Invasion windows for a global legume invader are revealed after joint examination of abiotic and biotic filters

• Successful alien plant invasion is influenced by both climate change and plant–plant interactions. We estimate the single and interactive effects of competition and extreme weather events on the performance of the global legume invader Lupinus polyphyllus (Lindl.).
• In three experimental studies we assessed (i) the stress tolerance of seedling and adult L. polyphyllus plants against extreme weather events (drought, fluctuating precipitation, late frost), (ii) the competitive effects of L. polyphyllus on native grassland species and vice versa, and (iii) the interactive effects of extreme weather events and competition on the performance of L. polyphyllus.
• Drought reduced growth and led to early senescence of L. polyphyllus but did not reduce adult survival. Fluctuating precipitation events and late frost reduced the length of inflorescences. Under control conditions, interspecific competition reduced photosynthetic activity and growth of L. polyphyllus. When subjected to competition during drought, L. polyphyllus conserved water while simultaneously maintaining high assimilation rates, demonstrating increased water use efficiency. Meanwhile, native species had reduced performance under drought.
• In summary, the invader gained an advantage under drought conditions through a smaller reduction in performance relative to its native competitors but was competitively inferior under control conditions. This provides evidence for a possible invasion window for this species. While regions of high elevation or latitude with regular severe late frost events might remain inaccessible for L. polyphyllus, further spread across Europe seems probable as the predicted increase in drought events may favour this non‐native legume over native species.

     

Release from belowground enemies and shifts in root traits as interrelated drivers of alien plant invasion success: a hypothesis

Our understanding of the interrelated mechanisms driving plant invasions, such as the interplay between enemy release and resource‐acquisition traits, is biased by an aboveground perspective. To address this bias, I hypothesize that plant release from belowground enemies (especially fungal pathogens) will give invasive plant species a fitness advantage in the alien range, via shifts in root traits (e.g., increased specific root length and branching intensity) that increase resource uptake and competitive ability compared to native species in the alien range, and compared to plants of the invader in its native range. Such root‐trait changes could be ecological or evolutionary in nature. I explain how shifts in root traits could occur as a consequence of enemy release and contribute to invasion success of alien plants, and how they could be interrelated with other potential belowground drivers of invasion success (allelopathy, mutualist enhancement). Finally, I outline the approaches that could be taken to test whether belowground enemy release results in increased competitive ability and nutrient uptake by invasive alien plants, via changes in root traits in the alien range.     

<Emphasis Type="Italic">Acacia</Emphasis><Emphasis Type="Italic">longifolia</Emphasis> invasion impacts vegetation structure and regeneration dynamics in open dunes and pine forests

Acacia spp. are among the most serious plant invaders worldwide, and Acacia longifolia specifically causes problems in Portugal. In this study, we evaluated the impacts of A. longifolia invasion on community structure, light climate, plant diversity and regeneration in pine forests and open stabilized dunes in northern and southern Portugal. Having the growth form of a small tree or shrub, between 1 and 8 m tall, A. longifolia tended to dominate the middle stratum of forests and to share dominance in the upper stratum of open dunes. The presence of A. longifolia was associated with a decreased canopy cover in the lower stratum of all studied habitats, and at some sites with a significantly increased leaf area index and reduced light intensity in the understory. Species number and diversity were reduced in some habitats by up to 50% in invaded compared to non-invaded areas. Furthermore, in forest habitats, A. longifolia seedlings were facilitated by proximity to an adult A. longifolia while the establishment and growth of native seedlings was negatively impacted. The replacement of drought tolerant native species by the water spending invader, A. longifolia, may have serious implications for ecosystem functioning, especially during the prolonged drought periods predicted to occur in Portugal in the future.     

