Responses of a native beetle to novel exotic plant species with varying invasion history

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Understanding plant drought resistance in a Mediterranean coastal sand dune ecosystem: differences between native and exotic invasive species

Aims Mediterranean coastal dunes are habitats of great conservation interest, with a distinctive and rich flora. In the last century, Acacia spp., native from Australia, have been introduced in Portugal, with the objective of stabilizing sand dunes, and since have become dominant in numerous sand dune habitats. This invasion process led to the reduction of native plant species richness, changed soil characteristics and modified habitat's microclimatic characteristics. The aim of this research was to typify and compare, in Mediterranean sand dune ecosystems, the ecophysiological responses to drought of Helichrysum italicum and Corema album, two native species, and Acacia longifolia, an exotic invasive species. We addressed the following specific objectives: (i) to compare water relations and water use efficiencies, (ii) to evaluate water stress, (iii) to assess water use strategies and water sources used by plants and (iv) to evaluate the morphological adaptations at leaf and phyllode level. Methods In order to obtain an integrative view of ecophysiological patterns, water relations and performance measuring methods have been applied: predawn (ψ PD) and midday (ψ MD) water potential, chlorophyll a fluorescence, oxygen isotopic composition of xylem, rain and groundwater (δ 18 O) and leaf carbon isotopic discrimination (Δ 13 C). The leaf characteristics of the three species, as well as the histochemistry of non-glandular trichome cell walls, were also studied to identify morpho-traits related to drought resistance.Important findings The results support our initial hypothesis: although A. longifolia clearly possesses a degree of resistance to water stress, such ability is provided by a different water strategy, when compared to native species. Natives relied on morphological adaptations to restrict water loss, whereas the invasive species adjusted the water uptake as a way to balance their limited ability of restricting water loss. We corroborate that woody native species (i) have a conservative water-saving strategy and minor seasonal variations relative to invasive species, (ii) use enriched water sources during drought periods, indicating different water sources and root systems comparing with invasive species and (iii) present drought leaf morpho-functional adaptations related with limiting water loss. Comparing the physiological performance of invasive and native species can offer causal explanations for the relative success of alien plant invasions on sand dunes ecosystems.     

Cave invertebrate assemblages differ between native and exotic leaf litter

Abstract Allochtonous leaf litter is an important source of energy and nutrients for invertebrates in cave ecosystems. A change to the quality or quantity of litter entering caves has the potential to disrupt the structure and function of cave communities. In this study, we adopted an experimental approach to examine rates of leaf litter decomposition and the invertebrate assemblages colonizing native and exotic leaf litter in limestone caves in the Jenolan Caves Karst Conservation Reserve, New South Wales, Australia. We deployed traps containing leaf litter from exotic sycamore (Acer pseudoplatanus) and radiata pine (Pinus radiata) trees and native eucalypts (Eucalyptus spp.) in twilight zones (near the cave entrance) and areas deep within the caves for 3 months. Thirty‐two invertebrate morphospecies were recorded from the litter traps, with greater richness and abundance evident in the samples from the twilight zone compared with areas deep within the cave. Sycamore litter had significantly greater richness and abundance of invertebrates compared with eucalypt and pine litter in samples from the twilight zone, but there was no difference in richness or abundance among litter samples placed deep within the cave. Relative rates of decay of the three litters were sycamore > eucalypt > pine. We discuss the potential for the higher decomposition rates and specific leaf area in sycamores to explain their higher invertebrate diversity and abundance. Our findings have important implications for the management of exotic plants and the contribution of their leaf litter to subterranean ecosystems.     

Responses of a native plant species from invaded and uninvaded areas to allelopathic effects of an invader

