High outcrossing in the annual colonizing species Ambrosia artemisiifolia (Asteraceae)

Background and Aims

Variation in mating patterns may be particularly evident in colonizing species because they commonly experience wide variation in plant density. Here, the role of density for the mating system of Ambrosia artemisiifolia (common ragweed), a wind-pollinated annual colonizing species previously reported as self-compatible, is explored.

Methods

The effect of population density on the proportion of self- and cross-fertilized seeds was examined using allozyme markers and experimental arrays conducted over two seasons in the field. Also the reproductive success of isolated plants located in diverse habitats was measured. The potential occurrence of a physiological mechanism preventing self-fertilization, i.e. self-incompatibility, following controlled self- and cross-pollinations in the glasshouse was examined.

Key Results

Outcrossing rates estimated using allozyme markers were uniformly high, regardless of the spacing between plants. However, when single plants were isolated from congeners they set few seeds. Observations of pollen-tube growth and seed set following controlled pollinations demonstrated that plants of A. artemisiifolia possess a strong self-incompatibility mechanism, contrary to earlier reports and assumptions.

Conclusions

The maintenance of high outcrossing rates in colonizing populations of A. artemisiifolia is likely to be facilitated by the prodigious production of wind-borne pollen, high seed production and extended seed dormancy.Key words: Self-incompatibility, outcrossing rate, density dependence, colonization, wind-pollination, Ambrosia artemisiifolia (ragweed), Asteraceae     

Distribution map of Ambrosia artemisiifolia L. (Asteraceae) in Italy

The spread of the invasive and allergenic Ambrosia artemisiifolia L. in Italy was analysed and mapped using distribution data from a wide range of sources. Ambrosia artemisiifolia occupies 1057 floristic quadrants which are mostly distributed in the Po plain. The distribution obtained represents the basis to implement urgent management strategies.     

Size-dependent variation of gender in high density stands of the monoecious annual,Ambrosia artemisiifolia (Asteraceae)

Summary We examined the relationship between size variability and the distribution of functional gender in stands of the monoecious, wind-pollinated annual Ambrosia artemisiifolia. Populations of 60 individuals were grown in the greenhouse at a density of 372 m^–2 and at two nutrient levels. Among the surviving plants, after self-thinning, variability in above-ground biomass and gender was higher in the high nutrient treatment. Among individuals there was a significant positive correlation between maleness and both height and biomass. Fecundity was also positively correlated with both measures of size. Based on the pattern of distribution of male and female flowers within the plant, it appears that the increase in maleness in larger plants is due to increased branching and axis elongation. These results demonstrate that competitive interactions, which lead to increased variability in biomass and fecundity, can also generate variability in gender within populations.     

Isolation of five polymorphic microsatellite loci in the invasive weed Ambrosia artemisiifolia (Asteraceae) using an enrichment protocol

We report the development of five microsatellite markers for the invasive North American native plant Ambrosia artemisiifolia (Asteraceae). An enrichment protocol was used to isolate microsatellite loci and polymorphism was explored with samples from 16 natural populations collected in the native range, across North America, and in an invaded area along the Rhône Valley in France. In these areas, we found a high level of polymorphism within population and significant heterozygote deficiencies.     

The effects of feeding damage in ragweed Ambrosia artemisiifolia (Asteraceae) on populations of Zygogramma suturalis (Coleoptera,Chrysomelidae)

Summary Field sampling indicated that the number of eggs laid by a Zygogramma suturalis female within a sampling plot correlated inversely with the mean degree of ragweed damage. Feeding on extensively damaged ragweed in the laboratory caused a drop in oviposition intensity, and a considerable proportion of females completely stopped oviposition. Feeding on slightly damaged ragweed had no significant effect on oviposition intensity. Ovipositing females preferred to feed on the intact ragweed and lay their eggs close to it. The locomotory activity of ovipositing females was significantly higher on highly damaged ragweed whereas non-ovipositing (diapausing) females and males were behaviourally indifferent to the extent of ragweed damage. Under natural conditions, ovipositing females more frequently left damaged host plants for less damaged one. If the degree of ragweed damage is high over a large area, the insects that were unable to find undamaged plants for several days oviposited less and some females entered diapause. The adaptive effect of these reactions is a decrease of population density in advance, before it might drop as a result of starvation. These results are in agreement with the second model of insect reaction to the damage-induced changes in a host plant (Edwards and Wratten 1987).     

