Allelopathy and exotic plant invasion

The primary hypothesis for the astonishing success of many exotics as community invaders relative to their importance in their native communities is that they have escaped the natural enemies that control their population growth – the `natural enemies hypothesis'. However, the frequent failure of introduced biocontrols, weak consumer effects on the growth and reproduction of some invaders, and the lack of consistent strong top-down regulation in many natural ecological systems indicate that other mechanisms must be involved in the success of some exotic plants. One mechanism may be the release by the invader of chemical compounds that have harmful effects on the members of the recipient plant community (i.e., allelopathy). Here, we provide an abbreviated compilation of evidence for allelopathy in general, present a detailed case study for Centaurea diffusa, an invasive Eurasian forb in western North America, and review general evidence for allelopathic effects of invasive plants in native communities. The primary rationale for considering allelopathy as a mechanism for the success of invaders is based on two premises. First, invaders often establish virtual monocultures where diverse communities once flourished, a phenomenon unusual in natural communities. Second, allelopathy may be more important in recipient than in origin communities because the former are more likely to be naïve to the chemicals possessed by newly arrived species. Indeed, results from experiments on C. diffusa suggest that this invader produces chemicals that long-term and familiar Eurasian neighbors have adapted to, but that C. diffusa's new North American neighbors have not. A large number of early studies demonstrated strong potential allelopathic effects of exotic invasive plants; however, most of this work rests on controversial methodology. Nevertheless, during the last 15 years, methodological approaches have improved. Allelopathic effects have been tested on native species, allelochemicals have been tested in varying resource conditions, models have been used to estimate comparisons of resource and allelopathic effects, and experimental techniques have been used to ameliorate chemical effects. We do not recommend allelopathy as a `unifying theory' for plant interactions, nor do we espouse the view that allelopathy is the dominant way that plants interact, but we argue that non-resource mechanisms should be returned to the discussion table as a potential mechanism for explaining the remarkable success of some invasive species. Ecologists should consider the possibility that resource and non-resource mechanisms may work simultaneously, but vary in their relative importance depending on the ecological context in which they are studied. One such context might be exotic plant invasion.     

Invader to invader facilitation: ice plant mats prompt increased densities and grazing impacts of white garden snails in a Pampean coastal dune system

Non-native species can facilitate other non-native species via their effects on the physical environment. Here, I evaluate whether the ice plant, Carpobrotus edulis, prompts increased densities and grazing impacts of the white garden snail, Theba pisana, in coastal dunes of the Argentinean Pampas, where both species are non-native. Snail densities were higher in ice plant patches than in native graminoid patches over four seasonal samplings. Snail activity and grazing impacts on two co-occurring forbs (the native Hydrocotyle bonariensis and the non-native Calystegia soldanella) were also higher within ice plant patches. Experimental ice plant transplant into graminoid-dominated areas led to increased snail densities and grazing impacts, which demonstrates cause-effect relationships linking ice plant cover with increasing snail densities and grazing damage to forbs. The findings here suggest that ice plant mats provide suitable microclimatic conditions for T. pisana, and thereby facilitate greater snail numbers in ice plant patches leading to increased grazing rates on other plants. On that basis, it can be expected that the cumulative effect of both invaders on co-occurring plants would be larger than their individual effects.

     

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.     

Differential responses of native and exotic coastal sage scrub plant species to N additions and the soil microbial community

Background and aims

Native shrub species of southern California have a long history of displacement by exotic annual herbs and forbs. Such invasions may be mediated by interactions with the microbial community and changes in the N cycle as a result of N pollution. However, the simultaneous effects of the soil microbial community status and N fertilization on dominant native and exotic plant species growth have not been thoroughly explored in this ecosystem.

Methods

Three species of native shrubs and of exotic annuals were grown in an orthogonal two-factor greenhouse experiment. To assess the importance of the soil microbial community pre-sterilized soils were inoculated with sterilized or non-sterilized field soil; to assess the importance of N type pots were fertilized with nitrate, ammonium or glycine solutions. Plant shoot and root biomass was measured after harvesting.

Results

The natives Artemisia californica and Eriogonum fasciculatum had lower growth in sterilized soil, suggesting microbial facilitation of these species, and E. fasciculatum higher growth with ammonia than either nitrate or glycine. Salvia apiana had equal growth under all conditions. The exotics Brassica nigra and Bromus madritensis grew equally in sterilized and unsterilized soil, and B. madritensis greater growth with ammonia fertilizer. Centaurea melitensis had greater growth in sterilized soil, and with either form of inorganic N.

