Natural hybridization between Senecio jacobaea and Senecio aquaticus: molecular and chemical evidence

Hybridization is known to be involved in a number of evolutionary processes, including species formation, and the generation of novel defence characteristics in plants. The genus Senecio of the Asteraceae family is highly speciose and has historically demonstrated significant levels of interspecific hybridization. The evolution of novel chemical defence characteristics may have contributed to the success of Senecio hybrids. Chemical defence against pathogens and herbivores has been studied extensively in the model species Senecio jacobaea, which is thought to hybridize in nature with Senecio aquaticus. Here, we use amplified fragment length polymorphisms (AFLPs) and pyrrolizidine alkaloid (PA) composition to confirm that natural hybridization occurs between S. jacobaea and the closely related species S. aquaticus. AFLPs are also used to estimate the ancestry of hybrids. We also demonstrate that even highly back-crossed hybrids can possess a unique mixture of defence chemicals specific to each of the parental species. This hybrid system may therefore prove to be useful in further studies of the role of hybridization in the evolution of plant defence and resistance.     

Occurrence of tetraploid and octoploid cytotypes in Senecio jacobaea ssp. jacobaea (Asteraceae) in Pannonia and the Carpathians

Four different cytotypes have been reported for Senecio jacobaea L. ssp. jacobaea throughout Europe, with the most common occurrence of tetraploids (2 n  = 40). Here we present a survey of previously published chromosome number data on this subspecies and its geographical distribution, and focus on populations from Pannonia and the Carpathians. Two ploidy levels have been determined in the study area, using chromosome counting and flow cytometry: tetraploid (2 n  = 40) and octoploid (2 n  = 80). Fifty-one populations originating from Slovakia, Austria, the Czech Republic, Hungary, Ukraine and Romania have been analysed. Multivariate morphometric analyses have been performed on 39 populations to study morphological differentiation between these two cytotypes. Despite slight morphological tendencies expressed on the level of populations, tetraploid and octoploids cannot be reliably distinguished morphologically and they are not taxonomically classified formally here.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 153 , 231–242.     

Somatic hybrids and cybrids of Senecio fuchsii Gmel. (x) jacobaea L.

Summary In situ hybridisation and restriction fragment length polymorphism (RFLP) analysis were used to determine the relative location of the translocation breakpoint and the size of the integrated chromatin segment in hexaploid wheat-Lophopyrum translocation stocks. Three 7el₂-7D recombinant stocks were Robertsonian translocations, 7DS.7el. The remaining recombinant stock (KS10-2) was 7elS.7el-7DL and contained only the distal one-half of the long arm of 7D. The recombinant stock with 7el₁ (K11695) could be designated 7DS.7DL-7el where approximately the distal one-half of 7DL was replaced. RFLP analysis indicated that on the 7DL RFLP map the breakpoints for K11695 and KS10-2 are in different locations and that the two recombinants contain an overlapping region (a common region) of the Lophopyrum chromosome 7 in which Lr19, a leaf-rust resistant gene, is located. RFLP analysis also indicated that RFLP markers which mapped to within 1.5 cm of the centromere of chromosome 7D are located in the distal half of the long arm.     

The genetic basis of floral variation in Senecio jacobaea (Asteraceae).

The self-incompatible composite Senecio jacobaea (ragwort) exhibits geographic variation in the frequency of rayed and discoid (rayless) individuals. Hybrid progenies from within- and between-morph crosses were established in a seminatural (garden) environment to determine whether patterns of segregation conform to single-gene predictions (as found in other Senecio species), whether the direction of dominance is conducive to rapid evolutionary change in ray morphology, and whether geographically distant populations of the discoid morph utilize the same or different genes to suppress ray development. Data from segregating F2 and BC families were consistent with a genetic model involving one major locus and an unknown number of modifiers. Analysis of F1 progenies from different intermorph crosses using the same rayed plant as a seed parent revealed a variable and incomplete pattern of dominance, with a trend toward partial dominance in some crosses. Hybridizations between discoid populations produced a few rayed progeny (4%), but there was no tendency for the frequency of rayed progeny to increase with the geographic distance separating the parent populations. Results of this study indicate that major mutations have been important for the evolution of discoid populations of ragwort, that ray-suppressing mutations should be directly available to selection in most populations, and that the suppression of rays is conditioned by the same or similar gene(s) in Atlantic and Baltic populations of the discoid taxon.     

