Genetically Based Trait Differentiation but Lack of Trade-offs between Stress Tolerance and Performance in Introduced Canada Thistle.

Trade-offs between performance and tolerance of abiotic and biotic stress have been proposed to explain both the success of invasive species and frequently observed size differences between native and introduced populations. Canada thistle seeds collected from across the introduced North American and the native European range were grown in benign and stressful conditions (nutrient stress, shading, simulated herbivory, drought, and mowing), to evaluate whether native and introduced individuals differ in performance or stress tolerance. An additional experiment assessed the strength of maternal effects by comparing plants derived from field-collected seeds with those derived from clones grown in the glasshouse. Introduced populations tended to be larger in size, but no trade-off of stress tolerance with performance was detected; introduced populations had either superior performance or equivalent trait values and survivorship in the treatment common gardens. We also detected evidence of parallel latitudinal clines of some traits in both the native and introduced ranges and associations with climate variables in some treatments, consistent with recent climate adaptation within the introduced range. Our results are consistent with rapid adaptation of introduced populations, but, contrary to predictions, the evolution of invasive traits did not come at the cost of reduced stress tolerance.

Impact of Invasive Bees on Plant-Pollinator Interactions and Reproductive Success of Plant Species in Mixed Nothofagus antarctica Forests.

Invasive social bees can alter plant-pollinator interactions with detrimental effects on both partners. However, most studies have focused on one invasive bee species, while the interactions among two or more species remain poorly understood. Also, many study sites had a history of invasive bees, being hard to find sites with historical low abundances. In Patagonia, Bombus ruderatus (F.) invasion begun in 1993 and B. terrestris (L.) in 2006. Though honey bees (Apis mellifera L.) introduction started in 1859, their density is still low in some parts. By experimentally increasing honey bee densities, we evaluated the effect of honey bees and bumblebees floral visitation on native pollinator floral visitation, pollen deposition, and reproductive success of three plant species in mixed Nothofagus antarctica forests of northern Patagonia: Oxalis valdiviensis, Mutisia spinosa and Cirsium vulgare. Our results show that exotic bees became the main floral visitors. No negative association was found between invasive bee and native pollinator visitation rates, but there was evidence of potential competition between honey bees and bumblebees. Floral neighborhood diversity played an important role in pollinator behavior. Conspecific pollen deposition was high for all species, while deposition of heterospecific pollen was very high in M. spinosa and C. vulgare. Not as expected, honey bees visitation rate had a negative effect on heterospecific pollen deposition in C. vulgare. For O. valdiviensis, exotic visitation rates increased conspecific pollen deposition, which was positively related to reproductive success. Although exotic bees became main floral visitors, their contribution to reproductive success was only clear for one species.

Experimental manipulation of floral scent bouquets restructures flower-visitor interactions in the field.

A common structural feature of natural communities is the non-random distribution of pairwise interactions between organisms of different trophic levels. For plant-animal interactions, it is predicted that both stochastic processes and functional plant traits that facilitate or prevent interactions are responsible for these patterns. However, unbiased manipulative field experiments that rigorously test the effects of individual traits on community structure are lacking. We address this gap by manipulating floral scent bouquets in the field. Manipulation of floral scent bouquets led to quantitative as well as qualitative restructuring of flower-visitor networks, making them more generalized. Olfactometer trials confirmed both positive and negative responses to scent bouquets. Our results clearly show that the distribution of insect visitors to the two abundant study plant species reflects the insects' species-specific preferences for floral scents, rather than for visual or morphological floral traits. Thus, floral scents may be of major importance in partitioning flower-visitor interactions. Integrating experimental manipulations of plant traits with field observations of interaction patterns thus represents a promising approach for revealing the processes that structure species assemblages in natural communities.

Infection by a foliar endophyte elicits novel arabidopside-based plant defence reactions in its host, Cirsium arvense.

Endophytic fungi live asymptomatically within plants. They are usually regarded as nonpathogenic or even mutualistic, but whether plants respond antagonistically to their presence remains unclear, particularly in the little-studied associations between endophytes and nongraminoid herbaceous plants. We investigated the effects of the endophyte Chaetomium cochlioides on leaf chemistry in Cirsium arvense. Plants were sprayed with spores; leaf material from both subsequent new growth and the sprayed leaves was analysed 2 wk later. Infection frequency was 91% and 63% for sprayed and new growth, respectively, indicating that C. cochlioides rapidly infects new foliage. Metabolomic analyses revealed marked changes in leaf chemistry with infection, especially in new growth. Changes in several novel oxylipin metabolites were detected, including arabidopsides reported here for the first time in a plant species other than Arabidopsis thaliana, and a jasmonate-containing galactolipid. The production of these metabolites in response to endophyte presence, particularly in newly infected foliage, suggests that endophytes elicit similar chemical responses in plants to those usually produced following wounding, herbivory and pathogen invasion. Whether endophytes benefit their hosts may depend on a complex series of chemically mediated interactions between the plant, the endophyte, other microbial colonists and natural enemies.

