Effects of nutrient addition and clipping on biomass production of invasive and native annual Asteraceae plants

Invasive plants dominate many habitats and cause significant harm to native plants, agriculture and the wider environment. Clipping by mowing machines has been a common method to control invasive plants, but whether it can reduce the performance of invaders and increase the performance of co‐occurring natives is not known. This study examined the effects of soil nutrient conditions and clipping on the performance of three invasive and three co‐occurring native annual Asteraceae plants under either isolated or competitive conditions. Clipping tended to reduce the aboveground biomass of the invasive plants more than that of the native plants under low, but not under high nutrient conditions. However, invasive plants still produced an aboveground biomass equal to, or larger than, that of native plants under the low nutrient and clipping treatment combination. Nutrient addition increased the aboveground biomass of the invasive plants more than that of the native plants, although this was only marginally significant under competitive conditions. Clipping did not reduce the performance advantage of invaders over natives under high soil nutrient conditions. The ability of invaders to benefit more from nutrient addition, along with its ability to perform equally well, or better than, the native plants after human mechanical control (i.e. clipping), may explain their high abundance and dominance in many habitats.     

Evidence for recent evolution in an invasive species,Microstegium vimineum,Japanese stiltgrass

The establishment and spread of invasive plants are often associated with a ‘general‐purpose genotype’, with a corresponding high degree of phenotypic plasticity when introduced to a new environment. Evolutionary potential of invasive species following introduction should also be considered, however, as such changes can facilitate rapid range expansion. In this study, we utilised seed from three geographically distinct mid‐Atlantic populations of an invasive species, Microstegium vimineum (Japanese stiltgrass), to assess whether populations varied in their biomass and reproductive responses to light and nitrogen under neutral glasshouse conditions. To assess evolutionary adaptation, we quantified and correlated the abiotic environment from each collection location with the final biomass and seed production of that population under common glasshouse conditions. For M. vimineum, growth and reproductive responses to a common set of abiotic parameters (e.g. light) under these conditions indicated considerable phenotypic variation between and within populations. Interestingly, the degree of variation of seed production and final biomass among populations in the glasshouse was strongly correlated with growing season length and cumulative degree days recorded near each population location. These latter data indicated that M. vimineum, introduced into this region <100 years ago, may be undergoing adaptive evolution. Although additional populations of M. vimineum need to be examined, this study suggests that local evolutionary adaptation may be occurring rapidly and could, potentially, be an important aspect in the establishment and spread of this invasive species.     

Performance of the invasive Eupatorium catarium and Ageratum conyzoides in comparison with a common native plant under varying levels of light and moisture

Invasive alien plants are often found to be more plastic than the natives, which enable the former to outperform the native plants under some, or in a wide range, of environmental conditions. In addition, their performance under non‐competitive conditions might not predict their performance under competitive conditions. The performance of the recent invader, Eupatorium catarium, was compared with that of a historical invader, Ageratum conyzoides, and a common native plant, Vernonia cinerea, under varying levels of light and moisture with and without competition. When grown alone and in response to decreases in light levels, E. catarium and V. cinerea were more plastic in their root‐to‐shoot ratio than A. conyzoides, while V. cinerea was more plastic in its specific leaf area (SLA) than E. catarium and A. conyzoides. Eupatorium catarium was more plastic than A. conyzoides and V. cinerea in the SLA in response to the drought treatment. However, these differences in trait plasticities did not lead to a difference in biomass production under each light and moisture treatment combination. When E. catarium was competing with either A. conyzoides or V. cinerea, the interacting species also did not differ in their biomass production and relative interaction intensity under each light and moisture treatment combination. These results indicate that the two invasive species and the native plant do not differ in their performance under relatively low soil nutrient conditions, regardless of the moisture, light and competition treatments that are imposed. Therefore, E. catarium and A. conyzoides cannot outperform V. cinerea under low nutrient conditions, regardless of the light and water availability.     