Natural hybrids in Senecio (Asteraceae): new records from Argentina

Abstract

Hybridisation is very common within Senecio. It is considered a fundamental mechanism of speciation and also a major feature causing systematic complexity. Morphological features are the preliminary characteristics used to identify hybrids; however this approach is usually not sufficient when this is the case chromosome studies represent a valuable tool for reinforcing the observations. Two natural homoploid interspecific hybrids are reported here: Senecio × lulioi M. G. López et Xifreda (S. brasiliensis (Spreng.) Less. × S. grisebachii Baker) and Senecio × piepaloensis M. G. López et Xifreda (S. ragonesei Cabrera × S. sectilis Griseb.). Classic cytogenetic analysis and pollen stainability were performed to evaluate hybrid origins and to designate the putative parents. S. × lulioi presented a low frequency of quadrivalents and heteromorphic bivalents, both strong evidences of hybridisation between closely related species. S. × piepaloensis exhibited a high frequency of meiotic irregularities, and reduced pollen stainability, certain characteristics of hybrids. This analysis is discussed in relation to the role of hybridisation in the evolution of the genus, and its systematic significance.     

Floral display and pollination success in Senecio jacobaea (Asteraceae): Interactive effects of head and corymb size

The self-sterile Senecio jacobaea (Asteraceae) presents its rayed heads in large, compound inflorescences (corymbs). I examined the role of head and corymb size in pollinator attraction, and whether the positive effect of intact rays (if any) depends on the size of the corymb. Using female fertility as a measure of pollination success, I assessed the performance of stems representing four experimentally produced character combinations: (1) few heads without rays, (2) many heads without rays, (3) few heads with rays, and (4) many heads with rays. The proportion of flowers setting fruit was higher for intact stems (treatments 2, 4) than for stems on which the majority of the heads had been removed (treatments 1, 3), suggesting selection for maximum inflorescence production. By contrast, experimental removal of all rays had a relatively weak negative effect on fruit set, with few-headed stems (treatment 1) experiencing a greater reduction than stems with many heads (treatment 2). These results suggest that clusters of heads produce a synergistic effect on pollinator attraction, allowing plants to maintain high visitation rates even if there are drastic reductions in the basic attraction units. Hence, the number of heads and the attractiveness of the individual heads interacted in their effect on pollination success. Fruit set per flower differed greatly between sites and was positively correlated with plant density.     

Taxonomìc analysis of Senecio pauciflorus and S. indecorus (Asteraceae)

Senecio pauciflorus and S. indecorus (= Packera pauciflora and P. indecora ) are autogamous, high polyploid species with similar geographic ranges in North America. Their east-west distribution pattern including a disjunction in California is unusual within the aureoid Senecio complex and may provide insights into the origins of the aureoid complex. Taxonomic treatment of the two taxa has varied in the past due to the similarity and variability of the taxa. Results of morphological analyses of 192 herbarium specimens show that the taxa are best treated as two distinct species and that infraspecifìc variation is not related to geographical distribution. Separation of taxa remains difficult. California specimens are the most difficult to classify and may represent the ancestral group. Based on morphology and chromosome number it is suggested that the 5. pauciflorus-S. indecorus group may have originated from the arctic-alpine aureoid group including S. cymbalaria .     

Chloroplast DNA and isozyme analysis of the progenitor-derivative species relationship between Senecio nebrodensis and S. viscosus (Asteraceae)

A comparison of chloroplast DNA (cpDNA) and isozyme diversity in Senecio nebrodensis L., a species restricted to certain mountain ranges in Spain, and S. viscosus L., a widespread ruderal species in Europe, revealed that S. viscosus possessed the more common of two cpDNA haplotypes resolved in S. nebrodensis and contained only a small subset of the alleles found in S. nebrodensis at enzyme coding loci. The two species shared one restriction length mutation and one site mutation in their cpDNA, which distinguished them from other European Senecio species examined previously. Taken overall, these results support the hypothesis that S. nebrodensis and S. viscosus are related as a progenitor-derivative pair of species. The fact that no novel alleles were found in S. viscosus would suggest a relatively recent origin of the species, most probably in late glacial or postglacial times.     

