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The biotic resistance hypothesis asserts that native species may hinder the invasion of exotic species, which can occur either directly or indirectly by influencing interactions between exotic and local species. Aphid-tending ants may play a key role in the indirect biotic resistance to plant invasion. Ants may protect aphids, thus increasing their negative effect on exotic plants, but may also deter chewing herbivores, thus benefiting exotic plants. We studied native aphid-tending ants (Dorymyrmex tener, Camponotus distinguendus, and Dorymyrmex richteri) on exotic nodding thistles (Carduus thoermeri), which are attacked by thistle aphids (Brachycaudus cardui) and thistle-head weevils (Rhinocyllus conicus). We evaluated the impact of ants, aphids, and weevils on thistle seed set. We compared ant species aggressiveness towards aphid predators and weevils and performed ant-exclusion experiments to determine the effects of ants on aphid predators and weevils. We analysed whether ant species affected thistle seed set through their effects on aphids and/or weevils. The ant D. tener showed the most aggressive behaviour towards aphid predators and weevils. Further, D. tener successfully removed aphid predators from thistles but did not affect weevils. Excluding D. tener from thistles increased seed set. Analyses supported a negative indirect pathway between the aggressive D. tener and thistle seed set through aphid populations, while the other ant species showed no indirect effects on thistle reproduction. Therefore, aggressive aphid-tending ants may enhance biotic resistance by increasing aphid infestation on exotic invasive plants. This study highlights the importance of indirect biotic resistance in modulating the success of invasive species. [ABSTRACT FROM AUTHOR]

Native bee abundance has long been assumed to be limited by floral resources. This paradigm has been established in large measure because more bees are often found in areas supporting greater floral abundance. This could result from attraction to resource-rich sites as well as greater local demographic performance in sites supporting high floral abundance; however, demographic performance is usually unknown. Factors other than floral resources such as availability of nest sites, pressure from natural enemies, or whether floral resources are from a mixed native or mostly monodominant exotic assemblage might influence survival or fecundity and hence abundance. We examined how the survival and fecundity of the native solitary bee Osmia lignaria varied along a gradient in floral resource abundance. We released bees alongside a nest block at 27 grassland sites in Montana (USA) that varied in floral abundance and the extent of invasion by exotic forbs. We monitored nest construction and the fate of offspring within each nest. The number of nests established was positively related to native forb abundance and was negatively related to exotic forb species richness. Fecundity was positively related to native forb species richness; however, offspring mortality caused by the brood parasite Tricrania stansburyi was significantly greater in native-dominated sites. These results suggest that native floral resources can positively influence bee populations, but that the relationship between native floral resources and bee population performance is not straightforward. Rather, bees may face a trade-off between high offspring production and low offspring survival in native-dominated sites. [ABSTRACT FROM AUTHOR]

Invasive plants have the potential to reduce native plant abundance through both direct and indirect interactions. Direct interactions, such as competition for soil resources, and indirect interactions, such as competition for shared pollinators, have been shown to influence native plant performance; however, we know much less about how these interactions influence native plant abundance in the field. While direct competitive interactions are often assumed to drive declines in native abundance, an evaluation of their influence relative to indirect mechanisms is needed to more fully understand invasive plant impacts. We quantified the direct effects of resource competition by the invasive perennial forb, Euphorbia esula (Euphorbiaceae), on the recruitment, subsequent performance, and ultimate adult abundance of the native annual, Clarkia pulchella (Onagraceae). We contrast these direct effects with those that indirectly resulted from competition for shared pollinators. Although E. esula dramatically reduced pollinator visitation to C. pulchella, plants were only weakly pollen-limited. Pollen supplementation increased the number of seeds per fruit from 41.28 to 46.38. Seed addition experiments revealed that the impacts of ameliorating pollen limitation only increased potential recruitment by 12.3 %. In contrast, seed addition experiments that ameliorated direct competition with E. esula resulted in an increase in potential future recruitment of 574 %. Our results show that, while the indirect effects of competition for pollinators can influence plant abundance, its effects are dwarfed by the magnitude of direct effects of competition for resources. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Effects of invasive plant species on native plant species are frequently assumed or inferred from comparisons, but rarely quantified experimentally. Such quantification is important to assessing risks and impacts of invasives. We quantified the effects of Lonicera maackii, an exotic shrub invasive in many eastern North American forests, on survival, growth, and reproduction of three perennial herbs: Allium burdickii, Thalictrum thalictroides , and Viola pubescens. We predicted that the spring ephemeral, A. burdickii , would be most impacted, due to early leaf expansion of L. maackii. Field experiments were carried out in two deciduous forest stands, one (Gregg’s Woodlot, GW) disturbed and the other (Western Woods, WW) relatively undisturbed. In each stand, individual herbs were transplanted into a blocked design of 60 plots where L. maackii was present, absent, or removed, and monitored for 5 growing seasons. Lonicera maackii did not affect survival of transplants, but reduced growth and final size of individuals of all three species. For two of the species, A. burdickii and V. pubescens, L. maackii reduced the proportion of live plants flowering in both stands, and reduced the seed or fruit number per flowering individual in GW. For T. thalictroides the proportion flowering was not affected, but seed number per flowering plant was reduced by L. maackii in both stands. For all three species, cumulative seed production over the course of the study was reduced by L. maackii. Overall, effects on the spring ephemeral, A. burdickii, were similar to effects on the other herbs. Because mortality of these established individuals was not affected, short-term studies might conclude forest herbs are unaffected by invasive shrubs. However, the growth and reproduction impacts documented here suggest that populations are impacted in the long-term. [ABSTRACT FROM AUTHOR]

