Potential selection in native grass populations by exotic invasion

Ecological impacts of invasive plant species are well documented, but the genetic response of native species to invasive dominance has been often overlooked. Invasive plants can drastically alter site conditions where they reach dominance, potentially exerting novel selective pressures on persistent native plant populations. Do native plant populations in old exotic invasions show evidence of selection when compared to conspecific populations in adjacent, noninvaded areas? We employ amplified fragment length polymorphism (AFLP) analysis to screen a large number of loci from two native grass species (Hesperostipa comata (Trin. & Rupr.) Barkworth and Sporobolus airoides Torr.) that occur in old infestations of the invasive forb Acroptilon repens. We then compare observed locus by locus F_ST values with distributions of F_ST estimated from simulation models under expectation of neutrality. We also compare the proportion of loci possibly linked to selection and those not linked to selection which exhibit parallel trends in divergence between two community types (invaded, noninvaded). Few loci (H. comata, 2.6%; S. airoides, 8.7%) in the two native grasses may be linked to genes under the influence of selection. Also, loci linked to selection showed a greater portion of parallel trends in divergence than neutral loci. Genetic similarities between community types were less than genetic similarity within community types suggesting differentiation in response to community alteration. These results indicate that a small portion of scored AFLP loci may be linked to genes undergoing selection tied to community dominance by an invasive species. We propose that native plants in communities dominated by exotic invasives may be undergoing natural selection.     

Impact of two shoot-galling biological control candidates on Russian knapweed, Acroptilon repens

Russian knapweed, Acroptilon repens, is one of the most serious exotic invaders of temperate grasslands in North America. Here we present results from a field experiment in which we quantified the impact of two potential biological control agents, the gall wasp Aulacidea acroptilonica V.Bel. (Hymenoptera, Cynipidae) and the gall midge Jaapiella ivannikovi Fedotova (Diptera, Cecidomyiidae), on A. repens under field conditions in the plant’s native range in Uzbekistan. Attack by A. acroptilonica reduced shoot length by 21%, above-ground biomass by 25% and seed output by 75%, while attack by J. ivannikovi reduced shoot length by 12%, above-ground biomass by 24%, and seed output by 92%. The results of these field experiments are likely to accurately reflect the potential of these two gall formers to reduce above-ground biomass and sexual reproduction of A. repens shoots, since the shoots were part of a clonal network. Despite this, the attacked shoots were not able to compensate for the reallocation of plant resources to gall formation. Moreover, the mean number of galls per shoot obtained in the experiments was within the range of observed gall incidences in the native range. The impact of these two gall-forming insects on Russian knapweed in North America will depend on the population size the species reach and on the timing of attack. The highest impact is likely to occur when the insects attack shoots that have not yet started producing flower-buds.     

Influence of an exotic species,Acroptilon repens (L.) DC. on seedling emergence and growth of native grasses

Our objective was to evaluate the effects of an invasive perennial forb, Acroptilon repens (Russian knapweed), on seed emergence and seedling survival of four native grass species that are important in semiarid grasslands of North America. Greenhouse experiments and field studies conducted at three sites in Colorado, USA were used to examine the response by four perennial grasses: Bouteloua gracilis, Koelaria cristata, Sporobolus cryptandrus, and Agropyon smithii to A. repens. In the greenhouse, seeds of each species were sown in pots that contained either live A. repens roots, A. repens litter on the soil surface, or both roots and litter. Field transects were placed inside stands of A. repens with adjacent control transects in the surrounding grass-dominated community. Emergence and initial survival were decreased by the presence of A. repens roots for K. cristata (35%), B. gracilis (31%), and S. cryptandrus (44%) in the greenhouse, and 57, 32, and 36%, respectively, in the field. Root weight was decreased by the presence of A. repens roots for both B. gracilis and K. cristata by more than 55% in the greenhouse. A. smithii survival and growth were unaffected by A. repens in either the greenhouse or the field. These species-specific responses to the presence of A. repens may explain, at least in part, differences in susceptibility and recovery of sites with different native grass species composition. This revised version was published online in August 2006 with corrections to the Cover Date.     