Sporophytic self-incompatibility in Senecio squalidus L (Asteraceae)--the search for S

Senecio squalidus (Oxford Ragwort) is being used as a model species to study the genetics and molecular genetics of self-incompatibility (SI) in the Asteraceae. S. squalidus has a strong system of sporophytic SI (SSI) and populations within the UK contain very few S alleles probably due to a population bottleneck experienced on its introduction to the UK. The genetic control of SSI in S. squalidus is complex and may involve a second locus epistatic to S. Progress towards identifying the female determinant of SSI in S. squalidus is reviewed here. Research is focused on plants carrying two defined S alleles, S(1) and S(2). S(2) is dominant to S(1) in pollen and stigma. RT-PCR was used to amplify three SRK-like cDNAs from stigmas of S(1)S(2) heterozygotes, but the expression patterns of these cDNAs suggest that they are unlikely to be directly involved in SI or pollen-stigma interactions in contrast to SSI in the Brassicaceae. Stigma-specific proteins associated with the S(1) allele and the S(2) allele have been identified using isoelectric focusing and these proteins have been designated SSP1 (Stigma S-associated Protein 1) and SSP2. SSP1 and SSP2 cDNAs have been cloned by 3' and 5' RACE and shown to be allelic forms of the same gene, SSP. The expression of SSP and its linkage to the S locus are currently being investigated. Initial results show SSP to be expressed exclusively in stigmas and developmentally regulated, with maximal expression occurring at and just before anthesis when SI is fully functional, SSP expression being undetectable in immature buds. Together these data suggest that SSP is a strong candidate for a Senecio S-gene.     

Reproductive isolation of a new hybrid species, Senecio eboracensis Abbott & Lowe (Asteraceae)

The nature and extent of reproductive isolation was examined between a new self-compatible hybrid species Senecio eboracensis (2n=40) and its parents, self-incompatible S. squalidus (2n=20) and self-compatible S. vulgaris (2n=40). The triploid F(1) of S. eboracensis x S. squalidus exhibited very low seed set (x=0.63%), and F(2) and F(3) progeny were able to recover nominal levels of fertility (x=23.9 and 9.7%), while F(1) and F(2) offspring of S. eboracensis x S. vulgaris showed reduced seed set (x=63.8 and 58.8%). In both cases, evidence from previous work indicates that reduced fertility is associated with meiotic chromosome mispairing, and is a likely consequence of recombining both parental genomes within this new taxon. No hybrid offspring between S. eboracensis and S. squalidus were found in the wild, and only one such hybrid was recorded among 769 progeny produced by S. eboracensis surrounded by S. squalidus on an experimental plot. Natural crossing between S. eboracensis and S. vulgaris was recorded to be very low (between 0 and 1.46%) in the wild, but rose to 18.3% when individuals of S. eboracensis were surrounded by plants of S. vulgaris. It was concluded that strong breeding barriers exist between the new hybrid species and its two parents. Prezygotic isolation between S. eboracensis and S. vulgaris is likely to be largely due to both species reproducing by predominant self-fertilisation. However, differences recorded for germination, seedling survival, time of flowering and characters associated with pollinator attraction, plus significant clumping of juvenile and adult conspecifics in the wild, probably also contribute to reproductive isolation and ecological differentiation.     

Taxonomic studies on Senecio aquaticus (Asteraceae). A recommendation for the taxonomic status of Aquaticus and Barbareifolius

Aquaticus and Barbareifolius are two subgroups of Senecio aquaticus s. l. (Asteraceae) that have been recognized at various taxonomic levels in European taxonomic literature. In this paper we present the results of morphological studies on S. aquaticus s. l. in order to establish the taxonomic status of these two subgroups. Using multivariate morphometric techniques (discriminant analysis and principal component analysis), we examined the diagnostic potential of 60 macro-morphological characters and studied the delimitation of Aquaticus and Barbareifolius. Seven of the characters studied were included in the discriminant function of the stepwise discriminant analysis and are therefore considered to be the most reliable diagnostic characters of Aquaticus and Barbareifolius. Barbareifolius has, amongst other distinguishing traits, a more divaricate inflorescence than Aquaticus. Because both subgroups are distinct but overlapping clusters in multivariate morphometric space, and because clear differences in their geographical distribution are absent, we recommend the taxonomic status of variety to both groups. Following the rules of nomenclature, Aquaticus and Barbareifolius should therefore be named S. aquaticus Hill var. aquaticus and S. aquaticus Hill var. barbareifolius (Krock.) Wimm. & Grab.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 489–498.     