On the efficiency of introduction of American insects feeding on the common ragweed (Ambrosia artemisiifolia L.) in the south of Russia

The explosive spread of the common ragweed induced by disturbance of the natural habitats in the postwar years created an ideal vacant niche for acclimation of North American phytophagous insects introduced to the South of Russia in the 1960–70s. The propagation of these species resulted in restoration of natural succession periodicity and promoted the subsequent acclimation of the predatory stink bug Perillus bioculatus, an extremely important agent of biological control of the Colorado potato beetle. Besides the economic effect, the studies of these introduction events were significant from the theoretical viewpoint, revealing the phenomenon of a solitary population wave (SPW) of the ragweed leaf beetle. The theoretical concept of an SPW as the key factor of efficiency of the biological control of weeds underlies the method of suppression of the common ragweed which consists in inducing SPWs by establishing local refuges for the initial buildup of the beetle population.     

Establishment of trophic associations of native leaf beetle species (Coleoptera,Chrysomelidae) with Ambrosia artemisiifolia (Asteraceae) in Primorskii Territory of Russia

The establishment of trophic associations between the native leaf beetle species and the adventive ragweed Ambrosia artemisiifolia in the territory of southern Primorskii Territory of Russia is considered. The native leaf beetles Neocrepidodera obscuritarsis (Motschulsky, 1859) and Chrysolina aurichalcea (Gebler in Mannerheim, 1825) feed on A. artemisiifolia despite their trophic preferences for native plants.     

The population genomic basis of geographic differentiation in North American common ragweed (Ambrosia artemisiifolia L.)

Common ragweed (Ambrosia artemisiifolia L.) is an invasive, wind‐pollinated plant nearly ubiquitous in disturbed sites in its eastern North American native range and present across growing portions of Europe, Africa, Asia, and Australia. Phenotypic divergence between European and native‐range populations has been described as rapid evolution. However, a recent study demonstrated major human‐mediated shifts in ragweed genetic structure before introduction to Europe and suggested that native‐range genetic structure and local adaptation might fully explain accelerated growth and other invasive characteristics of introduced populations. Genomic differentiation that potentially influenced this structure has not yet been investigated, and it remains unclear whether substantial admixture during historical disturbance of the native range contributed to the development of invasiveness in introduced European ragweed populations. To investigate fine‐scale population genetic structure across the species' native range, we characterized diallelic SNP loci via a reduced‐representation genotyping‐by‐sequencing (GBS) approach. We corroborate phylogeographic domains previously discovered using traditional sequencing methods, while demonstrating increased power to resolve weak genetic structure in this highly admixed plant species. By identifying exome polymorphisms underlying genetic differentiation, we suggest that geographic differentiation of this important invasive species has occurred more often within pathways that regulate growth and response to defense and stress, which may be associated with survival in North America's diverse climatic regions.     

Causes of differences in the distribution of the invasive plants Ambrosia artemisiifolia and Ambrosia trifida in the Yili Valley,China

