Effects of Soil Characteristics,Allelopathy and Frugivory on Establishment of the Invasive Plant Carpobrotus edulis and a Co-Occuring Native,Malcolmia littorea

Background

The species Carpobrotus edulis, native to South Africa, is one of the major plant invaders of Mediterranean coastal ecosystems around the world. Invasion by C. edulis exerts a great impact on coastal habitats. The low number of native species in invaded communities points to the possible existence of mechanisms suppressing their germination. In this study we assessed whether soil factors, endozoochory, competition and allelopathic effects of the invader affect its own early establishment and that of the native species Malcolmia littorea. We used laboratory solutions representing different chemical composition and moisture of the soil, herbivore feeding assays to simulate seed scarification and rainwater solutions to account for the effect of differently aged C. edulis litter.

Principal Findings

We show that unlike that of the native species, germination and early growth of C. edulis was not constrained by low moisture. The establishment of C. edulis, in terms of germination and early growth, was increased by scarification of seeds following passage through the European rabbit intestines; the rabbits therefore may have potential implications for plant establishment. There was no competition between C. edulis and M. littorea. The litter of the invasive C. edulis, which remains on the soil surface for several years, releases allelopathic substances that suppress the native plant germination process and early root growth.

Conclusions

The invasive species exhibits features that likely make it a better colonizer of sand dunes than the co-occurring native species. Allelopathic effects, ability to establish in drier microsites and efficient scarification by rabbits are among the mechanisms allowing C. edulis to invade. The results help to explain the failure of removal projects that have been carried out in order to restore dunes invaded by C. edulis, and the long-lasting effects of C. edulis litter need to be taken into account in future restoration projects.     

Physiological integration increases the survival and growth of the clonal invader <Emphasis Type="Italic">Carpobrotus edulis</Emphasis>

Clonal growth seems to be a common trait for many of the most aggressive invasive plant species. However, little research has been conducted to determine the role of clonality in the successful invasion of new areas by exotic species. Carpobrotus edulis (L.) N.E. Br. is a mat-forming succulent plant, native to South Africa that is invasive in coastal dunes of Australia, New Zealand, USA and Southern Europe. Although Carpobrotus edulis is a clonal plant, there is no information on the role of clonality for the invasion by this species, therefore the objective of this study was to test whether or not physiological integration improves the performance of C. edulis invading coastal sand dunes. To do that, a 6-month field experiment was designed in which the stolon connections between the apical ramets and the C. edulis mats were severed to prevent physiological integration. This treatment was applied to ramets growing under high and low competition with the native species. Apical ramets with intact stolon connections were used as control. Integration improved the survivorship and growth of apical ramets, both in high and low competition. Connected ramets showed a more pronounced increase of clonal growth (estimated as stolon length) during the experimental period and a higher total biomass and number of ramets at the completion of the experiment. In terms of survivorship, the benefit of integration was greater under high competition. Physiological integration can therefore be considered an important factor in the invasiveness of C. edulis, both in open space and in direct competition with the native plants.     

Understanding the influence of urbanization on invasibility: <Emphasis Type="Italic">Carpobrotus edulis</Emphasis> as an exemplar

Coastal dune areas are valuable ecosystems, generally impacted by habitat destruction and invasive alien species. In this study, we assessed how human disturbance and invasion by Carpobrotus edulis impact the soils and the establishment of native flora in the north-western coastal regions of Spain. We compared soil characteristics (pH, conductivity, water content, nutrients and enzymatic activities) and native plant as well as C. edulis fitness correlates (germination and early growth) between uninvaded and invaded soils from urban and natural coastal dune areas. We found that human disturbance impacts coastal soils by increasing organic matter and water content, modifying soil nutrients and cycles, and reducing the pH in urban soils. The presence of invasive C. edulis further increases these impacts. These changes in soil characteristics allow for the establishment of the native, but ruderal, Scolymus hispanicus and non-native C. edulis, both of which are not adapted to the typically limiting conditions of coastal dunes. In some instances, the coastal dune endemic, Malcolmia littorea, showed no fitness effects in response to urbanization or the presence of C. edulis. These results suggest that human disturbed coastal areas might be more easily invaded than natural areas. More broadly, our findings of differential responses of different native species to disturbance and invasion, illustrate the need for multi-taxon approaches when assessing the impacts of invasive species.     

