Invasion by Non-Native Annual Grasses: The Importance of Species Biomass, Composition, and Time Among California Native Grasses of the Central Valley

The Central Valley of California is noted for its dearth of remnant native grass populations and for low native grass seedling establishment within grasslands now dominated by non‐native annual species. In contrast, remnant populations are common along the coast, and studies have shown an ability for seedlings and adults to compete with non‐native annual grasses. The invasibility of well‐established populations of native grasses in the Central Valley remains unclear. The objectives of this study were to compare the invasibility of native grasses differing in density and species composition and, given the species in this study, to assess the ability of mixes with greater species richness to resist invasion relative to their abilities in monoculture. In the Sacramento Valley of California, six species of native grasses were planted at three densities in monospecific and mixed‐species plots. Percent cover of native perennial and non‐native annual grasses was measured in years 2 and 3, and biomass was sampled in year 5. Native grass biomass and, to a lesser extent, species composition were important in explaining variation in non‐native grass invasibility in the fifth year. Species‐rich treatments did not experience less invasion than would be expected by the proportional invasibility of each species in monoculture. However, invasibility of plots consisting of slower growing, shorter statured species decreased over time, suggesting a successional benefit to diverse communities. This study demonstrates that established stands of native grasses in the Sacramento Valley can resist invasion by non‐native annual grasses and that stand biomass is a particularly important factor in determining invasibility.     

The Mark of Zorro: Effects of the Exotic Annual Grass Vulpia myuros on California Native Perennial Grasses

Native perennial grasses were once common in California prairies that are now dominated by annual grasses introduced from Europe. Competition from exotics may be a principal impediment to reestablishment of native perennial grasses. Introduced annual grasses, such as Vulpia myuros (zorro fescue), are often included with native perennial species in revegetation seed mixtures used in California. To examine the potential suppressive effect of this graminoid, we evaluated the growth and performance of a mixture of California native perennial grasses and resident weeds when grown with varying densities of V. myuros. The annual fescue exhibited a strongly plastic growth response to plant density, producing similar amounts of above‐ground biomass at all seeding densities. Perennial grass seedling survival and above‐ ground biomass decreased and individuals became thinner (i.e., reduced weight‐to‐height ratio) with increasing V. myuros seeding density. V. myuros also significantly suppressed above‐ground biomass and densities of weeds and had a more negative effect on weed densities than on native perennial grass densities. Biomass of native grasses and weeds was not differentially affected by increasing densities of V. myuros. Overall, because V. myuros significantly reduced the survival and performance of the mixture of native perennial grasses and this effect increased with increasing V. myuros density, we conclude that including this exotic annual in native seed mixtures is counterproductive to restoration efforts.     

Effects of Established Perennial Grasses on Introduction of Native Forbs in California

Prairie restoration is not complete without the establishment of both grasses and forbs. However, if desirable forbs and grasses are seeded simultaneously, control of broadleaf weeds is problematic. If possible, a two‐step process of introducing forbs after establishing grasses would allow use of broadleaf‐specific herbicides at the critical early stages of grass growth. We conducted experiments to investigate methods for introducing forbs into previously restored native perennial grasslands on rural roadsides in the Sacramento Valley of California. In one experiment, we studied the effects of background vegetation (established perennial grasses or tilled ground) on seven native forb species planted from seed. In a second experiment, we evaluated the effects of background vegetation (existing perennial grasses or tilled ground) and container size (36 ml or 105 ml) with excavation technique (excavation by core removal [core] or by creating an impression [dibble]) on the growth of transplants of the native perennial forbs Asclepias fascicularis and Sisyrinchium bellum. The presence of established perennial grasses reduced the growth of seeded forbs, but did not affect transplants, indicating the vulnerability of seedling forbs to interference. When compared to control plots that had been tilled in the autumn, weed canopy cover was significantly lower in the presence of perennial grasses if seeded with forbs, but not in the presence of perennial grasses alone. Both transplanted species grew better in the large container/core treatment than the small container/dibble treatment; however, existing grasses eliminated these positive effects. Asclepias fascicularis performed better when grown in large containers than in small containers, but its growth was not affected by excavation method; S. bellum performed better when planted with the core method than the dibble method of excavation, but container size made no difference. We attribute differences in the responses of the species to interactions between phenological differences and expansive clay soils that naturally de‐compact upon drying.     