The relative importance of allelopathy and resource competition in plant-plant interactions has been vigorously debated but seldom tested. We used activated carbon to manipulate the effects of root exudates of Centaurea maculosa, a noxious weed in much of western North America, on root elongation rates and growth of the native bunchgrass Festuca idahoensis in order to investigate the relative importance of allelopathy in the total interference of Centaurea. In root observation chambers, Festuca root elongation rates decreased to ≈50% of the control, beginning 4 days before contacting Centaurea roots in silica sand. However, when activated carbon, which has a high affinity for adsorbing to organic compounds, was added to the sand the effects of Centaurea roots on Festuca root elongation were reduced. In other experiments, Festuca plants were 50% smaller when grown with Centaurea than with conspecifics in pure silica sand. However, Festuca grown with Centaurea in mixtures of sand and activated carbon were 85% larger than Festuca grown with Centaurea in silica sand without carbon. These results suggest that allelopathy accounts for a substantial proportion of the total interference of Centaurea on Festuca, shifting the balance of competition in favor of Centaurea. However, Centaurea outperformed Festuca even in the presence of activated carbon, demonstrating the importance of the combined roles of resource competition and allelopathy. [ABSTRACT FROM AUTHOR]

Release of exotic insects as biological control agents is a common approach to controlling exotic plants. Though controversy has ensued regarding the deleterious direct effects of biological control agents to non-target species, few have examined the indirect effects of a ”well-behaved” biological control agent on native fauna. We studied a grassland in west-central Montana infested with spotted knapweed (Centaurea maculosa) to examine the effects of knapweed invasion and two gall flybiological control agents (Urophora affinis and U. quadrifasciata) on the native deer mouse (Peromyscus maniculatus). Stomach-content analysis revealed that Urophora were the primary food item in Peromyscus diets for most of the year and made up 84–86% of the winter diet. Stomach contents indicated that wild-caught mice consumed on average up to 247 Urophora larvae mouse–1 day–1, while feeding trials revealed that deer mice could depredate nearly 5 times as many larvae under laboratory conditions. In feeding trials, deer mice selected knapweed seedheads with greater numbers of galls while avoiding uninfested seedheads. When Urophora larvae were present in knapweed seedheads, deer mice selected microhabitats with moderately high (31–45% cover) and high knapweed infestation (≥46% cover). After Urophora emerged and larvae were unavailable to Peromyscus, mice reversed habitat selection to favor sites dominated by native-prairie with low knapweed infestation (0–15%). Establishment of the biological control agent, Urophora spp., has altered deer mouse diets and habitat selection by effecting changes in foraging strategies. Deer mice and other predators may reduce Urophora populations below a threshold necessary to effectively control spotted knapweed. [ABSTRACT FROM AUTHOR]

Variation in timing and amount of rainfall in California has been noted to cause strong year-to-year differences in the composition of vernal pool communities. We explored the effects of possible variation of the “rainy season” in a seedbank germination study conducted with three factors: monthly timing of first soil moistening, length of moist period before inundation, and length of inundation. Monthly timing was the most important factor in determining the number of plants and number of species that germinated in the pots. Many species, mostly generalist wetland and pool-endemic species, showed sharp decreases in number in later trials. Five species did not germinate in the final trial, and two of these were abundant only in the first trial. Inundation was required for the germination of two species. Treatments with a combination of inundated and moist conditions resulted in higher numbers of species and numbers of plants compared to treatments with only moistness or only inundation. Our results indicate that vernal pool species have mechanisms that keep them from emerging under unfavorable conditions. The sensitivity of vernal pool species to out-of-season germination suggests that they could be negatively affected by climatic change, with generalist species like non-native Lythrum hyssopifolium having a potential advantage. [ABSTRACT FROM AUTHOR]