Influence of Microsite Disturbance on the Establishment of Two Congeneric Invasive Thistles

The successful establishment of invasive species has been shown to depend on aspects of the invaded community, such as gap characteristics. Biotic resistance may be particularly critical for stopping invaders at early life history stages, but new species can often invade following disturbances, which may create microsites with very different characteristics than are usually present. We examine the response of two invasive thistle species, Carduus nutans L. and C. acanthoides L., to three different microsite characteristics: disturbance type, size, and water availability. The two species initially responded differently to the type of disturbance: C. acanthoides had higher emergence and survival in plots with both above- and belowground disturbance, whereas C. nutans had better early performance in large microsites with above-ground disturbance only. Later in their life cycle, C. nutans performed better in plots that had been disturbed both above- and belowground, whereas C. acanthoides was largely unaffected by disturbance type. Increased emergence and survival, larger size and a higher proportion flowering were observed in larger gaps for both species throughout the life cycle. Watering had a negative impact on C. nutans emergence and fall survival and on C. acanthoides survival to the following summer. Overall, these results suggest that disturbance-generated microsite characteristics (disturbance type and size) may have large impacts on establishment of these two Carduus species, which in turn may persist well beyond the initial stages of growth. Studying invader responses to disturbance can help us to understand under what circumstances they are likely to establish and create persistent problems; avoiding or ameliorating such situations will have significant management benefits.     

Warming leads to divergent responses but similarly improved performance of two invasive thistles

Plants’ responses to climate change are complex. Even the same net performance changes may involve different responses of multiple life history traits. Here we show that two congeneric thistles, Carduus nutans and Carduus acanthoides, both grew taller under increased temperature, albeit following divergent response patterns. For C. nutans, warming advanced bolting more than flowering, leading to a longer growing period before flowering and ultimately taller plant height at the end of the growing season. Carduus acanthoides maintained the same length of growing period because of equally shifted events in the phenological sequence, however, post-flowering growth rate was increased, which also led to enhanced final plant height. As seeds from taller plants disperse farther, their responses imply that future invasion spread rates of these two species will increase. Similar consequences due to divergent responses in life history traits, as demonstrated in this study, suggest that considering only ultimate performance outcomes, and not the underlying processes generating such outcomes, is not enough to understand the impacts of climate change.     

Spatial Segregation of Congeneric Invaders in Central Pennsylvania, USA

Carduus acanthoides and Carduus nutans (plumeless and musk thistles) are among the most noxious weeds in the United States of America, presenting a serious challenge in cropping and pasture systems. Unfortunately, a lack of detailed spatial distribution information hampers both our ability to understand the factors affecting their invasive success, and the effectiveness of monitoring and management efforts. To examine patterns of distribution and co-occurrence at a local level, we sampled a 5000 km² area of central Pennsylvania that cut a transect across known areas of C. acanthoides and C. nutans infestation. A number of potential environmental explanatory variables were recorded and analyzed to examine whether they correlated with observed species distribution patterns. Patterns of forest density and spatial aggregation of the thistles were the primary covariates that significantly impacted both species’ distributions. The survey established that the frequency of sightings for each species diminished as the ranges converged, with only brief overlap: the two species are strongly negatively correlated in space. Understanding environmental correlates of infestation and the pattern of spatial dissociation of these two invasive species is an important step towards an improved understanding of the mechanisms underlying their invasive potential, and hence towards effective weed control.     

Can Soil Seed Banks Serve as Genetic Memory? A Study of Three Species with Contrasting Life History Strategies

