Bottom-up effects shift top-down community structure of the parasitoid guild that attacks Asphondylia borrichiae (Diptera: Cecidomyiidae) on Borrichia frutescens (Asteraceae)

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Genome size variation in the Artemisia arborescens complex (Asteraceae, Anthemideae) and its cultivars.

Different wild Mediterranean populations of Artemisia arborescens from diverse locations representing its geographical distribution, as well as some of its well-known cultivars and some specimens cultivated as ornamentals in gardens, streets, roads and nurseries, were analysed for genome size. Other closely related species endemic to Macaronesia, Artemisia canariensis, Artemisia argentea, and Artemisia gorgonum, were also analysed, and their nuclear DNA amount has been related to the biogeography of this group of species. Additionally, 5 populations of the closely related Artemisia absinthium were analysed to establish comparisons. Measurements acquired by flow cytometry ranged from 8.29 to 11.61 pg for 2C values. Statistically significant differences of 2C nuclear DNA amounts with respect to factors such as insularity or domestication have been detected. However, quite a low intraspecific genome size variation has been found in these species. Furthermore, the study also addressed the possible hybrid origins and possible misidentifications of some of the supposed cultivars of A. arborescens.     

Narrow hybrid zone between two subspecies of big sagebrush (Artemisia tridentata: Asteraceae)

Several species of gall-forming insects specialize on big sagebrush (Artemisia tridentata), a species that shows much clinal and subspecific variation throughout its geographic range. Two of those subspecies, basin big sagebrush (A. t. ssp. tridentata) and mountain big sagebrush (A. t. ssp. vaseyana), form a narrow hybrid zone at Salt Creek, Utah. Reciprocal transplant experiments have shown that the hybrid big sagebrush at Salt Creek are more fit than either parental subspecies, but only in the hybrid zone. Do genotype and environment influence the density and distribution of galls on big sagebrush? We counted galls on parental and hybrid big sagebrush in three reciprocal transplant gardens at Salt Creek. Gardens were in each of the two parental zones and in the hybrid zone. Transplanted seedlings came from five source populations: two parental and three hybrid populations. We identified seven kinds of gall-forming flies (Rhopalomyia midges and Eutreta fruitflies) that produced identifiable galls. Densities of galls varied among the three gardens and five source populations, and there was also a significant garden by source interaction in gall density. In general, variation in gall density among gardens (i.e., environments) was much greater than the variation among source populations (i.e., genotypes). Nevertheless, significant genotype-environment interactions were observed for five of the seven kinds of galls. Overall density of galls, mostly due to Rhopalomyia ampullaria, was greatest in the high-elevation (mountain) garden and least in the low-elevation (basin) garden. To the best of our knowledge, this is the first reciprocal transplant experiment addressing herbivore richness in a hybrid zone.     

Genetic components of heterocarpy inMicroseris hybrid B87 (Asteraceae,Lactuceae)

Microseris B87 is derived from a single hybrid specimen betweenM. pygmaea with few, weakly hairy peripheral achenes and aM. bigelovii with many, strongly hairy peripheral achenes. Offspring through the F4 and F5 generations obtained by spontaneous selfing were analyzed for the segregation of quantitative and qualitative characters relating to achene dimorphism. The phenotypic effects of two previously identified unlinked genes determining the relative number of outer achenes are characterized in partially and completely homozygous sublines. We show that two morphological markers genetically linked to one of these genes are themselves regulated by the system inducing heterocarpy. Not more than two more unlinked genes are involved in the genetic basis of the heterocarpic response. The interaction of these genes in determining the heterocarpy phenotypes is discussed in the framework of a model postulating genes for a morphogen gradient across the capitulum and genes responding to this gradient.     

Pollen competition among two species of Senecio (Asteraceae) that form a hybrid zone on Mt. Etna, Sicily

Hybridization between interfertile, sympatric or parapatric, plant species can be reduced significantly by conspecific pollen advantage (CPA), whereby conspecific pollen has an advantage over heterospecific pollen in terms of ovule fertilization. We examined CPA in two interfertile species of Senecio, S. aethnensis, and S. chrysanthemifolius (Asteraceae), which form a hybrid zone on Mt. Etna, Sicily. Individuals of both species were pollinated with pollen mixtures containing 0, 25, 50, 75, or 100% heterospecific pollen, and offspring were genotyped to determine if they were products of conspecific or heterospecific pollen fertilizing the ovules. The mean proportion of hybrid offspring produced on S. aethnensis plants was not significantly different to that expected based on the proportion of heterospecific pollen applied to the flower head. However, S. chrysanthemifolius mother plants showed moderate CPA, with the proportion of hybrid offspring significantly less than expected. Seed set or seed germination was not reduced, hence the CPA found for S. chrysanthemifolius acts before ovule fertilization. The consequences of asymmetry in CPA on the reproductive isolation of S. aethnensis are briefly discussed, along with other mechanisms that may play a role in the maintenance of the hybrid zone on Mt. Etna.     