Restoration of a Native Shrubland Impacted by Exotic Grasses, Frequent Fire, and Nitrogen Deposition in Southern California

Natural ecosystems globally are often subject to multiple human disturbances that are difficult to restore. A restoration experiment was done in an urban fragment of native coastal sage scrub vegetation in Riverside, California that has been subject to frequent fire, high anthropogenic nitrogen deposition, and invasion by Mediterranean annual weeds. Hand cultivation and grass‐specific herbicide were both successful in controlling exotic annual grasses and promoting establishment of seeded coastal sage scrub vegetation. There was no native seedbank left at this site after some 30 years of conversion to annual grassland, and the only native plants that germinated were the seeded shrubs, with the exception of one native summer annual. The city green‐waste mulch used in this study (C:N of 39:1) caused short‐term N immobilization but did not result in decreased grass density or increased native shrub establishment. Seeding native shrubs was successful in a wet year in this Mediterranean‐type climate but was unsuccessful in a dry year. An accidental spring fire did not burn first‐year shrubs, although adjacent plots dominated by annual grass did burn. The shrubs continued to exclude exotic grasses into the second growing season, suggesting that successful shrub establishment may reduce the frequency of the fire return interval.     

A Non-Native Prey Mediates the Effects of a Shared Predator on an Ecosystem Service

Non-native species can alter ecosystem functions performed by native species often by displacing influential native species. However, little is known about how ecosystem functions may be modified by trait-mediated indirect effects of non-native species. Oysters and other reef-associated filter feeders enhance water quality by controlling nutrients and contaminants in many estuarine environments. However, this ecosystem service may be mitigated by predation, competition, or other species interactions, especially when such interactions involve non-native species that share little evolutionary history. We assessed trophic and other interference effects on the critical ecosystem service of water filtration in mesocosm experiments. In single-species trials, typical field densities of oysters (Crassostrea virginica) reduced water-column chlorophyll a more strongly than clams (Mercenaria mercenaria). The non-native filter-feeding reef crab Petrolisthes armatus did not draw down chlorophyll a. In multi-species treatments, oysters and clams combined additively to influence chlorophyll a drawdown. Petrolisthes did not affect net filtration when added to the bivalve-only treatments. Addition of the predatory mud crab Panopeus herbstii did not influence oyster feeding rates, but it did stop chlorophyll a drawdown by clams. However, when Petrolisthes was also added in with the clams, the clams filtered at their previously unadulterated rates, possibly because Petrolisthes drew the focus of predators or habituated the clams to crab stimuli. In sum, oysters were the most influential filter feeder, and neither predators nor competitors interfered with their net effect on water-column chlorophyll. In contrast, clams filtered less, but were more sensitive to predators as well as a facilitative buffering effect of Petrolisthes, illustrating that non-native species can indirectly affect an ecosystem service by aiding the performance of a native species.     

Evaluating Ecosystem Services Provided by Non-Native Species: An Experimental Test in California Grasslands

The concept of ecosystem services – the benefits that nature provides to human''s society – has gained increasing attention over the past decade. Increasing global abiotic and biotic change, including species invasions, is threatening the secure delivery of these ecosystem services. Efficient evaluation methods of ecosystem services are urgently needed to improve our ability to determine management strategies and restoration goals in face of these new emerging ecosystems. Considering a range of multiple ecosystem functions may be a useful way to determine such strategies. We tested this framework experimentally in California grasslands, where large shifts in species composition have occurred since the late 1700''s. We compared a suite of ecosystem functions within one historic native and two non-native species assemblages under different grazing intensities to address how different species assemblages vary in provisioning, regulatory and supporting ecosystem services. Forage production was reduced in one non-native assemblage (medusahead). Cultural ecosystem services, such as native species diversity, were inherently lower in both non-native assemblages, whereas most other services were maintained across grazing intensities. All systems provided similar ecosystem services under the highest grazing intensity treatment, which simulated unsustainable grazing intensity. We suggest that applying a more comprehensive ecosystem framework that considers multiple ecosystem services to evaluate new emerging ecosystems is a valuable tool to determine management goals and how to intervene in a changing ecosystem.     

