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.     

Geographical dispersion of ragweed leaf beetle (Ophraella communa) based on climatic and biological characters in the Palearctic habitats

1. The accidentally introduced ragweed leaf beetle (Ophraella communa) is a most promising biological control agent for common ragweed (Ambrosia artemisiifolia), which herbivore has already emerged in several areas of the Palearctic region.
2. The aim of our study was to model the expansion of O. communa and the number of generations in the various Palearctic regions. Furthermore, our objective was to determine the effect of the prevailing wind on the direction of its spread and to ascertain the relationship between the green biomass production of ragweed and individual numbers of this leaf beetle.
3. According to our meta-analytical findings, the advancement of O. communa is continuous in the Palearctic areas. This phytophagous insect invades new habitats, which are occupied by A. artemisiifolia, and spreads quickly. The stable populations of O. communa seem to be strictly linked to the presence of its primary host, A. artemisiifolia.
4. We show that the rapid spread of this insect is due to the combination of wind direction and topography features, which was reinforced by our analysis. O. communa possesses uniformly multivoltine populations in its Palearctic habitats. So, insects possessing facultative diapause are able to colonize northern areas depending on the presence of their host, which statement is based on the processing of 143 East- and 68 West-Palearctic records
5. According to our investigations, the mass appearance of this phytophagous insect coincided with the assimilation peak of its main host, common ragweed.

     

Reevaluation of biosecurity of Ophraella communa against sunflower (Helianthus annuus)

Ophraella communa (Coleoptera: Chrysomelidae), originally from North America, has been used for biological control of common ragweed, Ambrosia artemisiifolia, in China since 2007. However, there is still a debate on whether O. communa can attack sunflowers under field conditions. To re-evaluate the biosecurity of O. communa against sunflower (Helianthus annuus), we investigated the population density of O. communa on three sunflower varieties that were intercropped with or planted in circumambience of A. artemisiifolia under field conditions. Our results showed that only very few O. communa eggs (<0.5 eggs/plant) were found on sunflower plants at the last two surveys when sunflowers were planted in circumambience of common ragweed. O. communa eggs were not found on sunflower plants at each survey when sunflowers were intercropped with common ragweed. The first–second instar larvae, third instar larvae, pupae and adults of O. communa were occasionally found on sunflower plants, but their densities were very low under either case of planting patterns. Based on these results, we conclude that sunflower is not a potential host plant for O. communa and the beetle is an effective host-specific biological control agent of common ragweed.     

Review of the biology and ecology of a ragweed leaf beetle,Ophraella communa (Coleoptera: Chrysomelidae), which is a biological control agent of an invasive common ragweed,Ambrosia artemisiifolia (Asterales: Asteraceae)

A ragweed leaf beetle, Ophraella communa (Coleoptera: Chrysomelidae), has been highlighted as a potential biological control agent of Ambrosia artemisiifolia. O. communa and A. artemisiifolia are native in North America and alien species in East Asia and Europe. As an invasive weed, A. artemisiifolia causes severe economic losses as reducing agricultural production as well as producing severe allergenic pollen. As an herbivore insect, O. communa has strong host preference on A. artemisiifolia. All the developmental stages of O. communa can be found on A. artemisiifolia and it attacks a single plant in repeated and extended manners. With few individuals on A. artemisiifolia, O. communa can completely defoliate before pollen production. Therefore, O. communa had been focused as a biological control of this invasive weed, but its introduction was denied because of possible damage on an important crop, Helianthus annuus. O. communa was accidentally introduced in East Asia and Europe in 1990s and 2010s, respectively. Fortunately, O. communa population was well established to suppress A. artemisiifolia in the introduced areas. Following detailed field surveys and host specificity tests of O. communa were conducted and proved a strong potential of O. communa as a biological control agent of A. artemisiifolia. Moreover, O. communa has been investigated in physiological and evolutionary studies. In this study, the potential of O. communa as a biological control agent and a study organism are reviewed.     

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.     

