Parasitism by Cuscuta gronovii mediated soil legacy effects and the competitive ability of invasive and native plant species by changing soil abiotic and biotic properties

Impact of invasive alien plants Gutenbergia cordifolia and Tagetes minuta on native taxa in the Ngorongoro crater, Tanzania

Invasive plant species often competitively displace native plant species but some populations of native plant species can evolve adaptation to competition from invasive plants and persist in invaded habitats. However, studies are lacking that examine how variation in abiotic conditions in invaded landscapes may affect fitness of native plants that have adapted to compete with invasive plants. I tested whether invasion by Parthenium hysterophorus in Nairobi National Park - Kenya may have selected for native plant individuals with greater competitive ability than conspecific naïve natives in nutrient-rich and mesic soil conditions. I compared vegetative growth and seed yields of invader-experienced and conspecific naïve individuals of seven native species. Invader-experienced natives grew shorter than naïve natives regardless of growth conditions. Nevertheless, the two groups of native plants also exhibited treatment-specific differences in competitive ability against P. hysterophorus. Invader-experienced natives displayed plasticity in seed yield under drought treatment, while naïve natives did not. Moreover, drought treatment enhanced competitive effects of invader-experienced natives on P. hysterophorus, while nutrient enrichment relaxed competitive effects of invader-experienced natives on the invader. The results suggest that P. hysterophorus may have selected for shorter native plant genotypes that also exhibit plasticity in competitive ability under drought conditions.

Insights into ecological drivers of alien plant invasions can be gained through comparative studies of growth and fecundity of invasive alien plants versus those of co‐occurring non‐invasive alien plants and native plants across environmental conditions in common garden settings. Habitats that harbour alien plant species in many ecosystems globally are presently experiencing light pollution resulting from artificial light at night (ALAN) and increased rates of nutrient enrichment of the soil. However, the potential interactive effects of ALAN and nutrient enrichment on invasiveness of alien plant species remain unknown. Here, we performed a common‐garden experiment to test the interactive effects of ALAN and soil nutrient enrichment on the growth of a random set of ten alien (five invasive and five non‐invasive naturalized) and seven co‐occurring native ornamental plant species that are commonly cultivated within urban and peri‐urban areas of Nairobi city in Kenya. We predicted that a simultaneous increase in photoperiod via ALAN and nutrient enrichment will favour growth of invasive alien plant species over that of non‐invasive alien and native plant species. We grew the 17 plant species under natural daylight (ALAN−) versus natural daylight followed by ALAN (ALAN+) and fully crossed with two levels of nutrient enrichment (low vs. high) and competition (competition vs. no‐competition against a native plant Ocimum gratissimum ) treatments. Under simultaneous high‐nutrient and no‐competition treatments, ALAN enhanced mean total biomass of invasive and non‐invasive naturalized alien species by 61.1% and 131.4%, respectively but decreased that of native plant species by 34%. In contrast, under simultaneous high‐nutrient and competition treatments, ALAN enhanced mean total biomass of invasive alien plant species by 68.6% and that of non‐invasive naturalized alien species by 51.9% and native species by 35.4%. High‐nutrient treatment enhanced flower formation more strongly in invasive and non‐invasive naturalized alien plants than in native plants. The invasive and non‐invasive naturalized alien species grew taller than native species across the light, nutrient, and competition treatments. Synthesis : The present findings suggest that light pollution and nutrient enrichment may jointly confer growth advantage to invasive alien plant species over that of co‐occurring native plant species and enhance invasiveness of alien plant species.

Human influence on the environment is so extensive that virtually all ecosystems on the planet are now affected by biological invasions. And, often, ecosystems are invaded by multiple co‐occurring non‐native species. Hence, it is important to understand the impacts these invasions are producing on biodiversity and ecosystem processes.Here, we present results of a 2‐year long field experiment where we tested the effects of co‐occurring invasive C4African grasses in a Cerrado area in central Brazil. We compared plant and arthropod communities, plant biomass, and soil nitrogen dynamics and soil chemical characteristics across five experimental treatments:Urochloa decumbensremoval;Melinis minutifloraremoval; bothU.decumbensandM.minutifloraremoval;U.decumbensandM.minutiflorainvaded plots; and uninvaded Cerrado. We hypothesized that selective removal of invasive grasses would have distinct effects on the native ecosystem structure and functioning. We expected that each invasive grass would produce a different type of impact on the native ecosystem and that their impacts would be synergistic when co‐occurring.Removal ofM.minutifloradoubled native plant diversity and biomass when compared to invaded plots, whereas removal ofU.decumbensdid not alter these parameters. Cerrado plots had four times more plant species than plots cleared of invasives. Removal of invasive grasses did not affect the species richness or community composition of soil epigeal fauna. Cerrado soils had lower fertility, organic matter content and pH than invaded soils. The effects were generally higher when both invasive grasses were removed, suggesting impacts were synergistic, butM.minutiflorahad greater effects on plants and soils thanU.decumbens. Both invasive species produced negative impacts, but a single species was the main driver. We also detected persistent effects of the invasive grass species on the ecosystem after 2 years of removal.We conclude that invasive species of the same functional group have similar types of effects in native ecosystems, but the magnitude of impact was largely dependent on invasive species biomass and cover. Where multiple invasive species are present, research and management of invaded ecosystems should tackle the interacting effects of co‐occurring invaders.