Radial Dispersion of Neighbors and the Small-Scale Competitive Impact of Two Annual Grasses on a Native Perennial Grass

In California's Mediterranean type grasslands, native perennial grasses such as Nassella pulchra are surrounded by introduced annual species and these annuals are thought to have displaced natives through much of their range. Amongst other invaders, two grasses Lolium multiflorum and Bromus hordeaceus, commonly dominate portions of the grassland with potential for N. pulchra restoration. We hypothesized that competitor species differences and small‐scale gaps (150 cm²) could be important determinants of N. pulchra survival and performance on these sites. Lolium multiflorum and B. hordeaceus were planted in 20 cm diameter circular plots at a constant rate of 1 seed per cm² surrounding newly transplanted N. pulchra plants. Nassella pulchra showed no significant effect of the species of competitor or from the distribution of the competitors. Both interspersion of patches of bare ground and separation of competitors into patches did not increase N. pulchra pre‐dawn water potential, basal area change, number of seeds produced, or average weight of seeds. The presence of L. multiflorum was associated with a decrease in N. pulchra survival compared with plots with only B. hordeaceus. Plants with increases in basal area of less than 0.75 cm² during the growing season had 74% mortality compared with no mortality in plants with more growth. However, initial N. pulchra plant size was not a good predictor of mortality. Limiting competition from annuals may increase survival of N. pulchra plantings, but 60% of the plants survived for at least 1 year, despite being transplanted into soil containing substantial annual grass seed.     

Control of Exotic Annual Grasses to Restore Native Forbs in Abandoned Agricultural Land

Exotic annual grasses are a major challenge to successful restoration in temperate and Mediterranean climates. Experiments to restore abandoned agricultural fields from exotic grassland to coastal sage scrub habitat were conducted over two years in southern California, U.S.A. Grass control methods were tested in 5 m² plots using soil and vegetation treatments seeded with a mix of natives. The treatments compared grass‐specific herbicide, mowing, and black plastic winter solarization with disking and a control. In year two, herbicide and mowing treatments were repeated on the first‐year plots, plus new control and solarization plots were added. Treatments were evaluated using percent cover, richness and biomass of native and exotic plants. Disking alone reduced exotic grasses, but solarization was the most effective control in both years even without soil sterilization, and produced the highest cover of natives. Native richness was greatest in solarization and herbicide plots. Herbicide application reduced exotics and increased natives more than disking or mowing, but produced higher exotic forb biomass than solarization in the second year. Mowing reduced grass biomass and cover in both years, but did not improve native establishment more than disking. Solarization was the most effective restoration method, but grass‐specific herbicide may be a valuable addition or alternative. Solarization using black plastic could improve restoration in regions with cool, wet summers or winter growing seasons by managing exotic seedbanks prior to seeding. While solarization may be impractical at very large scales, it will be useful for rapid establishment of annual assemblages on small scales.     

Native and Non-Native Supergeneralist Bee Species Have Different Effects on Plant-Bee Networks

Supergeneralists, defined as species that interact with multiple groups of species in ecological networks, can act as important connectors of otherwise disconnected species subsets. In Brazil, there are two supergeneralist bees: the honeybee Apis mellifera, a non-native species, and Trigona spinipes, a native stingless bee. We compared the role of both species and the effect of geographic and local factors on networks by addressing three questions: 1) Do both species have similar abundance and interaction patterns (degree and strength) in plant-bee networks? 2) Are both species equally influential to the network structure (nestedness, connectance, and plant and bee niche overlap)? 3) How are these species affected by geographic (altitude, temperature, precipitation) and local (natural vs. disturbed habitat) factors? We analyzed 21 plant-bee weighted interaction networks, encompassing most of the main biomes in Brazil. We found no significant difference between both species in abundance, in the number of plant species with which each bee species interacts (degree), and in the sum of their dependencies (strength). Structural equation models revealed the effect of A. mellifera and T. spinipes, respectively, on the interaction network pattern (nestedness) and in the similarity in bee’s interactive partners (bee niche overlap). It is most likely that the recent invasion of A. mellifera resulted in its rapid settlement inside the core of species that retain the largest number of interactions, resulting in a strong influence on nestedness. However, the long-term interaction between native T. spinipes and other bees most likely has a more direct effect on their interactive behavior. Moreover, temperature negatively affected A. mellifera bees, whereas disturbed habitats positively affected T. spinipes. Conversely, precipitation showed no effect. Being positively (T. spinipes) or indifferently (A. mellifera) affected by disturbed habitats makes these species prone to pollinate plant species in these areas, which are potentially poor in pollinators.     