Ecophysiological factors contributing to the invasion of Panicum maximum into native Miscanthus sinensis grassland in Taiwan

Panicum maximum (guinea grass), a native grass of Africa, has invaded and displaced the native Miscanthus sinensis grassland at Dadu terrace in central Taiwan. The Dadu terrace has distinct wet and dry seasons and is prone to fire during the dry seasons. We compared photosynthesis and growth, as well as the growth response to two water treatments, between P. maximum and M. sinensis plants to understand ecophysiological factors contributing to the successful invasion of the grass. In comparison with M. sinensis, P. maximum had a significantly higher photosynthetic rate, larger specific leaf area and a higher leaf area/total biomass ratio, and thus grew faster and higher, and accumulated more biomass under well‐watered conditions. The growth of P. maximum was more susceptible than M. sinensis to drought stress. Faster growth of seedlings and more biomass accumulation of mature plants would give P. maximum a highly competitive advantage over M. sinensis for resource acquisition in the Dadu terrace during the wet season, which might contribute to the successful invasion of P. maximum into the grassland of M. sinensis.     

Distribution and abundance of invasive Tamarix genotypes in South Africa

The exotic Tamarix chinensis and T. ramosissima, believed to have been introduced into South Africa in the early 1900s to control erosion on mine dumps, are invading riparian zones and have been proven to hybridise with T. usneoides, which is native to southern Africa. In this study, we document the abundance of invasive Tamarix genotypes in South Africa. Eleven riparian zones from the Northern, Eastern and Western Cape Provinces were surveyed. Three quadrats of 600 m² each were selected per site. Plant density, canopy cover and tree height were recorded to quantify invasiveness. Leaf samples were randomly collected from an average of eight individuals per site to record genotypes of the invaders. Tamarix density and canopy cover were significantly greater than those of co‐occurring trees and shrubs in Olifants River in De Rust (Western Cape Province). A linear correlation between percentage Tamarix spp. cover and other co‐occurring tree and shrub species showed a strong negative relationship (R² = 0.78). Genetic analysis showed that the Western and Eastern Cape Provinces have the highest proportion of the exotic Tamarix species and their hybrids. This suggests that these two provinces require urgent management intervention to contain the spread of the weed. The distinctions made between the native and the exotic Tamarix species and their hybrids should also facilitate the testing and future release of potential biological control agents.     

Plant invasiveness and target plant density: high densities of native Schima wallichii seedlings reduce negative effects of invasive Ageratina adenophora

Economically feasible strategies to cope with invasive species are urgently needed. Plant density can be increased to reduce competitive effects on target plants. This study indicates that increasing native plant density can be used to reduce the effect of invasive Ageratina adenophora. Seedlings of an indigenous tree species, Schima wallichii, were grown in pots containing uninvaded or invaded soil, with or without A. adenophora leaf litter on the soil surface. Schima wallichii seedlings were also grown at four densities under four levels of A. adenophora leaf litter. Root and shoot biomass and length were measured as response parameters in both bioassays. Schima wallichii growth was inhibited by A. adenophora leaf litter and invaded soil. High litter levels reduced S. wallichii root length and dry weight at low plant densities. The inhibition disappeared at high S. wallichii plant densities. As A. adenophora did not inhibit S. wallichii growth at high plant densities, adjustments of seedling density should be studied as a possible management strategy for invasion by A. adenophora and potentially by other exotic plant species. As density‐dependent growth inhibition is the key characteristic of chemical interference, we propose that phytotoxins contribute to A. adenophora invasion particularly at low densities of native seedlings.     

Growth and photosynthetic responses of invasive Bidens frondosa to light and water availability: A comparison with invasive and native congeners

Bidens frondosa is a new invasive species in China. However, the mechanisms underlying its invasiveness are still not understood. This study compared the new invader with its invasive congener, Bidens pilosa, and two native congeners, Bidens biternata and Bidens tripartita, in terms of their morphology, growth, biomass allocation and photosynthetic parameters across variant light and water conditions, aiming to explore the traits that are associated with the invasiveness of the invasive species. All these species are annual herbs. The results showed that the four congeners had a similar tendency of plasticity in their morphological and growth traits to varying light and water, but their response was quantitatively different. Most of the morphological and growing variables, as well as the photosynthetic parameters, were affected more strongly by manipulated light levels. Although B. frondosa had a lower total biomass and relative growth rate than the invasive congener, B. pilosa, the trait values related to growth (e.g. plant height, total biomass, relative growth rate and specific leaf area) and the mean value of the plasticity index were higher for the invasive B. frondosa than for the native B. tripartita. These could contribute to the invasiveness of the invasive species. In addition, the native species, B. biternata, is a vigorous weed with a strong adaptive capacity across various light and water gradients, suggesting that it has the potential to be invasive if introduced into areas with suitable conditions for growth.     