Conclusions

These results highlight the importance of the soil microbial community in contributing to relative success of native vs. exotic species, and could inform restoration approaches for these species.     

Arrival order among native plant functional groups does not affect invasibility of constructed dune communities

Different arrival order scenarios of native functional groups to a site may influence both resource use during development and final community structure. Arrival order may then indirectly influence community resistance to invasion. We present a mesocosm experiment of constructed coastal dune communities that monitored biotic and abiotic responses to different arrival orders of native functional groups. Constructed communities were compared with unplanted mesocosms. We then simulated a single invasion event by bitou (Chrysanthemoides monilifera ssp. rotundata), a dominant exotic shrub of coastal communities. We evaluated the hypothesis that plantings with simultaneous representation of grass, herb and shrub functional groups at the beginning of the experiment would more completely sequester resources and limit invasion than staggered plantings. Staggered plantings in turn would offer greater resource use and invasion resistance than unplanted mesocosms. Contrary to our expectations, there were few effects of arrival order on abiotic variables for the duration of the experiment and arrival order was unimportant in final community invasibility. All planted mesocosms supported significantly more invader germinants and significantly less invader abundance than unplanted mesocosms. Native functional group plantings may have a nurse effect during the invader germination and establishment phase and a competitive function during the invader juvenile and adult phase. Arrival order per se did not affect resource use and community invasibility in our mesocosm experiment. While grass, herb and shrub functional group plantings will not prevent invasion success in restored communities, they may limit final invader biomass.     

Spatially contingent interactions between an exotic and native plant mediated through flower visitors

Exotic plants can negatively impact the fitness of native plants by changing the behavior of flower visitors and thus affecting pollen transfer. The presence of an exotic plant may decrease the visitation rate to native plants and thus increase pollen limitation. Flower visitors may also switch between exotic and native plants and if pollen from an exotic plant is transferred to native plant stigmas this may impede siring by conspecific pollen. As flower visitors forage within a spatial context, the distribution of plants may affect the type and magnitude of pollinator‐mediated competition. In this study we examined two questions: 1) does the exotic plant, Carduus nutans (Asteraceae) interact with the native Monarda fistulosa (Lamiaceae) through flower visitors by changing visitation rate and/or through heterospecific pollen transfer, and does this affect seed set of the native plant? 2) Does spatial context affect how the native and exotic plants interact through flower visitors? We created plots containing potted M. fistulosa with and without the presence of potted C. nutans. In the presence of C. nutans, M. fistulosa stigmas had significantly fewer conspecific and more C. nutans pollen grains. Visitation rate and seed set tended to be lower in these invaded plots, however they were not significant. In a second experiment, we examined whether changes in visitation rate to M. fistulosa due to the presence of C. nutans was a function of M. fistulosa distance from C. nutans. We found that visitation rate did not decrease in the presence of C. nutans when M. fistulosa were adjacent to C. nutans or 15 meters from C. nutans. However, floral visitation rate to M. fistulosa decreased at 1 and 5 meters from C. nutans. Our results suggest interactions between plant species through flower visitors may depend on spatial scale.     

Eco-evolutionary litter feedback as a driver of exotic plant invasion

Many studies have examined positive feedbacks between invasive plant traits and nutrient cycling, but few have investigated whether feedbacks arise from introduction of pre-adapted species or from eco-evolutionary feedback that develops after introduction. Eco-evolutionary feedback could occur between an invader's leaf tissue C:N ratio and its response to litter accumulation. Previous modeling predicts that occurrence of this feedback would be reflected by: (1) field data showing higher litter:biomass ratios in the invasive range; (2) high C:N genotypes benefiting more from experimental litter additions than low C:N genotypes; (3) this beneficial effect on high C:N genotypes inducing a critical transition toward invader dominance when a critical amount of litter is added to a native species-dominated community experiencing low nutrient conditions. Here, we empirically tested these predictions for the invasive grass Phalaris arundinacea, which has undergone post-introduction evolutionary change toward attaining higher C:N ratios under high nutrient conditions. We performed a biogeographical comparison of litter:biomass ratios in the native (Europe) and invasive (USA) range, and an experiment with mesocosms from the invasive range under low nutrient conditions. Low and high C:N Phalaris genotypes were introduced into native-dominated and bare mesocosms, to which varying litter amounts were added. The biogeographical comparison revealed that litter:biomass ratios were higher in the invasive range. The mesocosm experiment showed that when grown in isolation, only high C:N genotypes responded positively to litter. This effect, however, was not strong enough to stimulate Phalaris when exposed to competition with native species. Our results suggest that eco-evolutionary feedback between Phalaris’ C:N ratio and litter accumulation could occur, but only under high nutrient conditions. Our experiments suggest that eco-evolutionary feedback may select for specialist rather than superior genotypes. Hence, genotypic variation induced by post-introduction admixture may be subject to context-dependent selection due to eco-evolutionary feedback, increasing trait variation within invasive populations.     