Drought response of a native and introduced Hawaiian grass

The alien grass, Pennisetum setaceum, dominates many of the lowland arid regions that once supported native Heteropogon contortus grassland on the island of Hawaii. Response to drought in a glasshouse was compared between these C₄ grasses to test if success as an invader is related to drought tolerance or plasticity for traits that confer drought tolerance. Pennisetum produced 51% more total biomass, allocated 49% more biomass to leaves, and had higher net photosynthetic rates (P _n) on a leaf area basis than Heteropogon. Plants of both species under drought produced less total biomass and increased their allocation to roots compared to well-watered plants, but there was no difference between the two species in the magnitude of these responses. The decline in P _n with decreasing leaf water potential (₁) was greater for Pennisetum compared to Heteropogon. Plasticity in the response of P _n to ₁, osmotic potentials, and the water potentials at turgor loss in response to drought were not different between the two species. Stomata were more responsive to w in Heteropogon than in Pennisetum and for well-watered plants compared to droughted plants. Plasticity for the stomatal response to w, however, was not different between the species. There was no evidence that the alien, Pennisetum, had greater plasticity for traits related to drought tolerance compared to the native, Heteropogon. Higher P _n and greater biomass allocation to leaves resulted in greater growth for Pennisetum compared to Heteropogon and may explain the success of Pennisetum as an invader of lowland arid zones on Hawaii.     

Competitive displacement of the native species <Emphasis Type="Italic">Bursaphelenchus mucronatus</Emphasis> by an alien species <Emphasis Type="Italic">Bursaphelenchus xylophilus</Emphasis> (Nematoda: Aphelenchida: Aphelenchoididae): a case of successful invasion

The presence of alien invasive species has serious negative impact on endemic biodiversity, especially on native species that occupy the same niche in the ecosystem. To study the influence of the alien invasive species Bursaphelenchus xylophilus on its native sister species B. mucronatus, the two nematode species were mix-cultured in a fungal mat and mix-inoculated into a susceptible host. By comparing the propagation parameters of both species under competitive and noncompetitive conditions it was shown that the propagation level of B. xylophilus was clearly higher than that of B. mucronatus under laboratory culture. Furthermore, the propagation capacity of B. xylophilus under competitive conditions was much higher than that under noncompetitive conditions, both in laboratory culture and with host inoculation. Bursaphelenchus xylophilus also excluded B. mucronatus when the two species were cultured as a mixture for a longer time. The relative abundance ratios of the two species in natural pinewoods were also determined by random sampling of dying pine trees from regions with different invasion histories. It was noted that with an increase in invasion years the distribution frequency of B. xylophilus increased while that of B. mucronatus decreased. Experimental tests verified our hypothesis that because of its high fecundity and strong competitive ability, the invasive species B. xylophilus out-competed the native species B. mucronatus and displaced it in natural ecosystems. The successful invasion of B. xylophilus is attributed to competitive displacement, which may be one of the ecological invasive mechanisms.     

Effects of growing conditions on the growth of and interactions between salt marsh plants: implications for invasibility of habitats

A common but often less tested explanation for the successful invasion of alien species is that invasive alien species outcompete their co-occurring natives, which may not always be the case. In this study, we established artificial environmental gradients in a series of pot experiments with controlled environments to investigate the effects of salinity, sediment type and waterlogging on the performance of and interactions between Phragmites australis (native) and Spartina alterniflora (alien), which generally co-exist in the saline intertidal zones of Chinese and American coasts. Significant effects of salinity and waterlogging were detected on biomass production and morphological characteristics of S. alterniflora and P. australis, and the competitive interactions between the two species were found to vary with all three environmental factors in our experiments. Relative Neighbor Effect (RNE) analyses indicate that competitive dominance of S. alterniflora occurred under the conditions of high salinity, sandy sediment and full immersion, whereas P. australis showed competitive dominance under the conditions of low salinity and non-immersion. Our results suggest that S. alterniflora might outcompete P. australis under conditions present in early salt marsh succession, which support the viewpoint that the outcomes of competition between co-occurring native and invasive alien plants depend on the growing conditions. The implication of this study is that in response to the environmental changes expected from seawater intrusion and sea-level rise, the range of S. alterniflora is expected to expand further in the Yangtze River estuary in the future.     