We attempted to confirm that seed banks can be viewed as an important genetic reservoir by testing the hypothesis that standing (aboveground) plants represent a nonrandom sample of the seed bank. We sampled multilocus allozyme genotypes from three species with different life history strategies: Amaranthus retroflexus, Carduus acanthoides, Pastinaca sativa. In four populations of each species we analysed the extent to which allele and genotype frequencies vary in consecutive life history stages including the summer seed bank, which has been overlooked up to now. We compared the winter seed bank (i.e., seeds collected before the spring germination peak), seedlings, rosettes, the summer seed bank (i.e., seeds collected after the spring germination peak) and fruiting plants. We found that: (1) All three species partitioned most of their genetic diversity within life history stages and less among stages within populations and among populations. (2) All genetic diversity parameters, except for allele frequencies, were similar among all life history stages across all populations in different species. (3) There were differences in allele frequencies among life history stages at all localities in Amaranthus retroflexus and at three localities in both Carduus acanthoides and Pastinaca sativa. (4) Allele frequencies did not differ between the winter and summer seed bank in most Carduus acanthoides and Pastinaca sativa populations, but there was a marked difference in Amaranthus retroflexus. In conclusion, we have shown that the summer seed bank is not genetically depleted by spring germination and that a majority of genetic diversity remains in the soil through summer. We suggest that seed banks in the species investigated play an important role by maintaining genetic diversity sufficient for recovery rather than by accumulating new genetic diversity at each locality.     

Local pre‐adaptation to disturbance and inbreeding–environment interactions affect colonisation abilities of diploid and tetraploid Centaurea stoebe

• Primary colonisation in invasive ranges most commonly occurs in disturbed habitats, where anthropogenic disturbance may cause physical damage to plants. The tolerance to such damage may differ between cytotypes and among populations as a result of differing population histories (adaptive differentiation between ruderal verus natural habitats). Moreover, founder populations often experience inbreeding depression, the effects of which may increase through physical damage due to inbreeding–environment interactions. We aimed to understand how such colonisation processes differ between diploid and tetraploid Centaurea stoebe populations, with a view to understanding why only tetraploids are invasive.
• We conducted a clipping experiment (frequency: zero, once or twice in the growing season) on inbred versus outbred offspring originating from 37 C. stoebe populations of varying cytotype, range and habitat type (natural versus ruderal). Aboveground biomass was harvested at the end of the vegetation period, while re‐sprouting success was recorded in the following spring.
• Clipping reduced re‐sprouting success and biomass, which was significantly more pronounced in natural than in ruderal populations. Inbreeding depression was not detected under benign conditions, but became increasingly apparent in biomass when plants were clipped. The effects of clipping and inbreeding did not differ between cytotypes.
• Adaptive differentiation in disturbance tolerance was higher among populations than between cytotypes, which highlights the potential of pre‐adaptation in ruderal populations during early colonisation on anthropogenically disturbed sites. While the consequences of inbreeding increased through clipping‐mediated stress, they were comparable between cytotypes, and consequently do not contribute to understanding the cytotype shift in the invasive range.

     

Using phenological niche separation to improve management in a Northern Glaciated Plains grassland

Many of the remaining patches of untilled (native) prairie in the Northern Glaciated Plains of North America are heavily invaded by the cool‐season grasses, Bromus inermis and Poa pratensis. However, the native vegetation in these patches contains many warm‐season species. This difference in phenology can be used to benefit restoration. We conducted an experiment to examine the efficacy of restoration treatments (mowing and prescribed fire) applied early in the growing season for consecutive years to decrease cool‐season invasive plant biomass without impacting the native warm‐season species. Our treatments were successful at significantly decreasing invasive cool‐season plant biomass and increasing native warm‐season plant biomass. No differences between treatments (mowing and prescribed fire) were found. Results suggest that incorporating differences in phenology between target and nontarget species into management may increase restoration success.     

Associations of dominant plant species with arbuscular mycorrhizal fungi during vegetation development on coal mine spoil banks

Among plants colonizing mine spoil banks in Northern Bohemia the first colonizers, mainly ruderal annuals fromChenopodiaceae andBrassicaceae were found not to be associated with arbuscular mycorrhizal fungi (AMF). These species cultivated in pots with soil from four sites in different succession stages of the spoil bank did not respond to the presence of native or non-native AMF. All grass species studied (Elytrigia repens, Calamagrostis epigejos andArrhenatherum elatius) were found moderately colonized in the field and they seemed to be facultative mycotrophs, because their response to mycorrhiza in the experiment was negligible.Carduus acanthoides was found to be highly colonized in the field, however, it did not show growth response to AMF in the pot experiment. The AMF native in four sites on the spoil banks showed high infectivity but low effectiveness in association with colonizing plants compared to the non-native isolateG. fistulosum BEG23. In general, dependence on AMF in the cultivation experiment was rather low, regardless of the fact that plants were found to be associated with AMF either in the field or in pots. Occurrence and effectiveness of mycorrhizal associations might relate primarily to the mycotrophic status of each plant species rather than to the age of the spoil bank sites studied.     