Fitness consequences of floral variation in Senecio jacobaea (Asteraceae): evidence from a segregating hybrid population and a resource manipulation experiment

The present study examines some of the ecological factors that might exert selection on floral morphology in Senecio jacobaea , a self-sterile composite which exhibits geographic variation in the frequency of rayed and discoid individuals. Regression analyses of phenotypic data from a large, segregating hybrid population, established in a semi-natural (garden) environment and studied over a 2-year period, revealed a negative relationship between the size of the rays and the average germination rate of the maternal seed crop, a pattern that can be attributed to the reduced germination speed of achenes from ray florets. There was no effect of ray size on the amount of cross-pollination achieved, the proportion of heads infested by larvae of seed flies ( Pegohylemyia ) and the amount of resources retained for the next flowering season. The lack of resource costs was also apparent in a manipulation experiment with greenhouse-grown plants of the rayed phenotype: artificial removal of all rays at the early bud or flowering stage had no detectable effect on subsequent flower and fruit development, regardless of whether the plants experienced high or low water stress. Given these and other observations, I hypothesize that plant-animal interactions and resource costs sometimes play a minor role in exerting selection on flower morphology and that spatially varying selection on germination behaviour accounts for some of the morph frequency variation in S. jacobaea.     

Senecio josei and S. superparamensis spp. nov. (Asteraceae: Senecioneae) from the Andes of Ecuador

Research on the phylogeny of a distinct group of Andean Senecio, previously considered as Lasiocephalus, resulted in the discovery of two new species, which are here described and illustrated. Senecio josei Sklená?, which grows in rocky habitats in grass páramo in southern Ecuador, is characterized by linear leaves and a racemose or paniculate synflorescence. Senecio superparamensis Sklená? is characterized by grayish–white pubescence, solitary capitula and leafy stems, and inhabits superpáramos in northern and central Ecuador.     

Phylogeny reconstruction in the gap between too little and too much divergence: the closest relatives of Senecio jacobaea (Asteraceae) according to DNA sequences and AFLPs

The species composition and molecular phylogeny of Senecio sect. Jacobaea (Asteraceae; Senecioneae) were studied to identify the closest relatives of Senecio jacobaea. Maximum parsimony and Bayesian inference analyses of DNA sequence data of the plastid (the trnT-L igs, the trnL intron, two parts of the trnK intron, and the psbA-trnH igs) and nuclear genome (ITS1, 5.8S, and ITS2) showed these markers to be suitable to assess the species composition of sect. Jacobaea, identifying 24 species as members of this section. Of these, nine species were not previously assigned to the section. The selected DNA sequence regions, however, showed too little sequence divergence to be optimal for phylogenetic inference within sect. Jacobaea. In contrast, AFLPs proved to be too variable to be used to study relationships between the basal lineages in sect. Jacobaea. Nonetheless, these markers are very useful to study the phylogeny of S. jacobaea and its closest relatives. The combined use of DNA sequence data and AFLPs allowed us to take a major step towards resolving phylogenetic relationships in sect. Jacobaea, identifying Senecio alpinus, Senecio pancicii and Senecio subalpinus (using DNA sequence data) or Senecio chrysanthemoides (using AFLPs) as the closest relatives of S. jacobaea.     

Dormancy and dispersal in dimorphic achenes of tansy ragwort,Senecio jacobaea L. (Compositae)

