in the species despite potential mate availability restrictions imposed by low numbers of S alleles. Surveys of other forms of genetic diversity in S. squalidus show that allozyme variation is reduced relative to that within the progenitor species, but Randomly Amplified Polymorphic DNA variation is relatively high. Both types of genetic variation show little or no pattern of isolation-by-distance between populations in keeping with the recent range expansion of the species. During its spread in the British Isles, S. squalidus has hybridized with the native self-compatible (SC) tetraploid species, S. vulgaris, which has led to the origin of three new SC hybrid taxa: a radiate form of S. vulgaris (var. hibernicus), a tetrapoid hybrid species (S. eboracensis) and an allohexaploid (S. cambrensis).     

The Effects of Infection by Rust (Puccinia lagenophorae Cooke) on the Growth of Groundsel (Senecio vulgaris L.) Cultivated under a Range of Nutrient Concentrations

Groundsel (Senecio vulgaris L.), healthy or infected with therust fungus Puccinia lagenophorae Cooke, was grown at a rangeof nutrient concentrations in sand culture. There were statisticallysignificant interactions between the effects of infection andnutrient supply upon the dry weights of stems, leaves, rootsand reproductive tissues, leaf area and cumulative capitulumproduction. This interaction occurred since infection causedsignificant inhibitions of growth only at moderate or high nutrientconcentrations. At low concentrations rusted plants were similarto or slightly larger than controls. Both in controls and rustedplants root: shoot ratios increased as nutrient supply declined.The ratio of root: shoot dry weight was consistently reducedby infection whilst root length: leaf area ratio was relativelyunchanged. More detailed investigations confirmed that infection had littleeffect on plant growth under nutrient deficient conditions despitesuppression of the host's ability to increase root: shoot ratiosin response to nutrient stress. This reflected the inhibitionof relative growth rates in rusted plants at high but not lownutrient concentrations, which in turn reflected reduced netassimilation rates (NAR). Increases in leaf-area ratio (LAR)often ameliorated the decline in NAR in rusted plants. Senecio vulgaris L., Puccinia lagenophorae Cooke, nutrient deficiency, growth, root: shoot ratio     

Nutrient Relations of Groundsel (Senecio vulgaris) Infected by Rust (Puccinia lagenophorae) at a Range of Nutrient Concentrations II. Uptake of N, P and K and Shoot-Root Interactions

Groundsel (Senecio vulgaris), healthy or infected with rust,Puccinia lagenophorae, was grown at a range of nutrient concentrationsin sand culture. Specific absorption rates calculated on thebasis of root dry weight (SAR_W) were greater in rusted thancontrol groundsel for nitrogen, potassium and phosphorus. Whilethe magnitudes of these stimulations varied, they occurred acrossthe whole range of nutrient concentrations. By contrast, specificabsorption rate on the basis of root length (SAR_L) were littlechanged by rust at any external nutrient concentration; SAR_Lfor phosphate and potassium were slightly reduced when nutrientswere freely available. Water flux per unit dry root weight and length was stimulatedby rust because transpiration per unit leaf area was more rapidin infected plants after fungal sporulation. However, water-fluxand the rate of uptake of nutrients were correlated only whenexpressed on the basis of root weight and increased transpirationdid not appear to be the mechanism underlying increased rootactivity. Rather, increased SAR_W for N, P and K could very largelybe attributed to increased shoot demand per unit root, whichresulted from the higher shoot: root (S: R) ratios of infectedindividuals. Changes in S: R accounted for 92, 81 and 57% oftotal variation in SAR_W for K, P and N respectively. Greatervalues for SAR_W were possible because specific root length (SRL)increased, producing more functional root per unit root weight.The lack of stimulation in SAR_L in response to rust could beexplained since the higher SRL of infected plants resulted instable values of shoot weight per unit root length, i.e. shootdemand was not increased by infection on this basis. Senecio vulgaris, Puccinia lagenophorae, rust infection, nutrient uptake, water uptake, shoot: root interactions     

Nutrient Relations of Groundsel (Senecio vulgaris) Infected by Rust (Puccinia lagenophorae) at a Range of Nutrient Concentrations I. Concentrations, Contents and Distribution of N, P and K

