Responses of invasive silverleaf nightshade (Solanum elaeagnifolium) populations to varying soil water availability

Aspects of the invasive potential of the noxious weed silverleaf nightshade (Solanum elaeagnifolium Cav.) were studied. Greenhouse experiments were conducted in order to assess several growth, production and fecundity traits of three silverleaf nightshade populations under differing levels of water availability and to correlate them with the invasive potential of each genotype. Our results showed that water availability certainly improves growth and fecundity of this serious invasive species. In ideal conditions, S. elaeagnifoilum shows a massive seed production which clearly promotes its invasiveness. However, the three populations show different phenotypic plasticity, which could explain their different adaptation and invasiveness in habitats with different water availability. Under low water availability, plants of the Etoloakarnania (ET) population allocate more than half of their total biomass to their roots (high R/S ratio), which serve as an important reproductive organ, while they maintain a relatively high seed production. These differences in phenotypic plasticity between weed populations mean that some of them (such as ET) would be more efficient in coping with changing water availability than others and therefore should be managed on high priority.     

Evolution and spread of glyphosate resistance in Conyza bonariensis in California and a comparison with closely related Conyza canadensis

Glyphosate‐resistant weeds are an increasing problem in perennial cropping systems in the Central Valley of California, USA. To elucidate the evolutionary origins and spatial spread of resistance, we investigated the geographical distribution of glyphosate resistance and the population genetic diversity and structure of Conyza bonariensis and compared the results with previously studied C. canadensis. Thirty‐five populations from orchards and vineyards across the Central Valley were sampled. Population genetic structure was assessed using microsatellite markers. Population‐level resistance was assessed in glasshouse screening of plants grown from field‐collected seed. Bayesian clustering and analyses of multilocus genotypes indicated multiple origins of resistance, as observed in C. canadensis. Pairwise F_ST analysis detected spatial spread of resistance in the south of the Central Valley, also similar to C. canadensis. The results strongly indicate that the southern valley was an environment markedly more suitable than the northern valley for resistance spread and that spread in Conyza species was driven by increased uniformity of strong selection in the southern valley, due to recent regulation on herbicides other than glyphosate. Accordingly, resistant C. canadensis individuals occurred at high frequencies only in the southern valley, but interestingly resistant C. bonariensis occurred at high frequencies throughout the valley. Expression of resistance showed varying degrees of plasticity in C. bonariensis. The lower selfing rate and substantially greater genotypic diversity in C. bonariensis, relative to C. canadensis, indicate greater evolutionary potential over shorter time periods. Interspecific hybridisation was detected, but its role in resistance evolution remains unclear.     

Fine‐tuning of a mowing regime,a method for the management of the invasive plant,Ambrosia artemisiifolia,at different population densities

Common ragweed (Ambrosia artemisiifolia) is an invasive annual plant with highly allergenic pollen. Its spread in introduced and native ranges often occurs on roadsides, where it builds stable and rapidly growing populations. The most sustainable way of controlling the population size of this species is to prevent seed production in order to deplete the soil seed bank. Populations on roadsides are submitted to regular mowing management, which can even exacerbate the situation by inducing resprouting after cutting or by accidentally spreading seeds along the road. The population density in the juvenile stages of development could play an important role in the success of cutting regimes, as it might influence the resprouting capacity of this plant. The influence of the juvenile population density and of seven cutting regimes, differing in the timing and frequency of cuts, on easily measurable reproductive traits was investigated in a glasshouse experiment. The cutting regimes had a strong influence on the reproductive success and on the phenology of the development stages of ragweed. The population density in the juvenile stages did not play a role in further phenological development, but did influence the reproductive traits. The reproduction of ragweed can be lowered by locally adapted combinations of the timing and frequency of mowing. As the optimal management option for the reduction of both the male and female flowers, the authors suggest a first cut just before the start of male flowering, followed by subsequent cuts every 3–4 weeks.     