No impact of a native beetle on exotic plant performance and competitive ability due to plant compensation

The novel associations between invasive plants and their natural enemies in the introduced range have recently received increasing attention; however, the effects of novel enemies on exotic plant performance and competition with native species remain poorly explored. Here, we tested the impact of herbivory by a native beetle, Cassida piperata, on the performance of the exotic species Alternanthera philoxeroides and competition with a native congener, Alternanthera sessilis, using common garden experiments in central China. We found A. philoxeroides was able to fully compensate for intense herbivory by C. piperata. Herbivory by C. piperata that released at the average density in this region had no impact on competition between the native and exotic plant species. Our results indicate that herbivory by novel enemies may not reduce exotic plant performance due to plant compensation. However, high tolerance to herbivory may not confer a competitive advantage for exotic species compared to less tolerant native competitors if the herbivore damage is below a certain threshold. Thus, it is necessary to assess the impact of novel enemies on exotic plant performance and competition with native plants along gradients of insect densities. This may lead to a better understanding of how best to exploit the role of native herbivores in facilitating or slowing plant invasions.     

Zonal plant distribution and edaphic and micrometeorological conditions on a coastal sand dune

Some edaphic and meteorological conditions were examined to detect environmental gradients from shoreline to inland at the Kado-ori coast, Ibaraki, Japan, in 1989. Zonal distribution patterns of coastal dune plant species, including three ubiquitous perennials,Calystegia soldanella, Carex kobomugi andIschaemum anthephoroides, were described in relation to the environmental gradients. Environmental gradients were found in water availability, evaporative demand and soil-water salinity. Water availability, evaluated by thickness of capillary water layer, increased from 10 cm at 20 m to 48 cm at 85 m from the shoreline, reflecting the percentage of fine sand. Evaporative demand, which was evaluated by the evaporation rate from a wet black filter paper, decreased with increasing distance from the shoreline. Soil-water salinity was lowest (15 mmol/L) at 85 m from the shoreline and highest (90 mmol/L) at 30m. On the coast,C. soldanella, a salt-tolerant perennial, was distributed mainly in the environmentally harsh area 40–60 m from the shoreline.Ischaemum anthephoroides andC. kobomugi, less salt-tolerant perennials, occurred mainly 70–80 m from the shoreline, where environmental conditions were more hospitable.     

Contrasting responses of native ant communities to invasion by an ant invader,Linepithema humile

Biological Invasions - Invasive alien species pose a serious threat to the integrity and function of natural ecosystems. Understanding how these invaders alter natural communities is therefore an...     

Relationship between plant cover type and soil properties on Syunkunitai coastal sand dune in eastern Hokkaido

A survey of soils and trees was conducted on Syunkunitai coastal sand dune in eastern Hokkaido to clarify the relationships between the soil properties and the plant cover type. A belt transect of 360m in length was established across the dune. Three community types, that is, a Picea glehnii forest, an Abies sachalinensis forest, and a salt marsh were recognized. Soil types at the study area were determined to be sandy immature soil and peat soil. Their horizon sequences were described as A₀–V–C or T–V–C layers (A₀, T, V, and C indicate layers of leaf litter, peat, volcanic deposit, and parent material, respectively). The Abies sachalinensis forest was characterized by a relatively high calcium concentration in the surface soil layer and a tendency for podzolization in the volcanic deposit layer. The Picea glehnii forest was characterized by peat accumulation because of the high ground water table, volcanic deposits in the soil profile, and the strong influence of sea salt on the soil chemistry. The roots in the Picea glehnii forest were distributed more shallowly than those in the Abies sachalinensis forest, thus avoiding the high water table level as well as the influence of seawater in the soil. The salt marsh showed an extremely high sodium concentration and base saturation, indicating that this area was directly affected by seawater. Recently, the periphery of the Picea glehnii forest on Syunkunitai sand dune has been declining because of seawater inundation caused by ground subsidence.     