Restoring Native Perennial Grasses to Rural Roadsides in the Sacramento Valley of California: Establishment and Evaluation

Along rural roadsides of the Sacramento Valley of California, we seeded native and non-native perennial grasses to gauge their potential value in roadside vegetation management programs. In trial I (polycultures), three seeded complexes and a control (resident vegetation only) were tested. Each seeded plant complex included a different mix of perennial grasses seeded into each of several roadside topographic zones. The seeded levels of plant complex were: native perennial grasses 1 (8 species); native perennial grasses 2 (13 species); and non-native perennial grasses (3 species). In trial II, plots were seeded to monocultural plots of 15 accessions of native Californian and three cultivars of non-native perennial grasses. Plots in both trials were seeded during January 1992 and evaluated for three successive years.     

Trophic Strategies of a Non-Native and a Native Amphibian Species in Shared Ponds

One of the critical factors for understanding the establishment, success and potential impact on native species of an introduced species is a thorough knowledge of how these species manage trophic resources. Two main trophic strategies for resource acquisition have been described: competition and opportunism. In the present study our objective was to identify the main trophic strategies of the non-native amphibian Discoglossus pictus and its potential trophic impact on the native amphibian Bufo calamita. We determine whether D. pictus exploits similar trophic resources to those exploited by the native B. calamita (competition hypothesis) or alternative resources (opportunistic hypothesis). To this end, we analyzed the stable isotope values of nitrogen and carbon in larvae of both species, in natural ponds and in controlled laboratory conditions. The similarity of the δ¹⁵N and δ¹³C values in the two species coupled with isotopic signal variation according to pond conditions and niche partitioning when they co-occurred indicated dietary competition. Additionally, the non-native species was located at higher levels of trophic niches than the native species and B. calamita suffered an increase in its standard ellipse area when it shared ponds with D. pictus. These results suggest niche displacement of B. calamita to non-preferred resources and greater competitive capacity of D. pictus in field conditions. Moreover, D. pictus showed a broader niche than the native species in all conditions, indicating increased capacity to exploit the diversity of resources; this may indirectly favor its invasiveness. Despite the limitations of this study (derived from potential variability in pond isotopic signals), the results support previous experimental studies. All the studies indicate that D. pictus competes with B. calamita for trophic resources with potential negative effects on the fitness of the latter.     

Restoration of Native Perennials in a California Annual Grassland after Prescribed Spring Burning and Solarization

Grasslands dominated by exotic annual grasses have replaced native perennial vegetation types in vast areas of California. Prescribed spring fires can cause a temporary replacement of exotic annual grasses by native and non‐native forbs, but generally do not lead to recovery of native perennials, especially where these have been entirely displaced for many years. Successful reintroduction of perennial species after fire depends on establishment in the postfire environment. We studied the effects of vegetation changes after an April fire on competition for soil moisture, a key factor in exotic annual grass dominance. As an alternative to fire, solarization effectively kills seeds of most plant species but with a high labor investment per area. We compared the burn to solarization in a study of establishment and growth of seeds and transplants of the native perennial grass Purple needlegrass (Nassella pulchra) and coastal sage species California sagebrush (Artemisia californica). After the fire, initial seed bank and seedling densities and regular percent cover and soil moisture (0–20 cm) data were collected in burned and unburned areas. Burned areas had 96% fewer viable seeds of the dominant annual grass, Ripgut brome (Bromus diandrus), leading to replacement by forbs from the seed bank, especially non‐native Black mustard (Brassica nigra). In the early growing season, B. diandrus dominating unburned areas consistently depleted soil moisture to a greater extent between rains than forbs in burned areas. However, B. diandrus senesced early, leaving more moisture available in unburned areas after late‐season rains. Nassella pulchra and A. californica established better on plots treated with fire and/or solarization than on untreated plots. We conclude that both spring burns and solarization can produce conditions where native perennials can establish in annual grasslands. However, the relative contribution of these treatments to restoration appears to depend on the native species being reintroduced, and the long‐term success of these initial restoration experiments remains to be determined.     