Disturbance influences the outcome of plant–soil biota interactions in the invasive Acacia longifolia and in native species

Interrelated causes of plant invasion have been gaining increasing recognition. However, research on this subject has mainly focused around conceptual models. Here we explore whether plant–soil biota feedbacks and disturbance, two major factors capable of facilitating invasive plants in introduced ranges, interact to preferentially benefit exotics compared to native plants. We investigated the influence of fire disturbance on plant–soil biota interactions for the invasive Acacia longifolia and two dominant natives (Cytisus striatus and Pinus pinaster) in Portuguese dune systems. In the first experiment, we grew exotic and native plants in soil inoculated with soil biota from unburned or recently burned soils collected in an area with small invasion intensity by A. longifolia. Soil biota effects on the exotic legume A. longifolia changed from neutral to positive after fire, whereas the opposite outcome was observed in the native legume C. striatus, and a change from negative to neutral effects after fire occurred in the native P. pinaster. Fire reduced mycorrhizal colonization in all species and rhizobial colonization in C. striatus but not in A. longifolia. In the second experiment, we grew the exotic and native plants with conspecific and heterospecific soil biota from undisturbed soils (area with low invasion intensity by A. longifolia), and from post‐fire soils (area affected by a fire ～12 years ago and currently heavily invaded by A. longifolia). The exotic benefited more from post‐fire than from undisturbed soil biota, particularly from those associated with natives. Natives did not experience detrimental effects with invasive‐associated soil biota. Our results show that fire disturbance affected the functional interactions between soil biota and plants that may benefit more the exotic than some native species. Disturbance may open a window of opportunity that promotes invader success by altering soil enemy and mutualistic impacts.     

Competitive impacts and responses of an invasive weed: dependencies on nitrogen and phosphorus availability

Changes in competitive interactions under conditions of enhanced resource availability could explain the invasion success of some problematic plant species. For one invader of North American grasslands, Centaurea diffusa (diffuse knapweed), we test three hypotheses: (1) under ambient (high resource) conditions, C. diffusa is better able to tolerate competition from the resident community (competitive response), (2) under ambient conditions, C. diffusa strong impacts the competitive environment (competitive effect), and (3) reductions in nitrogen and/or phosphorus availability diminish these advantages. In support of our first hypothesis, C. diffusa was the most tolerant to neighbor competition of the four focal species under current resource conditions. In opposition to our second hypothesis, however, neighborhoods that contained C. diffusa and those where C. diffusa had been selectively removed did not differ in their impact on the performance of target transplant individuals or on resource conditions. Reduction in resource availability influenced competitive tolerance but not competitive impact, in partial support of our last hypothesis. Reduction in soil nitrogen (via sucrose carbon addition) enhanced the degree of neighbor competition experienced by all species but did not change their relative rankings; C. diffusa remained the best competitor under low nitrogen conditions. Reduction of soil phosphorus (via gypsum addition) weakened the ability of C. diffusa to tolerate neighbor competition proportionately more than the other focal species. Consequently, under low phosphorus conditions, C. diffusa lost its competitive advantage and tolerated neighbor competition similarly to the other focal species. We conclude that C. diffusa invasion may be double-edged: C. diffusa is less limited by nitrogen than the other focal species and is better able to utilize phosphorus to its competitive advantage.     

Does disturbance,competition or resource limitation underlie Hieracium lepidulum invasion in New Zealand? Mechanisms of establishment and persistence,and functional differentiation among invasive and native species

The processes underlying plant invasions have been the subject of much ecological research. Understanding mechanisms of plant invasions are difficult to elucidate from observations, yet are crucial for ecological management of invasions. Hieracium lepidulum, an asteraceous invader in New Zealand, is a species for which several explanatory mechanisms can be raised. Alternative mechanisms, including competitive dominance, disturbance of resident vegetation allowing competitive release or nutrient resource limitation reducing competition with the invader are raised to explain invasion. We tested these hypotheses in two field experiments which manipulated competitive, disturbance and nutrient environments in pre‐invasion and post‐invasion vegetation. H. lepidulum and resident responses to environmental treatments were measured to allow interpretation of underlying mechanisms of establishment and persistence. We found that H. lepidulum differed in functional response profile from native species. We also found that other exotic invaders at the sites were functionally different to H. lepidulum in their responses. These data support the hypothesis that different invaders use different invasion mechanisms from one another. These data also suggest that functional differentiation between invaders and native resident vegetation may be an important contributing factor allowing invasion. H. lepidulum appeared to have little direct competitive effect on post‐invasion vegetation, suggesting that competition was not a dominant mechanism maintaining its persistence. There was weak support for disturbance allowing initial establishment of H. lepidulum in pre‐invasion vegetation, but disturbance did not lead to invader dominance. Strong support for nutrient limitation of resident species was provided by the rapid competitive responses with added nutrients despite presence of H. lepidulum. Rapid competitive suppression of H. lepidulum once nutrient limitation was alleviated suggests that nutrient limitation may be an important process allowing the invader to dominate. Possible roles of historical site degradation and/or invader‐induced soil chemical/microbial changes in nutrient availability are discussed.     

