An invasive aster (Ageratina adenophora) invades and dominates forest understories in China: altered soil microbial communities facilitate the invader and inhibit natives

Exotic plant invasion may alter underground microbial communities, and invasion-induced changes of soil biota may also affect the interaction between invasive plants and resident native species. Increasing evidence suggests that feedback of soil biota to invasive and native plants leads to successful exotic plant invasion. To examine this possible underlying invasion mechanism, soil microbial communities were studied where Ageratina adenophora was invading a native forest community. The plant–soil biota feedback experiments were designed to assess the effect of invasion-induced changes of soil biota on plant growth, and interactions between A. adenophora and three native plant species. Soil analysis showed that nitrate nitrogen (NO₃⁻-N), ammonium nitrogen (NH₄⁺-N), and available P and K content were significantly higher in a heavily invaded site than in a newly invaded site. The structure of the soil microbial community was clearly different in all four sites. Ageratina adenophora invasion strongly increased the abundance of soil VAM (vesicular-arbuscular mycorrhizal fungi) and the fungi/bacteria ratio. A greenhouse experiment indicated that the soil biota in the heavily invaded site had a greater inhibitory effect on native plant species than on A. adenophora and that soil biota in the native plant site inhibited the growth of native plant species, but not of A. adenophora. Soil biota in all four sites increased A. adenophora relative dominance compared with each of the three native plant species and soil biota in the heavily invaded site had greater beneficial effects on A. adenophora relative dominance index (20% higher on average) than soil biota in the non-invaded site. Our results suggest that A. adenophora is more positively affected by the soil community associated with native communities than are resident natives, and once the invader becomes established it further alters the soil community in a way that favors itself and inhibits natives, helping to promote the invasion. Soil biota alteration after A. adenophora establishment may be an important part of its invasion process to facilitate itself and inhibit native plants.     

A Panax-centric view of invasive species

Plant-centric sampling provides a novel approach to quantifying the potential impact of invasive species on native plant species. The aim of this study was to determine the level of exposure of individuals and populations of Panax quinquefolius to invasive plant species using this approach in thirty natural ginseng populations. A high level of invasion was found with 63–70% of ginseng populations containing at least one invasive species. Approximately one-third of all individuals were found in close proximity to invasive plants. The most prevalent invasive species were Rosa multiflora and Berberis thunbergii. The exposure to invasives of plants in different size classes varied among populations. Invasive species presence increased with greater ginseng population sizes and presence of harvest. The abundance of invasives plants within forest interiors near this valuable medicinal herb suggests that the economic and ecological costs of competitive interactions with native species could be high.     

Effects of invasive plants on fire regimes and postfire vegetation diversity in an arid ecosystem

We assessed the impacts of co‐occurring invasive plant species on fire regimes and postfire native communities in the Mojave Desert, western USA. We analyzed the distribution and co‐occurrence patterns of three invasive annual grasses (Bromus rubens, Bromus tectorum, and Schismus spp.) known to alter fuel conditions and community structure, and an invasive forb (Erodium cicutarium) which dominates postfire sites. We developed species distribution models (SDMs) for each of the four taxa and analyzed field plot data to assess the relationship between invasives and fire frequency, years postfire, and the impacts on postfire native herbaceous diversity. Most of the Mojave Desert is highly suitable for at least one of the four invasive species, and 76% of the ecoregion is predicted to have high or very high suitability for the joint occurrence of B. rubens and B. tectorum and 42% high or very high suitability for the joint occurrence of the two Bromus species and E. cicutarium. Analysis of cover from plot data indicated two or more of the species occurred in 77% of the plots, with their cover doubling with each additional species. We found invasive cover in burned plots increased for the first 20 years postfire and recorded two to five times more cover in burned than unburned plots. Analysis also indicated that native species diversity and evenness as negatively associated with higher levels of relative cover of the four invasive taxa. Our findings revealed overlapping distributions of the four invasives; a strong relationship between the invasives and fire frequency; and significant negative impacts of invasives on native herbaceous diversity in the Mojave. This suggests predicting the distributions of co‐occurring invasive species, especially transformer species, will provide a better understanding of where native‐dominated communities are most vulnerable to transformations following fire or other disturbances.     