Invasive Phragmites australis management outcomes and native plant recovery are context dependent

The outcomes of invasive plant removal efforts are influenced by management decisions, but are also contingent on the uncontrolled spatial and temporal context of management areas. Phragmites australis is an aggressive invader that is intensively managed in wetlands across North America. Treatment options have been understudied, and the ecological contingencies of management outcomes are poorly understood. We implemented a 5‐year, multi‐site experiment to evaluate six Phragmites management treatments that varied timing (summer or fall) and types of herbicide (glyphosate or imazapyr) along with mowing, plus a nonherbicide solarization treatment. We evaluated treatments for their influence on Phragmites and native plant cover and Phragmites inflorescence production. We assessed plant community trajectories and outcomes in the context of environmental factors. The summer mow, fall glyphosate spray treatment resulted in low Phragmites cover, high inflorescence reduction, and provided the best conditions for native plant recruitment. However, returning plant communities did not resemble reference sites, which were dominated by ecologically important perennial graminoids. Native plant recovery following initial Phragmites treatments was likely limited by the dense litter that resulted from mowing. After 5 years, Phragmites mortality and native plant recovery were highly variable across sites as driven by hydrology. Plots with higher soil moisture had greater reduction in Phragmites cover and more robust recruitment of natives compared with low moisture plots. This moisture effect may limit management options in semiarid regions vulnerable to water scarcity. We demonstrate the importance of replicating invasive species management experiments across sites so the contingencies of successes and failures can be better understood.     

Allergenicity of Acroptilon repens and Juglans regia pollen in rats

Pollen proteins that are located in the cytoplasm or on the surface of the exine can function as allergens and evoke immune system responses in sensitive patients, leading to allergic rhinitis and asthma. In this research, the pollen allergenicity and ability to induce IgE response of the pollen of two plant species were studied in rats. Acroptilon repens is an herbaceous, invasive plant with entomophilous pollen, while Juglans regia which is a tree crop produces anemophilous pollen. Immunoblot analysis using sera of sensitised rats revealed IgE reactivity to three protein bands including the 70, 41 and 25.12 kDa bands present in the A. repens pollen extract, while only one single immunogenic band of 11 kDa was detected in J. regia pollen extract. Both pollen extracts increased the eosinophil content and caused some clinical signs of allergy in treated rats. The results showed that both entomophilous and anemophilous pollen can be allergenic.     

Response of Acroptilon repens to simulated herbivory and soil disturbance

Acroptilon repens is an invasive weed in North America but also causes problems in disturbed habitats in its native range in Asia. In order to test the effect of simulated biological control and soil disturbance on established A. repens patches and the competing vegetation, two levels of shoot clipping as well as soil tillage were imposed on A. repens patches in an undisturbed meadow and at two fallowland sites in the native range of the weed. At the meadow site, 2 years of partial clipping of shoots and of soil tillage had no influence on A. repens performance, while soil tillage significantly reduced the above‐ground biomass of the competing vegetation. At the fallowland sites, which had been continuously cultivated for several years prior to the experiment, A. repens shoot density, biomass and number of seed heads were significantly higher in the undisturbed control than in the tillage plots. The total number of seed heads per unit area increased with shoot density up to 200 shoots m^?2. These results indicate that A. repens has considerable regrowth capacities that allow established patches to tolerate substantial losses of above‐ground biomass and that the competitive ability of A. repens is favoured both when soil disturbance is imposed on previously undisturbed sites, as well as when repeated soil disturbance is abandoned. The only promising nonchemical herbicide‐based approach to reduce the competitive ability and seed output of A. repens appears to be a long‐term management that enhances the interspecific plant competition by reducing soil disturbance and selectively damaging A. repens.     