Ambrosia artemisiifolia and Ambrosia trifida are two species of very harmful and invasive plants of the same genus. However, it remains unclear why A. artemisiifolia is more widely distributed than A. trifida worldwide. Distribution and abundance of these two species were surveyed and measured from 2010 to 2017 in the Yili Valley, Xinjiang, China. Soil temperature and humidity, main companion species, the biological characteristics in farmland ecotone, residential area, roadside and grassland, and water demand of the two species were determined and studied from 2017 to 2018. The area occupied by A. artemisiifolia in the Yili Valley was more extensive than that of A. trifida, while the abundance of A. artemisiifolia in grassland was less than that of A. trifida at eight years after invasion. The interspecific competitive ability of two species was stronger than those of companion species in farmland ecotone, residential, and roadside. In addition, A. trifida had greater interspecific competitive ability than other plant species in grassland. The seed size and seed weight of A. trifida were five times or eight times those of A. artemisiifolia. When comparing the changes under simulated annual precipitation of 840 mm versus 280 mm, the seed yield per m² of A. trifida decreased from 50,185 to 19, while that of A. artemisiifolia decreased from 15,579 to 530.

The differences in the distribution of the two species are mainly due to differences in interspecific competitive ability, seed size, and water dependence. The two species have stronger interspecific competitive ability than that of companion species, but A. artemisiifolia has a smaller seed size and stronger drought tolerance, which allows A. artemisiifolia to spread farther than A. trifida. The reason for wider distribution of A. trifida in grassland is that A. trifida has stronger interspecific competitive ability than A. artemisiifolia under sufficient water.     

Influence of target plant density on herbivorous insect oviposition choice: Ambrosia artemisiifolia L. (Asteraceae) and Zygogramma suturalis F. (Coleoptera: Chrysomelidae)

Zygogramma suturalis F. has been introduced to Russia for the biological control of the common ragweed Ambrosia artemisiifolia L. The functional response of Z. suturalis to the density of the target plant was studied at two levels: between plants and between 0.1 m² sampling plots. The number of Z. suturalis eggs per unit of plant weight was almost independent of plant weight, indicating that a functional response between plants is absent. However, tall and intermediate height plants received more eggs per phytomass unit than short plants. Between plots, correlation analysis showed that absolute Z. suturalis egg density is positively dependent on the ragweed density. Experiments showed that the locomotor activity of ovipositing females decreased when on the ragweed, suggesting orthokinesis. The number of leaf beetle eggs per unit of ragweed weight was negatively correlated with host plant density in May‐June, when the majority of eggs are laid, but was positively correlated in July and loosely, or not significantly, correlated in August. Seasonal changes in the functional response of a herbivorous insect to host plant population density have not been reported previously. A negative functional response of a herbivorous insect to target plant density can be a serious obstacle to the biological control of weeds.     

Multiple introductions,admixture and bridgehead invasion characterize the introduction history of Ambrosia artemisiifolia in Europe and Australia

Admixture between differentiated populations is considered to be a powerful mechanism stimulating the invasive success of some introduced species. It is generally facilitated through multiple introductions; however, the importance of admixture prior to introduction has rarely been considered. We assess the likelihood that the invasive Ambrosia artemisiifolia populations of Europe and Australia developed through multiple introductions or were sourced from a historical admixture zone within native North America. To do this, we combine large genomic and sampling data sets analysed with approximate Bayesian computation and random forest scenario evaluation to compare single and multiple invasion scenarios with pre‐ and postintroduction admixture simultaneously. We show the historical admixture zone within native North America originated before global invasion of this weed and could act as a potential source of introduced populations. We provide evidence supporting the hypothesis that the invasive populations established through multiple introductions from the native range into Europe and subsequent bridgehead invasion into Australia. We discuss the evolutionary mechanisms that could promote invasiveness and evolutionary potential of alien species from bridgehead invasions and admixed source populations.     