<p>Drivers of global change have direct impacts on the structure of communities and functioning of ecosystems, and interactions between drivers may buffer or exacerbate these direct effects. Interactions among drivers can lead to complex non-linear outcomes for ecosystems, communities and species, but are infrequently quantified. Through a combination of experimental, observational and modelling approaches, I address critical gaps in our understanding of the interactive effects of climate change and plant invasion, using Tongariro National Park (TNP; New Zealand) as a model. TNP is an alpine ecosystem of cultural significance which hosts a unique flora with high rates of endemism. TNP is invaded by the perennial shrub Calluna vulgaris (L.) Hull. My objectives were to: 1) determine whether species-specific phenological shifts have the potential to alter the reproductive capacity of native plants in landscapes affected by invasion; 2) determine whether the effect of invasion intensity on the Species Area Relationship (SAR) of native alpine plant species is influenced by environmental stress; 3) develop a novel modelling framework that would account for density-dependent competitive interactions between native species and C. vulgaris and implement it to determine the combined risk of climate change and plant invasion on the distribution of native plant species; and 4) explore the possible mechanisms leading to a discrepancy in C. vulgaris invasion success on the North and South Islands of New Zealand. I show that species-specific phenological responses to climate warming increase the flowering overlap between a native and an invasive plant. I then show that competition for pollination with the invader decreases the sexual reproduction of the native in some landscapes. I therefore illustrate a previously undescribed interaction between climate warming and plant invasion where the effects of competition for pollination with an invader on the sexual reproduction of the native may be exacerbated by climate warming. Furthermore, I describe a previously unknown pattern of changing invasive plant impact on SAR along an environmental stress gradient. Namely, I demonstrate that interactions between an invasive plant and local native plant species richness become increasingly facilitative along elevational gradients and that the strength of plant interactions is dependent on invader biomass. I then show that the consequences of changing plant interactions at a local scale for the slope of SAR is dependent on the pervasion of the invader. Next, I demonstrate that the inclusion of invasive species density data in distribution models for a native plant leads to greater reductions in predicted native plant distribution and density under future climate change scenarios relative to models based on climate suitability alone. Finally, I find no evidence for large-scale climatic, edaphic, and vegetative limitations to invasion by C. vulgaris on either the North and South Islands of New Zealand. Instead, my results suggest that discrepancies in invasive spread between islands may be driven by human activity: C. vulgaris is associated with the same levels of human disturbance on both islands despite differences in the presence of these conditions between then islands. Altogether, these results show that interactive effects between drivers on biodiversity and ecosystem dynamics are frequently not additive or linear. Therefore, accurate predictions of global change impacts on community structure and ecosystems function require experiments and models which include of interactions among drivers such as climate change and species invasion. These results are pertinent to effective conservation management as most landscapes are concurrently affected by multiple drivers of global environmental change.</p>