Native and exotic Amphipoda and other Peracarida in the River Meuse: new assemblages emerge from a fast changing fauna

Samples issued from intensive sampling in the Netherlands (1992–2001) and from extensive sampling carried out in the context of international campaigns (1998, 2000 and 2001) were revisited. Additional samples from artificial substrates (1992–2003) and other techniques (various periods) were analysed. The combined data provide a global and dynamic view on the Peracarida community of the River Meuse, with the focus on the Amphipoda. Among the recent exotic species found, Crangonyx pseudogracilisis regressing, Dikerogammarus haemobaphesis restricted to the Condroz course of the river, Gammarus tigrinusis restricted to the lowlands and seems to regress, Jaera istriis restricted to the ‘tidal’ Meuse, Chelicorophium curvispinumis still migrating upstream into the Lorraine course without any strong impact on the other amphipod species. After a rapid expansion Dikerogammarus villosushas continued its upstream invasion between 1998 and 2002 at a rate of 30–40 km per year, but no further progression was noticed in 2003. Locally and temporarily the native species (Gammarus fossarum and G. pulex) and naturalized species (G. roeseliand Echinogammarus berilloni)mayhave been excluded by the most recent invaders (mainly D. villosus), but none of the native and naturalized species has disappeared completely. Therefore, the number of amphipod species found in the River Meuse has increased. Moreover, the native and naturalized species keep on dominating the tributaries from which the recent invaders seem to be excluded. A changing Peracarida community structure is observed along the course of the River Meuse: four native or naturalized species inhabit the upstream (Lorraine) course, three invasive species dominate in the middle reach (Ardenne-Condroz zone), one exotic species is housed in the Border Meuse and three or four invasive species dominate the assemblages in the lowlands.     

An invasive dandelion unilaterally reduces the reproduction of a native congener through competition for pollination

The impact of invasive alien species on native species is of increasing global concern. Invasive plants have various negative effects on natives through competition; however, relatively little is known about competition for pollination. The relationship between Japanese native dandelions (Taraxacum spp.) and invasive congeners may be a typical case of such an interaction. For example, native dandelions are being replaced by invasive congeners, especially in urban and suburban areas of Japan. To explain this phenomenon, we hypothesized that when natives are mixed with attractive invasives, natives may suffer from reduced seed set because invasives deprive natives of pollinators or because pollinators frequently move between species, resulting in interspecific pollen transfer. To test this hypothesis, we studied the effect of the invasive dandelion T. officinale on the pollination and seed set of the native T. japonicum using artificial arrays of monospecific and mixed-species plots as well as natural populations. Taraxacum officinale attracted more pollinator visits, perhaps because it produced more nectar than T. japonicum. The number of pollinator visits to T. japonicum was reduced when the congeners were grown together, and pollinators moved frequently between the two species. The proportion of seed set for T. japonicum was reduced in the presence of T. officinale in both artificial arrays and natural populations. These results support our hypothesis that interspecific competition for pollination plays an important role in the recent replacement of native dandelions by invasive congeners in Japan. Because invasive dandelions are apomicts, negative effects are incurred only by sexual natives. Thus, this system can be recognized as a rare case of interspecific interaction through pollination.     

Interactions among invaders: community and ecosystem effects of multiple invasive species in an experimental aquatic system

With ecosystems increasingly supporting multiple invasive species, interactions among invaders could magnify or ameliorate the undesired consequences for native communities and ecosystems. We evaluated the individual and combined effects of rusty crayfish (Orconectes rusticus) and Chinese mystery snails [Bellamya (=Cipangopaludina) chinensis] on native snail communities (Physa, Helisoma and Lymnaea sp.) and ecosystem attributes (algal chlorophyll a and nutrient concentrations). Both invaders are widespread in the USA and commonly co-occur within northern temperate lakes, underscoring the importance of understanding their singular and joint effects. An outdoor mesocosm experiment revealed that while the two invaders had only weakly negative effects upon one another, both negatively affected the abundance and biomass of native snails, and their combined presence drove one native species to extinction and reduced a second by >95%. Owing to its larger size and thicker shell, adult Bellamya were protected from crayfish attack relative to native species (especially Physa and Lymnaea), suggesting the co-occurrence of these invaders in nature could have elevated consequences for native communities. The per capita impacts of Orconectes (a snail predator) on native snails were substantially greater than those of Bellamya (a snail competitor). Crayfish predation also had a cascading effect by reducing native snail biomass, leading to increased periphyton growth. Bellamya, in contrast, reduced periphyton biomass, likely causing a reduction in growth by native lymnaeid snails. Bellamya also increased water column N:P ratio, possibly because of a low P excretion rate relative to native snail species. Together, these findings highlight the importance of understanding interactions among invasive species, which can have significant community- and ecosystem-level effects.     