Shifts in dominance of native and invasive plants in experimental patches of vegetation

While it is widely accepted that disturbance often promotes the process of plant invasion, few data are available on specific relationships between different types of disturbance and success of plant invaders. Nor is much known about variation in invader behaviour in response to changing composition of the dominant native species in the recipient habitats. In the middle Main valley near Würzburg (Germany), two invasive Brassicaceae forbs, Bunias orientalis and Rorippa austriaca, are frequently associated with a small number of dominant native species in the herbaceous vegetation of productive, disturbed sites. Variation in the relative dominance of these species suggests high variability between sites in the factors influencing the local invasion process. In our study, we examined variation in regeneration mode, type and frequency of disturbance, and native species composition, and considered life history differences between the two invasive species as factors contributing to the distinct dominance patterns. These factors were tested in a controlled field experiment in which individuals of the two invasive species and four native species were grown in mixed stands for almost 3 years. The development of the plants was monitored by cover values and all plants were harvested at the end of the study. All factors investigated contributed considerably to the development of distinct dominance patterns in the species mixtures. Relative to the natives, both invasive species were promoted by regeneration from vegetative fragments as compared to development from sexual offspring. While both invasive species were generally promoted by disturbance relative to the natives, they differed in their response to distinct types of disturbance, and these differences were strongly affected by species composition. Our results indicate that invasion success of R. austriaca may depend more on soil disturbance, soil transport and deposition, while B. orientalis is expected to particularly expand at mown sites that do not have dense cover by meadow grasses. Whether the native resident vegetation is mainly composed of successional competitors that suffer from disturbance or of ruderal competitors that compete early and strongly with the invasive species may be of critical importance for invasion by the two species.     

Colonization and early succession on anthropogenic soils

Abstract. This paper reports on vegetation development on permanent experimental plots during five years of succession. Nine (1 m²) plots were filled with three typical substrates from man-made habitats of urban and industrial areas in the region of Berlin. The three substrates (a commercial ‘topsoil’, a ruderal ‘landfill’ soil and a sandy soil), differ in organic matter and nutrient contents. Relevés of species composition and percent cover of each species present were made monthly during the growing season from the start of vegetation development. This paper describes the different successional pathways on topsoil and ruderal soil and the colonization process on sandy soil. On topsoil, ruderal annuals are dominant in the first year and are replaced by short-lived perennials from the second year. Those species were replaced by long-lived perennial herbs (Ballota nigra or Urtica dioica) from the third year of succession onwards. On the ruderal land-fill soil the early successional stages are less sharp and the perennial Solidago canadensis is able to dominate within one year after the succession was initiated. On sandy soil there is still an ongoing colonization process, where pioneer tree species like Betula pendula and Populus nigra play a main role. The importance of ‘initial floristic composition’, the role of substrate for community structure and the peculiarities of successional sequences on anthropogenic soils in the context of primary and secondary successions are discussed.     

Establishment and spread of founding populations of an invasive thistle: the role of competition and seed limitation

Successful plant invasions require both the founding and local spread of new populations. High plant densities occur only when founding plants are able to disperse their seeds well locally to quickly colonize and fill the new patch. We test this ability in a 7-year field experiment with Carduus acanthoides, an invasive weed in several North American ecosystems. Founder plants were planted in the center of 64 m² plots and we monitored the recruitment, distribution pattern, mortality, and seed production of the seedlings that originated from these founding plants. Competing vegetation was clipped not at all, once, or twice each year to evaluate the importance of interspecific competition. More seedlings recruited in the intermediate once-clipped plots, and these seedlings also survived better. The control plots had fewer microsites for seedling recruitment; clipping a second time in September stimulated grasses to fill up the gaps. The number of C. acanthoides recruits and their median distances from the founder plants were also explained by the initial seed production of the founding plants. Overall, the experiment shows that the success of founder plants can fluctuate strongly, as 55% of the plots were empty by the sixth year. Our study suggests that the local invasion speed following initial establishment depends strongly on both the propagule pressure and availability of suitable microsites for seedling recruitment and growth.     