Control efficiency of leaf beetle,Ophraella communa,on the invasive common ragweed,Ambrosia artemisiifolia,at different growing stages

Ophraella communa LeSage (Coleoptera: Chrysomelidae), an effective biological control agent of A. artemisiifolia, was unintentionally introduced into China. To understand the biological control potential of O. communa on A. artemisiifolia, plant height, the number of branches and leaf control index of A. artemisiifolia injured by the beetle were determined. The results showed that plant height and number of branches decreased and leaf control index increased with the increasing initial release density of O. communa adults, when they were released at 90–100-cm plant height stage in 2008 and 60–70-cm stage in 2009. Even when O. communa was released at a low average density of 1.07 adults per plant at early growth stage of A. artemisiifolia in 2009, it could significantly suppress plant height and number of branches and led to a higher leaf control index compared with the herbicide treatment with Roundup. Prior to the fructicative period, the leaf control index of A. artemisiifolia by O. communa was up to 1.0 on the 47th day after the average initial release of 12 beetles per plant in 2008 or on the 85th day after the average release of 1.07 beetles per plant in 2009, which was significantly higher than that in the herbicide treatment. We suggest that the initial release average density of O. communa adults should be ≥1.07 beetles per plant at the early growth stage, or ≥12 beetles per plant at the late growth stage for an effective control of A. artemisiifolia in the field.     

Rapid evolution of photoperiodic response in a recently introduced insect Ophraella communa along geographic gradients

The introduced beetle Ophraella communa was first found in 1996 in Japan and has rapidly expanded its distribution to include regions that encompass a wide range of latitude and altitude and are dominated by different host‐plants. In this study, we investigated geographic variation in its photoperiodic response for the induction of reproductive diapause, with which the beetle adjusts its life cycle to local climate and host‐plant phenology. The beetle lines were collected from 18 sites in Japan. The diapause incidence under a photoperiodic condition of 13 h light : 11 h dark (LD 13:11) and the critical day length differed among the beetle lines. Analysis with the generalized linear model showed that latitude, altitude and host‐plant species (Ambrosia artemisiifolia vs. Ambrosia trifida) had significant effects on diapause incidence under LD 13:11. These results suggest that the O. communa populations have rapidly adapted to local environmental conditions after their colonization. However, the photoperiodic response of the O. communa population in Tomakomai, the northernmost part of its distribution range in Japan, deviated significantly from the general trend. We suggest that this deviation is attributed to either: (i) that this beetle has colonized Tomakomai more recently compared to the other sites; or (ii) that the Tomakomai population has adapted to local environments in a different way from other populations.     

Which role can arbuscular mycorrhizal fungi play in the facilitation of Ambrosia artemisiifolia L. invasion in France?

Ambrosia artemisiifolia L. (common ragweed), an annual invasive plant, was introduced more than 100 years ago from North America to Europe. Like the majority of other invasive plants in Europe, it develops in open, disturbed areas such as fields, wastelands, roadsides, and riverbanks. Recently, arbuscular mycorrhizal fungi (AMF) have been suspected to play a role in some plant invasion processes. As the common ragweed is known to be colonized by AMF in its native range, the intensity of mycorrhizal root colonization was studied in 35 natural populations in eastern France. About 94% of the A. artemisiifolia populations sampled were mycorrhizal. Root colonization levels varied from 1 to 40% depending on the ecological sites, with lower levels for agricultural habitats and higher levels in disturbed sites, such as wastelands or roadsides. A subsequent greenhouse experiment showed positive impacts of AMF on the growth and development of A. artemisiifolia. It is proposed that the spread of this invasive plant species could be facilitated by AMF, underlining the need to integrate symbiotic interactions in future work on invasive plant processes.     

Defoliation of common ragweed by Ophraella communa beetle does not affect pollen allergenicity in controlled conditions

Ragweed allergy is one of the primary causes of seasonal allergies in Europe and its prevalence is expected to rise. The leaf beetle Ophraella communa, recently and accidentally established in N-Italy and S-Switzerland, represents a promising approach to control ragweed, but negative side effects should be excluded before its use. Since biotic and abiotic stresses are known to influence the allergenicity of pollen, we set out to assess the effect of sub-lethal defoliation by O. communa on the quantity and quality of ragweed pollen. Seventeen sister pairs (including six clones) of ragweed plants were grown in controlled conditions. One of each pair was exposed to O. communa as soon as the plant started to produce reproductive structures. After 10 weeks of exposure, plant traits were measured as a proxy for pollen quantity. Pollen quality was assessed by measuring its viability and allergenicity. Generally, plants produced very few male flowers and little amount of pollen. Damage by the beetle was severe with most of the leaf tissue removed, but no treatment effect was found on any of the quantitative and qualitative traits assessed. In conclusion, O. communa did not increase the amount or allergenicity of ragweed pollen grains in our experimental conditions.     