Groundsel (Senecio vulgaris L.), healthy or infected with therust fungus Puccinia lagenophorae was grown in sand and fedwith a complete nutrient medium diluted to give a range of concentrations.Analysis of bulked, dried tissues of the plant showed that undernutrient-rich conditions rust infection resulted in increasedconcentrations of total (Kjeldahl) nitrogen and potassium buthad little effect on phosphorus concentration. Thus, despitereduced dry weight growth, total plant nitrogen contents wereno less in rusted than control plants. Although total contentsof phosphorus and potassium were reduced by rust, effects wereprobably related to loss of these nutrients in fungal spores. Interactions between rust infection and nutrient supply weresignificant but differed between nutrients: rust caused increasednitrogen concentrations only under nutrient-rich conditionsbut increased phosphorus concentrations only when nutrient supplywas limited. Increased concentrations were not confined to infectedtissues. Mechanisms underlying rust-nutrient interactions appearto be complex and to depend inter alia on the partitioning andrecycling of the particular nutrient within the plant. Rust-inducedincreases in potassium concentration occurred under both highand low nutrient conditions but were confined to infected tissues.Potassium accumulation in nutrient deficient conditions wasprobably due to increased transpirational flux into infectedtissues, but under nutrient-rich conditions reduced potassiumexport appeared to assume greater significance. The possible implications of the changed nutrient relationsfor the wider interactions of rust-infected plants in naturalvegetation are discussed. Senecio vulgaris, Puccinia lagenophorae, rust infection, nutrient deficiency, nutrient content, nutrient concentration, nutrient distribution     

The Effects of Nutrient Deficiency and Rust Infection on the Relationship Between Root Dry Weight and Length in Groundsel (Senecio vulgaris L.)

Groundsel (Senecio vulgaris L.) was grown in sand culture ata range of nutrient concentrations. Except when nutrient deficiencywas severe, infection by the rust fungus Puccinia lagenophoraeCooke substantially reduced root dry weight but had little effecton root length. Thus, specific root length (SRL, cm root mg^–1d. wt) was significantly increased in rust-infected plants.The inhibition of root dry weight caused by rust infection wasmost pronounced late in development, especially after floweringwhen, in control plants, root elongation but not dry weightaccumulation ceased. In rusted plants, and in all plants subjectedto severe nutrient deficiency, dry weight accumulation in theroots ceased concurrently with root elongation. Late in developmentat high nutrient concentration adventitious roots with low SRLswere produced. However, infection did not modify the productionof such roots and increases in SRL could not be attributed tochanges in any single type of root. There was an inverse relationship between SRL and root diameter.This relationship was unaffected by rust infection whilst nutrientdeficiency changed only its intercept: at a given SRL rootsof nutrient stressed plants were thinner than those of plantswith adequate nutrient supply. Thus, the smaller diameter ofroots of nutrient-stressed plants occurred independently ofmeasured changes in SRL but, in the absence of nutrient stress,the decrease in root diameter caused by rust was closely relatedto increases in SRL. Changes in the root: length relationships in rusted plants mayhave important implications for root activity in the field.In view of the reported changes in SRL, inhibition of root growthin terms of dry weight may be a poor indicator of potentialchanges in activity. Senecio vulgaris, rust infection, nutrient deficiency, root weight: length ratio, root diameter     

Early adjustment of patterns of metaphase association in the evolution of a polyploid species

Evidence is presented for the adjustment of chromosomal association at metaphase in the very early stages of establishment of a new level of polyploidy. The species concerned is S. cambrensis Rossner, the hexaploid hybrid between S. squalidus L. (2n=20) and S. vulgaris (2n=40). It is suggested that the perfect bivalent pairing of S. vulgaris is the basis for the success of S. cambrensis.     

Hybrid swarms: catalysts for multiple evolutionary events in Senecio in the British Isles

Background: Introgressive hybridisation is an evolutionary catalyst producing novel variants able to explore new ecological niches and evolve as new hybrid taxa. However, the role of ‘hybrid swarms’ – highly variable populations produced following interspecific hybridisation – in generating this evolutionary novelty has been poorly studied.

Aims: We examine the alternative origins of tetraploid hybrid derivatives of Senecio vulgaris and S. squalidus, via local polytopic formation or long-distance dispersal from a single perennial hybrid swarm around Cork, Ireland.