Summary Marginal and central florets of the capitula of tansy ragwort Senecio jacobaea yield different kinds of fruit. The central (disk) achenes are lighter (x±SE=199±5g), more numerous (x±SE=58±0.6 achenes per head), and are equipped with a pappus aiding wind transport and rows of trichomes aiding animal transport. The marginal (ray) achenes are heavier (x±SE=286±7g), less numerous (virtually invariant at 13 achenes per head), and lack dispersal structures. Whereas disk achenes are relased shortly after they mature, ray achenes are retained by the parent for a period of months following maturity.Germination at constant temperature (20°C) and with alternating light (12 h light: 12 h dark) demonstrated that disk and ray achenes exhibit different germination syndromes. Germination percentage increases linearly with achene fresh weight in both types; for a given weight, disk achenes have a higher germination percentage than ray achenes. Germination time decreases with increasing achene weight in disk achenes, but increases with achene weight in ray achenes.The germination percentages and germination times for disk and ray achenes diverge progressively with increasing achene weight. The divergence in behavior is a result of diverging patterns of dry matter allocation in the two achene types. Increase in the size of disk achenes favors the embryo fraction, thereby speeding germination while reducing protection. Increase in the size of the ray achenes favors the pericarp fraction, thereby increasing protection while delaying germination.Reduced germination percentage and germination speed of the ray achenes were shown by experimental manipulation to be caused by physical inhibition by their thicker pericarps.Dimorphism in ragwort likely speards germination out in space and time, thereby increasing the number of safe sites an individual parent can exploit in disseminating offspring. The syndrome in other heteromorphic composites resembles that of ragwort, generally combining reduced dispersal-delayed germination in the outer achenes and distance dispersal-quick germination in the central achenes. The outer achenes are generally less numerous and larger. Dispersal traits (large numbers, early release and light wieght) are the direct opposite of dormancy traits (small numbers, delayed release and heavier weight). Thus conflicts between the properties determining dormancy and dispersal appear to require separate dormancy and dispersal phenotypes.     

The sesquiterpenoids and their chemotaxonomic implications in Senecio L. (Asteraceae)

Sesquiterpenoids are important characteristic compounds in Asteraceae plants. These compounds have been proposed to be potential chemotaxonomic markers, but this application has not been comprehensively investigated. In this paper, sesquiterpenoids from 149 species of the genus Senecio were investigated to assess their taxonomic utility. The presence and absence were encoded as binary taxonomic characteristics and subsequently utilised in a clustering analysis. The previously biosynthesis pathway of sesquiterpenoids was taken into account to explore the relationship among these species contained different sesquiterpenoid types. In addition, the DNA phylogeny was also considered to explore the distribution and evolution of sesquiterpenoids in genus Senecio further. As a result, seven chemical major sections within the Senecio were recognised based on the chemical character. Although with several exceptions, the biosynthesis pathway, molecular phylogeny and the geographical origin of these sesquiterpenoid were found to be related to the chemical sections, which largely confirmed the validity of our chemo-classification for genus Senecio. In conclusion, we suggest that the sesquiterpenoid character in Senecio has some value for the genus taxonomy but should be analysed carefully and critically.     

Three new species of Senecio (Asteraceae) from Mexico

Senecio moranii, Senecio martirensis, andSenecio semperamatae, all from Mexico, are described as new.     

Taxonomic notes on Senecio tomentosus and its allies (Asteraceae)

Distribution maps, specimen citations and taxonomic commentary are provided for 14 species ofSenecio in temperate North America allied withS. tomentosus (the “Tomentosi”). These data support the author’s treatment of the group inNorth American Flora.     

Adaptation of Senecio vulgaris (Asteraceae) to ruderal and agricultural habitats

Adaptation of the annual plant Senecio vulgaris to ruderal and agricultural habitats was investigated. We expected S. vulgaris to be adapted to the agricultural habitat through nutrient-specific differentiation of relatively few genotypes responding to the generally high homogenous nutrient levels at the agricultural habitat caused by constant fertilization. To assess adaptation of S. vulgaris, vegetative and reproductive responses of seed families from various populations of agricultural and ruderal habitats, grown in the greenhouse at high and low nutrient levels, were compared. Data were analyzed with a three-level nested ANOVA based on the levels habitat, population, and family. A significant habitat effect indicated that S. vulgaris from ruderal and agricultural habitats were genetically different with plants from the agricultural habitat having larger leaves and a higher reproduction. A significant habitat by nutrient effect showed a stronger response of reproduction to nutrients at the agricultural habitat, suggesting that genetic differentiation among habitats is nutrient-specific. Contrary to expectations, only the agricultural habitat showed genetic diversity of S. vulgaris. Results suggest that nutrient levels at the agricultural habitat are more heterogeneous as generally proposed leading to a relatively high genetic variation.