Contrasting effects of invasive plants in plant–pollinator networks

The structural organization of mutualism networks, typified by interspecific positive interactions, is important to maintain community diversity. However, there is little information available about the effect of introduced species on the structure of such networks. We compared uninvaded and invaded ecological communities, to examine how two species of invasive plants with large and showy flowers (Carpobrotus affine acinaciformis and Opuntia stricta) affect the structure of Mediterranean plant–pollinator networks. To attribute differences in pollination to the direct presence of the invasive species, areas were surveyed that contained similar native plant species cover, diversity and floral composition, with or without the invaders. Both invasive plant species received significantly more pollinator visits than any native species and invaders interacted strongly with pollinators. Overall, the pollinator community richness was similar in invaded and uninvaded plots, and only a few generalist pollinators visited invasive species exclusively. Invasive plants acted as pollination super generalists. The two species studied were visited by 43% and 31% of the total insect taxa in the community, respectively, suggesting they play a central role in the plant–pollinator networks. Carpobrotus and Opuntia had contrasting effects on pollinator visitation rates to native plants: Carpobrotus facilitated the visit of pollinators to native species, whereas Opuntia competed for pollinators with native species, increasing the nestedness of the plant–pollinator network. These results indicate that the introduction of a new species to a community can have important consequences for the structure of the plant–pollinator network.     

Reduced performance of native infauna following recruitment to a habitat-forming invasive marine alga

Despite well-documented negative impacts of invasive species on native biota, evidence for the facilitation of native organisms, particularly by habitat-forming invasive species, is increasing. However, most of these studies are conducted at the population or community level, and we know little about the individual fitness consequences of recruitment to habitat-forming invasive species and, consequently, whether recruitment to these habitats is adaptive. We determined the consequences of recruitment to the invasive green alga Caulerpa taxifolia on the native soft-sediment bivalve Anadara trapezia and nearby unvegetated sediment. Initially, we documented the growth and survivorship of A. trapezia following a natural recruitment event, to which recruitment to C. taxifolia was very high. After 12 months, few clams remained in either habitat, and those that remained showed little growth. Experimental manipulations of recruits demonstrated that all performance measures (survivorship, growth and condition) were significantly reduced in C. taxifolia sediments compared to unvegetated sediments. Exploration of potential mechanisms responsible for the reduced performance in C. taxifolia sediments showed that water flow and water column dissolved oxygen (DO) were significantly reduced under the canopy of C. taxifolia and that sediment anoxia was significantly higher and sediment sulphides greater in C. taxifolia sediments. However, phytoplankton abundance (an indicator of food supply) was significantly higher in C. taxifolia sediments than in unvegetated ones. Our results demonstrate that recruitment of native species to habitat-forming invasive species can reduce growth, condition and survivorship and that studies conducted at the community level may lead to erroneous conclusions about the impacts of invaders and should include studies on life-history traits, particularly juveniles.     

3种菊科入侵植物不同生长时期的土壤酶活性和养分变化

【目的】黄顶菊、三叶鬼针草和豚草是我国危害较严重的3种菊科入侵植物。了解3种菊科植物生长过程中土壤养分和酶活性的变化,可以为研究其入侵机制提供依据。【方法】在中国农业科学院植物保护研究所廊坊中试基地开展同质园实验,比较分析了黄顶菊、三叶鬼针草和豚草3种菊科入侵植物在幼苗期、旺盛生长期和生殖生长期根际土壤养分和酶活性的变化。【结果】3种外来菊科植物的生长时期对土壤养分和酶活性存在显著影响。3种外来植物入侵域的土壤速效磷、硝态氮含量以及磷酸酶活性随生长时期变化都表现出先升高后降低的趋势,均在旺盛生长期达到最大值。入侵域土壤速效磷和硝态氮含量的变化趋势与狗尾草明显不同。3种外来植物入侵降低了土壤速效钾含量,提高了土壤脲酶、磷酸酶活性。【结论】3种外来菊科植物的入侵改变了土壤养分和酶活性,创造出更有利于自身生长和繁殖的条件,以利于其进一步扩张。     