Development of microsatellite markers for the endangered medicinal plant Launaea arborescens (Asteraceae)

• Premise of the study: Microsatellite loci were developed in Launaea arborescens, an endangered and medicinal Asteraceae species in North Africa, for further investigation of its conservation genetics. • Methods and Results: We isolated and characterized 10 polymorphic and nine monomorphic microsatellite loci from L. arborescens using the protocol of Fast Isolation by AFLP of Sequences COntaining repeats (FIASCO). For the 10 polymorphic loci, the number of alleles detected per locus varied from two to six, and the observed and expected heterozygosities ranged from 0.000 to 0.833 and 0.059 to 0.713, respectively. • Conclusions: The polymorphic markers provide a useful tool for conservation genetics studies of L. arborescens, including analysis of mating system, estimating gene flow, and identifying discrete genetic units within the species.     

The developmental floral morphology of Montrichardia arborescens (Araceae) revisited

The spadix of Montrichardia arborescens contains unisexual flowers without a perianth. The pistillate flowers are located in the basal portion of the inflorescence, and the staminate flowers are located in the apical portion. There is a narrow :zone between male flowers and female flowers consisting of atypical flowers. The portion of the atypical flowers facing the staminate zone exhibits staminate characters (stamens), and the portion facing the pistillate zone has an aborted gynoecium. The floral development of Montrichurdia is compared with that of Philodendron and a new interpretation of the morphology of atypical flowers of Montrichardia is proposed. Ontogenetic evidence supports relationships with Philodendron rather than Cercestis. 2001 The Linnean Society of London     

Narrow hybrid zone between two subspecies of big sagebrush (Artemisia tridentata: Asteraceae). II. Selection gradients and hybrid fitness

The dynamic equilibrium hypothesis proposes that hybrid zones are stabilized by a balance between dispersal and selection against hybrids. A key prediction of this hypothesis is that hybrids should have lower fitness than either parental taxon, regardless of habitat. Hybrid big sagebrush (Artemisia tridentata ssp. tridentata × ssp. vaseyana) in two narrow hybrid zones do show greatly decreased recruitment. Hybrids in one zone also show increased browsing by grasshoppers, while those in the other zone have increased numbers of aphids. Overall herbivore loads, however, are not greater on the hybrids than on the parental subspecies. Browsing by mule deer is greatest on ssp. vaseyana in both hybrid zones. Incidence of galls is also greatest on ssp. vaseyana in one hybrid zone. Moreover, browsing by Artemisia weevils is greatest on ssp. tridentata in one hybrid zone. Hybrids produce more flowers than either ssp. tridentata or ssp. vaseyana, while seed production rates of hybrids do not differ from those of the parental taxa. Finally, hybrid seeds germinate as well as those of ssp. tridentata and better than those of ssp. vaseyana. Thus, our data do not support the dynamic equilibrium hypothesis.     

Phylogenetic implications of corolla morphology in subfamily Barnadesioideae (Asteraceae)

The previously insignificant and small South American subtribe Barnadesiinae of tribe Mutisieae has been shown recently to be ancient within Asteraceae. Due to absence of a 22 kb diagnostic chloroplast inversion, plus other features, this subtribe has been elevated to subfamilial status as Barnadesioideae, now containing 90 species in nine genera. Recent cladistic analyses at the generic and specific levels based on morphology and DNA sequences (ITS and trnL intron) have revealed different relationships among taxa of the subfamily. To better understand these conflicts, we analyze specific morphological features of corollas (shape and vascularization) to develop trends (trees) in these features based on minimal structural change (i.e., morphological parsimony), and to compare these with relationships among genera derived from the two recent cladistic analyses. We define six principal types of corollas in Barnadesioideae (tubular, split, double split, ligulate, subbilabiate and bilabiate) and six principal types of corolla vascularization patterns (combinations of presence or absence of central bundles, fusion of adjacent bundles and fusion of bundles at the apex of corolla lobes). In all features we assume character states in Calyceraceae to be ancestral. In corolla shape, from tubular ancestry, we hypothesize two general evolutionary trends within the subfamily: (1) splitting of the tube, and (2) flattening of the tube. In vascularization of corollas, from an ancestral condition of one central and one lateral bundle in each corolla lobe and all traces fused at the apex, we hypothesize four basic trends: (1) gain or (2) loss of lateral vascular traces in each lobe, (3) loss of the central vein, and (4) loss of fusion of traces in lobe apices. These morphological trends allow tests of the two previous phylogenetic hypotheses by (1) counting step changes (following steps in the morphological network) in the two characters on the two competing phylogenetic trees and (2) constructing an index of morphological advancement for each genus (based on morphological trees of the two characters) and correlating these with cladistic distances within the two phylogenies. Results of both tests reveal the molecular phylogeny to be more compatible with evolutionary inferences from the two morphological features. With Fulcaldea excluded, a highly significant correlation is seen between morphological advancement and cladistic distance in the molecular phylogeny.     