Phenotypes of Two Generations of Sporobolus airoides Seedlings Derived from Acroptilon repens‐invaded and Non‐invaded Grass Populations

Although the ecological impacts of invasive species are well known, the evolutionary impacts on recipient native grass communities are not. We suggest that remnant native plants may provide desirable seed sources for restoration and native plant production. Native populations exposed to the selective pressures associated with exotic invasion may retain traits that increase their ability to coexist with invasive species. Two generations of Sporobolus airoides Torr. (Alkali sacaton) plants derived from lineages collected from within long‐term invaded areas of Acroptilon repens (L.) DC (Russian knapweed) and from adjacent non‐invaded areas were propagated in a greenhouse to evaluate generational changes in phenotypic traits from the production environment. Given the difference in invasion history of the two populations, we hypothesized that invaded and non‐invaded subpopulations would differ phenotypically. Phenotypic measurements revealed that invaded subpopulations had greater vegetative growth, whereas non‐invaded subpopulations had increased sexual reproduction. Phenotypic expression changed from the first to the second generation, predominantly in the invaded subpopulation. Generational phenotypic shifts are disadvantageous for native seed production which requires a standard product to sell commercially. However, phenotypic variation may improve field seed survival. This research demonstrates the potential value of targeting post‐invasion remnant grass populations for restoration.     

Presence of Megalorhipida leucodactyla (Lepidoptera Pterophoridae) in Sicily: new report for the Italian fauna and notes on the diffusion of its host plant Boerhavia repens L. ssp. viscosa (Choisy) Maire (Nyctaginaceae)

Despite its circum-tropical distribution, we report, for the first time in Italy and Sicily, the presence of Megalorhipida leucodactyla (Fabricius 1974), a species recorded only once before in Europe. Several adults, pupae and larvae of different instars were found on Boerhavia repens L. ssp. viscosa (Choisy) Maire (Nyctaginaceae), recorded for the first time in Italy (Sicily) in 1967. We provide both an update of Boerhavia repens ssp. viscosa’s distribution in Sicily and the results of our vegetational survey, based upon the phytosociological method by Braun-Blanquet Sigmatist School, on Boerhavia repens ssp. viscosa in Palermo city’s urban areas. This plant, naturalised especially in Palermo city, appears to have become an invasive plant, increasingly spreading in larger areas, by even replacing the native flora in urban ecosystems. This study also provides the results of laboratory bioethological observations on some specimen of both adults and larvae (the latter while feeding on Boerhavia repens ssp. viscosa’s fruits). Finally, using bibliographic data, we drew a map of the distribution of both Megalorhipida leucodactyla and Boerhavia repens ssp. viscosa in the Mediterranean basin.     

基于个体的空间显性模型和遥感技术模拟入侵植物扩张机制

基于个体的空间显性模型和遥感技术,以互花米草为例,模拟了自1997到2010年的种群扩张动态,揭示了土地利用变化与潮间带高程的影响;并通过全局敏感性分析揭示了种子扩散、成体存活率、有性和无性繁殖等种群统计学特征对互花米草种群扩张的相对重要性。研究结果发现:1)有性繁殖与无性繁殖共同决定互花米草种群快速扩张;2)潮间带高程和土地利用变化显著影响模型预测的精度,对互花米草种群扩张有非常重要的影响;3)成体存活率与种子长距离扩散是影响互花米草种群扩张速度最重要的因素;无性繁殖比有性繁殖对种群扩张的影响更大;种子长距离扩散比本地扩散更为重要,同时,小概率的种子长距离扩散事件对种群扩张有非常重要的影响。为了经济有效地控制外来入侵植物的扩张,应该抑制种子的长距离扩散和移除种子长距离扩散形成的位于入侵前沿的小斑块。     

An evaluation of the impact of Melaleuca quinquenervia invasion and management on plant community structure after fire