Plant-soil feedback as a mechanism of invasion by Carpobrotus edulis

Invasive plant species have been suggested to change the composition of the soil community in a way that results in a positive feedback for them and a negative feedback for the native plant community. Carpobrotus edulis, a species native to South Africa, is one of the most aggressive exotic species in Mediterranean Europe. Although several aspects of its invasion biology have been studied, the occurrence of plant-soil feedback has been scarcely investigated. We first checked for the existence of biotic resistance in soils from two invaded sites of Mediterranean Europe and one site in the native area. Secondly, we evaluated the effects of soil conditioning on the germination and plant growth of C. edulis and two key species of native dunes. Finally, we tested the effects of short- and long-term soil conditioning on the performance and reproductive effort of C. edulis. Our results show that at first there is a natural resistance to invasion by the soil biota. Later, biotic resistance in invaded soil is suppressed by the establishment of a soil community that enhances the growth of C. edulis and that negatively influences the growth and survival of the native plants. Long-term soil conditioning in the field resulted in shifts in the balance between vegetative growth and sexual reproduction. Long-term invasion was also reflected in high levels of endophyte colonization by chytrids in roots, although the physiological consequences of this colonization remain unknown. The results obtained illustrate a mechanism that explains how C. edulis breaks the initial biotic resistance of newly-invaded landscapes. Finally, this study highlights the importance of studying plant-soil interactions on different members of the plant community and temporal stages in order to fully understand invasion.     

Assessing the residual effects of Carpobrotus edulis invasion, implications for restoration

We examined whether the residual effects on soil caused by the invasion of Carpobrotus edulis, common iceplant, would inhibit the reestablishment of a native plant species. Carpobrotus edulis interacts both directly by suppressing the growth and establishment of other plants and indirectly by altering soil chemistry. We tested whether the residual effects of C. edulis resulted in lowered germination, survival, growth, and reproduction of Gilia millefoliata, a rare dune annual. We compared G. millefoliata planted in plots previously occupied by C. edulis to G. millefoliata planted in plots that previously had native vegetation. Each plot received three treatments: seed, transplant, and unplanted, and were censused every three weeks until senescence. Carpobrotus edulis had strong negative effects on the germination, survival, growth, and reproduction of G. millefoliata. C. edulis lowers soil pH and increases organic content due to the recalcitrance of tissue to decomposition, which may have evolved as a mechanism to facilitate recolonization and invasion.     

Responses to invasion and invader removal differ between native and exotic plant groups in a coastal dune

The spread of exotic, invasive species is a global phenomenon that is recognized as a major source of environmental change. Although many studies have addressed the effects of exotic plants on the communities they invade, few have quantified the effects of invader removal on plant communities, or considered the degree to which different plant groups vary in response to invasion and invader removal. We evaluated the effects of an exotic succulent, iceplant (Carpobrotus edulis), on a coastal dune plant community in northern California, as well as the community responses to its removal. To assess possible mechanisms by which iceplant affects other plants, we also evaluated its above- and belowground influences on the germination and growth of a dominant exotic annual grass, Bromus diandrus. We found that iceplant invasion was associated with reduced native plant cover as well as increased cover and density of some exotic plants—especially exotic annual grasses. However, iceplant removal did not necessarily lead to a reversal of these effects: removal increased the cover and density of both native and exotic species. We also found that B. diandrus grown in iceplant patches, or in soil where iceplant had been removed, had poorer germination and growth than B. diandrus grown in soil not influenced by iceplant. This suggests that the influence of iceplant on this dune plant community occurs, at least in part, due to belowground effects, and that these effects remain after iceplant has been removed. Our study demonstrates the importance of considering how exotic invasive plants affect not only native species, but also co-occurring exotic taxa. It also shows that combining observational studies with removal experiments can lead to important insights into the influence of invaders and the mechanisms of their effects.     