Restoration of California Native Grasses and Clovers: The Roles of Clipping,Broadleaf Herbicide,and Native Grass Density

One of the major challenges confronting grassland restoration of highly invaded communities is increasing the diversity of native species. There is surprisingly little research investigating how reconstructed native grasslands respond to common management techniques and how these techniques influence the relative establishment of both native grasses and forbs. Despite the diversity and wide distribution of native clovers in California, few practitioners incorporate them into grassland restoration plans. Conversely, non‐native clovers have been seeded extensively onto California rangelands. This study addresses the following questions: (1) Using readily available management tools, is there a strategy that can benefit the growth of both planted native bunchgrasses and seeded clovers? (2) Do native bunchgrasses compete with establishing clovers and non‐native grasses? (3) Do native and non‐native clovers differ in their response to management treatments or in their productivity? Plots were established to test three factors in different combinations over 3 years: (1) early spring clipping, (2) initial broadleaf herbicide, and (3) native bunchgrass planting density. Native and non‐native clovers were seeded in years 2 and 3. Early spring clipping did not have a significant effect on native bunchgrass cover, yet it did result in greater growth of native and non‐native clovers. The direction of the response to broadleaf herbicide changed between years for native bunchgrasses and was consistently negative for native clovers. Plots with higher native grass densities did not adversely affect the seeded clovers, yet non‐native grass cover was reduced. Native and non‐native clovers exhibited similar responses to clipping and established at similar densities.     

Reducing Competitive Suppression of a Rare Annual Forb by Restoring Native California Perennial Grasslands

Populations of the rare annual forb Amsinckia grandiflora may be declining because of competitive suppression by exotic annual grasses, and may perform better in a matrix of native perennial bunchgrasses. We conducted a field competition experiment in which Amsinckia seedlings were transplanted into forty 0.64‐m² experimental plots of exotic annual grassland or restored perennial grassland. The perennial grassland plots were restored using mature 3 cm‐diameter plants of the native perennial bunchgrass Poa secunda planted in three densities. The exotic annual grassland plots were established in four densities through manual removal of existing plants. Both grass types reduced soil water potential with increasing biomass, but this reduction was not significantly different between grass types. Both grass types significantly reduced the production of Amsinckia inflorescences. At low and intermediate densities (dry biomass per unit area of 20–80 g/m²), the exotic annual grasses reduced Amsinckia inflorescence number to a greater extent than did Poa, although at high densities (>90 g/m²) both grass types reduced the number of Amsinckia inflorescences to the same extent. The response of Amsinckia inflorescence number to Poa biomass was linear, whereas the same response to the annual grass biomass is logarithmic, and appeared to be related to graminoid cover. This may be because of the different growth forms exhibited by the two grass types. Results of this research suggest that restored native perennial grasslands at intermediate densities have a high habitat value for the potential establishment of the native annual A. grandiflora.     

Impact of Non-Native Birds on Native Ecosystems: A Global Analysis

Introduction and naturalization of non-native species is one of the most important threats to global biodiversity. Birds have been widely introduced worldwide, but their impacts on populations, communities, and ecosystems have not received as much attention as those of other groups. This work is a global synthesis of the impact of nonnative birds on native ecosystems to determine (1) what groups, impacts, and locations have been best studied; (2) which taxonomic groups and which impacts have greatest effects on ecosystems, (3) how important are bird impacts at the community and ecosystem levels, and (4) what are the known benefits of nonnative birds to natural ecosystems. We conducted an extensive literature search that yielded 148 articles covering 39 species belonging to 18 families -18% of all known naturalized species. Studies were classified according to where they were conducted: Africa, Asia, Australasia, Europe, North America, South America, Islands of the Indian, of the Pacific, and of the Atlantic Ocean. Seven types of impact on native ecosystems were evaluated: competition, disease transmission, chemical, physical, or structural impact on ecosystem, grazing/ herbivory/ browsing, hybridization, predation, and interaction with other non-native species. Hybridization and disease transmission were the most important impacts, affecting the population and community levels. Ecosystem-level impacts, such as structural and chemical impacts were detected. Seven species were found to have positive impacts aside from negative ones. We provide suggestions for future studies focused on mechanisms of impact, regions, and understudied taxonomic groups.     

Root Inputs Influence Soil Water Holding Capacity and Differentially Influence the Growth of Native versus Exotic Annual Species in an Arid Ecosystem

Invasion by exotic annual species is increasingly impacting Southern California arid lands, altering ecosystem processes and plant community composition. With climate change, the Southwestern United States is expected to experience increasingly variable rainfall. Larger rainfall events could facilitate invasion by exotic species that can capitalize on high resource conditions. Exotic annual species also have dense shallow root systems that could create positive feedbacks to further invasion by increasing soil organic matter and water holding capacity. Alternatively, fine root inputs could create negative feedbacks to exotic plant growth if they stimulate microbial nutrient immobilization. The dual influences of rainfall regime and fine root inputs on species performance were evaluated in an experiment where native and exotic species were grown individually and in combination under varying watering regimes (large infrequent or small frequent pulses, holding total rainfall constant) and root additions (with or without sterilized exotic roots). Mean soil moisture increased with larger infrequent watering events, and also with root addition. Plant growth (both native and exotic) increased with larger watering events, but declined with root addition. Exotic species growth declined more than native species growth with root additions. Mechanistically, root addition lowered inorganic nitrogen (N) availability, and microbial N immobilization increased with soil moisture content. Together these results show that increased fine root production promotes negative feedbacks to growth of exotic species via microbial N immobilization, especially under conditions of high soil moisture. Further, our results suggest that organic carbon additions are a potentially effective strategy for suppressing growth of problematic desert invaders.     