Methods: Morphometric, isozyme and chloroplast DNA analysis.

Results: The Cork hybrid swarm and UK hybrid swarms exhibited a broad range of morphological variation and contained individuals similar to the stable tetraploid hybrid derivatives; S. eboracensis and S. vulgaris var. hibernicus. Chloroplast DNA analysis shows that S. eboracensis did not evolve from the Cork hybrid swarm. However, UK S. vulgaris var. hibernicus populations exhibit a broad range of variation for both chloroplast and isozyme markers, but were not distinguishable from Cork material.

Conclusions: Our study confirms that S. eboracensis did not evolve from the Cork hybrid swarm, and while our analyses could not demonstrate this conclusively for S. vulgaris var. hibernicus the ease with which hybrid swarms have been generated in the past makes a polytopic origin for S. vulgaris var. hibernicus the most likely scenario.     

Comparison of Atrazine-Resistant and -Susceptible Biotypes of Senecio vulgaris L.: Effects of High and Low Temperatures on the in vivo Photosynthetic Electron Transfer in Intact Leaves

The effects of temperature on the yield of in vivo modulatedchlorophyll fluorescence were measured in intact leaves of atrazineresistantand -susceptible biotypes of the weed Senecio vulgaris L. At25 ?C, the photochemical quenching (q_Q of steady-state chlorophyllfluorescence was reduced by around 30% in the atrazine-resistantmutant as compared to the susceptible wild type, indicatinga higher reduction state of the primary electron acceptor Q_Aof photosystem II in the former biotype. Moderately elevatedtemperatures (above 30 ?C) further increased the steady-stateconcentration of reduced in the mutant. Analysis of the temperature dependence of both the photochemicalquenching q_Q and the initial fluorescence level F_o clearly indicateda drastically enhanced heat-sensitivity of the photochemicalapparatus in the atrazine-resistant Senecio leaves. The heat-inducedchanges in F₀ and q_Q were closely correlated, suggesting thatthe phenomenon responsible for the rise in F₀ was also involvedin the inhibition of the photosynthetic electron flux. Low temperaturesalso affected reoxidation but, in contrast to heating, no apparent differences were observed in the behaviourof the two biotypes exposed to chilling stress. Key words: Atrazine-resistance, chlorophyll fluorescence, heat stress, Senecio vulgaris     

Members of the<Emphasis Type="Italic"> S</Emphasis>-receptor kinase multigene family in<Emphasis Type="Italic"> Senecio squalidus</Emphasis> L. (Asteraceae), a species with sporophytic self-incompatibility

While the molecular basis of sporophytic self-incompatibility (SSI) has been investigated extensively in the Brassicaceae, almost nothing is known about the molecular regulation of SSI in other families, such as the Asteraceae. In species of Brassica and in Arabidopsis lyrata, a stigma-specific serine-threonine receptor kinase (SRK) and its cognate ligand, a pollen coating-borne cysteine-rich protein (SCR/SP11), determine the female and male sides of the SSI response, respectively. Here we have used RT-PCR with degenerate primers to conserved regions of SRK to amplify three SRK-like gene fragments expressed in stigmas of Senecio squalidus (Asteraceae). The Senecio S-receptor-like kinase (SSRLK) sequences share ~43% amino acid sequence identity with Brassica SRK₃ but higher amino acid sequence identity (~50%) with two Solanum bulbocastanum receptor-like kinase genes of unknown function. Despite expression in stigmas, all three SSRLKs were expressed at varying levels in floral and vegetative tissues. No allelic polymorphism was detected for the three SSRLKs in two S homozygous lines of S. squalidus or three other lines of S. squalidus carrying different S alleles. A full-length cDNA clone was obtained for SSRLK1 and its predicted amino acid sequence revealed significant structural differences to Brassica SRKs, most notably a major N-terminal truncation of 169 amino acids and the presence of just 8 conserved cysteine residues within the putative receptor domain instead of 12. Together, the sequence characteristics and expression characteristics of SSRLKs suggest that they are unlikely to be directly involved in the SSI response of S. squalidus. These findings are discussed in terms of the evolution of the SRK multigene family and the molecular basis of SSI in S. squalidus and the Asteraceae.     

