Differences in arbuscular mycorrhizal fungal communities associated with sugar maple seedlings in and outside of invaded garlic mustard forest patches

Garlic mustard (Alliaria petiolata) is a Eurasian native that has become invasive in North America. The invasive success of A. petiolata has been partly attributed to its production of allelopathic compounds that can limit the growth of arbuscular mycorrhizal fungi (AMF). Although such effects are well known, specific effects on the richness and community composition of AMF associated with woody species have not been explored. We collected sugar maple (Acer saccharum) seedlings from eight natural forest sites in Ohio and Massachusetts, containing areas either invaded or uninvaded by A. petiolata. We measured AMF root colonization of seedlings, isolated DNA from the roots and performed PCR-TRFLP analysis to assess the richness and community composition of AMF. As expected, we found reduced AMF colonization in A. petiolata invaded patches. A. petiolata did not alter the detected TRF richness, but was associated with significant changes in the composition of AMF communities in half of the sites monitored in each region. Our results suggest that although AMF colonization was reduced in A. petiolata patches, many indigenous AMF communities include AMF that are tolerant to allelopathic effects of A. petiolata.     

Reduced competitive ability in an invasive plant

One explanation for successful plant invaders is that they evolved to be more competitive. An intuitive prediction of this Evolution of Increased Competitive Ability (EICA) hypothesis never previously tested is that invasive populations should outcompete their native ‘ancestors’ in a common environment. We tested this idea in a diallel competition experiment with Alliaria petiolata where offspring from native and invasive populations were grown alone or in all pairwise combinations. While without competition, there were no differences between native and invasive populations, native populations outperformed invasive ones when competing against each other. Our results contradict the EICA hypothesis and we conclude that it does not not hold for Alliaria petiolata. Instead, we formulate a new ERCA (Evolutionary Reduced Competitive Ability) hypothesis: if there is less competition in the invasive range and competitive ability involves traits that have a fitness cost, then selection might act against it, thereby reducing intraspecific interactions too.     

Seed survival and periodicity of seedling emergence in eight species of Cruciferae

Seeds of eight species of Cruciferae were collected, usually in each of 3 years, and mixed with the top 7-5 cm of sterilised soil confined in cylinders sunk in the ground outdoors and cultivated three times yearly. The seedlings emerging were recorded for 5 yr and the numbers of viable seeds remaining then determined. Emergence of Alliaria petiolata was almost entirely restricted to February and March. That of Erysimum cheiranthoides, Lepidium campestre, Sinapis arvensis and Raphanus raphanistrum took place mainly in March and April, but seedlings continued to appear until late autumn. Some seedlings of Arabidopsis thaliana, Cardamine hirsuta and Sisymbrium officinale appeared in spring, but most did so in summer (C. hirsuta and S. officinale) or early autumn (A. thaliana). Seed survival in A. petiolata was of short duration and there were few seedlings after the second year, whereas 18% of the seeds of R. raphanistrum were still present and viable after 5 yr. Seed survival at this time for the other six species ranged from 1 -6% (C. hirsuta) to 6-1% (S. arvensis).     

Pre-release impact assessment of two stem-boring weevils proposed as biological control agents for Alliaria petiolata

Experimental studies can be useful tools to test plant responses to herbivory and to quantify the impact of potential biological control agents prior to their release. We evaluated the per-capita effect of Ceutorhynchus alliariae and C. roberti, two stem-boring weevils currently being investigated as potential biological control agents for garlic mustard, Alliaria petiolata, in North America. Weevils were released at three different densities in individual and mixed-species treatments onto potted plants of A. petiolata. Damage by C. roberti alone and by both weevils combined caused an increase in the numbers of inflorescences produced per plant. Although plants could compensate for low levels of damage, moderate to high levels of damage by both C. alliariae and C. roberti, individually and in combination, caused a decrease in plant height and a reduction in seed output per plant. The damage inflicted by both weevil species is similar so the overall impact of both species combined can be predicted by summing the impact of each species alone. Provided they are sufficiently host specific, both weevils could be released as biocontrol agents. Because reduced seed production is necessary to suppress A. petiolata populations, both species have the potential to contribute to control of A. petiolata in North America.     