Competitive traits of the invasive grass Arundo donax are enhanced by carbon dioxide and nitrogen enrichment

Increased atmospheric carbon dioxide (CO₂) and soil nitrogen (N) may confer competitive advantages to invasive species over native plant species. We conducted a two‐way factorial experiment, growing the model invasive plant Arundo donax in CO₂ growth chambers to test how CO₂ and N availability interact to affect plant growth and biomass allocation. CO₂ was supplied at c. 400 mg L^?1 and c. 750 mg L^?1. N was supplied as ammonium nitrate at 640, 320 and 80 mg L^?1. We hypothesised that the fertilisation effect of CO₂ enrichment would offset limitations from N deficiency. The results indicated that A. donax plants grown with enriched CO₂ and abundant N accumulated approximately 50–100% more biomass and allocated approximately 50% more biomass to rhizomes than plants grown under ambient CO₂ conditions. Neither treatment affected the leaf area per unit mass (specific leaf area; SLA). Greater growth will likely increase A. donax's competitive potential, because increased rhizome biomass has been associated with increased stress tolerance and post‐disturbance resprouting capacity. The consistent SLA under all treatments suggests that A. donax has a morphological strategy that prioritises increasing leaf quantity over increasing individual leaves' photosynthetic potential. These results reveal the ecological strategies that contribute to the successful establishment, dominance and persistence of this invasive plant species.     

Increased free amino acid contents of the invasive weed Ageratina adenophora and the effects on its specialist herbivore Procecidochares utilis

In order to verify the escape‐from‐enemy hypothesis from the changes of nutrient substance and fitness of natural enemies on alien plants, contents of free amino acids in native and invasive plant populations of Ageratina adenophora and life history parameters of specialist herbivore Procecidochares utilis reared on these plants were investigated. Our results showed that the contents of glycine, valine, γ‐aminobutyric acid, proline, serine, alanine, and arginine in the invasive plants were higher than those in the native plants of A. adenophora. There was a shorter developmental duration and higher fecundity of P. utilis when fed on the invasive plants. The results indicated a possible fitness tradeoff of natural enemies between invasive and native plants arose from nutrient substance changes.     

Shade-induced plasticity in invasive Impatiens glandulifera populations

Phenotypic plasticity, which confers a fitness advantage under heterogeneous and novel environments, has been commonly suggested to contribute to the success of invasive plants in their introduced range. For example, plasticity in response to changes in light availability could facilitate invasiveness by allowing plants to both rapidly establish in unshaded, disturbed habitats, and tolerate shaded or crowded environments. The plastic responses of invasive plants to shade were mostly studied in morphological traits. However, plasticity in physiological traits might provide more rapid and reversible responses and thus be more effective in environments with rapid temporal variations. Here, we compared plasticity in a range of morphological and physiological traits that provide shade avoidance or tolerance between two native and six introduced populations of Impatiens glandulifera. In a common garden, we subjected second-generation siblings of native and invasive plants to two light availability treatments and measured their morphological, physiological and performance responses. Impatiens glandulifera from invasive populations exhibited greater phenotypic plasticity in response to light availability in four out of 12 measured traits. Moreover, this greater plasticity was mostly limited to physiological traits associated with photosynthetic acclimation. These results suggest high phenotypic plasticity in response to light availability could have facilitated I. glandulifera's spread in both disturbed habitats and woodlands or under intense light competition. The results of this study highlight the importance of considering physiological traits when studying the role of plasticity in the success of invasive plants.     

Assessing wind and mammals as seed dispersal vectors in an invasive legume

While some plants have modified seed structures to facilitate dispersal, many lack such specialised adaptations, making their mode of dispersal unclear. This can be particularly problematic for predicting shifts in species ranges or tracking the spread of invasive plants. As an example, the seed size and shape of the invasive legume, Lespedeza cuneata, suggest that wind and attachment to animals are not important for dispersal, yet populations can spread surprising distances within a few years. Using a series of experiments conducted in the laboratory and field, we tested wind and mammal fur as mediators of seed dispersal. To test wind dispersal, seed traps were arranged radially around a patch of L. cuneata and seeds were collected following dispersal. Attachment to mammal fur was tested by fitting pelts of a deer, coyote and raccoon to artificial torsos and determining seed retention in both the field and the laboratory. Laboratory trials also examined the influence of wet versus dry conditions. Our results showed that wind direction strongly influenced dispersal distance and seeds were readily dispersed by mammal fur. The number of seeds retained was species specific, depending on fur depth and mammal size, with seed retention increasing under wet conditions. Together, these results suggest that both wind and mammal fur contribute to the movement of L. cuneata across grasslands. Consequently, both dispersal vectors should be considered when designing and implementing control strategies.     