Compensation of seed production after severe injury in the short-lived herb Barbarea vulgaris

A pot experiment with the common ruderal herb Barbarea vulgaris (Brassicaceae) was set up to elucidate to what extent short-lived species sprouting from roots regenerate and compensate for seed production after damage. We tested if sprouting from roots ensures survival after severe aboveground biomass damage, but the number of seeds produced declines with increasing severity of injury, decreasing nutrient availability and progress in the life cycle at the time of injury.Plants of B. vulgaris were cultivated in a 3-year garden experiment at two nutrient levels (high vs. low). During the experiment, two levels of injury severity were applied: high (removal of all aboveground biomass) and low (removal of aboveground biomass leaving basal axillary buds intact). Damage was applied at four life-cycle phases: young rosette, overwintered rosette, flowering plant and fruiting plant. All injured plants survived and resprouted irrespective of life-cycle phase, severity of injury and nutrient availability. Injury significantly affected seed production and also the plants’ life cycle. Plants injured in the second year of their life (overwintered rosette, flowering plant and fruiting plant) postponed reproduction to the third season (in the case of high injury severity) or their seed production was lower than in intact plants (in the case of low injury severity). In plants injured in the first life year, seed production and life cycle were not influenced. Nutrient level only marginally affected resprouting after injury and seed production.The experiment showed that the ability to sprout from roots enables plants to survive a 100% loss of aboveground biomass, and to keep some seed production or even compensate it. The short-lived ruderal species B. vulgaris successfully copes with severe disturbance by resprouting and does not rely only on its seed bank.     

Maternal warming affects early life stages of an invasive thistle

Maternal environment can influence plant offspring performance. Understanding maternal environmental effects will help to bridge a key gap in the knowledge of plant life cycles, and provide important insights for species’ responses under climate change. Here we show that maternal warming significantly affected the early life stages of an invasive thistle, Carduus nutans. Seeds produced by plants grown in warmed conditions had higher germination percentages and shorter mean germination times than those produced by plants under ambient conditions; this difference was most evident at suboptimal germination temperatures. Subsequent seedling emergence was also faster with maternal warming, with no cost to seedling emergence percentage and seedling growth. Our results suggest that maternal warming may accelerate the life cycle of this species via enhanced early life‐history stages. These maternal effects on offspring performance, together with the positive responses of the maternal generation, may exacerbate invasions of this species under climate change.     

Integrating multiple disturbance aspects: management of an invasive thistle,Carduus nutans

Background and Aims

Disturbances occur in most ecological systems, and play an important role in biological invasions. We delimit five key disturbance aspects: intensity, frequency, timing, duration and extent. Few studies address more than one of these aspects, yet interactions and interdependence between aspects may lead to complex outcomes.

Methods

In a two-cohort experimental study, we examined how multiple aspects (intensity, frequency and timing) of a mowing disturbance regime affect the survival, phenology, growth and reproduction of an invasive thistle Carduus nutans (musk thistle).

Key Results

Our results show that high intensity and late timing strongly delay flowering phenology and reduce plant survival, capitulum production and plant height. A significant interaction between intensity and timing further magnifies the main effects. Unexpectedly, high frequency alone did not effectively reduce reproduction. However, a study examining only frequency and intensity, and not timing, would have erroneously attributed the importance of timing to frequency.

Conclusions

We used management of an invasive species as an example to demonstrate the importance of a multiple-aspect disturbance framework. Failure to consider possible interactions, and the inherent interdependence of certain aspects, could result in misinterpretation and inappropriate management efforts. This framework can be broadly applied to improve our understanding of disturbance effects on individual responses, population dynamics and community composition.     