Comparative performance of the highly invasive Mimosa invisa and its less invasive subspecies under different nutrient and light conditions

Invasive plants may be more plastic than non‐invasive plants and maintain high fitness under various environmental conditions. Previous studies mainly focused on the comparisons between invasive and native plants, and comparisons between highly invasive and less invasive exotic species are still relatively rare, especially for comparisons at the subspecies level. This study examined the effects of nutrient addition and shading on the performance of the highly invasive Mimosa invisa and its less invasive subspecies M. invisa var. inermis under either isolated or competitive conditions. Nutrient addition increased biomass and plant height and decreased root‐to‐shoot ratio (R/S). Shading decreased biomass and R/S and increased plant height. Under isolated conditions, the two invaders did not differ in R/S, plant height and plasticity of these traits in response to nutrient addition or shading, and the two invaders also did not differ in biomass production under each of the nutrient and light treatments. When the two invaders competed with each other, M. invisa outcompeted M. invisa var. inermis under high soil nutrient conditions, and the two invaders did not differ in performance under other growth conditions. Thus, only considering competition may we find out the difference between highly invasive species and their closely related, less invasive subspecies. Management of M. invisa should focus on habitats with high soil nutrient availability, in which M. invisa is more likely to dominate.     

Assessing wind and mammals as seed dispersal vectors in an invasive legume

While some plants have modified seed structures to facilitate dispersal, many lack such specialised adaptations, making their mode of dispersal unclear. This can be particularly problematic for predicting shifts in species ranges or tracking the spread of invasive plants. As an example, the seed size and shape of the invasive legume, Lespedeza cuneata, suggest that wind and attachment to animals are not important for dispersal, yet populations can spread surprising distances within a few years. Using a series of experiments conducted in the laboratory and field, we tested wind and mammal fur as mediators of seed dispersal. To test wind dispersal, seed traps were arranged radially around a patch of L. cuneata and seeds were collected following dispersal. Attachment to mammal fur was tested by fitting pelts of a deer, coyote and raccoon to artificial torsos and determining seed retention in both the field and the laboratory. Laboratory trials also examined the influence of wet versus dry conditions. Our results showed that wind direction strongly influenced dispersal distance and seeds were readily dispersed by mammal fur. The number of seeds retained was species specific, depending on fur depth and mammal size, with seed retention increasing under wet conditions. Together, these results suggest that both wind and mammal fur contribute to the movement of L. cuneata across grasslands. Consequently, both dispersal vectors should be considered when designing and implementing control strategies.     

Seed germination response to temperature for a range of international populations of Conyza canadensis

Conyza canadensis is a surface‐germinating ruderal facultative winter annual with recruitment that is highly susceptible to changes in microsite conditions. A key adaptive characteristic for a facultative winter annual species, like C. canadensis, is germination response to temperature. The objective of this study was to determine the germination response to temperature for C. canadensis seed sourced from regions around the world with differing climates and, by doing so, gain insight into the role that seed germination biology plays in the adaptiveness and weediness of facultative winter annual weeds. Seed was sourced from populations in Málaga, Spain, Hertfordshire, UK, Shiraz, Iran and southern Ontario, Canada, and grown out in a common garden under controlled conditions to produce seed for this study. These seeds were then subjected to temperatures from 6.5 to 20°C at 1.5°C increments using a thermogradient plate. Cumulative daily germination counts for 30 days were recorded. Results indicated that temperature and source location had a significant effect on germination response. Estimated base germination temperature ranges were significantly different among the populations [Ontario (8–9.5°C), Iran (9.5–11°C), Spain (12.5–14°C), UK (11–12.5°C)], as were accumulated growing degree days (GDDs; d°C) required to reach 50% germination. For three of the four populations, estimated base germination temperature range values were below those previously reported in the literature. These differences are most likely rapid evolutionary adaptations to local climate and highlight the potential C. canadensis has to be problematic as a native and invasive species.     

Site‐specific management is crucial to managing Mikania micrantha

Increasingly, weeds have been taking on global distributions. With the proliferation of invasive weeds has come the challenge of managing these species over broad geographical regions, with diverse habitats and political jurisdictions. Here, we review the management of Mikania micrantha Kunth (Asteraceae; mile‐a‐minute) throughout its invaded range, extending through most of the Pacific islands and southern and south‐east Asia. Context matters when determining the best course of action for managing M. micrantha, as it has invaded a large variety of agricultural and natural systems. In Queensland, Australia and Florida, USA, M. micrantha has been targeted in relatively successful eradication campaigns, highlighting the importance of early detection and rapid response methods, while elsewhere in its invaded range, populations are either still increasing or showing limited signs of decline. An inter‐regional approach to research and management should incorporate successful management strategies employed throughout the invaded range including, but not limited to, chemical and cultural control practices, manual and mechanical control, classical biological control using the rust fungus Puccinia spegazzinii, plant–plant competition and integrated approaches utilising two or more control methods concurrently. Additional knowledge of M. micrantha genetics is required to determine if management approaches could be fine‐tuned for particular populations. Countries bordering the Mekong River formed a network in 2011 to co‐ordinate the management of invasive species such as M. micrantha. Expanding such a collaborative approach to other regions could further reduce populations of M. micrantha and limit its spread.     