Variation in floral morphology within a population of Aster hispidus var. tubulosus (Asteraceae,Astereae)

The family Asteraceae has a particular inflorescence, the capitulum, consisting of ray florets and disc florets. The ray florets function as petals that attract pollinators. Marked variation in the ray floret morphology is known in a natural population of Aster hispidus var. tubulosus (Asteraceae). We analyzed the variation and found two distinct types in the ray florets, the long tubular ray floret and the ligulate ray floret. In this species, therefore, the variation in floral morphology among capitula, each of which is the basic pollination unit, is caused by the variation in the composition of the two ray floret types among capitula. We evaluated the sources of the observed variation in the floral morphology among capitula within a population using a hierarchical analysis that separated within‐individual (i.e. among capitula within each individual) and between‐individual components of the variation. We found that the main source of the variation lay at the between‐individual level, not at the between‐capitulum level nested within individuals. This finding will provide the basic knowledge that enables future study exploring whether the between‐individual variation in floral morphology caused by the compositional variation of the ray floret types leads to differential pollination success of individual plants in species of Asteraceae.     

Narrow hybrid zone between two subspecies of big sagebrush,Artemisia tridentata (Asteraceae). III. Developmental instability

Stable hybrid zones are believed to occur because gene dispersal is opposed by selection, but the nature of this selection remains unclear. The dynamic equilibrium model postulates that hybridization disrupts coadapted gene complexes, leading to alterations in development and subsequent hybrid unfitness. Alternatively, the bounded hybrid superiority model assumes that selection is due to exogenous factors. Here, we examine the developmental instability of sagebrush across a narrow hybrid zone. The results show no difference among the parental and hybrid taxa for the majority of comparisons examined. When significant differences in developmental instability were observed, one of the parental taxa (mountain big sagebrush) was typically the least developmentally stable. Hybrids were found to be the least developmentally stable for only two of the 28 comparisons studied. Interestingly, for some comparisons, hybrids were the most developmentally stable. These results contradict predictions of the dynamic equilibrium model. The sagebrush hybrid zone does not appear to be maintained by endogenous selection, as there is little indication that development is disrupted by hybridization.     

Recent, allopatric, homoploid hybrid speciation: the origin of Senecio squalidus (Asteraceae) in the British Isles from a hybrid zone on Mount Etna, Sicily

Homoploid hybrid speciation occurs through stabilization of a hybrid segregate (or segregates) isolated by premating and/or postmating barriers from parent taxa. Theory predicts that ecological and spatial isolation are of critical importance during homoploid hybrid speciation, and all confirmed homoploid hybrid species are ecologically isolated from their parents. Until recently, such species have been identified long after they originated, and consequently it has not been possible to determine the relative importance of spatial and ecological isolation during their origin. Here we present evidence for the recent origin (within the past 300 years) of a new homoploid hybrid species, Senecio squalidus (Asteraceae), in the British Isles, following long-distance dispersal of hybrid material from a hybrid zone between S. aethnensis and S. chrysanthemifolius on Mount Etna, Sicily, Italy. Historical records show that such hybrid material from Sicily was introduced to the Oxford Botanic Garden in Britain in the early part of the 18th century and that S. squalidus began to spread from there after approximately 90 years. A survey of randomly amplified polymorphic DNA/intersimple sequence repeats (RAPD/ISSR) marker variation demonstrated that S. squalidus is a diploid hybrid derivative of S. aethnensis and S. chrysanthemifolius that grow at high and low altitudes, respectively, on Mount Etna and that form a hybrid zone at intermediate altitudes. Senecio squalidus contained 11 of 13 RAPD/ISSR markers that were recorded at high frequency in S. chrysanthemifolius but were absent or occurred at low frequency in S. aethnensis, and 10 of 13 markers for which the reverse was true. Bayesian admixture analysis showed that all individuals of S. squalidus surveyed were of mixed ancestry with relatively high mean proportions of ancestry derived from both S. chrysanthemifolius and S. aethnensis (0.644 and 0.356, respectively). We argue that long-distance isolation of hybrid material from its parents on Mount Etna would have helped favor the origin and establishment of S. squalidus in the British Isles, regardless of whether the initial hybrid material introduced to Britain was preadapted to local conditions.     