Realized niche shift associated with Galinsoga quadriradiata (Asteraceae) invasion in China

粗毛牛膝菊在中国的入侵与生态位漂移有关 在外来物种入侵和扩散过程中,生态位的漂移可能起到了重要作用。粗毛牛膝菊(Galinsoga quadriradiata) 在中国已造成了较为严重的入侵,占据了许多与其原产地不同的气候区。为此,本研究力图揭示粗毛牛膝菊入侵过程中的气候生态位漂移,分析其在该物种入 侵中国过程中可能发挥的作用。本研究结合粗毛牛膝菊原 产地和入侵地的分布点与气候数据, 采用Maxent模型预测了其在中国潜在的分布,并采用主成分分析的方法评估 了在入侵中国过程中粗毛牛膝菊气候生态位的漂移。模型结果显示,该物种原产地种群和入侵地种群之间只 有32.7%的生态位重叠,两个种群的生态位相似性较低(Schoener's D = 0.093, P < 0.005),这暗示了在其入侵过程中发生了生态位漂移。相比于其原产地种群,其在中国的入侵种群气候生态位的整体范围和中心都明 显地漂移向了温度更低、降水更少的区域;中国南方大部分区域属于粗毛牛膝菊的稳定适生区,而位于入侵 前沿的北方地区则存在局域适应和潜在拓殖区域。这些研究结果说明,粗毛牛膝菊在中国的入侵种群仍处于准平衡阶段,未来有可能继续向新的适生区扩散入侵,其生态位的变化有力地解释了为什么该物种在中国的入侵性强、危害范围大。     

野外条件下广聚萤叶甲对豚草的产卵选择

为探究影响广聚萤叶甲Ophraella communa LeSage选择豚草产卵的环境因素, 在南京城郊于2009年6月23日至9月1日豚草生长季节, 跟踪调查了60个豚草斑块上落卵数量动态, 运用跨栏模型分析了影响叶甲产卵以及产卵数量的因素。结果表明： （1）产卵发生随季节延长、 豚草被取食程度增大、 株高增长、 暴露环境中斑块增大而增加, 但随遮蔽环境中豚草斑块增大、 开花、 开花后季节延长、 季节延长情况下被取食程度增大、 开花后被取食程度增大等因素而降低; （2）落卵数量随季节延长、 豚草被取食程度增大、 暴露环境、 开花期被取食程度增大、 开花期株高增长等因素而增加, 但随豚草开花、 季节延长情况下被取食程度增大而减小; （3）落卵量随季节延长而增大的幅度在豚草营养生长期明显大于开花期, 在暴露环境中随季节延长而呈明显直线增大, 但在遮阴环境中先直线增加, 于10周后趋缓。研究结果说明, 影响产卵发生与落卵数量的因素不完全一致, 产卵发生与落卵数量主要受因子之间互作的影响, 而较少受到单因素的独立影响。     

Genetic differentiation in life-history traits of introduced and native common ragweed (Ambrosia artemisiifolia) populations

Introduced species represent opportunities to observe evolution over contemporary time scales, and as exotics encounter new environments, adaptive responses can occur, potentially contributing to invasion. Here, we compare 22 native North American populations and 12 introduced European populations of common ragweed (Ambrosia artemisiifolia) in five common gardens (control, herbivory, light stress, nutrient stress and drought). We found evidence for improved growth and reproduction of the introduced populations in most environments, particularly in the light stress. However, under drought conditions, the introduced plants experienced more rapid wilting and mortality than their native counterparts, evidence consistent with a life-history trade-off between rapid growth and drought tolerance. Moreover, we found parallel latitudinal clines in flowering time and correlations between fitness components and the local climate of the source populations in both ranges. Together these data provide evidence for adaptation to local environmental conditions in the native and introduced range of common ragweed.     

CO2 enrichment reduces reproductive dominance in competing stands of Ambrosia artemisiifolia (common ragweed)