Recovery of aSasa tsuboiana population after mass flowering and death

The recovery process of aSasa tsuboiana population after a mass flowering and death in 1977 was investigated by 15 years of observation in the Hira Mountains, Kinki district, western Japan. Seed production was high (6600–13 800 seeds m⁻² inSasa plots and 3900 seeds m⁻² in a forest plot) but emergent seedling density was low (14–21 seedlings m⁻²), probably because of seed predation byMicrotus montebelli occurring between seed shedding and the next spring. The seedling density had decreased further by the next year and theS. tsuboiana population recovered from only a limited number of seedlings. In spite of such a low initial density, theS. tsuboiana population was able to regenerate successfully and attained the previous full stand height in 7–16 years.Miscantbus sinensis invaded and delayed the recovery ofS. tsuboiana in one plot, butS. tsuboiana became dominant as it caught up with the height ofM. sinensis. Seedling growth patterns, such as frequent tillering, the onset of rhizome extension in the early stage of seedling growth and frequent culm production from rhizomes, played important roles in the successful regeneration ofS. tsuboiana.     

Exotic consumers interact with exotic plants to mediate native plant survival in a Midwestern forest herb layer

Consumer-facilitated invasions have been proposed as an alternative mechanism to direct competitive exclusion to explain the replacement of native plants by exotics. In a factorial field experiment manipulating competition from the exotic plant Alliaria petiolata and herbivory by exotic mollusks, we documented that mollusk herbivory significantly reduced the survival of two species of native palatable plants, but found minimal direct herbivore effects on less palatable species, including the invasive A. petiolata. These effects were evident after one growing season on younger juvenile plants of Aster cordifolius, but only after two growing seasons on older transplants of the same species, suggesting a greater vulnerability of young plants. In contrast to our expectations, A. petiolata competition alone had no effect on any of the six native species we tested. However, competition from A. petiolata did affect the survival of the most palatable native plant when mollusks were also present. While not significant for any other single species, this same pattern was observed for three of the five remaining native species tested. The selective grazing on palatable plants that we document provides novel evidence contributing to our understanding of observed shifts in the forest herbaceous layer towards the dominance of exotic plants and unpalatable species. More broadly, our results highlight the importance of the interactive effect of consumers and inter-specific competition in forest understories via its contribution to differential survival among regenerating species.     

Population stage structure,survival and recruitment in the endangered East African forest herb <Emphasis Type="Italic">Saintpaulia</Emphasis>

Climatic seasonality, local habitat quality, and edge effects created by forest fragmentation due to human activity may affect the performance of endangered rain forest understory herbs. Viability and seasonal dynamics of the populations of the endangered Saintpaulia confusa, S. difficilis, and S. grotei were studied in a protected seasonal submontane forest in NE Tanzania by examining plant life-history traits, population stage structure, and the effects of habitat quality on plant performance. The population stage structures were of the dynamic type. There was a lower frequency of seedlings (57.9%) and higher frequency of juvenile (13.3%) and adult plants (28.8%) in S. confusa than in S. difficilis (74%, 10.7% and 15.3%, respectively). Seedling recruitment occurred from May to August with an average of 54, 103 and 38 emerged seedlings per 1 m² study plot (S. confusa, S. difficilis, and S. grotei, respectively). Presence of the seed bank was also an indication of the regeneration potential of the populations. Mortality was high during the dry and hot season from December to March. Survival was lowest in juvenile plants, higher in sterile adult plants and the highest in fertile adult plants. Because survival was the lowest on dry substrates under open canopy, our data suggest that forest fragmentation, by reducing shade and humidity, will increase mortality in Saintpaulia. Furthermore, since the mortality was highest in seedlings and juveniles, forest fragmentation is likely to impede the regeneration of the Saintpaulia populations.     