Fine-Grained Distribution of a Non-Native Resource Can Alter the Population Dynamics of a Native Consumer

New interactions with non-native species can alter selection pressures on native species. Here, we examined the effect of the spatial distribution of a non-native species, a factor that determines ecological and evolutionary outcomes but that is poorly understood, particularly on a fine scale. Specifically, we explored a native butterfly population and a non-native plant on which the butterfly oviposits despite the plant’s toxicity to larvae. We developed an individual-based model to describe movement and oviposition behaviors of each butterfly, which were determined by plant distribution and the butterfly''s host preference genotype. We estimated the parameter values of the model from rich field data. We simulated various patterns of plant distributions and compared the rates of butterfly population growth and changes in the allele frequency of oviposition preference. Neither the number nor mean area of patches of non-native species affected the butterfly population, whereas plant abundance, patch shape, and distance to the nearest native and non-native patches altered both the population dynamics and genetics. Furthermore, we found a dramatic decrease in population growth rates when we reduced the distance to the nearest native patch from 147 m to 136 m. Thus changes in the non-native resource distribution that are critical to the fate of the native herbivore could only be detected at a fine-grained scale that matched the scale of a female butterfly’s movement. In addition, we found that the native butterfly population was unlikely to be rescued by the exclusion of the allele for acceptance of the non-native plant as a host. This study thus highlights the importance of including both ecological and evolutionary dynamics in analyses of the outcome of species interactions and provides insights into habitat management for non-native species.     

Specific Daylength Effects on Leaf Growth and Dry-matter Production in High-latitude Grasses

Plants of Phleum pratense L. cv. Engmo (origin 69° N) andBromus inermis Leyss. cv. Löfar (origin 64° N) raisedin short days gave large and significant increases in d. wt,height and leaf area upon exposure to continuous light, comparedwith 8 h short days, at essentially identical daily inputs ofradiant energy (8 h summer daylight ± low intensity extension).The increases in both plant height and leaf area were causedmainly by increases in the dimensions of the leaf sheaths andblades, which, in turn, were the result of increased cell sizeand number. However, there was a pronounced interaction betweentemperature and daylength such that the greatest photoperiodicstimulation occurred at lower growing temperatures (     

Direct and Indirect Influence of Non-Native Neighbours on Pollination and Fruit Production of a Native Plant

Background

Entomophilous non-native plants can directly affect the pollination and reproductive success of native plant species and also indirectly, by altering the composition and abundance of floral resources in the invaded community. Separating direct from indirect effects is critical for understanding the mechanisms underlying the impacts of non-native species on recipient communities.

Objectives

Our aims are: (a) to explore both the direct effect of the non-native Hedysarum coronarium and its indirect effect, mediated by the alteration of floral diversity, on the pollinator visitation rate and fructification of the native Leopoldia comosa and (b) to distinguish whether the effects of the non-native species were due to its floral display or to its vegetative interactions.

Methods

We conducted field observations within a flower removal experimental setup (i.e. non-native species present, absent and with its inflorescences removed) at the neighbourhood scale.

Results

Our study illustrates the complexity of mechanisms involved in the impacts of non-native species on native species. Overall, Hedysarum increased pollinator visitation rates to Leopoldia target plants as a result of direct and indirect effects acting in the same direction. Due to its floral display, Hedysarum exerted a direct magnet effect attracting visits to native target plants, especially those made by the honeybee. Indirectly, Hedysarum also increased the visitation rate of native target plants. Due to the competition for resources mediated by its vegetative parts, it decreased floral diversity in the neighbourhoods, which was negatively related to the visitation rate to native target plants. Hedysarum overall also increased the fructification of Leopoldia target plants, even though such an increase was the result of other indirect effects compensating for the observed negative indirect effect mediated by the decrease of floral diversity.