Invader presence disrupts the stabilizing effect of species richness in plant community recovery after drought

Higher biodiversity can stabilize the productivity and functioning of grassland communities when subjected to extreme climatic events. The positive biodiversity–stability relationship emerges via increased resistance and/or recovery to these events. However, invader presence might disrupt this diversity–stability relationship by altering biotic interactions. Investigating such disruptions is important given that invasion by non‐native species and extreme climatic events are expected to increase in the future due to anthropogenic pressure. Here we present one of the first multisite invader × biodiversity × drought manipulation experiment to examine combined effects of biodiversity and invasion on drought resistance and recovery at three semi‐natural grassland sites across Europe. The stability of biomass production to an extreme drought manipulation (100% rainfall reduction; BE: 88 days, BG: 85 days, DE: 76 days) was quantified in field mesocosms with a richness gradient of 1, 3, and 6 species and three invasion treatments (no invader, Lupinus polyphyllus, Senecio inaequidens). Our results suggest that biodiversity stabilized community productivity by increasing the ability of native species to recover from extreme drought events. However, invader presence turned the positive and stabilizing effects of diversity on native species recovery into a neutral relationship. This effect was independent of the two invader's own capacity to recover from an extreme drought event. In summary, we found that invader presence may disrupt how native community interactions lead to stability of ecosystems in response to extreme climatic events. Consequently, the interaction of three global change drivers, climate extremes, diversity decline, and invasive species, may exacerbate their effects on ecosystem functioning.     

The invasion of plant communities following extreme weather events under ambient and elevated temperature

Although the problem of plant invasions is expected to increase with climate change, there is as yet little experimental evidence, in particular, for the effects of extreme weather events. We established communities of European meadow species, which were subjected to warming and extreme event (drought and deluge) treatments in a factorial design at an experimental garden in Zurich, Switzerland. Phylogenetically matched pairs of native and alien species (Bromus erectus, B. inermis, Trifolium pratense, T. hybridum, Lactuca serriola, and Conyza canadensis) were introduced into the communities to test if invader performance is favored by warming and extreme events, and if alien invaders perform better than native colonizers. With a warming of on average 0.3?°C, a higher cover of native plant communities was observed, while drought decreased cover in the short-term and lowered biomass. Germination, survival, and growth of the introduced species were lower under elevated temperature. Survival of all pairs and growth of Trifolium was greater in drought pots, while deluge had no effect. While the alien species showed a faster rate of increase in the number of leaves, mortality of alien species was greater than of native species. Overall, the performance of the focal species varied much more among taxonomic groups than native/alien provenances. The results suggest that with climate change, different types of extreme events will differ in the severity of their effects on native plant communities. Meanwhile, the effects of climate change on plant invasions are more likely to operate indirectly through the impacts on native vegetation.     

From plant traits to invasion success: Impacts of the alien Fallopia japonica (Houtt.) Ronse Decraene on two native grassland species

Alien invasive plants threaten biodiversity, productivity and ecosystem functioning throughout the world. We examined the effect of Fallopia japonica on two native grassland species (Trifolium repens, Lolium perenne). We hypothesized that its negative effects on the native species are dependent on three mechanisms: (i) allelochemicals released and accumulated in soil with a history of invasion, (ii) altered soil biota and (iii) direct resource competition. We measured the response of the native species as the difference in their functional traits when grown under the three conditions. Our results demonstrate that neither allelochemicals nor soil biota from soil with history of F. japonica invasion had measurable effects on either species. Competition with the invader strongly reduced height, biomass and specific leaf area (SLA) of T. repens, while it had a lower effect on L. perenne. Furthermore, our results reveal that F. japonica took advantage of a positive plant–soil and plant–plant interaction. The results show that the prominent mechanism underpinning the invasion success of F. japonica in the grassland was the direct resource competition. This prominent role is also confirmed by the significant interactions between competition, allelochemicals and soil biota from soils with history of invasion of F. japonica on SLA of the native species.     