Genetic variation and structure in native and invasive Solidago canadensis populations

Solidago canadensis is native to North America, but has become a noxious invasive plant in China. We know only a little about its invasion history and the effects of introductions on its genetic composition. Here, we investigated genetic variation and structure between 15 North American and 13 Chinese populations of S. canadensis using AFLP makers. Four AFLP loci suggested relatively high genetic diversity of this weed and similar genetic variation between the invasive range and the native range. Most genetic variation was within populations across two ranges, but the Chinese range had a higher degree of among‐population variation than the North American range. Multiple tests, including Bayesian assignment, UPGMA analysis, PCoA and analysis of ‘isolation by distance’, showed that the Chinese populations originated from at least two distinct native sources and that secondary introduction or dispersal should be common in China. Also, North American populations were possibly a single genetic group. Overall, S. canadensis in China was probably founded from multiple introductions and then spread through long‐distance dispersal associated with human activities. Genetic variability in the species in the invaded range appears to have favoured establishment and spread and may well provide a challenge to successful control.     

Seed germination and viability of herbicide resistant and susceptible Chenopodium album populations after ensiling,digestion by cattle and manure storage

Chenopodium album became a problem weed in sugar beet production, due to resistance to metamitron, a key herbicide in this crop. Dispersal of the seeds from resistant biotypes may occur due to spread by wind, animals, agricultural machinery or manure. This study examined the effect of ensiling, digestion by cattle and storage in slurry and farmyard manure on the germination and viability of the seeds of one susceptible and three resistant C. album populations. After 4 weeks in a maize silo, seed viability of C. album populations was reduced drastically to 0–5%. Incubation for 24 h in the rumen followed by a post‐ruminal digestion in vitro of intact seeds only resulted in a small reduction in viability in one C. album population. Storage in a slurry cellar for 16 weeks reduced the viability of intact seeds of the C. album populations to 25–60%. Only 0–1% of the seeds remained viable after storage in a farmyard manure heap for 4 weeks. An accelerated ageing experiment showed seed persistence to be population specific and less related to seed weight. Keeping a fresh maize silo closed for at least 4 weeks and heaping farmyard manure are excellent preventive measures to limit the spread of resistant C. album seeds between fields.     

Weed seed decay in conventional and diversified cropping systems

Diversified cropping systems can have high soil microbial biomass and thus strong potential to reduce the weed seedbank through seed decay. This study, conducted in Iowa, USA, evaluated the hypothesis that weed seed decay is higher in a diversified 4‐year maize–soyabean–oat/lucerne–lucerne cropping system than in a conventional 2‐year maize–soyabean rotation. Mesh bags filled with either Setaria faberi or Abutilon theophrasti seeds and soil were buried at two depths in the maize phase of the two cropping systems and sampled over a 3‐year period. Setaria faberi seed decay was consistently greater at 2 cm than at 20 cm burial depth and was higher in the more diverse rotation than in the conventional rotation in 1 year. Abutilon theophrasti seeds decayed very little in comparison with seeds of S. faberi. Separate laboratory and field experiments confirmed differences in germination and seed decay among the seed lots evaluated each year. Fusarium, Pythium, Alternaria, Cladosporium and Trichoderma were the most abundant genera colonising seeds of both species. A glasshouse experiment determined a relationship between Pythium ultimum and S. faberi seed decay. Possible differences in seed susceptibility to decay indicate the need to evaluate weed seedbank dynamics in different cropping systems when evaluating overall population dynamics and formulating weed management strategies.     