Effects of inoculation with native arbuscular mycorrhizal fungi on clonal growth of Potentilla reptans and Fragaria moschata (Rosaceae)

Many clonal plants live in symbiosis with ubiquitous arbuscular mycorrhizal (AM) fungi, however, little is known about their interaction with respect to clonal reproduction and resource acquisition. The effects of arbuscular mycorrhiza on the growth and intraclonal integration between ramets of two stoloniferous species were studied experimentally in a nutritionally homogenous soil environment. Two species coexisting at the same field site, Potentilla reptans and Fragaria moschata, were selected as model plants for the study. Pairs of their ramets were grown in neighbouring pots with each ramet rooted separately. Four inoculation treatments were established: (1) both mother and daughter ramets remained non-inoculated, (2) both ramets were inoculated with a mixture of three native AM fungi from the site of plant origin, (3) only mother or (4) daughter ramet was inoculated. The stolons connecting the ramets were either left intact or were disrupted. Despite the consistent increase in phosphorus concentrations in inoculated plants, a negative growth response of both plant species to inoculation with AM fungi was observed and inoculated ramets produced fewer stolons and fewer offspring ramets and had lower total shoot dry weights as compared to non-inoculated ones. A difference in the extent of the negative mycorrhizal growth response was recorded between mother and daughter ramets of P. reptans, with daughter ramets being more susceptible. Due to AM effect on ramet performance, and thereby on the source-sink relationship, inoculation also significantly influenced biomass allocation within clonal fragments. Physiological integration between mother and daughter ramets was observed when their root systems were heterogeneous in terms of AM colonization. These results hence indicate the potential of mycorrhizal fungi to impact clonal growth traits of stoloniferous plant species, with possible consequences for their population dynamics.     

Conspecific plant–soil feedback scales with population size in Lobelia siphilitica (Lobeliaceae)

Plant–soil interactions directly affect plant success in terms of establishment, survival, growth and reproduction. Negative plant–soil feedback on such traits may therefore reduce the density and abundance of plants of a given species at a given site. Furthermore, if conspecific feedback varies among population sites, it could help explain geographic variation in plant population size. We tested for among-site variation in conspecific plant–soil feedback in a greenhouse experiment using seeds and soils from 8 natural populations of Lobelia siphilitica hosting 30–330 plants. The first cohort of seeds was grown on soil collected from each native site, while the second cohort was grown on the soil conditioned by the first. Our goal was to distinguish site-specific effects mediated by biotic and/or abiotic soil properties from those inherent in seed sources. Cohort 1 plants grown from seeds produced in small populations performed better in terms of germination, growth, and survival compared to plants produced in large populations. Plant performance decreased substantially between cohorts, indicating strong negative feedback. Most importantly, the strength of negative feedback scaled linearly (i.e., was less negative) with increasing size of the native plant population, particularly for germination and survival, and was better explained by soil- rather than seed-source effects. Even with a small number of sites, our results suggest that the potential for negative plant–soil feedback varies among populations of L. siphilitica, and that small populations were more susceptible to negative feedback. Conspecific plant–soil feedback may contribute to plant population size variation within a species’ native range.     