Native woody legumes respond more negatively to extreme drought than invasive ones

本地木本豆科植物比外来入侵木本豆科植物对极端干旱事件的负面响应更强 植物入侵可能会受到极端干旱事件增加和大气氮沉降的影响。然而,极端干旱、氮沉降及其相互作用如何影响外来木本豆科植物的成功入侵尚不清楚。本研究选取3种入侵木本豆科植物和3种本地木本豆科植物,开展温室控制实验。分别将这些植物种植在不同干旱处理(极端干旱和对照)和不同氮水平(低氮和高氮)下,然后对比分析两类植物的株高、叶片数、生物量以及根质量分数的差异。研究结果表明,极端干旱对本地木本豆科植物生长的抑制效应强于入侵木本豆科植物。尽管土壤氮素有效性的增加降低了植物的根质量分数,但这并不影响植物的整体表现。入侵木本豆科植物比本地木本豆科植物更能耐受长期极端干旱的负面效应。本研究加深了我们对气候变化导致的干旱事件如何影响外来木本豆科植物入侵的了解。     

Competition strength of two significant invasive species in coastal dunes

To investigate the effect of increased nutrient availability on competition amongst invasive and native plants, I measured changes in above and below ground biomass of Chrysanthemoides monilifera spp. rotundata (bitou bush) and Asparagus aethiopicus (asparagus fern) competing with two native species, Banksia integrifolia and Ficinia nodosa, under high- and low-nutrient regimes. Bitou bush, as a primary invader, was competitive under all conditions lowering the growth of native species in both high and low nutrients. Asparagus fern as a secondary invader, did not influence growth of native species but responded, like bitou bush, to high nutrients. Native species were generally negatively affected by increases in nutrients. With bitou bush soils often providing higher nutrients, the chance of secondary invasion by asparagus fern is more likely, although asparagus fern is unlikely to invade low nutrient soils quickly. The invasive species, therefore, differed in their competitive ability in these coastal dune communities.     

Tolerance to drought and flooding events provides a competitive advantage for an invasive over a native plant species

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Competitive reversals and environment-dependent resource partitioning in Erodium

Summary Effects of drought and varying plant density on the competitive coexistence of two winter annual Erodium species were studied using multiple regression analysis. Significant indications of resource partitioning were detected for interspecific mixtures under spring drought. Competitive superiority also was environment-dependent with E. botrys dominating with drought in autumn, while E. brachycarpum dominated with drought in spring. The results suggest that competitive coexistence in Erodium is promoted by processes both equilibrial (e.g., resource partitioning) and nonequilibrial (e.g. competitive reversals).     

Invasion strategies of Cynodon dactylon: Competitive ability under low-nutrient conditions

Cynodon dactylon is one of the five most important invasive alien species worldwide. It is the invasive alien species with the broadest distribution range in Uruguay, and its expansion is frequently associated with disturbances. Since natural grasslands are facing processes of productive intensification, C. dactylon represents a threat as it could displace native species. However, the mechanisms that explain its invasion success remain unclear. The objective of this study was to analyse interspecific interactions under low nutrient conditions between C. dactylon and two species that are native to the Campo grasslands in Uruguay. Specifically, we assessed differences in the components of competitive ability effects and responses (or tolerance) as possible mechanisms involved in C. dactylon invasiveness. We performed a greenhouse experiment in pots with low-nutrient substrate assessing pair-wise interactions between C. dactylon, Mnesithea selloana and Paspalum notatum plus control pots consisting of single individual of each species. The invasive species showed greater competitive ability than both native grasses, as it reduced their below and above-biomass. Conversely, the size of C. dactylon plants interacting with native species was similar to that of single C. dactylon plants growing alone (controls). This reveals that the greater competitive ability of the invasive species was due to a greater tolerance to grow with neighbouring plants. The reason underlying this tolerance was a marked increase in biomass allocation towards stolons and leaves, at the expense of roots. Conversely, native species barely changed their shoot-root allocation pattern when interacting with neighbours. Furthermore, C. dactylon induced reproductive development solely when interacting with neighbours. Along with the fact that the potential growth rate of the invasive and native species was quite similar, these results suggest that sensitive and rapidly triggered shade avoidance responses could be one mechanism involved in the invasion success of C. dactylon.     