Differences in growth and herbicide sensitivity among Cyperus esculentus clones found in Belgian maize fields

Cyperus esculentus is an invasive troublesome neophyte in many arable crops in Belgium. Applied weed control varies from field to field. One of the possible reasons for this variability might be a differential vegetative and reproductive behaviour among Belgian C. esculentus clones. In this study, growth characteristics and herbicide sensitivity of C. esculentus clones collected in Belgian maize (Zea mays) fields were evaluated. In a morphology Experiment, 25 clones were screened for growth characteristics and ability to set viable seeds under outdoor conditions. Dose – response experiments were conducted in the glasshouse to evaluate the effectiveness of two foliar‐applied herbicides (bentazon and glyphosate) and two pre‐sowing soil‐incorporated herbicides (S‐metolachlor and dimethenamid‐P) for controlling 14 C. esculentus clones. Response variables were aboveground dry biomass, tuber number, tuber dry biomass and individual tuber dry weight. Clones exhibited large differences in shoot number (up to 3.1‐fold), tuber dry biomass (up to 4.7‐fold), tuber number (up to 3.4‐fold), individual tuber dry weight (up to 4.8‐fold), inflorescence number and capacity to set viable seeds. Large interclonal differences in herbicide sensitivity (up to 8.3‐ and 4.0‐fold for aboveground dry biomass and tuber dry biomass, respectively) were observed. Contrary to foliar‐applied herbicides, soil‐incorporated herbicides were very effective and provided season‐long C. esculentus control at doses below the recommended maximum field dose. However, low doses stimulated tuber formation. Future C. esculentus management strategies should take into account differential growth characteristics and herbicide sensitivity of C. esculentus clones.     

Endophytic fungi in the invasive weed Impatiens glandulifera: a barrier to classical biological control?

The rust fungus, Puccinia komarovii var. glanduliferae, has been introduced into the UK for biological control of the invasive weed, Impatiens glandulifera (Himalayan balsam). However, establishment of the pathogen has differed across the country, which may be partly explained by variation in plant genotype. The aim of this study was to examine whether there is a further layer of phenotypic resistance, provided by indigenous foliar endophytic fungi. Culturable endophytes were isolated from a number of different balsam populations, and the commonest species were inoculated into ‘clean’ balsam plants, to test their interactions with the rust. We found that endophyte communities within balsam are low in diversity and become more dissimilar with increasing distance between populations. Three endophytes (Colletotrichum acutatum, Alternaria alternata and Cladosporium oxysporum) were common and appeared to be antagonistic to the rust, reducing pustule number and mitigating the effect of the pathogen on plant biomass. I. glandulifera thus partially conforms to the endophyte-enemy release hypothesis, in that as an introduced species, it has an impoverished endophyte complement, acquired from the local environment. However, these endophytes represent a potential barrier to effective biological control and future weed control strategies need to find strains of rust that can overcome plant genetic resistance and the overlaying phenotypic resistance, conferred by endophytes. Future classical biological control programmes of weeds must therefore take into account the fungal bodyguards that invasive species may acquire in their introduced ranges.     

Current situation with invasive Erigeron annuus (l.) Pers. (daisy fleabane) in the Republic of Macedonia

Detailed surveys of populations of Erigeron annuus (L.) Pers., an invasive plant species native to Eastern North America, were made in three locations in north‐western, northern and eastern parts of the Republic of Macedonia. The population densities were not quantified, but several stands of different sizes were found. A rapid ecological risk assessment, mainly based on knowledge of invasion histories in South‐Eastern and Central European countries, showed that this species is a serious threat to Macedonian biodiversity, particularly in the north‐western part, where monospecific stands of E. annuus were recorded. Biological invasions of E. annuus affect biodiversity worldwide through its rapid growth and high seed production, phenotypic plasticity in the native range with regard to the availability of soil nutrients and release of compounds to the soil over the period of plant growth. Consequently, invaded ecosystems suffer from significant loss in economic and cultural value.     