Phenological dynamics of the invasive plant Acacia longifolia in Portugal

Phenological shifts may play an important role in the success of invasive species. In the coastal regions of continental Portugal, Acacia longifolia is one of the most widespread invasive plants, but there are significant gaps in our understanding and interpretation of its phenology. This study is the first to investigate the phenological variation of A. longifolia and its association with climate in populations representing a temperature and precipitation gradient in mainland Portugal. The results highlighted significant variations in the vegetative (leaf initiation) and reproductive phenophases (bud formation, flowering and fruiting) among populations. Overall, leafing was observed throughout the year, bud formation occurred between summer and spring, anthesis between late winter and the beginning of spring, and fruiting started in late winter. Most of the reproductive phenophases varied across elevation or latitudinal gradients, with populations that were subjected to more stressful climatic conditions advancing their flowering and fruiting phenophases. The occurrence of each phenophase was strongly associated with the climatic conditions in the previous 5–12 months, suggesting that plants receive their phenological cues well in advance of their phenological response. Among climatic factors, temperature and irradiance were the most significant predictors of the phenological cycle of A. longifolia, but precipitation also exerted a greater influence on budding and flowering of the species. The phenological response of A. longifolia in a variety of sites represents an important step towards understanding how its phenology may react to the projected climate change in the Mediterranean region.     

Impact of <Emphasis Type="Italic">Sugarcane yellow leaf virus</Emphasis> on growth and sugar yield of sugarcane

A survey revealed that Sugarcane yellow leaf virus (SCYLV) is found on all Hawaiian sugarcane plantations including those where no yellow leaf symptoms were observed. In a comparison of growth and yield between SCYLV-infected and SCYLV-free plants of the cultivar H87-4094, germination and early shoot growth of infected plants were retarded. The number of stalks per stool was reduced by 30%, biomass was reduced by 29%, and sugar yield by 26% when plants were harvested after 11 months. Yields did not decrease when plants were harvested after 2 years. Thus, SCYLV could reduce yield, even when the plants were asymptomatic. In a field test of SCYLV-susceptible (infected) and -resistant cultivars to compare growth and yield, 10 commercial cultivars (six susceptible and four resistant to SCYLV) were grown in eight fields with different climates and soils. Primary stalk length, biomass and sugar yield did not differ between susceptible and resistant cultivars under any field conditions. Thus, harmful effects of SCYLV on yield cannot be deduced by comparing different cultivars.     

Pathogenicity and host‐parasite relationships of the root‐knot nematode Meloidogyne incognita on celery

Pathogenicity and host‐parasite relationships in root‐knot disease of celery (Apium graveolens ) caused by Meloidogyne incognita race 1 were studied under glasshouse conditions. Naturally and artificially infected celery cv. D’elne plants showed severe yellowing and stunting, with heavily deformed and damaged root systems. Nematode‐induced mature galls were spherical and/or ellipsoidal and commonly contained more than one female, males and egg masses with eggs. Feeding sites were characterized by the development of giant cells that contained granular cytoplasm and many hypertrophied nuclei. The cytoplasm of giant cells was aggregated along their thickened cell walls and consequently the vascular tissues within galls appeared disrupted and disorganized. The relationship between initial nematode population density (Pi) and growth of celery plants was tested in glasshouse experiments with inoculum levels that varied from 0 to 512 eggs and second‐stage juveniles (J₂) mL^?1 soil. Seinhorst's model y = m + (1 – m)z^P–T was fitted to height and top fresh weight data of the inoculated and control plants. The tolerance limit with respect to plant height and fresh top weight of celery to M. incognita race 1 was estimated as 0·15 eggs and J₂ mL^?1 soil. The minimum relative values (m) for plant height and top fresh weight were 0·37 and 0·35, respectively, at Pi ≥ 16 eggs and J₂ mL^?1 soil. The maximum nematode reproduction rate (Pf/Pi) was 407·6 at an initial population density (Pi) of 4 eggs and J₂ mL^?1 soil.     