毛竹浸提液对苦槠幼苗生长的化感效应

为探讨毛竹(Phyllostachys edulis)扩张过程中潜在的化感作用,选择苦槠(Castanopsis sclerophylla(Lindl)Schott)为研究对象。采用水浸提的方法,用毛竹茎叶、枯落物和土壤3部分浸提液浇灌苦槠幼苗,以蒸馏水处理作为对照,对比分析质量浓度分别为0.1、0.05、0.02 g/mL的3个浓度梯度浸提液处理下苦槠幼苗生长指标及各项光合生理指标的差异。结果表明,毛竹浸提液对苦槠幼苗苗高、地径和叶绿素相对含量的影响大体上呈现高浓度抑制低浓度促进的双重浓度效应。不同来源毛竹浸提液的化感效应不尽相同,土壤浸提液对苦槠幼苗生长和光合生理均呈现抑制作用,而茎叶、枯落物浸提液低浓度时为促进作用。毛竹潜在的化感作用,在其扩张过程中可能会干扰森林主要树种更新,从而对森林群落产生威胁。     

Phenotypic plasticity of invasive Carpobrotus edulis modulates tolerance against herbivores

The anthropogenic movement of species has favoured the introduction of invasive plants worldwide. Invasive plants are frequently released from their natural enemies; however, new associations with generalist herbivores may induce defence mechanisms of non-native plants. Defensive traits are often directly related to the highly competitive ability, but also to potential antagonisms and mutualisms that they can establish with soil microorganisms. Here, we examined whether the intraspecific competition and soil microorganisms influence the morphological and physiological traits of Carpobrotus edulis when is being attacked by the native generalist snail Theba pisana. To achieve this, we grew two C. edulis individuals in separate and same pots filled with live or sterile sand, and with or without T. pisana. Our results indicated that herbivory induced an increase of shoot biomass in attacked C. edulis individuals (i.e., treated donor plants), as well as in un-attacked neighbouring individuals co-growing in the same pot (i.e., untreated recipient plants). Nevertheless, intraspecific competition nor soil microorganisms did not affect the growth of C. edulis despite reduced physiological activity and damage caused by the herbivore. Overall, our findings revealed that C. edulis individuals tolerate snail attack by inducing a compensatory growth response. We conclude that phenotypic plasticity of invasive C. edulis favours tolerance against herbivores, but we also suggest that plant-plant interactions probably determine the plant growth of un-attacked neighbouring C. edulis individuals, thus favouring their invasion mechanisms.

     

Allelopathy and its coevolutionary implications between native and non‐native neighbors of invasive Cynara cardunculus L.

Invasive plants apply new selection pressures on neighbor plant species by different means including allelopathy. Recent evidence shows allelopathy functions as remarkably influential mediator for invaders to be successful in their invaded range. However, few studies have determined whether native and non‐native species co‐occurring with invaders have evolved tolerance to allelopathy. In this study, we conducted germination and growth experiments to evaluate whether co‐occurring native Juncus pallidus and non‐native Lolium rigidum species may evolve tolerance to the allelochemicals induced by Cyanara cardunculus in Australian agricultural fields. The test species were germinated and grown in pots filled with collected invaded and uninvaded rhizosphere soil of C. cardunculus with and without activated carbon (AC). Additionally, a separate experiment was done to differentiate the direct effects of AC on the test species. The soil properties showed invaded rhizosphere soils had higher total phenolic and lower pH compared with uninvaded soils. We found significant reduction of germination percentage and seedling growth in terms of above‐ and belowground biomass, and maximum plant height and root length of native in the invaded rhizosphere soil of C. cardunculus, but little effect on non‐native grass species. Even soil manipulated with AC showed no significant differences in the measured parameters of non‐native except aboveground biomass. Taken together, the results indicate allelochemicals induced by C. cardunculus exert more suppressive effects on native than non‐native linking the coevolved tolerance of those.     