How novel are the chemical weapons of garlic mustard in North American forest understories?

The Novel Weapons Hypothesis predicts that invasive plants excel in their new ranges because they produce novel metabolites to which native species possess little resistance. We examined the novelty of the phytochemistry of the Eurasian invader, Alliaria petiolata, in North America by comparing its phytochemical profile with those of closely related Brassicaceae native to North America. We examined the profile and/or concentrations of glucosinolates, alliarinoside, flavonoids, cyanide, and trypsin inhibitors in cauline leaves of field-collected A. petiolata, Arabis laevigata, Cardamine concatenata, C. bulbosa, and C. douglassii. Cyanide and the glucosinolates and flavonoids produced by A. petiolata were detected only in A. petiolata. Trypsin inhibitor activity was highest in A. laevigata, intermediate in the Cardamine species, and lowest in A. petiolata. The phytochemical profile of A. petiolata was distinct from those of four closely related and/or abundant Brassicaceaeous species native to North America, providing support for the Novel Weapons Hypothesis.     

Intraspecific variation in allelochemistry determines an invasive species’ impact on soil microbial communities

Invasive species can benefit from altered species interactions in their new range, and by interfering with species interactions among native competitors. Since exotic invasions are generally studied at the species level, relatively little is known about intraspecific variation in the traits that determine an invader’s effect on native species. Alliaria petiolata is a widespread and aggressive invader of forest understories that succeeds in part by interfering with mutualistic interactions between native plants and soil fungi. Here, I show that the impact of A. petiolata on soil microbial communities varied among individuals due to variation in their allelochemical concentrations. The differential impacts translated into varied effects on native tree growth, partly because A. petiolata’s allelochemicals preferentially affected the most mutualistic fungal taxa. These results highlight the importance of considering the spatial and temporal variation in an invasive species’ impacts for understanding and managing the invasion process.     

Aspects of the Ecology of an Invasive Plant, Garlic Mustard (Alliaria petiolata), in Central Illinois

Alliaria petiolata (garlic mustard), an exotic plant species, has invaded woodlands in several areas in mid-western and northeastern United States and adjacent Canada, and it is displacing the indigenous under-story flora. This study was conducted to provide information about the species' biology that might be useful in controlling its spread in native woodlands. The plant is a strict biennial in North America, spending the first year of growth as a basal rosette. This period of relatively slow growth is followed by a period of rapid shoot elongation (1.9 cm/day) during early spring of the second growing season. The plant is capable of cross- and self-pollination, although pollination and stigma receptivity occur before the flower is open, so autogamy is the most likely breeding system. Garlic mustard invests 20.4% of its biomass in reproductive effort, with an annual seed rain of 15,000 seeds/m². Seed dispersal from fruits begins in early July and continues into October. Most seeds germinate in the spring following the year in which they were produced. Seedling recruitment is high (8.3–18.0 seedlings/cm²), but only about 7.5% of the plants survive to maturity. The success of the plant in invading woodlands appears to be related to (1) its autogamous breeding system that allows a single individual, or a few individuals, to establish populations of genetically similar but interfertile individuals; (2) high seed production, permitting establishment of large numbers of individuals; and (3) rapid growth during the second growing season, which increases its competitive ability. Because of garlic mustard's ability to occupy understory habitats successfully, it may be unrealistic to expect to eliminate the plant from many habitats it has already invaded.     

PCR–RFLP assays for discerning three weevil stem feeders (Ceutorhynchus spp.) (Col.: Curculionidae) on garlic mustard (Alliaria petiolata)

Polymerase chain reaction–restriction fragment length polymorphism (PCR–RFLP) assays were developed to rapidly and cost-effectively differentiate the morphologically indistinguishable larval stages of three species, Ceutorhynchus alliariae, C. roberti and C. scrobicollis, proposed for the biological control of Alliaria petiolata. A PCR–RFLP using TaqαI restriction enzyme can differentiate the three target species only, while a PCR–RFLP double digestion using the restriction enzymes AluI and TaqαI should enable, based on virtual digestion, distinguishing them from 12 other Ceutorhynchus species associated with A. petiolata.     