Alkaloid patterns and biosynthetic capacity of root cultures from some pyrrolizidine alkaloid producing Senecio species

Root cultures of Senecio vulgaris, S. vernalis, S. erucifolius and S. squalidus were established. The patterns of pyrrolizidine alkaloids found in these root cultures were analyzed by high-resolution GC and GC-MS and compared with the alkaloids present in the respective plants. In vitro cultured roots produce alkaloid patterns and accumulate quantities which are comparable to those found in soil grown plants. With the exception of the otonecine derivative senkirkine all pyrrolizidines accumulate as N-oxides. Only senkirkine is partially released into the medium. The cultures incorporate biosynthetic precursors, e.g. ¹⁴C-labelled putrescine or spermidine with high efficiency into the alkaloids. Senecionine N-oxide was found to be the main product of biosynthesis. Evidence is presented that senecionine N-oxide is directly transformed into senkirkine, the main alkaloid of S. vernalis root cultures.Abbreviations GC Gas chromatography - MS Mass spectroscopy - PND Phosphorous-Nitrogen-Detector - FID Flame Ionization Detector - fr.wt Fresh weight     

Isolation and characterization of microsatellite loci in Senecio

Interspecific hybridization has resulted in the recent origin of several hybrid Senecio taxa at diploid, tetraploid and hexaploid levels. As part of research aimed at constructing and comparing genomic maps of each of these taxa and their parents, we have isolated microsatellite loci from genomic DNA libraries of S. vulgaris and S. squalidus. Primers of 35 loci amplified microsatellites resolved in agarose gels from one or more of S. vulgaris, S. squalidus, S. aethnensis and S. chrysanthemifolius. Approximately 71% of primers amplified a product in all four species. A survey of microsatellite variation in S. chrysanthemifolius over a subset of 14 loci resolved 2–11 alleles per locus in polyacrylamide gels with expected heterozygosity (H_E) ranging from 0.26 to 0.87.     

Changes in Tissue Freezing in Senecio vulgaris infected by Rust (Puccinia lagenophorae)

Freezing of healthy and rust (Puccinia lagenophorae) infectedleaves of Senecio vulgaris was compared calorimetrically bythermal analysis. In fully expanded leaves the threshold freezingtemperature was in the range –6.8 to –8.4 °Cin controls but –3.0 to –5.1 °C in leaves withsporulating rust sori. Comparable values in expanding leaveswere –5.0 to –8.9 °C and –3.9 to –6.7°C for healthy and rusted tissues, respectively. The bulktissue freezing point was between –1.0 and –4.0°C in both fully expanded and expanding healthy leaves,and was increased by infection by between +0.2 and 2.5 °C.Whereas healthy leaves supercooled by 3.1–5.8 °C,rusted leaves supercooled by only 1.8–4.9 °C Supercoolingof control leaves was reduced by dusting with aeciospores, particularlywhen leaves were wounded to simulate the rupture of the surfacecaused by sporulation, but wounding alone had no significanteffect. Supercooling of distilled water was also significantlyreduced by aeciospores, suspended at a concentration of 10⁵spores ml^–1. It is concluded that rust-induced changes in leaf freezing inS. vulgaris grown in controlled environments were due to anincrease in the number of sites for ice nucleation, caused bythe presence of the aeciospores, and increased penetration ofice into internal tissues, resulting from damage to the cuticleand epidermis. Although data for frost resistance obtained inthe growth-room are similar to previous field observations,the role of the above mechanisms under field conditions remainsunproven. Senecio vulgaris (groundsel), Puccinia lagenophorae (rust), low temperature, freezing resistance     

Palatability to a generalist herbivore,defence and growth of invasive and native <Emphasis Type="Italic">Senecio</Emphasis> species: testing the evolution of increased competitive ability hypothesis