Micropropagation and seed cryopreservation of the critically endangered species Tennessee yellow-eye grass, Xyris tennesseensis Kral

Xyris tennesseensis is a critically endangered species native to the southeastern USA. A micropropagation protocol was developed which may assist in the safeguarding and augmentation of dwindling natural populations of this ecologically and medically valuable plant. Four different batches of seeds were sterilized using hydrogen peroxide and germinated in vitro on modified one third-strength Murashige and Skoog medium. Shoot multiplication from seedling tissue was obtained using modified one third-strength Murashige and Skoog medium containing 1?mg/l kinetin and 0.1?C0.5?mg/l ??-naphthaleneacetic acid. Optimal shoot size and sustainable multiplication rates of three to five per 2-mo subculture occurred on medium containing 0.3?C0.4?mg/l ??-naphthaleneacetic acid. Shoots rooted successfully when placed on growth regulator-free medium for 10?d followed by transfer to greenhouse soil under high humidity. Use of seed cryopreservation resulted in significant increases in germination compared to control treatments with average germination rates of 97%. Shoot tip cultures from soil-grown plants of X. tennesseensis and Xyris spathifolia were also developed using the above protocols. Plant tissue culture tools will assist in the multiplication, long-term storage, and conservation of these rare and valuable plants as well as provide a template for the micropropagation of other Xyris species.     

Developmentally-programmed division of labour in the clonal invader Carpobrotus edulis

Invasive species are one of the main causes for the loss of global biodiversity. However, the mechanisms that explain the success of invasive species remain unsolved. Clonal growth has been pointed out as an attribute that could contribute to the invasiveness of plants, however little research has been conducted to determine the importance of clonal traits in successful invaders. One of the most interesting attributes related to clonal growth is the capacity for division of labour. In this experiment we investigated the capacity for division of labour in the aggressive invader Carpobrotus edulis, and how clonal integration can contribute to the expansion of this species. Division of labour was determined by studying the degree of morphological and physiological specialization of individual ramets to a specific activity: acquisition of soil or aboveground resources and aboveground expansion. Our results showed that there is division of labour in the clonal fragments, with older ramets increasing the biomass allocated to roots (specialization in the uptake of belowground resources) and younger ramets increasing the chlorophyll content and aboveground biomass (specialization in the uptake of aboveground resources). Physiological integration allows division of labour, and as consequence the overall performance of the clonal fragment was enhanced, with connected clonal fragments showing a higher total biomass than severed clonal fragments. Division of labour increased the aboveground growth of apical ramets of C. edulis, and therefore could contribute to an effective colonization of the surrounding area by this aggressive invader. Our study is the first exploring the role of division of labour in the expansion of an invader, and supports the idea that clonal traits could increase the invasiveness of plant species.     

Soil influence on the performance of 26 native herbaceous plants suitable for sustainable Mediterranean landscaping

Native herbaceous plants have the potential for renaturalizing and recovering derelict soils, such as urban or anthropized soils.Ecological restoration following the establishment of a native wildflower meadow should lead to a reduction in management costs and to the preservation of native plant populations. This study was aimed at determining the ecological characteristics and the cultivation needs of 26 herbaceous species native to Italy and southern Europe in order to identify their landscape potential in low-maintenance conditions. The species were selected on the basis of their adaptation to unproductive soils in semi-natural and rural areas, and on their ornamental value, including their ability to attract insects. Mono-specific plots were set up in three different soils. Seed germination, seedling emergence, flowering dynamics, and plant growth were determined. Dormancy-breaking treatments were effective in improving the germination of most species. The percentage of field establishment and biomass appeared to be affected by the physical and chemical characteristics of the soil. Soil texture slightly affected seedling emergence, whereas soil texture and the C and N levels affected plant growth, the number of flowers and the duration of flowering. Dianthus carthusianorum, Verbascum blattaria, Matricaria chamomilla and Hypochoeris radicata developed a higher biomass per plant in the soils with a low nutrient content, indicating their adaptability to infertile soils. Daucus carota, Papaver rhoeas, Verbascum sinuatum, Coleostephus myconis produced a higher biomass per plant in the most fertile soil, where they appeared to show a higher potential when competing with other species. The ecological characteristics shown by the native plants are extremely important in terms of combining seeds of different species to create and to maintain semi-natural herbaceous communities in low-maintenance landscapes.     