Impact of simulated herbivory on Alliaria petiolata survival, growth, and reproduction

In weed biological control, insect damage to target weeds can be simulated in invaded habitats to study potential responses of the plant to introduced natural enemies. In the present study, we investigated the impact of two levels of manual flower-shoot damage (shoots cut at tip or base) on Alliaria petiolata (garlic mustard) survival, size, and reproduction. Experiments were conducted in 2002 and 2003 using invasive field populations of A. petiolata under naturally varying plant densities. Plant survival was recorded, and size and reproduction parameters were measured. Manual flower-shoot damage had a significant effect on plant survival. In both years, fewer plants survived in the basal-cut treatment than in either the control (un-cut) or tip-cut treatment. Plant size and reproductive output were likewise reduced in the basal-cut treatment. In both years, total seed production was significantly lower in the basal-cut treatment than either the control or tip-cut treatment. When combined, increased mortality and reduced seed production of basal-cut plants greatly reduced the contribution these plants made to the seed bank. Plant density did not affect reproduction or plant size. The impacts of cutting were consistent across years and sites with distinct biotic and abiotic conditions, and A. petiolata densities. We anticipate that herbivore damage to A. petiolata populations by introduced biological control agents will likewise remain consistent under varying biotic and abiotic conditions if the agents are equally adapted to these.     

Flexible N uptake and assimilation mechanisms may assist biological invasion by Alliaria petiolata

Although nitrogen has historically limited terrestrial plant productivity in the northern hemisphere, accelerated industrial activity is changing the availability of N, with consequences for ecosystem properties including altered susceptibility to biological invasion. Alliaria petiolata (Bieb.) Cavara & Grande is an increasingly problematic invader in forests of eastern North America. Population growth rate of this species is especially high in N-rich habitats, and it produces a variety of N-based compounds that have been shown to interfere with the growth and reproduction of native plants. To investigate how increases and shifts in forms of N will impact A. petiolata, seedlings were transplanted to the greenhouse from the field and grown in sand culture. We applied three concentrations of N (0.25, 1 and 2 mM) using five different ratios of NH₄ ⁺ and NO₃ ⁻ (100/0, 75/25, 50/50, 25/75, 0/100) in a crossed design to yield fifteen different treatments. Plants were measured throughout the growing season and a final harvest yielded measures of biomass and tissue quality. Plant growth increased significantly in response to increased concentration of total N. These increases were similar for all combinations of N. This flexibility in uptake ability may facilitate the invasion of this species, not only by increasing the range of habitats A. petiolata can occupy but also by enhancing N uptake that can lead to the production of secondary compounds disrupting other species’ belowground mutualisms. We suggest that this species’ ability to respond rapidly to changes in N availability, regardless of its form, may modify competitive interactions with natives and intensify its negative impacts.     

Conditions for seedling establishment and probable function of seed in the clonal sedge Carex rugulosa Kük. in riverside marshes

Carex rugulosa Kük. forms large meadows in moist zones along estuarine riversides. These meadows are usually maintained by rhizomatous ramet production (clonal growth), and the appearance of seedlings is uncommon. We studied the conditions required for seedling establishment. In areas in which clonal ramets were dense (906–1050/m²), and the relative light intensity at ground level was low (0.8–3.8%), seed germination was entirely suppressed. In contrast, many seedlings (288–513/m²) were observed in areas in which clonal ramets had been considerably reduced (13–269/m²) by continuous inundation in the previous year. In these areas, the relative light intensity had increased (20.5–79.3%). It was inferred that seeds resisted the prolonged inundation that killed the ramets, and that germination was induced under these open conditions. These results suggest that the main functions of sexual reproduction in C. rugulosa are recovery after severe damage to vegetation using seeds from the seed bank and the dispersal of seeds to areas without dense vegetation. Most seedlings died when the marshy meadow was temporarily inundated by heavy rain. However, transplanted seedlings survived better at artificially elevated sites that were not submerged. This shows that inundation during the seedling stage impedes seedling survival. Therefore, seeds can contribute to recovery after ramet decline or colonization to open area when (i) the rainfall is not heavy to cause seedling submergence or (ii) seeds are dispersed to higher area which is not inundated.     