Population genetic analysis reveals a homoploid hybrid origin of Stephanomeria diegensis (Asteraceae)

Homoploid hybrid speciation has generally been viewed as a rare evolutionary phenomenon, with relatively few well-documented cases in nature. Here, we investigate the origin of Stephanomeria diegensis , a diploid flowering plant species that has been proposed to have arisen as a result of hybridization between S. exigua and S. virgata . Across the range of S. diegensis , all individuals share a common chloroplast haplotype with S. virgata while showing a greater affinity for S. exigua in terms of nuclear genetic diversity. A prinicipal coordinates analysis (PCO) based on the nuclear data revealed that S. diegensis is most similar to each parent along different axes. Moreover, a Bayesian clustering analysis as well as a hybrid index-based analysis showed evidence of mixed ancestry, with approximately two thirds of the S. diegensis nuclear genome derived from S. exigua . These results provide strong support for a homoploid hybrid origin of S. diegensis . Finally, contrary to the finding that homoploid hybrid species are typically multiply-derived, our results were most consistent with a single origin of this species.     

Origin(s) of the diploid hybrid species Helianthus deserticola (Asteraceae)

Homoploid hybrid speciation has traditionally been considered a rare event, dependent on the establishment of both a novel, balanced genotype and reproductive isolating barriers between the new species and its progenitors. However, more recent studies have shown that synthetic hybrids converge toward the chromosomal structure of natural hybrids after only a few generations, suggesting that this phenomenon may be more frequent than previously assumed. Here, the possibility that the diploid hybrid species Helianthus deserticola arose from more than one hybrid speciation event was investigated using patterns of variation from cpDNA, 18 nuclear microsatellite loci, and population interfertility. Helianthus deserticola contains cpDNA haplotypes characteristic of both parental species, is polyphyletic with one parental species based on nine microsatellite loci, and has a high degree of interfertility among populations. The data are consistent with either a single origin followed by introgression with the parental species or multiple origins. Analysis of microsatellite variation places the origin of H. deserticola between 170?000 and 63?000 years before present, making it unlikely that anthropogenic disturbances influenced its origin. Finally, the hybrid species generally has lower levels of genetic diversity but higher levels of differentiation among populations than either parental species.     

The comparative pollen morphology of genera Matricaria L. and Tripleurospemum Sch. Bip. (Asteraceae) in Turkey

Pollen morphology of four Matricaria species and 28 Tripleurospermum species was investigated with light microscopies (LM) and scanning electron microscopies (SEM). Pollen slides were prepared using Wodehouse technique. Measurements were based on 20 or more pollen grains per specimen. For SEM studies, dried pollen grains were transferred on aluminum stubs and coated with gold for 4 min in a sputter-coater. The pollen grains of Matricaria and Tripleurospermum are radially symmetric and isopolar. The pollen grains of the Matricaria are oblate-spheroidal with the polar axes 16.6–31.2 μm and the equatorial axes 18.7–23.9 μm. Tripleurospermum is oblate-spheroidal, suboblate and prolate-spheroidal with the polar axes 15.6–32.2 μm and the equatorial axes 17.7-38.5 μm. The pollen grains of Tripleurospermum are operculate and tricolporate. Matricaria is operculate and usually tricolporate or rarely syncolporate, tricolpate and tetracolporate. The pollen grain of both taxa shows echinate ornamentation. The spines are commonly conical with a broadened base and a tapered apical portion. The spine length varies between 1.8–4 μm in Tripleurospermum and 2.3–3.3 μm in Matricaria. The width of spines varies between 2.8–4.6 μm in Tripleurospermum and 2.4–3.6 μm in Matricaria. Inter-spinal area shows granulate–perforate, reticulate–perforate, rugulate–perforate ornamentations and the tectum surrounding the spine base is micro perforate. Overall exine thickness ranges from 2.8 to 4.8 μm in Tripleurospermum, 3.6 to 5.2 μm in Matricaria. Intine is thicker under pores in Tripleurospermum (0.3–0.62 μm) than in Matricaria (0.6–0.8 μm). Inter-spinal ornamentations, pollen shape and the numbers of perforations at the spin base have been observed as important morphological characters.     

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

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