<p>Drivers of global change have direct impacts on the structure of communities and functioning of ecosystems, and interactions between drivers may buffer or exacerbate these direct effects. Interactions among drivers can lead to complex non-linear outcomes for ecosystems, communities and species, but are infrequently quantified. Through a combination of experimental, observational and modelling approaches, I address critical gaps in our understanding of the interactive effects of climate change and plant invasion, using Tongariro National Park (TNP; New Zealand) as a model. TNP is an alpine ecosystem of cultural significance which hosts a unique flora with high rates of endemism. TNP is invaded by the perennial shrub Calluna vulgaris (L.) Hull. My objectives were to: 1) determine whether species-specific phenological shifts have the potential to alter the reproductive capacity of native plants in landscapes affected by invasion; 2) determine whether the effect of invasion intensity on the Species Area Relationship (SAR) of native alpine plant species is influenced by environmental stress; 3) develop a novel modelling framework that would account for density-dependent competitive interactions between native species and C. vulgaris and implement it to determine the combined risk of climate change and plant invasion on the distribution of native plant species; and 4) explore the possible mechanisms leading to a discrepancy in C. vulgaris invasion success on the North and South Islands of New Zealand. I show that species-specific phenological responses to climate warming increase the flowering overlap between a native and an invasive plant. I then show that competition for pollination with the invader decreases the sexual reproduction of the native in some landscapes. I therefore illustrate a previously undescribed interaction between climate warming and plant invasion where the effects of competition for pollination with an invader on the sexual reproduction of the native may be exacerbated by climate warming. Furthermore, I describe a previously unknown pattern of changing invasive plant impact on SAR along an environmental stress gradient. Namely, I demonstrate that interactions between an invasive plant and local native plant species richness become increasingly facilitative along elevational gradients and that the strength of plant interactions is dependent on invader biomass. I then show that the consequences of changing plant interactions at a local scale for the slope of SAR is dependent on the pervasion of the invader. Next, I demonstrate that the inclusion of invasive species density data in distribution models for a native plant leads to greater reductions in predicted native plant distribution and density under future climate change scenarios relative to models based on climate suitability alone. Finally, I find no evidence for large-scale climatic, edaphic, and vegetative limitations to invasion by C. vulgaris on either the North and South Islands of New Zealand. Instead, my results suggest that discrepancies in invasive spread between islands may be driven by human activity: C. vulgaris is associated with the same levels of human disturbance on both islands despite differences in the presence of these conditions between then islands. Altogether, these results show that interactive effects between drivers on biodiversity and ecosystem dynamics are frequently not additive or linear. Therefore, accurate predictions of global change impacts on community structure and ecosystems function require experiments and models which include of interactions among drivers such as climate change and species invasion. These results are pertinent to effective conservation management as most landscapes are concurrently affected by multiple drivers of global environmental change.</p>

Herbivores can profoundly influence plant species assembly, including plant invasion, and resulting community composition. Population increases of native herbivores, e.g. white-tailed deer (Odocoileus virginianus), combined with burgeoning plant invasions raise concerns for native plant diversity and forest regeneration. While individual researchers typically test for the impact of deer on plant invasion at a few sites, the overarching influence of deer on plant invasion across regional scales is unclear. We tested the effects of deer on the abundance and diversity of introduced and native herbaceous and woody plants across 23 white-tailed deer research sites distributed across the east-central and north-eastern USA and representing a wide range of deer densities and invasive plant abundance and identity. Deer access/exclusion or deer population density did not affect introduced plant richness or community-level abundance. Native and total plant species richness, abundance (cover and stem density) and Shannon diversity were lower in deer-access vs. deer-exclusion plots. Among deer-access plots, native species richness, native and total cover, and Shannon diversity (cover) declined as deer density increased. Deer access increased the proportion of introduced species cover (but not of species richness or stem density). As deer density increased, the proportion of introduced species richness, cover and stem density all increased. Because absolute abundance of introduced plants was unaffected by deer, the increase in proportion of introduced plant abundance is likely an indirect effect of deer reducing native cover. Indicator species analysis revealed that deer access favoured three introduced plant species, including Alliaria petiolata and Microstegium vimineum, as well as four native plant species. In contrast, deer exclusion favoured three introduced plant species, including Lonicera japonica and Rosa multiflora, and 15 native plant species. Overall, native deer reduced community diversity, lowering native plant richness and abundance, and benefited certain invasive plants, suggesting pervasive impacts of this keystone herbivore on plant community composition and ecosystem services in native forests across broad swathes of the eastern USA.

Native plant communities are often invaded by multiple alien species. It is still unclear how increasing diversity of alien invasive species suppresses the growth of native species and thus contributes to invasion success. In the subtropical monsoon region of southeast China, we experimentally created a native plant community with 18 herbaceous species. One week later, we let it be invaded by either zero (controls without invasion), one, two, four or eight alien plant species, with either high or low species evenness. After a four‐month growth period we harvested the aboveground biomass of each species. We found that increasing invasive species richness significantly increased invasive plant biomass, the biomass of all invasive and native plant species within the community, and invasion success (the ratio of invasive plant biomass to the biomass of all native and invasive plants), but it did not significantly reduce native plant biomass. Experimentally manipulating invasive species evenness did not influence invasion success and did not show any differential suppression effects on native plants. One invasive species, Sesbania cannabina , became dominant in terms of plant biomass, irrespective of its proportion in the alien plant mixtures. Throughout this experiment, effects of invasive species richness on invasion success were mainly due to such selection effects among the invasive species. On the other hand, the unchanged biomass of native species under increasing invasive plant richness suggests the presence of at least partly complementary resource niches between invasive and native species.