Germination ecology of Emex spinosa and Emex australis,invasive weeds of winter crops

Emex spinosa and Emex australis are invasive dicotyledonous weeds. The effects of various environmental factors on the germination of these weeds were investigated under laboratory and glasshouse conditions. Germination response of both species was lower at warmer temperature, and maximum germination was recorded at 20/12°C (day/night). Light stimulated germination in both species, but considerable germination also occurred under darkness. More than 80% of E. spinosa seeds germinated at pH between 6 and 9, whereas E. australis seeds germination was considerably decreased at pH 9. Emex spinosa was fairly tolerant to salinity as compared with E. australis and germination (21%) of E. spinosa occurred even at 200 mm NaCl. Both species were sensitive to osmotic stress, but E. spinosa tolerated more osmotic stress than E. australis. Temperature above 20/12°C (day/night) and low osmotic potential increased time to start germination and mean germination time (MGT), as well as decreased germination index (GI) of both species. Darkness resulted in increased MGT and decreased GI in both species when compared with 10 h photoperiod. Salt stress strongly increased time to obtain 50% germination and reduced GI of both species. In both species, an increasing burial depth decreased emergence percentage and emergence index and increased time to start emergence, although some seed emerged even at 10 cm burial depth. It was concluded that both species can germinate over a wide range of environmental conditions. However, E. australis was more sensitive under adverse environmental conditions compared with E. spinosa. This information on germination ecology may aid in developing tools and strategies for management.     

Comparative seed germination study across alien grapes (Vitis,Vitaceae) in Europe

Alien grape (Vitis, Vitaceae) species and recently described hybrids are recognised as major environmental weeds in southern Europe, based on their tendency to spread into natural and semi‐natural habitats, often behaving as invasive species. Despite a high number of agronomic, genetic and phytopathological studies on Vitis cultivars, regeneration from seed has been poorly investigated, although it plays a key role in plant distribution and evolution. To this end, in this study we exposed seeds of eight wild grapes in Europe (three species and five nothospecies, including the native Vitis vinifera), to different temperature treatments in the laboratory. Fresh seeds from all studied taxa were dormant and germinated only after cold and/or warm plus cold stratification, but showing significant differences between the species. Seeds of V. vinifera showed an intermediate complex morphophysiological dormancy, while all other tested Vitis taxa exhibited a deep complex morphophysiological dormancy. Dark conditions reduced the germination percentage in Vitis labrusca, Vitis riparia and V. vinifera and their hybrids, and alternating temperatures elicited the highest germination percentages in all populations. Our results demonstrated for the first time that Vitis hybrids are capable of regeneration by seed, even in the absence of dispersal by animals. The germination processes studied here help understand the current expansion of alien Vitis taxa in Europe outside the viticultural areas. Consequently, germination requirements contribute to a better understanding of the mechanisms underlying alien Vitis taxa establishment and invasion.     

Predicting the dynamics of local adaptation in invasive species

An invasive plant species may restrict its spread to only one type of habitat,or,after some time,may continue to spread into a different,often stressful,secondary,habitat.The question of whether evolution is required for an invasive species to spread from one habitat to another is currently hotly debated.In order for local adaptation to occur,genetic variation must be present within invasive populations.In this paper,I focus on the effect of habitat on the maintenance of genetic variation during the lag phase,the phase of population stability prior to expansion.Genetic diversity in invasive plant populations accumulates through multiple introductions,gene flow,mutation,and hybridization,but diversity is maintained by population level processes influencing effective population size(Ne).I show that when the plastic response to the environment results in little variation in reproductive output among individuals,Ne is maximized and genetic variation is maintained.Established models of plant competition show that below-ground competition reduces the variation in reproductive output,whereas competition for light increases variation in reproductive output.The same environments that maintain high Ne also reduce the opportunity for selection and minimize the response to selection,and thus the effects of the environment are synchronized to prevent genetic purges.When the primary invasion habitat supports high Ne,conditions are ripe for local adaptation to a secondary habitat,particularly if the secondary habitat has high opportunity for selection.When the primary invasion habitat supports low Ne,genetic diversity is less likely to be sufficient for local adaptation to secondary habitat to occur.     

A novel species-specific satellite DNA family in the invasive root-knot nematode <Emphasis Type="Italic">Meloidogyne mayaguensis</Emphasis> and its potential use for diagnostics