Classical Biological Control of Nodding and Plumeless Thistles

Nodding (musk) thistle (Carduus thoermeri Weinmann in the Carduus nutans L. group) and plumeless thistle (Carduus acanthoides L.) are introduced noxious weeds of Eurasian origin. Both weeds are problematic in pastures, rangelands, and croplands and along state highways in many parts of the United States. The success of both species of thistles is largely due to their prolific seed production, seed longevity, competitive ability, and lack of natural enemies. Classical biological control of nodding thistle in Virginia has been achieved with three exotic thistle herbivores, Rhinocyllus conicus Froelich (Coleoptera: Curculionidae), Trichosirocalus horridus (Panzer) (Coleoptera: Curculionidae), and Cassida rubiginosa Müller (Coleoptera: Chrysomelidae). T. horridus also effectively controls plumeless thistle. These insect herbivores complement each other. Nodding thistle biological control is achieved in about 5–6 years in Virginia, Missouri, and Montana. In addition, a rust fungus (Puccinia carduorum Jacky) (Uredinales: Pucciniaceae) has been introduced and established for control of nodding thistle in Virginia. Development and reproduction of the three thistle herbivores are not adversely affected by the rust. The rust hastens plant senescence and reduces seed production. Control of plumeless thistle with R. conicus and T. horridus takes approximately twice as long as control of nodding thistle.     

Plant spatial arrangement affects projected invasion speeds of two invasive thistles

The spatial arrangement of plants in a landscape influences wind flow, but the extent that differences in the density of conspecifics and the height of surrounding vegetation influence population spread rates of wind dispersed plants is unknown. Wind speeds were measured at the capitulum level in conspecific arrays of different sizes and densities in high and low surrounding vegetation to determine how these factors affect wind speeds and therefore population spread rates of two invasive thistle species of economic importance, Carduus acanthoides and C. nutans. Only the largest and highest density array reduced wind speeds at a central focal thistle plant. The heights of capitula and surrounding vegetation also had significant effects on wind speed. When population spread rates were projected using integrodifference equations coupling previously published demography data with WALD wind dispersal models, large differences in spread rates resulted from differences in average horizontal wind speeds at capitulum height caused by conspecific density and surrounding vegetation height. This result highlights the importance of spatial structure for the calculation of accurate spread rates. The management implication is that if a manager has time to remove a limited number of thistle plants, an isolated thistle growing in low surrounding vegetation should be targeted rather than a similar sized thistle in a high density population with high surrounding vegetation, if the objective is to reduce spread rates.     

Selective and Non-Selective Control of Invasive Plants: The Short-Term Effects of Growing-Season Prescribed Fire, Herbicide, and Mowing in Two Texas Prairies

Conservation of North American grasslands is hampered by the impact of invasive herbaceous species. Selective control of these plants, although desirable, is complicated by the shared physiology and phenology of the invader and the native components of the invaded plant community. Fortunately, there is evidence that some management practices, such as prescribed fire, herbicide, and mowing, can cause differential responses in native and invasive grassland species. However, timing of treatment is critical, and fire has been shown to increase rates of invasion when implemented during the dormant season. Bothriochloa ischaemum, an introduced C4 Eurasian grass is an increasing problem in grasslands, particularly in southern and central regions of North America. To date, there has been little success in effective selective control. Two invaded grassland sites representative of Blackland Prairie and Edwards Plateau ecoregions were subjected to two growing‐season prescribed fire treatments, single and double herbicide applications, and single and double mowing treatments. Mowing had no effect on either B. ischaemum or other dominant species at either site one‐year posttreatment. However, growing‐season fire and herbicide were both effective at reducing the abundance of B. ischaemum, with other codominant species responding either negatively to herbicide or neutrally or positively to fire. The vulnerability of B. ischaemum to growing‐season fire may be associated with the ecology of its native range. The negative growth response to growing‐season fire, combined with its lower implementation costs, indicates that this method warrants further investigation as a selective management tool for other problematic species in invaded grasslands.     