Effects of apical meristem mining on plant fitness, architecture, and flowering phenology in Cirsium altissimum (Asteraceae).

UNLABELLED: • PREMISE OF THE STUDY: Interactions that limit lifetime seed production have the potential to limit plant population sizes and drive adaptation through natural selection. Effects of insect herbivory to apical meristems (apical meristem mining) on lifetime seed production rarely have been quantified experimentally. We studied Cirsium altissimum (tall thistle), whose meristems are mined by Platyptilia carduidactyla (artichoke plume moth), to determine how apical damage affects plant maternal fitness and evaluate both direct and indirect mechanisms underlying these effects.• METHODS: In restored prairie, apical mining was manipulated on tall thistles by applying insecticide, water, or no spray to apical meristems. We quantified effects on lifetime seed production, plant architecture, and flowering phenology. Seed germinability and seedling mass were evaluated in a greenhouse.• KEY RESULTS: Apical meristem miners decreased lifetime seed production of C. altissimum, but not seed quality. Higher mortality rates of damaged plants contributed to reduced seed production. Apical damage reduced plant height and increased the proportion of blooming flower heads in axial positions on branches. Apical damage delayed flowering and shortened flowering duration.• CONCLUSIONS: Apical meristem mining reduced plant maternal fitness. The shift in the identity of blooming flower heads from terminal to axial positions contributed to this reduction because axial heads are less fecund. Shorter, meristem-mined plants may have been more susceptible to competition, and this susceptibility may explain their higher mortality rates. The kinds of changes in architecture and phenology that resulted from apical damage to C. altissimum have been shown to affect floral visitation in other plant species.

Native insect herbivory limits population growth rate of a non-native thistle.

The influence of native fauna on non-native plant population growth, size, and distribution is not well documented. Previous studies have shown that native insects associated with tall thistle (Cirsium altissimum) also feed on the leaves, stems, and flower heads of the Eurasian congener C. vulgare, thus limiting individual plant performance. In this study, we tested the effects of insect herbivores on the population growth rate of C. vulgare. We experimentally initiated invasions by adding seeds at four unoccupied grassland sites in eastern Nebraska, USA, and recorded plant establishment, survival, and reproduction. Cumulative foliage and floral herbivory reduced C. vulgare seedling density, and prevented almost any reproduction by C. vulgare in half the sites. The matrix model we constructed showed that this herbivory resulted in a reduction of the asymptotic population growth rate (λ), from an 88% annual increase to a 54% annual decline. These results provide strong support for the hypothesis that indigenous herbivores limit population invasion of this non-native plant species into otherwise suitable grassland habitat.

Marsh thistle in New York: early detection and rapid response to a recent invader.

Case studies implementing early detection and rapid response (EDRR) protocols are crucial to our understanding of how to evaluate the threat of a new invasive organism. Most EDRR schematics depict a linear approach to EDRR beginning with detection, then assessment, and concluding with the rapid response. Our case study exemplified the need to adjust this protocol by interlacing all three phases. Our target species is a recent invader to central New York: Cirsium palustre (marsh thistle). We observed 192 populations and reported them on iMapInvasives.org. These locations were used to generate risk maps of neighboring counties. Six test plots evaluated the plant's bienniality and monocarpism. A life-stage analysis allowed us to delineate the invasion front to target control and management solutions where they are most effective for marsh thistle. An adaptive approach to EDRR can expedite response strategies, develop risk maps, and train partners for better control.

Insect herbivory and propagule pressure influence Cirsium vulgare invasiveness across the landscape.

A current challenge in ecology is to better understand the magnitude, variation, and interaction in the factors that limit the invasiveness of exotic species. We conducted a factorial experiment involving herbivore manipulation (insecticide-in-water vs. water-only control) and seven densities of introduced nonnative Cirsium vulgare (bull thistle) seed. The experiment was repeated with two seed cohorts at eight grassland sites uninvaded by C. vulgare in the central Great Plains, USA. Herbivory by native insects significantly reduced thistle seedling density, causing the largest reductions in density at the highest propagule inputs. The magnitude of this herbivore effect varied widely among sites and between cohort years. The combination of herbivory and lower propagule pressure increased the rate at which new C. vulgare populations failed to establish during the initial stages of invasion. This experiment demonstrates that the interaction between biotic resistance by native insects, propagule pressure, and spatiotemporal variation in their effects were crucial to the initial invasion by this Eurasian plant in the western tallgrass prairie.