The root-knot nematode (RKN) Meloidogyne mayaguensis is considered as one of the most damaging RKN species because of its extremely wide host range. Recent surveys have shown the rapid spread of this parasite in agro-ecosytems, often making crop cultivation not viable in the heavily infested areas. Here, we report the identification, molecular cloning, genomic organisation and sequence analysis of a new satellite DNA (satDNA) family from M. mayaguensis (named pMmPet). It is comprised of two groups of A+T rich, tandemly repeated units of 174 and 180 bp, respectively. Using these sequences as targets, hybridisation and PCR experiments performed on a wide collection of 44 populations belonging to 15 RKN species showed that the pMmPet family could only be detected in the 16 M. mayaguensis populations tested. In addition, because of their repetitive nature, positive detection of pMmPet sequences was achieved in single individual nematodes. Therefore, the repeated sequence described here possesses features that make it an excellent candidate for use as a specific and extremely sensitive tool for the accurate detection and identification of this invasive pest on a routine basis. Clearly, monitoring the occurrence and spread of M. mayaguensis at the domestic and international levels are needed to avoid wholesale loss of agricultural resources in the infested regions.     

Recruitment potential from asexual and sexual reproduction in Polymeria longifolia

Sexual and asexual reproduction in the perennial Australian weed Polymeria longifolia (polymeria take-all), was assessed using glasshouse and field studies. These studies aimed at elucidating the contribution that each reproductive mechanism makes to the success of the species as a weed in cotton in Australia. Asexual reproduction, including vegetative fragment movement because of cultivation and regeneration from in situ rhizome material, was the predominant means of dispersal and recruitment of the species. Sexual reproduction had a relatively minor but significant role in the reproductive biology of P. longifolia. Seedling establishment was <5% from buried seed in both the glasshouse and field. In the field, seedlings comprised <1% of the total number of new shoots recruited at the start of two consecutive seasons, with <3% of these reaching reproductive maturity. Vegetative fragments were more likely to survive and lead to subsequent shoot production when they were large and intact. Intensive cultivation, where plants of P. longifolia are cut into small fragments, may reduce the survival of fragments moved by cultivation and reduce recruitment from existing populations. Long-term management strategies should focus on reducing the large vegetative flushes of this weed, as well as controlling the small number of seedlings recruited.     

Vegetative propagation of Solidago canadensis – do fragment size and burial depth matter?

Clonal species may benefit from human disturbance because their vegetative fragments may be distributed via soil. Solidago canadensis is an invasive rhizomatous perennial frequently found in ruderal environments. When creating new infrastructure, digging and cutting are two main factors that may influence the spread of S. canadensis into new areas. To have a better understanding of the invasive potential of S. canadensis, we investigated whether S. canadensis was able to survive and grow from stem cuttings as well as from rhizomes. Rhizomes and cuttings were collected from three populations in Eastern Norway. The rhizomes and cuttings were planted in a pot experiment to assess their vegetative ability to propagate. Rhizome fragments (5 and 10 cm long) were buried at 0.5, 10 and 30 cm depths. The cuttings were planted as 15 cm stems, with the bottom 5 cm pushed into the soil. The results showed that rhizome length did not have an effect on survival. Although some sprouting occurred at all burial depths, increasing depth had a negative effect on rhizome survival. In general, development of the cuttings was good, but there were differences between population performance and survival. These results imply that care must be taken when (i) constructing new sites, because digging and transport of soil masses may spread S. canadensis into new areas by rhizomes or cuttings, and (ii) mowing road verges and other ruderal areas to prevent the spread of stem cuttings from one area to another.     

Biology,distribution and impacts of silverleaf nightshade (Solanum elaeagnifolium Cav.)*

Silverleaf nightshade, Solanum elaeagnifolium, is listed as a noxious weed in its native range (Americas) and as an invasive alien plant in many countries across the world. Its local pattern of distribution in the EPPO region indicates that it is still in an establishment phase. S. elaeagnifolium invasiveness could be related to several biological traits: copious production of sexual and asexual propagules, its facility for long‐distance dispersion, its ability to endure considerable drought and to dominate shallow‐rooted vegetation, especially during summer dry periods, and its capacity to suppress more valuable species because it is unpalatable and toxic to cattle. Invasion of the Mediterranean semi‐arid region by S. elaeagnifolium was mainly facilitated by irrigation and animal production. Low moisture stress may favour this invasive species and sheep are an important pathway for its long‐distance dispersal. Once established, it is one of the most difficult weeds to eradicate because of its network of creeping horizontal and deep vertical roots. Its phenotypic plasticity enables it to adopt a rosette‐like growth pattern to escape control by repeated slashing. Thus, S. elaeagnifolium monitoring in the EPPO region is vital in order to contain established populations and prevent invasion.