Measurement of volatile terpene emissions in 70 dominant vascular plant species in Hawaii: aliens emit more than natives

Aim Alien plant invasion is prominent in the Hawaiian Islands. There are many factors involved in invader success. To date, there is a general lack of information about one of them, which we aim to study here: the terpene emission capacity of both Hawaiian native and alien plants. Location Oahu (Hawaii). Methods We screened 35 alien and 35 native dominant plant species on Oahu Island for monoterpene emissions. The emission rates were measured from field‐grown plants under standardized conditions of temperature and quantum flux density in the laboratory. Results The emission rates of total terpenes ranged from 0 µg g^?1 h^?1 to 55 µg g^?1 h^?1, and altogether 15 different terpenes were emitted in detectable amounts by the overall set of species. A phylogenetic signal was observed for total terpene emissions. Total terpene emission rates were higher in aliens than in native species (12.8 ± 2.0 vs. 7.6 ± 1.9 µg g^?1 h^?1, respectively). Main conclusions The greater terpene emission capacity may confer protection against multiple stresses and may partly account for the success of the invasive species, and may make invasive species more competitive in response to new global change‐driven combined stresses. These results are consistent with aliens coming from very diverse ecosystems with generally higher biotic and abiotic stress pressures, and having higher nutrient concentrations. On the contrary, these results are not consistent with the ‘excess carbon’ hypotheses. These results indicate changes in vegetation terpene emissions brought about by alien plant invasions.     

Native species richness buffers invader impact in undisturbed but not disturbed grassland assemblages

Many systems are prone to both exotic plant invasion and frequent natural disturbances. Native species richness can buffer the effects of invasion or disturbance when imposed in isolation, but it is largely unknown whether richness provides substantial resistance against invader impact in the face of disturbance. We experimentally examined how disturbance (drought/burning) influenced the impact of three exotic invaders (Centaurea stoebe, Linaria dalmatica, or Potentilla recta) on native abundance across a gradient of species richness, using previously constructed grassland assemblages. We found that invaders had higher cover in experimentally disturbed plots than in undisturbed plots across all levels of native species richness. Although exotic species varied in cover, all three invaders had significant impacts on native cover in disturbed plots. Regardless of disturbance, however, invader cover diminished with increasing richness. Invader impacts on native cover also diminished at higher richness levels, but only in undisturbed plots. In disturbed plots, invaders strongly impacted native cover across all richness levels, as disturbance favoured invaders over native species. By examining these ecological processes concurrently, we found that disturbance exacerbated invader impacts on native abundance. Although diversity provided a buffering effect against invader impact without disturbance, the combination of invasion and disturbance markedly depressed native abundance, even in high richness assemblages.     

Disturbance governs dominance of an invasive forb in a temporary wetland

Dominance of invasive species is often assumed to be due to a superior ability to acquire resources. However, dominance in plant communities can arise through multiple interacting mechanisms, including disturbance. Inter-specific competition can be strongly affected by abiotic conditions, which can determine the outcome of competitive interactions. We evaluated competition and disturbance as mechanisms governing dominance of Phyla canescens (hereafter lippia), an invasive perennial forb from South America, in Paspalum distichum (perennial grass, hereafter water couch) meadows in floodplain wetlands of eastern Australia. Water couch meadows (in the study area) are listed under the Ramsar Convention due to their significance as habitat for migratory waterbirds. In the field, we monitored patterns of vegetation boundaries between the two species in response to flooding. Under controlled glasshouse conditions, we explored competitive interactions between the native water couch and lippia subject to different soil moisture/inundation regimes. We did this using a pairwise factorial glasshouse experiment that manipulated neighbor density (9 treatments) and soil moisture/inundation (4 treatments). In the field trial, inundation increased the cover of water couch. Under more controlled conditions, the invader had a competitive effect on the native species only under dry soil conditions, and was strongly inhibited by inundation. This suggests that dry conditions favor the growth of the invader and wetter (more historical) conditions favor the native grass. In this system, invader dominance is governed by altered disturbance regimes which give the invader a competitive advantage over the native species.