Biological control of the invasive alien weed Solidago canadensis: combining an indigenous fungal isolate of Sclerotium rolfsii SC64 with mechanical control

Solidago canadensis L. is a major invasive weed that is highly tolerant to disturbances and difficult to control in China. In order to develop a rapid non-chemical control strategy for this weed in heterogeneous environments, we investigated different treatments including mechanical control (cutting and hoeing) and inoculation with an indigenous pathogen, Sclerotium rolfsii SC64, which was isolated from S. canadensis and applied by means of a solid formulation. Greenhouse and field trials were conducted to test how the control regimes (i.e. individual treatment methods, combination of these methods and different treatment timing) influence control efficiency. The fungal isolate S. rolfsii SC64 caused 70% plant mortality and fresh weight reduction of S. canadensis under 150 cm growth stage, and efficacy increased to 80% when the above-ground material was removed. However, the use of cutting, hoeing or treating with S. rolfsii SC64, on its own, did not provide sufficient control of S. canadensis. Cutting treatments performed in July and September only eliminated sexual reproduction of S. canadensis. Combination of cutting, hoeing and treating with isolate SC64 during the growing season in May, July and September was able to kill more than 90% of the ramets. This combination of methods not only eliminated sexual reproduction of S. canadensis, but also destroyed its underground stems and prevented its regrowth. Therefore, this integrated approach may provide an optima control strategy for S. canadensis.     

Impact of the belowground herbivore and potential biological control agent, Ceutorhynchus scrobicollis, on Alliaria petiolata performance

We studied the influence of the root-crown weevil Ceutorhynchus scrobicollis on its host plant Alliaria petiolata, a European biennial herb that is currently invading much of temperate North America. Varying timing of attack and herbivore densities in a common garden allowed to assess seasonality of plant response, density-dependence of impact, and the effect of intraspecific competition on C. scrobicollis recruitment (number of F₁ generation adults emerged). Data collected in the common garden were compared with data collected at field sites. C. scrobicollis is a common weevil in Europe, frequently attaining high attack levels on its host plant. In the common garden, weevil attack decreased plant survival by up to 43%, reduced plant height by 54%, increased the number of shoots by up to four–fold and delayed seed ripening, but had no significant negative effect on seed production. Plants infested in spring allocated less biomass to aboveground plant parts, and remained smaller than plants attacked in autumn, indicating that the latter were able to partly compensate for weevil attack. Increasing weevil density rarely had an effect on A. petiolata performance, and did not increase F₁ recruitment, suggesting strong intraspecific competition. At field sites, C. scrobicollis attack is spread over a long time period, which probably alleviates intraspecific competition. In summary, attack by the root-crown feeding weevil, C. scrobicollis, can substantially reduce growth and survival of A. petiolata. If introduced as a biological control agent into North America, C. scrobicollis is likely to decrease the fitness and competitive superiority of A. petiolata.     