The Efficacy of Grafting on Alkali Stressed Watermelon Cultivars Under Hydroponic Conditions

The goal of the present study was to determine whether grafting of watermelon on gourd rootstocks could improve alkalinity tolerance and to investigate the physiological and morphological response mechanisms of the grafted plants under different pH levels. The experiment was carried out in a climate chamber to investigate growth, leaf chlorophyll content (SPAD), leaf area, stem length, shoot and root dry weight, root length, electrolyte leakage, leaf mineral composition, total chlorophyll and carotenoid contents. Two watermelon cultivars (Crimson tide, CT, and Crisby) were grafted onto three commercial Cucurbita maxima?×?C. moschata hybrid rootstocks under climate-chamber conditions (Strong tosa, ST, Ercole and Nun 9075). The grafted seedlings were transplanted onto 8?L continuously aerated pots containing nutrient solution with two different pH levels (8.5 and 6.5) and replicated three times. The results showed that in both grafted and non-grafted plants, there was a substantial reduction in shoot and root biomass production at high pH levels. At high pH level, the significantly highest leaf area, stem length, SPAD, concentration of P, Ca, S and Mn in leaf tissues were recorded in graft combination ‘Crisby/Ercole’, whereas the significantly highest concentration of Fe in leaf tissues, shoot dry weight were recorded in graft combination ‘Crisby/Nun 9075’. Moreover, at high pH, the significantly highest concentration of Mg and Cu in shoot under high pH levels was significantly found in graft combination of ‘CT/ST’. These results suggest that the use of interspecific Cucurbita maxima?×?C. moschata hybrid rootstocks can improve crop performance in watermelon plants under alkaline conditions.

     

Germination ecology of Emex spinosa and Emex australis,invasive weeds of winter crops

Emex spinosa and Emex australis are invasive dicotyledonous weeds. The effects of various environmental factors on the germination of these weeds were investigated under laboratory and glasshouse conditions. Germination response of both species was lower at warmer temperature, and maximum germination was recorded at 20/12°C (day/night). Light stimulated germination in both species, but considerable germination also occurred under darkness. More than 80% of E. spinosa seeds germinated at pH between 6 and 9, whereas E. australis seeds germination was considerably decreased at pH 9. Emex spinosa was fairly tolerant to salinity as compared with E. australis and germination (21%) of E. spinosa occurred even at 200 mm NaCl. Both species were sensitive to osmotic stress, but E. spinosa tolerated more osmotic stress than E. australis. Temperature above 20/12°C (day/night) and low osmotic potential increased time to start germination and mean germination time (MGT), as well as decreased germination index (GI) of both species. Darkness resulted in increased MGT and decreased GI in both species when compared with 10 h photoperiod. Salt stress strongly increased time to obtain 50% germination and reduced GI of both species. In both species, an increasing burial depth decreased emergence percentage and emergence index and increased time to start emergence, although some seed emerged even at 10 cm burial depth. It was concluded that both species can germinate over a wide range of environmental conditions. However, E. australis was more sensitive under adverse environmental conditions compared with E. spinosa. This information on germination ecology may aid in developing tools and strategies for management.     

Environmental and irrigation conditions can mask the effect of Magnaporthiopsis maydis on growth and productivity of maize

Maize production in temperate countries is threatened by late wilt, caused by Magnaporthiopsis maydis. Plant infection occurs early after sowing, but symptoms appear from flowering onwards. The disease is mainly controlled by genetic resistance, which is often partially expressed in the field. Development of disease symptoms is also highly dependent on environmental conditions. This study looked at whether production and growth of susceptible maize are affected by M. maydis under environmental conditions that are suboptimal for disease development. In addition, the effect of water availability on disease development under optimal conditions was determined. Pot experiments were conducted in an open-air enclosure in 2013, 2015 and 2016. Under unfavourable conditions for disease (low air temperature and relatively high air humidity), aboveground symptoms did not appear in the plants despite growth and production variables being clearly altered by the fungus. When air temperatures and humidity were optimal for disease development (air temperatures relatively high and humidity rather low), leaf symptoms on inoculated plants became apparent but with secondary importance compared to decreases in growth and production. The pathogen also affected the root:aboveground biomass ratio to a greater extent when the plants were under good water conditions than under deficit irrigation. Under optimal conditions and with good soil water content, the infected crop may end its cycle without symptoms, with the disease undetected, although reductions in yield and aboveground biomass can occur.     