Impacts of invasive plant species on riparian plant assemblages: interactions with elevated atmospheric carbon dioxide and nitrogen deposition

Resource competition is commonly invoked to explain negative effects of invasive plants on native plant abundance. If invasives out-compete natives, global changes that elevate resource availability may interact with invasives to exacerbate impacts on native communities. Indeed, evidence is accumulating that elevated CO₂ and N deposition decrease native biomass and simultaneously increase invasive biomass. However, superior competitive ability, and a relative increase in the magnitude of invasive impacts under elevated resource availability, remain to be definitively proven. Using model, multi-species, multi-individual riparian plant communities, where planting density was maintained by replacement of native with exotic individuals, we conducted a greenhouse, competition experiment using native (to the UK) and invaded communities exposed to ambient and elevated CO₂ (CO₂ experiment) or N availability (N experiment). We tested two hypotheses: (1) invasives are superior competitors to natives at ambient atmospheric CO₂ and N deposition; (2) negative effects of invasives on natives are exacerbated under elevated CO₂ or N availability. Our results provide some support for the first hypothesis: in the CO₂ experiment native biomass was significantly lower in invaded communities. In the N experiment, native biomass was unaffected by the presence of exotics but other characteristics (e.g. root:shoot ratios) were altered. Differences in light availability between the experiments may have modified the effects of the invasives on the native assemblages but our design did not permit us to determine this definitively. The hypothesis that elevated CO₂ and N availability benefit invasives at the expense of natives was not supported by our results. This may be explained either because the invasives showed minor responses to the resource manipulations or because native and exotic species were differentially limited by CO₂ and N. Our results confirm the expectation that invasives alter the characteristics of native assemblages but lead us to question whether elevated resource availability will magnify these effects.     

Apparent facilitation of an invasive mealybug by an invasive ant

Summary In the southeast United States, the invasive ant Solenopsis invicta is known to derive important carbohydrate (honeydew) resources from mealybugs utilizing grasses. Most important appears to be an invasive mealybug, Antonina graminis. We studied whether this mealybug and a similar native species also benefit from association with S. invicta. We found that mealybug occurrence increases significantly with increasing proximity to S. invicta mounds, suggesting that mealybugs benefit as well. Mutual benefits derived by S. invicta and A. graminis are consistent with a hypothesis proposing that associations among invasive species can be important in their success at introduced locations.     

Do Invasive Trees have a Hydraulic Advantage over Native Trees?

The hypothesis was tested that invasive trees have hydraulic traits that contribute to their invasive nature. Five pairs of co-occurring invasive and native trees, in mesic habitats, were selected: (1) Tamarix ramosissima and Salix amygdaloides; (2) Robinia pseudoacacia and Alnus rhombifolia (3) Schinus terebinthifolius and Myrica cerifera; (4) Ligustrum sinense and Acer negundo; and (5) Sapium sebiferum and Diospyros virginiana, respectively. Resistance to cavitation (the water potential [Ψ _x ] at 75% loss of hydraulic conductivity [Ψ₇₅]) was not consistently greater for invasive compared to native species (Ψ₇₅=−1.91 and −1.67 MPa, respectively). Xylem specific conductivity (K _s), a measure of xylem efficiency, was not different between native and invasive species (K _s = 3.50 and 3.70 kg s⁻¹ MPa⁻¹ m⁻¹, respectively). The lack of difference for resistance to cavitation among invasive and native species suggests that the sampled invaders are not more tolerant to water stress than co-occurring native species. Apparently the spread and invasive nature of the sampled species cannot be explained by hydraulic traits alone.     