Combined effects of plant competition and insect herbivory hinder invasiveness of an introduced thistle.

The biotic resistance hypothesis is a dominant paradigm for why some introduced species fail to become invasive in novel environments. However, predictions of this hypothesis require further empirical field tests. Here, we focus on evaluating two biotic factors known to severely limit plants, interspecific competition and insect herbivory, as mechanisms of biotic resistance. We experimentally evaluated the independent and combined effects of three levels of competition by tallgrass prairie vegetation and two levels of herbivory by native insects on seedling regeneration, size, and subsequent flowering of the Eurasian Cirsium vulgare, a known invasive species elsewhere, and compared its responses to those of the ecologically similar and co-occurring native congener C. altissimum. Seedling emergence of C. vulgare was greater than that of C. altissimum, and that emergence was reduced by the highest level of interspecific competition. Insect leaf herbivory was also greater on C. vulgare than on C. altissimum at all levels of competition. Herbivory on seedlings dramatically decreased the proportion of C. vulgare producing flower heads at all competition levels, but especially at the high competition level. Competition and herbivory interacted to significantly decrease plant survival and biomass, especially for C. vulgare. Thus, both competition and herbivory limited regeneration of both thistles, but their effects on seedling emergence, survival, size and subsequent reproduction were greater for C. vulgare than for C. altissimum. These results help explain the unexpectedly low abundance recorded for C. vulgare in western tallgrass prairie, and also provide strong support for the biotic resistance hypothesis.

The foliar endophytic fungal community composition in Cirsium arvense is affected by mycorrhizal colonization and soil nutrient content.

Foliar fungal endophytes are ubiquitous, but understudied symbionts of most plant species; relatively little is known about the factors affecting their occurrence, diversity and abundance. We tested the effects of soil nutrient content and arbuscular mycorrhizal (AM) colonization on the occurrence of foliar endophytic fungi in Cirsium arvense in two field studies. In the first study, we assessed relationships between soil moisture, organic matter, carbon and nitrogen content and plant water, nitrogen and carbon content and AM colonization and the occurrence of foliar endophytic fungal species. In the second study, we manipulated soil nutrient content and AM colonization of potted seedlings and identified differences in endophytic fungal species composition of the leaves and stems. The results reveal that endophytes can occur either more or less frequently, depending on soil nutrient and plant water content and AM colonization. We propose that these patterns were the result of differences in fungal growth responses to nutrient availability in the leaves, which can be affected by resources obtained from the soil or symbiotic fungi in the roots.

Seed availability and insect herbivory limit recruitment and adult density of native tall thistle.

Understanding spatial and temporal variation in factors influencing plant regeneration is critical to predicting plant population growth. We experimentally evaluated seed limitation, insect herbivory, and their interaction in the regeneration and density of tall thistle (Cirsium altissimum) across a topographic ecosystem productivity gradient in tallgrass prairie over two years. On ridges and in valleys, we used a factorial experiment manipulating seed availability and insect herbivory to quantify effects of: seed input on seedling density, insect herbivory on juvenile density, and cumulative impacts of both seed input and herbivory on reproductive adult density. Seed addition increased seedling densities at three of five sites in 2006 and all five sites in 2007. Insect herbivory reduced seedling survival across all sites in both years, as well as rosette survival from the previous year's seedlings. In both years, insecticide treatment of seed addition plots led to greater adult tall thistle densities in the following year, reflecting the increase in juvenile thistle densities in the experimental year. Seedling survival was not density dependent. Our analytical projection model predicts a significant long-term increase in adult densities from seed input, with a greater increase under experimentally reduced insect herbivory. While plant community biomass and water stress varied significantly between ridges and valleys, the effects of seed addition and insect herbivory did not vary with gradient position. These results support conceptual models that predict seedling and adult densities of short-lived monocarpic perennial plants should be seed limited. Further, the experiment demonstrates that even at high juvenile plant densities, at which density dependence potentially could have overridden herbivore effects on plant survival, insect herbivory strongly affected juvenile thistle performance and adult densities of this native prairie species.

No interaction between competition and herbivory in limiting introduced Cirsium vulgare rosette growth and reproduction.