Two questions were asked in this study: after a fire, does the choice of invasive plant management strategy, namely herbicidal or biological, alter (1) plant community assemblages and (2) the re-invasion potential of the Australian tree Melaleuca quinquenervia? Plant species richness was highest in the non-invaded and herbicide sites compared to the biological site with 10.5, 10.8, and 8.25 species m⁻² found in each site, respectively. Although the total count of live and dead seedlings was highest in the biologically controlled site at 22.8 and 13.6 plants m⁻², respectively, M. quinquenervia seedlings were recruited in all sites. While the ultimate goal of management programs is to restore ecosystem integrity, this work provides evidence that passive restoration may not be enough to restore plant community structure in this system.     

Propagule pressure and native species richness effects drive invasibility in tropical dry forest seedling layers

Understanding the factors that encourage or inhibit plant invasions is vital to focusing limited invasive control efforts within areas where they are most practical and cost-effective. To extend the range of contexts in which invasibility is studied and aid the development of practical strategies to limit damaging plant invasions, we set out to test the relative importance of native species richness, native seedling density, and invasive propagule pressure, on the invasibility of artificial assemblages of naturally occurring tropical woody seedling communities. Our greenhouse mesocosms included a species pool of twelve trees and woody shrubs native to South Florida's tropical hardwood hammocks, and an increasingly prevalent noxious woody invader of this system, Ardisia elliptica. We found that invader propagule pressure was the single most important factor determining community invasibility. We also revealed a positive relationship between invasibility and native species richness in our polyculture mesocosms. Because A. elliptica biomass production significantly differed among different native monocultures and was not related to overyielding in native polycultures, we suggest that the effect of species richness on invasibility in this experiment was the result of sampling effects rather than a true effect of diversity.Three broad findings hold potential for application in preventing and controlling plant invasions, especially in the seedling layers of tropical dry forests: (1) effective invasive control efforts will likely benefit from measures to minimize propagule pressure; (2) managers might do well to prioritize invasive monitoring and removal efforts on the most diverse habitats within a management region; and (3) while more data are necessary to further understand our finding of a lack of association between productivity and invasibility, management regimes aimed at maximizing primary productivity might not increase invasibility, and in fact, strategies for controlling invasive plants via the management of ecosystem productivity may be ineffective.     

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.     

Evaluation of elemental allelopathy in Acroptilon repens (L.) DC. (Russian Knapweed)

Although Acroptilon repens (L.) DC. (Russian knapweed) is known to concentrate zinc (Zn) in upper soil layers, the question of whether the elevated Zn has an allelopathic effect on restoration species has not been addressed. Experiments were conducted to investigate whether soils collected from within infestations of A. repens (high-Zn) inhibit the germination or growth and development of desirable restoration species, compared to soils collected adjacent to an A. repens infestation (low-Zn). Four bioassay species [Sporobolus airoides (Torrey) Torrey (alkali sacaton), Pseudoroegneria spicata (Pursh) A. Love (bluebunch wheatgrass), Psathyrostachys juncea (Fischer) Nevski (Russian wildrye) and A. repens] were germinated in a growth chamber and grown in a greenhouse in both soils and received treatments for the alleviation of Zn toxicity (P, Fe, Fe-oxide, and soil mixing) to isolate the effects of elevated soil Zn on plant performance. Percent germination, total plant biomass, tiller and stem number, inflorescence number, and tissue metal levels were compared among soil types and treatments for each species. There was no evidence from any of the indicators measured that high-Zn soils reduced plant performance, compared to low-Zn soils. Tissue Zn levels barely approached the lower range of phytotoxic levels established for native grasses. Older plants with longer exposure times may accumulate higher Zn concentrations. S. airoides and A. repens both had higher biomass in the high-Zn soil, most likely due to increased macronutrient (N and P) availability. As the Zn levels in the soils used in this study were much higher than any levels previously reported in soils associated with A. repens, it is unlikely that the elevation of soil Zn by A. repens will hinder germination or growth and development of desirable grasses during establishment.     