Soil seed densities and emergence patterns in pastures in the seasonally dry tropics of northeastern Australia

Germinable seed densities in the surface (0–10 cm) soil of pasture communities growing at Lansdown, near Townsville, were measured during the late dry season before the first germinating rain and again during the following wet season after germination but before the input of new seed. Seedlings emerging in the field were counted at approximately weekly intervals during this period to determine emergence patterns. Twelve communities were sampled in 1980–81 and six were re-sampled in 1981–82. During the late dry season germinable seed densities ranged from 5000 to 40 000 seeds m^-2. Seeds of the introduced legume, Stylosanthes hamata, were present in all pastures. There were many seeds of annual grasses (Digitaria ciliaris and Brachiaria miliiformis) and sedges (Cyperus and Fimbristylis spp.) but only few seeds of perennial grasses (both native and introduced). Soil seed densities were much lower during the wet season than during the preceding dry season, particularly for the grasses. Emergence commenced and approximately 70% of all seedlings emerged on the first major rainfall of the wet season. The subsequent emergence pattern varied between years. In 1980–81 there was a gradual and continuous increase in seedling numbers under the continuously moist conditions which prevailed. In 1981–82 further emergence occurred in discrete events related to rainfall and intervening dry periods. Maximum seedling densities exceeded 34 000 seedlings m^-2 including 29 000 grass seedlings (mainly annual species). The implications of these results for species survival and pasture composition are discussed.     

Observations of herbivore attack on garlic mustard (Alliaria petiolata) in Southwestern Ontario, Canada

We have observed what appears to be the first significant report of herbivory of a population of Alliaria petiolata (M. Bieb.) Cavara & Grande (Garlic Mustard) in Canada. Ceutorhynchus erysimi Fabr., Plutella xylostella L. (Diamondback Moth), and Philaenus spumarius (Meadow Spittlebug) were identified attacking, and successfully proceeding through different stages of development on A. petiolata. From the field observations, we hypothesize that these species have recognized A. petiolata as a host and therefore may be candidates for the management of the invasive species.     

The effects of arbuscular mycorrhizal (AM) fungal and garlic mustard introductions on native AM fungal diversity

Introduced, non-native organisms are of global concern, because biological invasions can negatively affect local communities. Arbuscular mycorrhizal (AM) fungal communities have not been well studied in this context. AM fungi are abundant in most soils, forming symbiotic root-associations with many plant species. Commercial AM fungal inocula are increasingly spread worldwide, because of potentially beneficial effects on plant growth. In contrast, some invasive plant species, such as the non-mycorrhizal Alliaria petiolata, can negatively influence AM fungi. In a greenhouse study we examined changes in the structure of a local Canadian AM fungal community in response to inoculation by foreign AM fungi and the manipulated presence/absence of A. petiolata. We expected A. petiolata to have a stronger effect on the local AM fungal community than the addition of foreign AM fungal isolates. Molecular analyses indicated that inoculated foreign AM fungi successfully established and decreased molecular diversity of the local AM fungal community in host roots. A. petiolata did not affect molecular diversity, but reduced AM fungal growth in the greenhouse study and in a in vitro assay. Our findings suggest that both introduced plants and exotic AM fungi can have negative impacts on local AM fungi.     

Genets of dwarf bamboo do not die after one flowering event: evidence from genetic structure and flowering pattern

Dwarf bamboos in the genus Sasa are believed to be long-lived, synchronously flowering, and monocarpic plants. However, the monocarpy of dwarf bamboo has not been confirmed, because whether all ramets within one genet flower at the same time cannot be determined without differentiating the genetic structure among ramets. This study aims to evaluate the reproductive traits of Sasa pubiculmis by verifying the monocarpy and physiological integration between flowering ramets and non-flowering ramets during a 4-year flowering period. One genotypically identified genet, which covered an area of approximately 3 ha, had both flowering and non-flowering patches of ramets during the 4-year flowering period (2004–2007). A fraction of the flowering genet remained non-flowering during the 4 years of observation, and did not die after mass flowering. Flowering ramets were physically connected to non-flowering ramets via rhizomes, and assimilated ¹³C was allocated from non-flowering ramets to flowering ramets. Consequently, we clarified that this dwarf bamboo potentially has polycarpic reproductive traits rather than monocarpic, and a genet can keep rhizomes and non-flowering patches alive to sustain the organism after mass flowering.