Contrasting effects of plant invasion on pollination of two native species with similar morphologies

Invasive plants may decrease native plant density and disrupt interactions between native plants and their pollinators. We hypothesized that invasive Solidago canadensis (Asteraceae) competes for pollination services with two confamilial species, Ixeris chinensis and Sonchus arvensis. Breeding-system studies revealed that both native species are self-incompatible. In plots with all three species we found that Solidago received the highest visitation rates. To test the hypothesis of competition for pollination in the context of reduced native density, we established 3 plots for both native species with three Solidago densities (uninvaded, 50 and 75 % invaded) and corresponding decreases in native density. We investigated the effects of varying densities of Solidago on honeybee visitation rates, number of successive visits within individual ramets, pollen-load composition on bees, and seed set. For both native species, increasing Solidago density and decreasing native density resulted in bees carrying higher ratios of Solidago pollen and in bees visiting fewer capitula prior to departing from a plant. However, for other aspects of pollination, the native species responded very differently to Solidago. With increasing Solidago and decreasing native density, Ixeris received fewer honeybee visits and produced fewer seeds, demonstrating competition for pollination, but Sonchus attracted more honeybee visits and showed a non-significant trend toward setting more seeds, suggesting facilitation. These opposing effects occurred despite similarities in the native species’ floral morphology, suggesting that the effects of invasive plants are difficult to predict. In this case the different effects may relate to Sonchus being a taller plant with larger flowers.     

Differential influence of clonal integration on morphological and growth responses to light in two invasive herbs

Background and aims

In contrast to seeds, high sensitivity of vegetative fragments to unfavourable environments may limit the expansion of clonal invasive plants. However, clonal integration promotes the establishment of propagules in less suitable habitats and may facilitate the expansion of clonal invaders into intact native communities. Here, we examine the influence of clonal integration on the morphology and growth of ramets in two invasive plants, Alternanthera philoxeroides and Phyla canescens, under varying light conditions.

Methods

In a greenhouse experiment, branches, connected ramets and severed ramets of the same mother plant were exposed under full sun and 85% shade and their morphological and growth responses were assessed.

Key results

The influence of clonal integration on the light reaction norm (connection×light interaction) of daughter ramets was species-specific. For A. philoxeroides, clonal integration evened out the light response (total biomass, leaf mass per area, and stem number, diameter and length) displayed in severed ramets, but these connection×light interactions were largely absent for P. canescens. Nevertheless, for both species, clonal integration overwhelmed light effect in promoting the growth of juvenile ramets during early development. Also, vertical growth, as an apparent shade acclimation response, was more prevalent in severed ramets than in connected ramets. Finally, unrooted branches displayed smaller organ size and slower growth than connected ramets, but the pattern of light reaction was similar, suggesting mother plants invest in daughter ramets prior to their own branches.

Conclusions

Clonal integration modifies light reaction norms of morphological and growth traits in a species-specific manner for A. philoxeroides and P. canescens, but it improves the establishment of juvenile ramets of both species in light-limiting environments by promoting their growth during early development. This factor may be partially responsible for their ability to successfully colonize native plant communities.     

Post-maturation treatment improves and synchronizes somatic embryo germination of three species of Japanese pines

Somatic embryos of three Japanese pines, Pinus thunberghii, P. densiflora, and P. armandii var. amamiana, were subjected to different post-maturation treatments to improve both germination and plant conversion frequencies. Slow desiccation of somatic embryos at high relative humidity resulted not only in a marked increment in germination frequencies but also subsequently improved plant conversion rates in all genotypes of the three species tested. Overall three species, germination and plant conversion frequency of somatic embryos was improved by more than fourfold (19?C81%) and more than fivefold (15?C77%), respectively, compared to those of untreated control. In addition, this treatment resulted in a considerable improvement of synchronization of the germination period. The time required for root emergence was synchronized over a half period (28?C14?days) compared with the control. Somatic plants were acclimatized and their growth was monitored in the field.     