Impacts of the emerald ash borer (EAB) eradication and tree mortality: potential for a secondary spread of invasive plant species

Since the discovery of the emerald ash borer in 2002, eradication efforts have been implemented in an attempt to eliminate or contain the spread of this invasive beetle. The eradication protocol called for the removal of every ash tree within a 0.8 km radius around an infested tree. In 2005 this study was established to identify environmental changes attributed to the eradication program and measure subsequent shifts in forest community composition and structure. We conducted this study in Ohio and compared areas that received the eradication treatment (ash trees cut down), to areas that were left uncut, (ash still standing). The goal of this project was to identify how the plant community is responding in these two areas. The eradication protocol accelerated the formation and size of gaps within the forest and thus increased the duration and intensity of light penetrating through to the forest floor. In addition, the use of track vehicles for removal of cut trees resulted in significant soil compaction. The resultant plant community had greater species diversity (H′). When specific species composition differences were compared, an increase in the establishment of invasive plant species was detected in areas that received eradication efforts compared to those that did not. Invasive species accounted for 18.7% of the total herbaceous cover in this highly disturbed environment which included Cirsium arvense, Rhamnus cathartica and 2 species of Lonicera. In contrast, invasive species accounted for <1% of the total herbaceous cover in the undisturbed uncut areas.     

Models of Experimentally Derived Competitive Effects Predict Biogeographical Differences in the Abundance of Invasive and Native Plant Species

Mono-dominance by invasive species provides opportunities to explore determinants of plant distributions and abundance; however, linking mechanistic results from small scale experiments to patterns in nature is difficult. We used experimentally derived competitive effects of an invader in North America, Acroptilon repens, on species with which it co-occurs in its native range of Uzbekistan and on species with which it occurs in its non-native ranges in North America, in individual-based models. We found that competitive effects yielded relative abundances of Acroptilon and other species in models that were qualitatively similar to those observed in the field in the two ranges. In its non-native range, Acroptilon can occur in nearly pure monocultures at local scales, whereas such nearly pure stands of Acroptilon appear to be much less common in its native range. Experimentally derived competitive effects of Acroptilon on other species predicted Acroptilon to be 4–9 times more proportionally abundant than natives in the North American models, but proportionally equal to or less than the abundance of natives in the Eurasian models. Our results suggest a novel way to integrate complex combinations of interactions simultaneously, and that biogeographical differences in the competitive effects of an invader correspond well with biogeographical differences in abundance and impact.     

The importance of roads, nutrients, and climate for invasive plant establishment in riparian areas in the northwestern United States

Natural and anthropogenic site characteristics play a role in determining the current distribution of invasive plant species. An understanding of these characteristics can be used to prioritize areas for monitoring and control efforts and to determine appropriate management actions to lower site invasion risk. We used species distribution models to look for attributes associated with invasion and to determine the extent to which these attributes varied across a suite of species. We modeled the presence-absence of 11 invasive plant species along riparian areas in the northwestern United States using the model Random Forests. We found that climate variables were most important for predicting species distributions across the large study area and factors related to nutrients, land cover, and disturbance had moderate importance. We also found that there was a general pattern related to invasion for almost all species. Invasion was more likely to occur at hotter, drier sites near roads in unforested areas. In addition, high nutrient levels and proximity to streams with lower baseflow values also generally increased the likelihood that at least one invasive species would be present. Examining patterns across a broad range of regions can help suggest general mechanisms of invasion as well as provide region-specific management recommendations.     