Contemporary evolution of host plant range expansion in an introduced herbivorous beetle Ophraella communa

Host range expansion of herbivorous insects is a key event in ecological speciation and insect pest management. However, the mechanistic processes are relatively unknown because it is difficult to observe the ongoing host range expansion in natural population. In this study, we focused on the ongoing host range expansion in introduced populations of the ragweed leaf beetle, Ophraella communa, to estimate the evolutionary process of host plant range expansion of a herbivorous insect. In the native range of North America, O. communa does not utilize Ambrosia trifida, as a host plant, but this plant is extensively utilized in the beetle's introduced range. Larval performance and adult preference experiments demonstrated that native O. communa beetles show better survival on host plant individuals from introduced plant populations than those from native plant populations and they also oviposit on the introduced plant, but not on the native plant. Introduced O. communa beetles showed significantly higher performance on and preference for both introduced and native A. trifida plants, when compared with native O. communa. These results indicate the contemporary evolution of host plant range expansion of introduced O. communa and suggest that the evolutionary change of both the host plant and the herbivorous insect involved in the host range expansion.     

The influence of nitrogen,water and competition on the vegetative and reproductive growth of common ragweed (<Emphasis Type="Italic">Ambrosia</Emphasis><Emphasis Type="Italic">artemisiifolia</Emphasis> L.)

As common ragweed (Ambrosia artemisiifolia L.) spreads across Europe and other regions, it is becoming both a health and an economic threat. To better understand which environmental conditions facilitate the spread of the invasive species, in 2010, a greenhouse experiment was conducted determining the effects of various nitrogen levels (10, 50 and 100 kg N/ha), soil moisture level (low and high) and competition levels (no competition, medium competition and high competition) on the growth parameters of ragweed. Single-grown ragweed responded favourably to the medium nitrogen and water increase, whereas the ragweed growth parameters in competition stands increased only when high levels of nitrogen and water were added. High competition reduced the total dry matter of ragweed by up to 83%, but the ragweed continued to increase its relative growth rate during the full-flowering stage and allocate its dry matter to reproductive parts, producing up to 70 seeds per plant. Ragweed is a poor competitor when there is high resource availability; however, under disturbance and in the shortage of nutrients and water conditions, the intensity of competition decreases and the ragweed performance is minimally affected. The addition of medium levels of nitrogen to promote the growth of competitive species, prevention of disturbance and establishment of plant communities with stress-tolerant species is measures that should help to prevent the further spread of ragweed.     

An Experimental Test of Trade-Offs Associated with the Adaptation to Alternate Host Plants in the Introduced Herbivorous Beetle, <Emphasis Type="Italic">Ophraella communa</Emphasis>

The adaptation to alternate host plants of introduced herbivorous insects can be vital to agriculture due to the emergence of crop pests. Historically, it is assumed that there are trade-offs associated with the adaptation to new host plants; a generalist genotype that adapts to an alternate host is expected to have a relatively lower fitness on the ancestral host than a specialist genotype (physiological cost) or a relatively lower host-searching ability for the ancestral host plant (behavioral cost). In this study, we tested the costs of adaptation to a new host plant in the introduced herbivorous insect, Ophraella communa LeSage (Coleoptera: Chrysomelidae). In its native range (United States), O. communa feeds mostly on Ambrosia artemisiifolia L. (Asterales: Asteraceae) and cannot utilize the related species, Ambrosia trifida L. (Asterales: Asteraceae), as a host plant. On the other hand, the introduced O. communa population in Japan utilizes A. trifida extensively, and is adapting to it, both physiologically and behaviorally. We compared larval performance on the ancestral and alternate plants and adult host-searching ability between the native and introduced beetle populations. The introduced O. communa showed higher larval survival and adult feeding preference for the alternate host plant A. trifida than did the native O. communa, indicating that the introduced O. communa has rapidly adapted to the alternate host plant. However, there are no differences in either larval performance on the ancestral host A. artemisiifolia or host-searching accuracy between the native and introduced O. communa.     

Is Puccinia xanthii a suitable biological control agent of Ambrosia artemisiifolia?