Subsequent to escape from intense herbivory in the native range, invasive plants are expected to reduce allocation to costly anti‐herbivory defences and have greater competitive ability than co‐occurring native species. Given that invasive alien plants often occur in open habitats where light is less limited, it is reasonable to hypothesize that invasive plants should express high concentrations of gibberellins that enable them to allocate more biomass to roots, and thus have higher competitive ability than native plants. To test this prediction, we grew five congeneric pairs of invasive alien plants and native plants under two levels of nutrient availability (low versus high) and treated a half of the plants with a gibberellin biosynthesis inhibitor, paclobutrazol. Paclobutrazol significantly decreased aboveground, belowground and total biomass of the test plants. Interestingly, the effects on belowground biomass were significantly stronger for invasive plants than for native plants. A similar pattern was found for total biomass (marginally significant effect p = 0.0592). Additionally, paclobutrazol decreased root mass fraction for invasive plants, but tended to increase it for native plants. Our findings suggest that plant hormones can differently regulate biomass allocation of invasive and native plants, and thus contribute to greater growth of invasive plants compared to native plants.

SummaryConcerns over the ecological impacts of invasive alien plant species have generated great research interest in understanding the mechanisms that underlie the capacity of such plants to occupy a broad range of habitats. It has been repeatedly suggested that rapid evolution of local adaptation to novel environments may enable invasive plants to occupy a broad range of habitats. However, the classical Darwinian view on evolution by natural selection is that the process is slow and gradual, occurring over thousands of years. Invasive plants typically have a relatively short residence time in their introduced ranges (decades or just a few centuries). Besides the time constraint, founder effects (reduction in population size and genetic diversity) may also limit the capacity of invasive plants to rapidly evolve local adaption. Thus, invasive plants may be less likely than native plants to evolve local adaptation. Interestingly, however, an expanding body of literature documents the existence of local adaptation in invasive plant species within their exotic ranges.Here, we did a phylogenetically controlled meta‐analysis to compare invasive and native plant species for differences in the frequency and magnitude of local adaptation. The meta‐analysis was based on different experiments performed in various habitats including grasslands, steppes, deserts, forests, mountains, wetlands and dunes, and used a total of 134 plant species in 52 families. Forty seven of these species (in 24 families) are alien invaders in the region where the studies were undertaken, while the other 91 species (in 38 families) are native.On average, local plants performed better than foreign plants, and invasive plant species expressed local adaptation just as frequently, and at least as strongly as that exhibited by native plant species. An analysis performed while taking into account different plant life‐history traits showed that self‐incompatible invasive plants exhibited significantly higher frequencies of local adaptation than native plants characterized by the same breeding system.Synthesis. The present results support the suggestion that rapid evolution of local adaptation may enable invasive plant species to occupy a broad range of novel habitats.

Large parts of the Earth are experiencing environmental change caused by alien plant invasions, rising atmospheric concentration of carbon dioxide (CO2 ), and nutrient enrichments. Elevated CO2 and nutrient concentrations can separately favour growth of invasive plants over that of natives but how herbivory may modulate the magnitude and direction of net responses by the two groups of plants to simultaneous CO2 and nutrient enrichments remains unknown. In line with the enemy release hypothesis, invasive plant species should reallocate metabolites from costly anti-herbivore defences into greater growth following escape from intense herbivory in the native range. Therefore, invasive plants should have greater growth than native plants under simultaneous CO2 and nutrient enrichments in the absence of herbivory. To test this prediction, we grew nine congeneric pairs of invasive and native plant species that naturally co-occurred in grasslands in China under two levels each of nutrient enrichment (low-nutrient vs. high-nutrient), herbivory (with herbivory vs. without herbivory) and under ambient (412.9 ± 0.6 ppm) and elevated (790.1 ± 6.2 ppm) levels of CO2 concentrations in open top chambers in a common garden. Elevated CO2 and nutrient enrichment separately increased total plant biomass, while herbivory reduced it regardless of the plant invasive status. High-nutrient treatment caused the plants to allocate a significantly lower proportion of total biomass to roots, while herbivory induced an opposite pattern. Herbivory suppressed total biomass production more strongly in native plants than invasive plants. The plants exhibited significant interspecific and intergeneric variation in their responses to the various treatment combinations. Overall, these results suggest that elevated CO2 and nutrients and herbivory may separately, rather than synergistically, impact productivity of the invasive and co-occurring native plant species in our study system. Moreover, interspecific variation in resource-use strategies was more important than invasive status in determining plant responses to the various treatment combinations.