Independent effects of arbuscular mycorrhiza and earthworms on plant diversity and newcomer plant establishment

Questions: How do arbuscular mycorrhiza and earthworms affect the structure and diversity of a ruderal plant community? Is the establishment success of newcomer plants enhanced by these soil organisms and their interactions? Methods: We grew a native ruderal plant community composed of different functional groups (grasses, legumes and forbs) in the presence and absence of arbuscular mycorrhizal fungi (AMF) and endogeic earthworms in mesocosms. We introduced seeds of five, mainly exotic, plant species from the same functional groups after a disturbance simulating mowing. The effects of the soil organisms on the native ruderal plant community and seedling establishment of the newcomer plants were assessed. Results: After disturbance, the total above‐ground regrowth of the native plant community was not affected by the soil organisms. However, AMF increased plant diversity and shoot biomass of forbs, but decreased shoot biomass of grasses of the native plant community. Earthworms led to a reduction in total root biomass. Establishment of the introduced newcomer plants increased in the presence of AMF and earthworms. Especially, seedling establishment of the introduced non‐native legume Lupinus polyphyllus and the native forb Plantago lanceolata was promoted in the presence of AMF and earthworms, respectively. The endogeic earthworms gained more weight in the presence of AMF and led to increased extraradical AMF hyphal length in soil. However, earthworms did not seem to modify the effect of AMF on the plant community. Conclusion: The present study shows the importance of mutualistic soil organisms in mediating the establishment success of newcomer plants in a native plant community. Mutualistic soil organisms lead to changes in the structure and diversity of the native plant community and might promote newcomer plants, including exotic species.     

Direct and indirect effects of land‐use intensification on ant communities in temperate grasslands

Land‐use intensification is a major driver of local species extinction and homogenization. Temperate grasslands, managed at low intensities over centuries harbored a high species diversity, which is increasingly threatened by the management intensification over the last decades. This includes key taxa like ants. However, the underlying mechanisms leading to a decrease in ant abundance and species richness as well as changes in functional community composition are not well understood. We sampled ants on 110 grassland plots in three regions in Germany. The sampled grasslands are used as meadows or pastures, being mown, grazed or fertilized at different intensities. We analyzed the effect of the different aspects of land use on ant species richness, functional trait spaces, and community composition by using a multimodel inference approach and structural equation models. Overall, we found 31 ant species belonging to 8 genera, mostly open habitat specialists. Ant species richness, functional trait space of communities, and abundance of nests decreased with increasing land‐use intensity. The land‐use practice most harmful to ants was mowing, followed by heavy grazing by cattle. Fertilization did not strongly affect ant species richness. Grazing by sheep increased the ant species richness. The effect of mowing differed between species and was strongly negative for Formica species while Myrmica and common Lasius species were less affected. Rare species occurred mainly in plots managed at low intensity. Our results show that mowing less often or later in the season would retain a higher ant species richness—similarly to most other grassland taxa. The transformation from (sheep) pastures to intensively managed meadows and especially mowing directly affects ants via the destruction of nests and indirectly via loss of grassland heterogeneity (reduced plant species richness) and increased soil moisture by shading of fast‐growing plant species.     

Mechanisms for Dominance in An Early Successional Old Field by the Invasive Non-Native Lespedeza Cuneata (Dum. Cours.) G. Don

Researchers studying invasive plants often concentrate their efforts on predictive models thought to allow invasive plants to dominate native landscapes. However, if an invasive is already well established then experimental research is necessary to provide the information necessary to effectively manage the species. Prescribing appropriate management strategies without prior experimental research may not only be ineffective but also may squander limited resources or have the unintended consequence of furthering spread. Lespedeza cuneata(Dum. Cours.) G. Don. is a well-established invasive plant of old fields and tall-grass prairie in the US. Managers suspect this species shades-out native plants and this is proposed as its primary mechanism for dominance. Using field experiments we tested probable factors allowing the species to establish itself and, once established, interfere in old field plant communities. We also examined the effects of two common anthropogenic disturbances (mowing and nutrients) on L. cuneata growth and establishment. When L. cuneata was treated (clipping, herbicide and stem pull-back) there was a significant increase in species richness and native species cover. Stem density and canopy cover of L. cuneata increased significantly with mowing frequency but decreased with nutrient input. We suggest that mowing benefits L. cuneata while also hindering woody competition. Results also indicate L. cuneata is less prevalent on nutrient enriched soils than on unamended soil. Lespedeza cuneata appears to suppress native plants by shading them out and it can subsequently take over grassland communities. Since it has a varying response to human induced disturbances and may actually benefit from mowing, land managers should be cautious when utilizing this as a management tool.