Both competition and herbivory have been shown to reduce plant survival, growth, and reproduction. Much less is known about whether competition and herbivory interact in determining plant performance, especially for introduced, weedy plant species in the invaded habitat. We simultaneously evaluated both the main and interactive effects of plant neighbors and insect herbivory on rosette growth and seed reproduction in the year of flowering for Cirsium vulgare (bull thistle, spear thistle), an introduced Eurasian species, in tallgrass prairie in 2 years. Effects of insect herbivory were strong and consistent in both years, causing reduced plant growth and seed production, whereas the effects of competition with established vegetation were weak. The amount of herbivore damage inflicted on rosettes did not depend on the presence of neighbor plants. We also found no interaction between competition and herbivory on key parameters of plant growth and fitness. The results of this study contradict the hypothesis that competitive context interacts with insect herbivory in limiting the invasiveness of this introduced thistle. Further, the results provide additional, experimental evidence that high levels of herbivory on established rosettes by native insects exert significant biotic resistance to the invasiveness of C. vulgare in western tallgrass prairie.

The relative importance of resources and natural enemies in determining herbivore abundance: thistles, tephritids and parasitoids.

1. The relative importance of host-plant resources and natural enemies in influencing the abundance of insect herbivores was investigated in potted plant and natural population experiments, using tephritid (Diptera: Tephritidae) flies, their host plant, creeping thistle Cirsium arvense, and their Hymenoptera parasitoids. 2. Experimental manipulation of host-plant quality (i.e. levels of host-plant nutrients) and resource availability (i.e. the number of buds) increased tephritid abundance. There was no evidence that the seed-feeding tephritid fly Xyphosia miliaria preferentially oviposited on fertilized C. arvense. 3. At low thistle densities, X. miliaria showed a constant rate of resource exploitation. At higher thistle densities, a threshold was detected, above which additional buds were not attacked. 4. Parasitism attack was variable across host (tephritid) densities but levels of parasitism were consistently higher on the fertilized thistles. 5. Experimental manipulation of host-plant quality and resource availability (quantity) not only directly affects the tephritid population but also, indirectly, leads to high rates of parasitism. Both chemical and physical characteristics of host plants affect the performance of natural enemies. 6. Both top-down and bottom-up forces act to influence tephritid abundance, with bottom-up influences appearing to be the most important.

Fragrance of Canada thistle (Cirsium arvense) attracts both floral herbivores and pollinators.

The evolution of floral scent as a plant reproductive signal is assumed to be driven by pollinator behavior, with little attention paid to other potential selective forces such as herbivores. I tested 10 out of the 13 compounds emitted by dioecious Cirsium arvense, Canada thistle, including 2-phenylethanol, methyl salicylate, p-anisaldehyde, benzaldehyde, benzyl alcohol, phenylacetaldehyde, linalool, furanoid linalool oxides (E and Z), and dimethyl salicylate. Single compounds (and one isomer) set out in scent-baited water-bowl traps trapped over 10 species of pollinators and 16 species of floral herbivores. The two dominant components of the fragrance blend of C. arvense, benzaldehyde and phenylacetaldehyde, trapped both pollinators and florivores. Other compounds attracted either pollinators or florivores. Florivores of C. arvense appear to use floral scent compounds as kairomones; by advertising to pollinators, C. arvense also attracts its own enemies.

The effect of pollination on floral fragrance in thistles.

We investigated postpollination changes in fragrance composition and emission rates, as well as pollinator discrimination in hand-pollinated flower heads of two thistle species: Canada thistle (Cirsium arvense) and sandhill thistle (C. repandum). Following pollination, neither species emitted any novel compounds that could function as repellents. Scent emission rates declined in pollinated plants of both species by approximately 89% within 48 hr. This decline was evident in all 13 scent components of C. arvense. Apis mellifera, the dominant pollinator in the study population of C. arvense, was nearly three times more likely to visit an unpollinated rather than a pollinated flower head. A more complex pattern was observed for C. repandum, whose scent comprised 42 compounds. Quantities of aromatic and sesquiterpenoid volatiles declined after pollination, whereas two classes of scent compounds, fatty acid derivatives and monoterperpenoids, continued to be emitted. In C. repandum, discrimination against pollinated flower heads by Papilio palamedes (its primary pollinator) was not as marked. Unpollinated control plants of both species maintained moderate levels of scent production throughout this experiment, demonstrating that senescence and floral advertisement may be delayed until pollination has occurred. We expect postpollination changes in floral scent contribute to communication between plants with generalized pollinator spectra and their floral visitors. This study provides the first field study of such a phenomenon outside of orchids.

Arbuscular mycorrhizal colonization of the dominant plant species in primary successional volcanic deserts on the Southeast slope of Mount Fuji.