Exotic vertebrate and invertebrate herbivores differ in their impacts on native and exotic plants: a meta-analysis

Herbivores modify various ecological processes including interactions between native and exotic plants that may affect invasion success by the exotic plants. It is unknown whether different types of exotic herbivores have similar effects on native and exotic plants. Using two distinct data sets, we ran meta-analyses to compare exotic vertebrate and invertebrate herbivore preferences for, and effects on performance and population sizes of native and exotic plants. We found that exotic vertebrate herbivores have positive effects on exotic plant performance and population sizes, and no significant effects on native plants. Exotic invertebrates have significant negative effects on performance and population sizes of both exotic and native plants. Vertebrates prefer to feed on native plants relative to exotic plants, while invertebrates prefer the exotic plants to native plants. Thus the exotic vertebrate herbivores may aid invasiveness of exotic plants, in accordance with the invasional meltdown hypothesis, while exotic invertebrate herbivores probably have no net effect on invasion process of the exotic plants. Invertebrate herbivore preferences for exotic plants support the biotic resistance hypothesis, as the native plants probably resist the invertebrate herbivory. We also tested an evolutionary logic that posits that herbivores with similar evolutionary history as plants will affect the plants less negatively than plants with which they have not co-evolved. Our results indicate that there is no consistent pattern in effects of exotic vertebrate and invertebrate herbivores on exotic plants with or without which they have co-evolved.     

Phytophagous insect fauna tracks host plant responses to exotic grass invasion

The high dependence of herbivorous insects on their host plants implies that plant invaders can affect these insects directly, by not providing a suitable habitat, or indirectly, by altering host plant availability. In this study, we sampled Asteraceae flower heads in cerrado remnants with varying levels of exotic grass invasion to evaluate whether invasive grasses have a direct effect on herbivore richness independent of the current disturbance level and host plant richness. By classifying herbivores according to the degree of host plant specialization, we also investigated whether invasive grasses reduce the uniqueness of the herbivorous assemblages. Herbivorous insect richness showed a unimodal relationship with invasive grass cover that was significantly explained only by way of the variation in host plant richness. The same result was found for polyphagous and oligophagous insects, but monophages showed a significant negative response to the intensity of the grass invasion that was independent of host plant richness. Our findings lend support to the hypothesis that the aggregate effect of invasive plants on herbivores tends to mirror the effects of invasive plants on host plants. In addition, exotic plants affect specialist insects differently from generalist insects; thus exotic plants affect not only the size but also the structural profile of herbivorous insect assemblages.     

Responses of photosynthesis,leaf conductance and growth to different salinities in three coastal dune plants

The effects of soil-water salinity on growth and photosynthesis of three coastal dune plants were examined by salt-treatment in order to clarify the causal relationship between salinity and plant distribution in a dune habitat. Plants were cultivated hydroponically at three salinity levels: 0, 10 and 100 mM NaCl. With the 100 mM salt treatment,Calystegia soldanella (C₃ species) had the highest relative growth rate (RGR) (0.085 g g⁻¹ d⁻¹), followed byCarex kobomugi (C₃) (0.066), andIschaemum anthephoroides (C₄) (0.060). This order coincides with the distribution pattern of the three species on coastal dunes;Calystegia soldanella is generally distributed in more seaward areas whereasI. anthephoroides occurs further inland. The order of RGR was determined exclusively by leaf area ratio (LAR) among the three species. Due to its C₄ pathway,I. anthephoroides had higher net photosynthetic rate (Pn) and net assimilation rate (NAR) than the two C₃ plants at all NaCl concentrations, despite its low RGR. This apparent discrepancy is explainable by differences of LAR among the three species; LAR ofI. anthephoroides was lowest, and about half that ofCalystegia soldanella. These results suggest that LAR is one of the main determinants of salt tolerance based on RGR, whereas Pn or NAR may not be significant. This article is dedicated to Professor Hideo Iwaki, University of Tsukuba, in appreciation of the sincere encouragement he has given to the authors.     

California native and exotic perennial grasses differ in their response to soil nitrogen,exotic annual grass density,and order of emergence