Plants growing in dense stands may not equally acquire or utilize extra carbon gained in elevated CO₂. As a result, reproductive differences between dominant and subordinate plants may be altered under rising CO₂ conditions. We hypothesized that elevated CO₂ would enhance the reproductive allocation of shaded, subordinate Ambrosia artemisiifolia L. (Asteraceae) individuals more than that of light-saturated dominants. We grew stands of A. artemisiifolia at either 360 or 720 μL L⁻¹ CO₂ levels and measured the growth and reproductive responses of competing individuals. To test whether elevated CO₂ altered size and reproductive inequalities within stands, we compared stand-level coefficients of variation (CV) in height growth and final shoot, root, and reproductive organ biomasses. Elevated CO₂ enhanced biomass and reduced the CV for all aspects of plant growth, especially reproductive biomass. Allocation to reproduction was higher in the elevated CO₂ than in the ambient treatment, and this difference was more pronounced in small, rather than large plant positive relationships between the CV and total stand productivity declined under elevated CO₂, indicating that growth enhancements to smaller plants diminished the relative biomass advantages of larger plants in increasingly crowded conditions. We conclude that elevated CO₂ stimulates stand-level reproduction while CO₂-induced growth gains of subordinate A. artemisiifolia plants minimize differences in the reproductive output of small and large plants. Thus, more individuals are likely to produce greater amounts of seeds and pollen in future populations of this allergenic weed.     

Effects of mating system on adaptive potential for leaf morphology in Crepis tectorum (Asteraceae)

Background and Aims

A shift from outcrossing to selfing is thought to reduce the long-term survival of populations by decreasing the genetic variation necessary for adaptation to novel ecological conditions. However, theory also predicts an increase in adaptive potential as more of the existing variation becomes expressed as homozygous genotypes. So far, relatively few studies have examined how a transition to selfing simultaneously affects means, variances and covariances for characters that might be under stabilizing selection for a spatially varying optimum, e.g. characters describing leaf morphology.

Methods

Experimental crosses within an initially self-sterile population of Crepis tectorum were performed to produce an outbred and inbred progeny population to assess how a shift to selfing affects the adaptive potential for measures of leaf morphology, with special emphasis on the degree of leaf dissection, a major target of diversifying selection within the study species.

Key Results

Three consecutive generations of selfing had a minor impact on survival, the total number of heads produced and the mean leaf phenotype, but caused a proportional increase in the genetic (co)variance matrix for foliar characters. For the degree of leaf dissection, the lowest 50th percentile of the inbred progeny population showed a disproportionate increase in the genetic variance, consistent with the recessive nature of the weakly lobed phenotype observed in interpopulation crosses. Comparison of inbreeding response with large-scale patterns of variation indicates a potential for selection in a (recently) inbred population to drive a large evolutionary reduction in degree of leaf dissection by increasing the frequency of particular sibling lines.

Conclusions

The results point to a positive role for inbreeding in phenotypic evolution, at least during or immediately after a rapid shift in mating system.     

Dysfunction of CYC2g is responsible for the evolutionary shift from radiate to disciform flowerheads in the Chrysanthemum group (Asteraceae: Anthemideae)

Evolutionary shifts among radiate, disciform and discoid flowerheads have occurred repeatedly in a number of major lineages across the Asteraceae phylogeny; such transitions may also appear within evolutionarily young groups. Although several studies have demonstrated that CYC2 genes partake in regulating floral morphogenesis in Asteraceae, the evolution of capitulum forms within a recently diverging lineage has remained poorly understood. Here, we study the molecular regulation of the shift from a radiate to a disciform capitulum within the Chrysanthemum group. This is a recently radiating group mainly comprising two genera, Chrysanthemum and Ajania, that are phylogenetically intermingled but distinct in flowerhead morphology: Chrysanthemum spp. with radiate capitula and Ajania spp. with disciform capitula. We found that the morphogenesis of zygomorphy in the marginal floret in Ajania was disrupted soon after floral primordium emergence; CYC2g, one of the CYC2 copies that was expressed prominently in the ray floret of Chrysanthemum was not expressed in flowerheads of Ajania. Weakening the expression of ClCYC2g in Chrysanthemum lavandulifolium led to the gradual transition of a ray flower toward the disc-like form. Molecular evolutionary analyses indicated that the disciform capitulum might have evolved only once, approximately 8 Mya, arising from dysfunction of the CYC2g orthologs. A 20-nt deletion, including a putative TATA-box of the Ajania-type CYC2g promoter, appeared to inhibit the expression of the gene. Considering the divergent habitats of Chrysanthemum and Ajania, we propose that the shift from radiate to disciform capitulum must have been related to changes in pollination strategies under selective pressure.     