A rapid method for constructing precaution maps based on a simple virtual ecology model: a case study on the range expansion of the invasive aquatic species Limnoperna fortunei

Before invasion, or in its early stages, information on the invader in target areas is generally extremely limited. In such situations, managers must select focal areas in which to concentrate control and mitigation efforts. Here, we discuss a rapid method for selecting areas in which to control invasive aquatic species based on limited information. We used a simple cellular automata model that does not require species-specific information, but simulates the process of invasive species expansion and includes observed expansion progress to detect keystone areas. As a case study, we simulated the expansion of an invasive aquatic mussel, Limnoperna fortunei, in Ibaraki Prefecture, Japan, and detected the areas in which control efforts should be concentrated. To some extent, our model was able to predict the expansion of L. fortunei from the initial detected invasion to the current distribution. We predicted areas with a high potential of spreading and areas that would suffer from high propagule pressure. Results revealed a mismatch between areas with high spread potential and those with high propagule pressure. Managers should concentrate their invasion prevention efforts in the former because these are likely to have a greater long-term influence. Additionally, we predicted future expansion from the current distribution and showed that current scattered populations could merge naturally. Our approach is useful for establishing a management plan before or in the early stages of invasion.     

The Effects of Habitat on Dispersal Patterns of an Invasive Thistle, Cynara Cardunculus

Dispersal of offspring is a critical step in the spread of invasive species, yet dispersal patterns are seldom well studied, inhibiting effective management and ecological understanding of invasions. Dispersal patterns can be affected by characteristics of the parent plant and by climatic or site characteristics, including community vegetation structure, but these factors have not been studied in the context of plant invasion processes. Cynara cardunculus (L.) is a polycarpic perennial thistle invasive in coastal grasslands in California that produces large numbers of wind-dispersed seeds. This study quantified and compared C. cardunculus dispersal patterns in an exotic grassland (vegetated site) and an agricultural field (non-vegetated site). Seed size variation in C. cardunculus was also quantified within inflorescences, between inflorescences, and between years of production, and compared across dispersal distances. Results indicate that dispersal distance increased dramatically from less than 20 m in the vegetated site to more than 40 m in the non-vegetated site. Plants producing fewer seeds produced heavier seeds and dispersal distance decreased over time, but seed size was not related to dispersal distance, which may serve to spread the risk to seeds or seedlings across the environment. C. cardunculus has great potential for dispersal in open areas such as agricultural fields or disturbed sites, but may be limited in highly structured natural communities. Management of C. cardunculus and other wind-dispersed plant invasions may be improved by prioritizing populations with open or disturbed areas downwind and minimizing the removal of vegetation during dispersal.     

Risk Assessment of Transgenic Virus-resistant White Clover: Non-target Plant Community Characterisation and Implications for Field Trial Design

One of the most difficult elements of the ecological risk assessment of transgenic plants is investigation of their potential impact on biodiversity in complex non-target communities. This problem is particularly acute for pasture plants, since many have a track record of invasiveness. In this paper we develop an understanding of some aspects of the ecology of Trifolium repens L. (white clover) in montane vegetation communities in southeastern Australia as part of a larger project investigating potential ecological risks associated with commercial release of newly-developed transgenic virus-resistant T. repens cultivars. We use a combination of floristic surveys and biomass sampling to determine the habitat affinity of T. repens, the structure and composition of associated communities, and the scale at which different abiotic and biotic factors correlate with T. repens abundance. We also compare the abundance of native and exotic species within specific morpho-functional groups and use this to determine the relative significance of T. repens as a community constituent, and to identify native species that would be most at risk by expanding populations of T. repens. We found that T. repens comprises a relatively small component of the total community cover and biomass, but is one of the most abundant herbaceous species in mesic Poa – dominated grasslands and Poa-Eucalyptus woodlands in the study area, and that T. repens abundance is correlated at the within-community scale primarily with soil moisture and fertility. At smaller scales T. repens is limited by the hierarchical dominance of native graminoids and we conclude that competition for inter-tussock space in mesic communities is the most likely mechanism by which expanding populations of T. repens would influence populations of associated native species. These results have significant implications for the manner in which future analysis and risk quantification stages of the risk assessment of virus-resistant T.␣repens and other transgenic pasture plants in complex plant communities are performed.