This paper tests the prediction that introduced plants may become successful invaders because they experience evolutionary changes in growth and defence in their new range [evolution of increased competitive ability hypothesis (EICA)]. Interspecific and intraspecific binary feeding choices were offered to the snail Helix aspersa. The choices were between: (1) plants of the invasive Senecio inaequidens and Senecio pterophorus derived from populations in the introduced range (Europe) and plants of three indigenous species (Senecio jacobea, Senecio vulgaris and Senecio malacitanus) from populations in Europe; (2) plants of the invasive S. inaequidens and S. pterophorus from populations in the introduced range (Europe) and from populations in the native range (South Africa). We did not find a clear pattern of preference for indigenous or alien species of Senecio. However, we found that European invasive populations of S. inaequidens and S. pterophorus were less palatable than South African native populations. Moreover, in contrast to the predictions of the EICA hypothesis, the invasive genotypes of both species also showed a higher total concentration of pyrrolizidine alkaloids, and in the case of S. inaequidens we also found higher growth than in native genotypes. Our results are discussed with respect to the refinement of the EICA hypothesis that takes into account the difference between specialist and generalist herbivores and between qualitative and quantitative defences. We conclude that invasive populations of S. inaequidens and S. pterophorus are less palatable than native populations, suggesting that genetic differentiation associated with founding may occur and contribute to the plants’ invasion success by selecting the best-defended genotypes in the introduced range.     

Quantitative assessment of heteroplasmy levels in <Emphasis Type="Italic">Senecio vulgaris</Emphasis> chloroplast DNA

Heteroplasmy in coding chloroplast DNA was only recently shown to occur and was so far not quantitatively assessed. We present a quantitative analysis of cpDNA heteroplasmy levels at a triazine-resistance determining site within and between individual Senecio vulgaris plants. Detectable levels of heteroplasmic haplotypes were observed in all tested plants. As expected, the levels of heteroplasmy vary greatly between plants. However, even within individual plants, the fraction of one haplotype may cover a range from below 1 to well over 90. Our results suggest that heteroplasmy may be a common phenomenon in S. vulgaris.Possible consequences for molecular diagnostics of chloroplast encoded traits as well as evolutionary consequences of chloroplast heteroplasmy are discussed.     

Sporophytic self-incompatibility in Senecio squalidus (Asteraceae): S allele dominance interactions and modifiers of cross-compatibility and selfing rates

Understanding genetic mechanisms of self-incompatibility (SI) and how they evolve is central to understanding the mating behaviour of most outbreeding angiosperms. Sporophytic SI (SSI) is controlled by a single multi-allelic locus, S, which is expressed in the diploid (sporophyte) plant to determine the SI phenotype of its haploid (gametophyte) pollen. This allows complex patterns of independent S allele dominance interactions in male (pollen) and female (pistil) reproductive tissues. Senecio squalidus is a useful model for studying the genetic regulation and evolution of SSI because of its population history as an alien invasive species in the UK. S. squalidus maintains a small number of S alleles (7–11) with a high frequency of dominance interactions. Some S. squalidus individuals also show partial selfing and/or greater levels of cross-compatibility than expected under SSI. We previously speculated that these might be adaptations to invasiveness. Here we describe a detailed characterization of the regulation of SSI in S. squalidus. Controlled crosses were used to determine the S allele dominance hierarchy of six S alleles and effects of modifiers on cross-compatibility and partial selfing. Complex dominance interactions among S alleles were found with at least three levels of dominance and tissue-specific codominance. Evidence for S gene modifiers that increase selfing and/or cross-compatibility was also found. These empirical findings are discussed in the context of theoretical predictions for maintenance of S allele dominance interactions, and the role of modifier loci in the evolution of SI.     

Lectotypification of the Bertoloni's names in the genus Senecio L. (Asteraceae)

Original material of the four Senecio taxa described by A. Bertoloni has been examined, and the typification of their names is discussed. A specimen from the Monti Herbarium (kept in BOLO) and iconographies from Bonanni and Bertoloni are designated as lectotypes, whereas another specimen from the Monti Herbarium is selected as neotype. The nomenclature of the names Senecio erucifolius L., Senecio tenuifolius Jacq., Senecio delphinifolius Vahl and Senecio rupestris Waldst. & Kit. is also analysed. The names S. crinitus Bertol. and S. laciniatus Bertol. belong to S. delphinifolius [now Jacobaea delphinifolia (Vahl) Pelser & Veldkamp] and S. rupestris [now Senecio squalidus subsp. rupestris (Waldst. & Kit.) Greuter], respectively, whereas S. erraticus Bertol. and S. praealtus Bertol. are accepted as separate taxa as Jacobaea erratica (Bertol.) Fourr. and Jacobaea erucifolia subsp. praealta (Bertol.) Greuter & B.Nord., respectively.