An invasive and native plant differ in their effects on the soil food-web and plant-soil phosphorus cycle

Summary 1. A large proportion of the world's land surface is extensively managed for livestock production. In areas where livestock systems are becoming more intensive, a major challenge is to predict those plant species likely to decline, persist or increase as a result of agricultural intensification. 2. Most analyses develop inferences for frequent or abundant species, or rely on intensive studies of single species. A promising approach is to identify plant traits related to disturbance to enable inference to be made about changes in plant community composition. We used a Bayesian hierarchical model to analyse the response to agricultural intensification of 494 plant species of pastures and woodlands in southern Australia, and to identify how simple species' traits (life form, growth form and species origin) influence those responses. 3. The probability of occurrence of most species declined along the two intensification gradients, grazing intensity and soil phosphorous concentration, although the occurrence of a greater proportion of species was negatively correlated with soil phosphorous. Responses could be broadly predicted from both plant origin and plant traits, in particular growth form. 4. Native perennial geophytes, ferns and shrubs were most negatively affected by both gradients, while exotic annual grasses and forbs were more tolerant. Along the phosphorous gradient, 24 of the 30 most negatively affected plant species were native geophytes. Mean within-group responses masked considerable within- and between-species variation, particularly for the exotic species group which included species that responded both negatively and positively to intensification. 5. Synthesis and applications. The hierarchical model described here provides a powerful method for estimating individual plant responses and identifying how species' traits influence those responses. Plant species native to southern Australia are sensitive to grazing and phosphorous apparently due to a shared evolutionary history of low grazing intensity and low phosphorous soils. Invading exotic plants have faced strongly contrasting ecological filters, leading to a greater diversity of responses. Where grazing systems have been most intense, a small suite of exotics dominate. Maintaining native and functional plant diversity will necessitate limits being placed on intensive livestock management systems.

The Hawaiian Islands have lost nearly all their native seed dispersers, but have gained many frugivorous birds and fleshy-fruited plants through introductions. Introduced birds may not only aid invasions of exotic plants but also may be the sole dispersers of native plants. We assessed seed dispersal at the ecotone between native- and exotic-dominated forests and quantified bird diets, seed rain from defecated seeds, and plant distributions. Introduced birds were the primary dispersers of native seeds into exotic-dominated forests, which may have enabled six native understory plant species to become reestablished. Some native plant species are now as common in exotic forest understory as they are in native forest. Introduced birds also dispersed seeds of two exotic plants into native forest, but dispersal was localized or establishment minimal. Seed rain of bird-dispersed seeds was extensive in both forests, totaling 724 seeds of 9 native species and 2 exotics with over 85% of the seeds coming from native plants. Without suitable native dispersers, most common understory plants in Hawaiian rainforests now depend on introduced birds for dispersal, and these introduced species may actually facilitate perpetuation, and perhaps in some cases restoration, of native forests. We emphasize, however, that restoration of native forests by seed dispersal from introduced birds, as seen in this study, depends on the existence of native forests to provide a source of seeds and protection from the effects of ungulates. Our results further suggest that aggressive control of patches of non-native plants within otherwise native-dominated forests may be an important and effective conservation strategy.

Invasive, non-native plant species often outcompete native species and reduce biodiversity. Invasive plants frequently begin growth before native plants, yet few studies have examined whether invasives win in competition partly by colonizing disturbed sites more quickly or by beginning growth earlier in the season than native plants (i.e. due to priority effects). We hypothesized that invasive plant species would benefit more from priority effects than would comparable native species and that earlier growth of invasive species would decrease plant biodiversity. To test this hypothesis, we grew three pairs of invasive and native plant species from three different functional groups/plant families (C3 grasses/Poaceae, non-leguminous forbs/Asteraceae, and legumes/Fabaceae). We seeded each of the species 3 weeks before seeding the other five species into large pots in a greenhouse. Consistent with our hypothesis, we found much stronger priority effects with invasive than native species. Each invasive species formed a near-monocultures when seeded first (97.5 % of total biomass, on average) whereas native species did not similarly dominate (29.8 % of total biomass, on average). Similarly, Simpson’s species diversity was 81 % higher when the initially sown species was native rather than invasive. The literature suggests that invasive species in the field often begin growth earlier in the spring than native species and that climate change may increasingly allow invasives to begin growth before native species, indicating invasive priority effects may become increasingly common.