Status of invasive alien plants included in EPPO Lists in Turkey

Turkey is one of the richest areas in the middle latitudes in terms of plant diversity. It has approximately 12 000 plant species and a great number of new species are being added each year. However, invasive alien species have not yet been fully considered in Turkey and only two plant species are in Turkey's quarantine list (Arceuthobium spp. and Eichhornia crassipes). EPPO is an intergovernmental organization responsible for cooperation in plant protection in the European and Mediterranean region and maintains the A1 and A2 Lists of pests recommended for regulation (A1 species are absent from the EPPO region, A2 species are present but of limited distribution), the List of Invasive Alien Plants, the Observation List of invasive alien plants and the Alert List. Eichhornia crassipes, Heracleum persicum, Heracleum sosnowskyi, Ludwigia peploides, Polygonum perfoliatum and Solanum elaeagnifolium are among the species which were recorded in Turkey and are included in the EPPO A2 List. Species recorded in the EPPO List of invasive alien plants which are present in Turkey are: Acroptilon repens, Ailanthus altissima, Ambrosia artemisiifolia (= A. elatior), Carpobrotus edulis, Cortaderia selloana, Cyperus esculentus, Paspalum distichum (= P. paspalodes), Oxalis pes‐caprae and Sicyos angulatus. Azolla filiculoides and Rhododendron ponticum are listed in the EPPO Observation List of Invasive alien plants and Miscanthus sinensis, listed in the EPPO Alert List, are also recorded in the Turkish flora.     

Effect of phosphate‐based seed priming on strigolactone production and Striga hermonthica infection in cereals

Strigolactones, plant‐secreted underground signalling molecules, play an important role in agricultural ecosystems, because they mediate the interaction of crops with symbiotic AM fungi and parasitic weeds like Striga hermonthica. Cereal host plants secret these signalling molecules particularly under nutrient‐deficient conditions and especially when phosphate (P) is limiting. The objective of the present study was to see the potential of P seed priming for S. hermonthica management in cereals in relation to strigolactone production. It has been demonstrated that P fertiliser application down‐regulates the production of these signalling molecules in the rhizosphere, which results in lower S. hermonthica infection of cereals. The laboratory study showed maximum production of strigolactones from dry and water‐soaked seeds, while seed soaking in P solution reduced their production. Similarly, maximum S. hermonthica infection was observed under control treatments with dry sowing or water soaking, while P seed soaking decreased S. hermonthica germination, emergence and dry biomass in all cereal crops. Our study shows that P seed priming resulted in lower exudation of strigolactones, which induced less S. hermonthica seeds germination and hence may lead to lower S. hermonthica infection. P‐based seed priming could prove to be an effective and affordable strategy to reduce S. hermonthica infection in cereals. Further research for practical field application is needed.     

Considering the preferences for nitrogen forms by invasive plants: a case study from a hydroponic culture experiment

Although preference for NH₄⁺, NO₃^? or a combination of the two often differs among species, we know little about the responses of invasive plants to different inorganic N forms. Furthermore, many studies have suggested that an increase in N availability may facilitate further invasions. However, most of these studies predicted the positive feedback without considering the preference for N forms of invasive plants. Therefore, we cultivated four common invasive species (Mikania micrantha, Ipomoea cairica, Wedelia trilobata and Bidens pilosa) in South China with hydroponic media containing different forms of N (i.e. NO₃^?, NH₄NO₃ and NH₄⁺) at equimolar concentrations. Our results showed that the N forms significantly affected the growth, biomass allocation and physiological traits of the plants. All four invasive plants supplied with NO₃^? alone had better performance and greater allocation to root biomass than did plants that were supplied with NH₄⁺ alone. Moreover, the photosynthetic rate, pigment content and photosystem II activity of plants supplied with NO₃^? or NH₄NO₃ were significantly higher than those of plants supplied with NH₄⁺alone. The results suggested that all four invasive plants preferred NO₃^? rather than NH₄⁺, and changes in NO₃^? played an important role in furthering the invasions of these plants than did changes in NH₄⁺. Our results implied that decreasing NO₃^? may be a useful tool for controlling and managing invasive plants preferring NO₃^?. In addition, this study highlighted the importance of considering plant N form preference to better understand plant invasions.