Context dependency of the allelopathic effects of Lonicera maackii on seed germination

Allelopathic effects of invasive plants on native flora may be mitigated by the abiotic and biotic environment into which the allelochemicals are released. Lonicera maackii (Amur honeysuckle), an invasive plant of the eastern deciduous forest, suppresses seed germination in laboratory assays. We investigated how L. maackii leachate interacts with abiotic conditions and with the soil microbial community. First, we tested the effects of leaf extract from L. maackii on germination of the native woodland herb, Blephilia hirsuta, under different light and soil conditions. We found that germination of Blephilia hirsuta was reduced by L. maackii extract, but abiotic conditions did not interact with this effect. We also tested the effects of leaf extract on germination of five native woodland species and L. maackii placed in sterile or live soil. There was an overall suppressive effect of L. maackii extract on itself and the other five native species tested. However, L. maackii extract interacted with live soil in ways that differed with the species being tested and, in some cases, changed over time. Our results indicate that allelopathic potential of L. maackii shows context dependency with respect to soil microorganisms and native species identity but not to light conditions or soil type. Our results imply that restoration of invaded areas may require active reintroduction of species sensitive to allelopathy in live soil. Further, laboratory assays of allelopathy should consider the interaction of allelochemicals with biotic and abiotic conditions to more accurately predict the impacts of allelopathy on plant communities.     

The effect of soil legacy on competition and invasion by Acacia dealbata Link

Plant–soil feedbacks can exacerbate competition between invasive and native species, although the net effect of the interaction between soil biota and competition is likely to be species-specific. Very few studies have addressed the combined effect of soil and competition on plant performance and invasion by exotic woody species. This study explores plant growth and competition between Acacia dealbata and Pinus pinaster in three different soils—native, disturbed and invaded—in Portugal. The invasion of native P. pinaster forests by A. dealbata can be explained by the stronger competition ability of the exotic tree species. Competition is stronger in the native soil, allowing the establishment of A. dealbata in this soil and the displacement of P. pinaster. During invasion, A. dealbata changes soil conditions and establishes positive plant–soil feedbacks that promote its own germination and growth and increase P. pinaster mortality. Soil disturbance by the introduction of a different exotic species, Eucalyptus globulus, did not promote invasion by A. dealbata. We found a significant effect of soil legacy on both growth and competitive ability of the invasive A. dealbata. The ability of A. dealbata to outcompete the native P. pinaster in its own soil and the positive plant–soil feedbacks established after invasion are important mechanisms for A. dealbata invasion.     

An allelopathic investigation of the domination of the introduced invasive Conyza canadensis L.

The introduced, invasive species Conyza canadensis L. covers large areas of the sandy levees next to the River Tamiš (Serbia), forming dense microcomplexes and dominating the other herbaceous species in the ruderal phytocoenosis with its aboveground mass and abundance. In addition to this species, a further 28 plant species were found, but the abundance and cover of these was significantly lower. The allelopathic influence of the species C. canadensis was investigated through analyzing the total phenolics and phenolic acids, as the main allelochemicals, in dead and vegetative parts and the soil beneath them. Seed germination and seedling growth of the target plants (Dactylis glomerata L. and Trifolium repens L.), which grow in this community, served as a measure of the inhibitory capacity of this species. It was established that the content of total phenolics and phenolic acids (p-coumaric, ferulic, p-hydroxybenzoic, vanillic and syringic) varies, following the order: vegetative plant parts > dead plant parts > sandy soil under C. canadensis. Water leachate and soils inhibited seed germination and seedling growth of the test plants to varying degrees, following the order: vegetative parts > dead parts > sandy soil, which is directly related to the content of total phenolics and phenolic acids in them. It was concluded that the pioneer species C. canadensis plays a decisive role in the first phases of vegetation succession and the process of soil formation on the barren sandy levees, owing to the synthesis of secondary phenolic metabolites.