Arbuscular mycorrhizal (AM) colonization was observed on four plant species in primary successional volcanic deserts on the Southeast slope of Mount Fuji. The AM colonization of the dominant species, Polygonum cuspidatum, contradicts the conclusion that Polygonaceae are often regarded as being non-mycorrhizal species. The secondary dominant species, Polygonum weyrichii var. alpinum, formed no mycorrhizas. The roots of Cirsium purpuratum, Clematis stans and Campanula punctata ssp. hondoensis, showed a higher percentage of AM colonization than P. cuspidatum. AM colonization and spore density in the rhizosphere soil of P. cuspidatum significantly decreased as elevation increased. AM colonization in roots of Cirsium purpuratum and Clematis stans also tended to decrease with increased altitudes. Cirsium purpuratum and Campanula punctata ssp. hondoensis formed single structural types of Arum- and Paris-type, respectively, whereas P. cuspidatum and Clematis stans formed both Arum- and Paris-type morphologies.

Above- and below ground trophic interactions on creeping thistle (Cirsium arvense) in high- and low-diversity plant communities: potential for biotic resistance?

The capacity of local communities to control introduced plants is called biotic resistance. Biotic resistance has been almost exclusively tested for plant competition and above ground herbivores and pathogens, while neglecting root herbivores and soil pathogens. Here, we present biotic resistance by above- and below ground herbivores in concert, and relate the abundance of the plant enemies to the species diversity of the local plant communities. The study was carried out in a 7-year-old biodiversity field experiment. We used creeping thistle (Cirsium arvense) as a model, and quantified sap-sucking herbivores: above ground aphids, their antagonists, and root-feeding nematodes. As plant diversity treatments, we used field plots sown with high (15 plant species, HSD) or low (4 plant species, LSD) diverse seed mixtures in 1996 and that were not weeded. Creeping thistle became established spontaneously at the start of the experiment. In 2002, in HSD plots, 90 % of the plant community was made up by 11 species, compared to seven species in LSD plots. No differences were found for C. arvense abundance or biomass. Above ground, three aphid species were found on C. arvense-Uroleucon cirsii, Aphis fabae, and Macrosiphum euphorbiae, but the latter was found only in low densities. Significantly more aphid species were found on individual plants in HSD plots. Moreover, in HSD plots, on average 10 % of aphids were parasitized, while no parasitism was observed in LSD plots. In the root zone of C. arvense, significantly more nematodes were found in HSD than in LSD plots, and a significantly higher proportion of those nematodes were plant parasites. The dominant plant parasitic nematode in both treatments was Paratylenchus. We conclude that biotic resistance by natural enemies may be enhanced by plant species diversity, but that above- and below ground sap-sucking herbivores do not necessarily have to respond similarly to the diversity of the surrounding plant community.

Phylogeny and ecological radiation of New World thistles (Cirsium, Cardueae - Compositae) based on ITS and ETS rDNA sequence data.

Sequence data from a portion of the external transcribed spacer (ETS) and internal transcribed spacers (ITS-1 and ITS-2) of 18S-26S nuclear ribosomal DNA were used to resolve historical biogeography and ecology of true thistles (Cirsium, Cardueae, Compositae) in the New World. The 650 base-pair, 3' portion of the ETS examined here showed a level of variation across taxa similar to that of the ITS sequences included. A maximum-likelihood tree based on combined ETS and ITS sequences leads us to suggest that the New World species of true thistles constitute a major lineage, which in turn comprises several smaller lineages. A western North American lineage shows weak quartet-puzzling support, but includes a well-supported lineage of species endemic to the California Floristic Province. Comparisons of this Californian lineage with other neoendemic angiosperm groups of the region show that the Californian Cirsium lineage exhibits unusually high ecological diversity for a group displaying such low levels of rDNA sequence divergence across taxa. Similarly low levels of sequence divergence were found throughout the New World Cirsium lineage. These results indicate either that Cirsium underwent a rapid ecological radiation in North America, or that rDNA evolution in North American Cirsium has been highly conservative.

Dynamics of a mutualism in a multi-species context.

Despite recent findings that mutualistic interactions between two species may be greatly affected by species external to the mutualism, the implications of such multi-species interactions for the population dynamics of the mutualists are virtually unexplored. In this paper, we ask how the mutualism between the shoot-base boring weevil Apion onopordi and the rust fungus Puccinia punctiformis is influenced by the dynamics of their shared host plant Cirsium arvense, and vice versa. In particular, we hypothesized that the distribution of the weevil's egg load between healthy and rust-infected thistles may regulate the abundance of the mutualists and their host plant. In contrast to our expectations we found that the dynamics of the mutualists are largely determined by the dynamics of their host. This is, to our knowledge, the first demonstration that the dynamics of a mutualism are driven by a third, non-mutualistic species.