Early emergence of plant seedlings can offer strong competitive advantages over later-germinating neighbors through the preemption of limiting resources. This phenomenon may have contributed to the persistent dominance of European annual grasses over native perennial grasses in California grasslands, since the former species typically germinate earlier in the growing season than the latter and grow rapidly after establishing. Recently, European perennial grasses have been spreading into both non-native annual and native perennial coastal grass stands in California. These exotic perennials appear to be less affected by the priority effects arising from earlier germination by European annual grasses. In addition, these species interactions in California grasslands may be mediated by increasing anthropogenic or natural soil nitrogen inputs. We conducted a greenhouse experiment to test the effects of order of emergence and annual grass seedling density on native and exotic perennial grass seedling performance across different levels of nitrogen availability. We manipulated the order of emergence and density of an exotic annual grass (Bromus diandrus) grown with either Nassella pulchra (native perennial grass), Festuca rubra (native perennial grass), or Holcus lanatus (exotic perennial grass), with and without added nitrogen. Earlier B. diandrus emergence and higher B. diandrus density resulted in greater reduction in the aboveground productivity of the perennial grasses. However, B. diandrus suppressed both native perennials to a greater extent than it did H. lanatus. Nitrogen addition had no effect on the productivity of native perennials, but greatly increased the growth of the exotic perennial H. lanatus, grown with B. diandrus. These results suggest that the order of emergence of exotic annual versus native perennial grass seedlings could play an important role in the continued dominance of exotic annual grasses in California. The expansion of the exotic perennial grass H. lanatus in coastal California may be linked to its higher tolerance of earlier-emerging annual grasses and its ability to access soil resources amidst high densities of annual grasses.     

Phenological and biogeographical aspects of coastal dune plant communities in southern Brazil

The coastline of southern Brazil is characterized by vast sand dunes. From March 1982 to April 1984 the dune vegetation was sampled along a transect between the Atlantic shore and an inland freshwater marsh, and phenological cycles of plant species were observed at monthly intervals.The occurrence of 23 annual species in the local dunes and peak germination, growth and flowering of 21 perennial species during spring, summer and fall, suggest that unfavourable temperatures and inundation of slacks during winter are seasonally limiting factors for this flora.Differences in the floristic composition between this region and its southern and northern extremes in the southwestern Atlantic together with a marked seasonality of the total species number and of germination and flowering periods, suggest that this area corresponds to a warm temperate biogeographic transition zone between northern tropical and southern cold temperate regions.     

Non-random co-occurrence of native and exotic plant species in Mediterranean grasslands

Invasion by exotic species in Mediterranean grasslands has determined assembly patterns of native and introduced species, knowledge of which provides information on the ecological processes underlying these novel communities. We considered grasslands from Spain and Chile. For each country we considered the whole grassland community and we split species into two subsets: in Chile, species were classified as natives or colonizers (i.e. exotics); in Spain, species were classified as exclusives (present in Spain but not in Chile) or colonizers (Spanish natives and exotics into Chile). We used null models and co-occurrence indices calculated in each country for each one of 15 sites distributed along a precipitation gradient and subjected to similar silvopastoral exploitation. We compared values of species co-occurrence between countries and between species subsets (natives/colonizers in Chile; exclusives/colonizers in Spain) within each country and we characterised them according to climatic variables. We hypothesized that: a) the different coexistence time of the species in both regions should give rise to communities presenting a spatial pattern further from random in Spain than in Chile, b) the co-occurrence patterns in the grasslands are affected by mesoclimatic factors in both regions. The patterns of co-occurrence are similar in Spain and Chile, mostly showing a spatial pattern more segregated than expected by random. The colonizer species are more segregated in Spain than in Chile, possibly determined by the longer residence time of the species in the source area than in the invaded one. The segregation of species in Chile is related to water availability, being species less segregated in habitat with greater water deficit; in Spain no relationship with climatic variables was found. After an invasion process, our results suggest that the possible process of alteration of the original Chilean communities has not prevented the assembly between the native and colonizer species together.     

Impact of exotic fish removal on native communities in farm ponds

Introduced largemouth bass (Micropterus salmoides spp.) and bluegill (Lepomis macrochirus spp.) are thought to threaten native aquatic organisms worldwide and hence their eradication has recently begun in Japan. Our previous studies suggested that the removal of largemouth bass increases native fish, shrimp, dragonflies, and exotic crayfish, but decreases macrophytes. To test this prediction, we removed the exotic fishes by draining farm ponds and compared the numbers of these organisms before and after the drain, as well as between drained and undrained ponds. The number of dragonfly Pseudothemis zonata, crayfish, shrimp, and goby increased rapidly after the drain, but the coverage of macrophyte declined. The reduction in macrophyte is assumed to be caused by increased herbivory by crayfish. The number of exuviae of damselfly Cercion calamorum and the total number of species of odonate also decreased after the drain. These decreases can be due to the reduction of macrophyte because reduced odonate species are known to use macrophytes as oviposition sites. Therefore, the removal of largemouth bass has a potential to cause negative effects on some native organisms. We propose that reduction of exotic crayfish should be considered when eradicating the exotic fishes.