Consistent performance of invasive plant species within and among islands of the Mediterranean basin

Since the success of an invasive species depends not only upon its intrinsic traits but also on particular characteristics of the recipient habitat, assessing the performance of an invader across habitats provides a more realistic analysis of risk. Such an analysis will not only provide insights into the traits related to invasiveness, but also the habitat characteristics that underpin vulnerability to invasion that, taken together, will facilitate the selection of management strategies to mitigate the invader’s effect. In the present study, we considered the Mediterranean basin islands as an excellent study region to test how the same invasive species perform in different habitats within a single island, and to scale up differences among islands with similar climate. We tested how the performance of three widespread plant invaders with clonal growth but contrasting life-history traits, a deciduous tree Ailanthus altissima, a succulent subshrub Carpobrotus spp., and an annual geophyte Oxalis pes-caprae, varied depending upon the species identity, habitat, and invaded island. The environmental parameters considered were habitat type, elevation, species diversity in the invaded plot, and several soil traits (% C, % N, C/N, pH, and relative humidity). The study documents that the performance of these three important and widespread plant invaders is dependent mainly on species identity, and less upon the invaded island’s general features. Likewise, differences in performance among habitats were only significant in the case of Ailanthus, whereas Carpobrotus and Oxalis appear to perform equally well in different environments. Ailanthus thus appears to have a broader spectrum of invasiveness, being able to invade a larger number of habitat types. On the contrary, Carpobrotus spp. have not yet invaded habitats different from those where the species have been originally introduced and where they are still commonly spread by humans. Oxalis distribution is mainly related to agricultural activities and disturbed sites, and the total area infested by this geophyte may be more reflection of the extent of suitable habitats than of invasiveness or ecological impact. Our results confirm the potential for these species to significantly alter the functioning of ecosystems in the Mediterranean islands and highlight the risk to other islands not yet invaded.     

Experimental evidence for an alkali ecotype of <Emphasis Type="Italic">Lolium multiflorum</Emphasis>, an exotic invasive annual grass in the Central Valley,CA, USA

One of the characteristics of highly invaded ecosystems is that exotic species are often poor invaders of edaphically severe sites, which become refuges for native flora. To investigate the invasive potential of Lolium multiflorum (Per.) into alkali sites in California, an ex-situ reciprocal transfer experiment was carried out using seeds from populations of L. multiflorum taken from three sites differing in alkalinity (and inundation), including alkali sink soils (pH 8.5) and sink matrix soils (pH 7.4) located within meters of each other, and non-sink soils (pH 5.0) located several km away. Survivorship, plant height, leaf number and seed production were assessed. In addition, a native composite, Hemizonia pungens (Hick.), commonly found on alkali sinks was also sampled at the sink and sink matrix microsites. Lolium multiflorum plants grown from alkali sink and sink matrix seeds produced fewer leaves and seeds but were taller than plants grown from non-alkali seeds, the latter perhaps an adaptation to frequently inundated soils. Non-alkali genotypes fared poorly in sink soils for all traits, both in comparison to their growth on non-sink soils, and in comparison to the sink and sink edge genotypes. This suggests the existence of L. multiflorum ecotypes adapted to inundated alkali sinks, a genotypic difference that occurs on a broad spatial scale (kilometers), but not so obviously on the micro-site scale (meters) between sink and sink matrix populations. These data suggest that the absence of exotic invasives from alkali sites may be temporary if they are evolving tolerance for these severe sites, and this may threaten the future of the native alkali specialists that currently find refuge in these sites.     

Influence of woody invader control methods and seed availability on native and invasive species establishment in a Hawaiian forest

When invasive woody plants become dominant, they present an extreme challenge for restoration of native plant communities. Invasive Morella faya (fire tree) forms extensive, nearly monospecific stands in wet and mesic forests on the Island of Hawai’i. We used logging, girdling, and selective girdling over time (incremental girdling) to kill stands of M. faya at different rates, with the objective of identifying a method that best promotes native forest re-establishment. We hypothesized that rapid canopy opening by logging would lead to establishment of fast-growing, non-native invaders, but that slower death of M. faya by girdling or incremental girdling would increase the establishment by native plants adapted to partial shade conditions. After applying the M. faya treatments, seed banks, seed rain, and plant recruitment were monitored over 3 years. Different plant communities developed in response to the treatments. Increased light and nitrogen availability in the logged treatment were associated with invasion by non-native species. Native species, including the dominant native forest tree, (Metrosideros polymorpha) and tree fern (Cibotium glaucum), established most frequently in the girdle and incremental girdle treatments, but short-lived non-native species were more abundant than native species. A diverse native forest is unlikely to develop following any of the treatments due to seed limitation for many native species, but girdling and incremental girdling promoted natural establishment of major components of native Hawaiian forest. Girdling may be an effective general strategy for reestablishing native vegetation in areas dominated by woody plant invaders.     