Common ragweed, Ambrosia artemisiifolia, is a highly allergenic North American plant that has become invasive in some parts of Europe, Asia and Australia following its introduction to many places in the world. Some earlier works suggested that a microcyclic autoecious rust fungus, Puccinia xanthii, known to infect A. artemisiifolia in the USA only, can be considered as a potential classical biocontrol agent (BCA) of this noxious weed in Europe and elsewhere. However, an extensive field survey did not reveal the presence of either P. xanthii or any other rusts on common ragweed in 14 US states and two Canadian provinces in 2002 and 2003. Moreover, P. xanthii infecting A. artemisiifolia has never been recorded in Canada, although it is known to occur on A. trifida and Xanthium spp. there. Nevertheless, herbarium specimens collected between 1855 and 1963 in five states of the USA confirmed the presence of P. xanthii on A. artemisiifolia. It is concluded that currently P. xanthii cannot be regarded as a promising BCA of A. artemisiifolia, although it did occur on common ragweed at least a few decades ago in the USA and some forms of this rust species have already been evaluated as effective BCAs of Xanthium in Australia.     

Performance of two monophagous leaf feeding beetles (Coleoptera: Chrysomelidae) on each other's host plant: Do intrinsic factors determine host plant specialization?

There has been much recent debate on whether physiological tradeoffs in performance across hosts or ecological factors such as predation are the primary determinants of host plant specialization in plant-herbivore interactions. This paper examines the relative role of intrinsic behavioral and physiological factors in host specialization of two species of leaf-feeding beetles (Coleoptera: Chrysomelidae). Ophraella notulata and Ophraella slobodkini are sister taxa that feed exclusively on the asteraceous plants Iva frutescens and Ambrosia artemisiifolia, respectively. Ambrosia is the ancestral host plant for this pair of beetles. I performed full-sib breeding experiments in both the laboratory and the field to assess mean responses of each species to both its native host and the host of its congener, genetic variation within each species for traits associated with using each host, and tradeoffs in performance across hosts. I reared each beetle species on each host plant and measured larval consumption, survival, development time and growth. I measured only survival and growth in the field. Genetic correlations were calculated to assess tradeoffs in performance across hosts. In the laboratory experiment, larval survival of O. slobodkini on I. frutescens was lower and development time longer than on A. artemisiifolia. Survival of O. notulata on A. artemisiifolia was marginally lower than on I. frutescens while development time did not differ. There was little genetic variation among families in host use traits for either species. None of the estimates of genetic correlations were negative. The results of the field experiment support the results of the laboratory experiment. I conclude that O. notulata, the species with the derived host association, retains considerable ability to utilize the ancestral host plant, while O. slobodkini, the species witht he ancestral host association, does not show a similar ability to ut ilize the derived host. Tradeoffs in performance across hosts were not documented for either species of Ophraella. That notulata performs so well on A. artemissifolia suggests that intrinsic factors may not provide a sufficient explanation for the host specialization of this species on I. frutescens.     

Nitrogen homeostasis in a willow leaf beetle, Plagiodera versicolora, is independent of host plant quality

While foliar nitrogen (N) content of host plants depends on environmental conditions, N content of herbivorous insects may remain relatively constant due to homeostasis. However, it is unknown to what extent insects can maintain their body elemental composition against natural variation in host plant quality. The present study examined the performance and N content of a willow leaf beetle, Plagiodera versicolora Laicharting (Coleoptera: Chrysomelidae), when fed leaves of host willow, Salix eriocarpa Franchet et Savatier (Salicaceae), with varying nutritional status. Water content, toughness, and N content of willow leaves varied seasonally, and they affected performance of the leaf beetle. The leaf beetle achieved high performance when fed young leaves. On the other hand, the N content of the leaf beetle changed little, and it was independent of that of willow leaves, indicating strong N homeostasis of the leaf beetle. We discussed the function of N homeostasis in herbivorous insects in tritrophic level interactions.     

恶性入侵植物豚草的繁育系统特性

豚草(Ambrosia artemisiifolia L.)是原产于北美的一年生恶性入侵植物,目前已广泛分布在我国东北、华北、华中和华东等地,威胁农业、生态和民众健康。该种雌、雄头状花序同株,以种子繁殖,因而其有性繁殖特性对其扩散和爆发具有重要影响。运用套袋授粉和联苯胺-过氧化氢法等实验方法,对入侵江苏常熟的豚草种群的繁育系统特性进行了研究。结果表明,豚草花粉活力在开花后第4天开始出现,第8天到第10天花粉活力比较高;柱头可授性在开花第2天开始出现,第5天到第8天柱头可授性较高;同植株上同一时期开花的雌花的柱头可授性和雄花的花粉活力有5d左右的重叠期。不套袋处理(自然条件)和异株授粉处理下,豚草的结实率都比较高,分别达48.4%和44.4%,两者间无显著差异(P0.05);而同株授粉处理的结实率较低,仅3.4%,极显著低于不套袋处理和异株授粉的结实率(P0.01)但显著大于0(P0.05)。分析表明,豚草属于自交不亲和种,但又可部分自交亲和,不具有无融合生殖特性。因此,对在新分布区的零星豚草植株进行及时防除,将取得事半功倍的效果。     

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).     