广聚萤叶甲和豚草卷蛾对 江西南昌豚草的联合控制作用

【背景】豚草是一种世界性的重要害草,在江西南昌地区普遍发生。广聚萤叶甲和豚草卷蛾是很有利用前景的豚草天敌,而有关这2种天敌在江西南昌地区田间对豚草的联合控制效果尚不明确。【方法】2009年6月25日分别按广聚萤叶甲0.7头.株-1和豚草卷蛾0.4头.株-1的密度在江西南昌豚草入侵地同时释放2种天敌,并于8月4日～9月13日,每隔10d调查广聚萤叶甲各虫态的数量、豚草卷蛾虫瘿数、豚草的株高和死亡株数。【结果】释放前期广聚萤叶甲和豚草卷蛾种群数量增长较快,但释放70d后,豚草植株死亡率达90.14%,天敌种群数量急剧下降;释放80d后,豚草植株死亡率达到100%。此外,在整个调查期间,天敌释放区豚草株高增长缓慢,均极显著低于对照区。【结论与意义】广聚萤叶甲和豚草卷蛾在江西南昌地区田间对豚草的联合控制效果显著,宜在江西大规模推广应用。     

Variation in cold hardiness among geographic populations of the ragweed beetle, Ophraella communa LeSage (Coleoptera: Chrysomelidae), a biological control agent of Ambrosia artemisiifolia L. (Asterales: Asteraceae), in China

The ragweed beetle, Ophraella communa, is a potential biological control agent of common ragweed that appeared in Nanjing City in 2001 and has since dispersed throughout southeast China. We compared the cold hardiness of five different O. communa populations by measuring the supercooling point (SCP), water and glycerol contents of adult beetles. All indices of cold hardiness varied significantly among the sampled populations. Male beetles from the most northerly population (Nanjing) had the lowest water content of any sampled and, although female beetles from Nanchang and Miluo had water content similar to those from Nanjing, they were still lower than those of females from Fuzhou and Wuchang. Beetles from the most southerly population (Fuzhou) had the highest SCP, although Nanchang males were not significantly different from Fuzhou males. Glycerol content yielded resolution of populations as follows: Nanjing > Wuchang = Miluo = Nanchang > Fuzhou, with beetles from Nanjing yielding twice the glycerol content of Fuzhou beetles. These results suggest that overwintering O. communa use freeze avoidance to survive winter cold and that geographically separated populations of O. communa are diverging with respect to their baseline cold hardiness in accordance with the severity of low temperatures experienced during the coldest winter months in each locality. The apparent ability of O. communa to rapidly adapt to different climatic conditions is predicted to facilitate its continued range expansion across mainland China, with consequent benefits in terms of fortuitous biological control of common ragweed.     

Development of a simple PCR-based assay for the identification of triazine resistance in the noxious plant common ragweed (Ambrosia artemisiifolia) and its applicability in higher plants

Bidirectional allele-specific PCR (Bi-PASA) was used to detect a point mutation causing triazine resistance in common ragweed (Ambrosia artemisiifolia). The external primers amplified a 278 bp standard DNA fragment in all genotypes. In the susceptible S264S genotypes, a 208 bp fragment was expected while in resistant S264G common ragweed genotypes a 109 bp band was expected. In resistant plants, both the wild and mutant type fragments were detected, indicating that the original triazine sensitive cpDNA is maintained in a heteroplasmic state in the resistant S264G genotypes. Additionally, in silico analysis confirmed the potential applicability of our diagnostic assay for other plant species. In 24 out of 74 taxa (32%), the assay could be used without any change, while in the others some of the primers should be redesigned before use.