Soil biota and invasive plants

Interactions between plants and soil biota resist invasion by some nonnative plants and facilitate others. In this review, we organize research and ideas about the role of soil biota as drivers of invasion by nonnative plants and how soil biota may fit into hypotheses proposed for invasive success. For example, some invasive species benefit from being introduced into regions of the world where they encounter fewer soil-borne enemies than in their native ranges. Other invasives encounter novel but strong soil mutualists which enhance their invasive success. Leaving below-ground natural enemies behind or encountering strong mutualists can enhance invasions, but indigenous enemies in soils or the absence of key soil mutualists can help native communities resist invasions. Furthermore, inhibitory and beneficial effects of soil biota on plants can accelerate or decelerate over time depending on the net effect of accumulating pathogenic and mutualistic soil organisms. These 'feedback' relationships may alter plant-soil biota interactions in ways that may facilitate invasion and inhibit re-establishment by native species. Although soil biota affect nonnative plant invasions in many different ways, research on the topic is broadening our understanding of why invasive plants can be so astoundingly successful and expanding our perspectives on the drivers of natural community organization.     

Competitive abilities of introduced and native grasses

Differencesin competitive ability may explain the maintenance of existing plantpopulationsand the invasion of new areas by plant species. We used field experiments toexamine the competitive responses of Agropyron cristatum(L.) Gaertn., an introduced C₃ grass, and Boutelouagracilis (HBK.) Lag., a native C₄ grass, and thecompetitive effects of Agropyron-dominated vegetation andsuccessional prairie. We also tested whether the outcome of competitiveinteractions varied with water availability. In each vegetation type,transplants of each species were grown under two levels of competition(presenceor absence of neighboring vegetation) and three levels of water availability(high, medium, or low). Transplant survival, growth, and biomass allocationpatterns were measured. Water availability had no effect on the measuredvariables, suggesting that both species were limited by another resource.Growthrates were affected more by competition, while survival and root: shoot ratiowere affected more by transplant species identity. In the successional prairie,neighboring vegetation suppressed the growth of Agropyrontransplants less than that of Bouteloua transplants,suggesting that Agropyron has a stronger ability to resistcompetitive suppression in that vegetation type. The spread ofAgropyron into surrounding vegetation may relate to itsability to resist competitive suppression. In theAgropyron-dominated vegetation, neighboring vegetationsuppressed the growth of both species by the same extent. However, competitionaccounted for more variation in transplant growth inAgropyron-dominated vegetation than in successionalprairie, suggesting that Agropyron has strong competitiveeffects which hinder plant growth and prevent other species from establishinginAgropyron fields.     

Intra and interspecific competition among invasive and native species during early stages of plant growth

Plant competition is a primary ecological process limiting grassland restoration success. Appropriate restoration techniques require an understanding of the degree to which intra and interspecific competition control invasive and native plant growth. The objective of this study was to determine how the intensity of intra and interspecific competition changes during early stages of plant growth. Two invasive (Bromus tectorum and Taeniatherum caput-medusae) and two native (Pseudoroegneria spicata and Poa secunda) species were grown in a diallel competition experiment, either alone or in 1:1 binary combinations and exposed to two levels of N (no N or 400 mg N kg⁻¹ soil added) in a greenhouse. Total biomass for each species was quantified over four harvests and competitive effects were calculated. Our results show that the relative magnitude of intra and interspecific competition changes through time. Intraspecific competition was intense for native species at the initial harvests and therefore important in contributing to the outcome of final size of native species seedlings. Interestingly, bluebunch wheatgrass imposed interspecific competition on annual grasses at the first two harvests and appeared to be a better competitor than Sandberg’s bluegrass. We found that fast growing invasive species became more competitive compared to slow growing native species with increasing N and appear to establish a positive feedback mechanism between size and resource uptake. Opportunities to improve restoration success exist from determining the optimum combination of density, species proportion, and their spatial arrangement in various ecosystems and environments.     