Early distribution and spread of Ambrosia artemisiifolia in Central and Eastern Europe

The introduction and early spread of common ragweed (Ambrosia artemisiifolia L.) in Central and Eastern Europe were reconstructed based on a study of all herbarium specimens of common ragweed deposited at six herbaria in Austria, Hungary and Serbia. More than 450 specimens were examined and the oldest ones, collected from 1907 to 1927, were used to map the historical spread of this highly allergenic invasive weed in this region. The herbarium records back-date the first known introduction of this noxious weed to Central and Eastern Europe as well as its early spread on the Danube-Tisza Plain, and in the North-Eastern part of Hungary. The data also confirm that the introduction of common ragweed to Central and Eastern Europe took place later than its introduction to the Western part of the continent.     

Leaf traits of congeneric host plants explain differences in performance of a specialist herbivore

1. Within the host range of herbivorous insects, performance hierarchies are often correlated with relatedness to a primary host plant, as plant traits are phylogenetically conserved. Therefore, it was hypothesised that differences in herbivore performance on closely related plant species are due to resistance traits that vary in magnitude, rather than in the nature of the traits. 2. This hypothesis was tested by manipulating putative resistance traits of three congeneric thistle species (Cirsium arvense, Cirsium palustre, and Cirsium vulgare) and assessing the performance of the oligophagous, leaf‐feeding beetle, Cassida rubiginosa. Measurements were done of survival, weight gain, and development time of the beetle on its primary host, C. arvense, and two alternative hosts under low and high nutrient availability, and on shaved and unshaved leaves. 3. Survival of C. rubiginosa was strongly dependent on plant species with final mean survival rates of 47%, 16%, and 8% on C. arvense, C. palustre, and C. vulgare, respectively. Survival was primarily explained by leaf trichome densities, and to a lesser extent by specific leaf area. Leaf flavonoid concentrations did not explain differences in beetle survival, and there were no differences in beetle weight gain or development time of individuals that survived to adulthood. 4. No beetles survived on unshaved (hairy) C. vulgare plants, but manipulating leaf trichome densities of the thistle species by shaving the leaves moderated the plant‐specific resistance, and equalised the survival rates. Survival of C. rubiginosa on alternative congeneric hosts was explained by a common physical resistance trait that varied in magnitude.     

Explaining variability in the production of seed and allergenic pollen by invasive <Emphasis Type="Italic">Ambrosia artemisiifolia</Emphasis> across Europe

To better manage invasive populations, it is vital to understand the environmental drivers underlying spatial variation in demographic performance of invasive individuals and populations. The invasive common ragweed, Ambrosia artemisiifolia, has severe adverse effects on agriculture and human health, due to its vast production of seeds and allergenic pollen. Here, we identify the scale and nature of environmental factors driving individual performance of A. artemisiifolia, and assess their relative importance. We studied 39 populations across the European continent, covering different climatic and habitat conditions. We found that plant size is the most important determinant in variation of per-capita seed and pollen production. Using plant volume as a measure of individual performance, we found that the local environment (i.e. the site) is far more influential for plant volume (explaining 25% of all spatial variation) than geographic position (regional level; 8%) or the neighbouring vegetation (at the plot level; 4%). An overall model including environmental factors at all scales performed better (27%), including the weather (bigger plants in warm and wet conditions), soil type (smaller plants on soils with more sand), and highlighting the negative effects of altitude, neighbouring vegetation and bare soil. Pollen and seed densities varied more than 200-fold between sites, with highest estimates in Croatia, Romania and Hungary. Pollen densities were highest on arable fields, while highest seed densities were found along infrastructure, both significantly higher than on ruderal sites. We discuss implications of these findings for the spatial scale of management interventions against A. artemisiifolia.