Variation in the establishment of a non-native annual grass influences competitive interactions with Mojave Desert perennials

Competition between native and non-native species can change the composition and structure of plant communities, but in deserts, the highly variable timing of resource availability also influences non-native plant establishment, thus modulating their impacts on native species. In a field experiment, we varied densities of the non-native annual grass Bromus madritensis ssp. rubens around individuals of three native Mojave Desert perennials—Larrea tridentata, Achnatherum hymenoides, and Pleuraphis rigida—in either winter or spring. For comparison, additional plots were prepared for the same perennial species and seasons, but with a mixture of native annual species as neighbors. Growth of perennials declined when Bromus was established in winter because Bromus stands had 2–3 months of growth and high water use before perennial growth began. However, water potentials for the perennials were not significantly reduced, suggesting that direct competition for water may not be the major mechanism driving reduced perennial growth. The impact of Bromus on Larrea was lower than for the two perennial grasses, likely because Larrea maintains low growth rates throughout the year, even after Bromus has completed its life cycle. This result contrasts with the perennial grasses, whose phenology completely overlaps with (Achnatherum) or closely follows (Pleuraphis) that of Bromus. In comparison, Bromus plants established in spring were smaller than those established in winter and thus did not effectively reduce growth of the perennials. Growth of perennials with mixed annuals as neighbors also did not differ from those with Bromus neighbors of equivalent biomass, but stands of these native annuals did not achieve the high biomass of Bromus stands that were necessary to reduce perennial growth. Seed dormancy and narrow requirements for seedling survivorship of native annuals produce densities and biomass lower than those achieved by Bromus; thus, impacts of native Mojave Desert annuals on perennials are expected to be lower than those of Bromus.     

Mycorrhizae transfer carbon from a native grass to an invasive weed: evidence from stable isotopes and physiology

Invasive exotic weeds pose one of the earth's most pressing environmental problems. Although many invaders completely eliminate native plant species from some communities, ecologists know little about the mechanisms by which these exotics competitively exclude other species. Mycorrhizal fungi radically alter competitive interactions between plants within natural communities, and a recent study has shown that arbuscular mycorrhizal (AM) fungi provide a substantial competitive advantage to spotted knapweed, Centaurea maculosa, a noxious perennial plant that has spread throughout much of the native prairie in the northwestern U.S. Here we present evidence that this advantage is potentially due to mycorrhizally mediated transfer of carbon from a native bunchgrass, Festuca idahoensis, to Centaurea. Centaurea maculosa, Festuca idahoensis (Idaho fescue, C₃), and Bouteloua gracilis (blue gramma, C₄) were grown in the greenhouse either alone or with Centaurea in an incomplete factorial design with and without AM fungi. Centaurea biomass was 87–168% greater in all treatments when mycorrhizae were present in the soil (P < 0.0001). However, Centaurea biomass was significantly higher in the treatment with both mycorrhizae and Festuca present together than in any other treatment combination (P < 0.0001). This high biomass was attained even though Centaurea photosynthetic rates were 14% lower when grown with Festuca and mycorrhizae together than when grown with Festuca without mycorrhizae. Neither biomass nor photosynthetic rates of Centaurea were affected by competition with the C₄ grass Bouteloua either with or without mycorrhizae. The stable isotope signature of Centaurea leaves grown with Festuca and mycorrhizae was more similar to that of Festuca, than when Centaurea was grown alone with mycorrhizae (P = 0.06), or with Festuca but without mycorrhizae (P = 0.09). This suggests that carbon was transferred from Festuca to the invasive weed. We estimated that carbon transferred from Festuca by mycorrhizae contributed up to 15% of the aboveground carbon in Centaurea plants. Our results indicate that carbon parasitism via AM soil fungi may be an important mechanism by which invasive plants out compete their neighbors, but that this interaction is highly species-specific.