Invaders exert new selection pressures on the resident species, for example, through competition for resources or by using novel weapons. It has been shown that novel weapons aid invasion but it is unclear whether native species co‐occurring with invaders have adapted to tolerate these novel weapons. Those resident species which are able to adapt to new selective agents can co‐occur with an invader while others face a risk of local extinction. We ran a factorial common garden experiment to study whether a native plant species, Anthriscus sylvestris, has been able to evolve a greater tolerance to the allelochemicals exerted by the invader, Lupinus polyphyllus. Lupinus polyphyllus produces allelochemicals which potentially act as a novel, strong selective agent on A. sylvestris. We grew A. sylvestris seedlings collected from uninvaded (naïve) and invaded (experienced) sites growing alone and in competition with L. polyphyllus in pots filled with soil with and without activated carbon. Because activated carbon absorbs allelochemicals, its addition should improve especially naïve A. sylvestris performance in the presence of the invader. To distinguish the allelochemicals absorption and fertilizing effects of activated carbon, we grew plants also in a mixture of soil and fertilizer. A common garden experiment indicated that the performances of naïve and experienced A. sylvestris seedlings did not differ when grown with L. polyphyllus. The addition of activated carbon, which reduces interference by allelochemicals, did not induce differences in their performances although it had a positive effect on the aboveground biomass of A. sylvestris. Together, these results suggest that naïve and experienced A. sylvestris plants tolerated equally the invader L. polyphyllus and thus the tolerance has not occurred over the course of invasion.     

Phylogenetic patterns differ for native and exotic plant communities across a richness gradient in Northern California

Aim Increasingly, ecologists are using evolutionary relationships to infer the mechanisms of community assembly. However, modern communities are being invaded by non‐indigenous species. Since natives have been associated with one another through evolutionary time, the forces promoting character and niche divergence should be high. On the other hand, exotics have evolved elsewhere, meaning that conserved traits may be more important in their new ranges. Thus, co‐occurrence over sufficient time‐scales for reciprocal evolution may alter how phylogenetic relationships influence assembly. Here, we examined the phylogenetic structure of native and exotic plant communities across a large‐scale gradient in species richness and asked whether local assemblages are composed of more or less closely related natives and exotics and whether phylogenetic turnover among plots and among sites across this gradient is driven by turnover in close or distant relatives differentially for natives and exotics. Location Central and northern California, USA. Methods We used data from 30 to 50 replicate plots at four sites and constructed a maximum likelihood molecular phylogeny using the genes: matK, rbcl, ITS1 and 5.8s. We compared community‐level measures of native and exotic phylogenetic diversity and among‐plot phylobetadiversity. Results There were few exotic clades, but they tended to be widespread. Exotic species were phylogenetically clustered within communities and showed low phylogenetic turnover among communities. In contrast, the more species‐rich native communities showed higher phylogenetic dispersion and turnover among sites. Main conclusions The assembly of native and exotic subcommunities appears to reflect the evolutionary histories of these species and suggests that shared traits drive exotic patterns while evolutionary differentiation drives native assembly. Current invasions appear to be causing phylogenetic homogenization at regional scales.     

An exotic invader drives the evolution of plant traits that determine mycorrhizal fungal diversity in a native competitor

The symbiosis between land plants and arbuscular mycorrhizal fungi (AMF) is one of the most widespread and ancient mutualisms on the planet. However, relatively little is known about the evolution of these symbiotic plant–fungal interactions in natural communities. In this study, we investigated the symbiotic AMF communities of populations of the native plant species Pilea pumila (Urticaceae) with varying histories of coexistence with a nonmycorrhizal invasive species, Alliaria petiolata (Brassicaceae), known to affect mycorrhizal communities. We found that native populations of P. pumila with a long history of coexistence with the invasive species developed more diverse symbiotic AMF communities. This effect was strongest when A. petiolata plants were actively growing with the natives, and in soils with the longest history of A. petiolata growth. These results suggest that despite the ancient and widespread nature of the plant–AMF symbiosis, the plant traits responsible for symbiotic preferences can, nevertheless, evolve rapidly in response to environmental changes.     

Recolonizing native wildlife facilitates exotic plant invasion into Singapore's rain forests

Halting biological invasions and rewilding extirpated native fauna are conservation interventions to bolster biodiversity, species interactions, and ecosystems. These actions are often considered separately and the potential for reintroduced wildlife to facilitate invasive plants has been largely overlooked. Here, we investigated the role of Singapore's recolonizing native wild pigs (Sus scrofa) in facilitating an invasive weed Miconia crenata into tropical rainforests, which are normally highly resistant to invasion. We conducted line-transect surveys in 11 Singaporean rain forests and used generalized linear mixed models to consider the contribution of pigs' soil disturbances, human forest paths, and other environmental covariates, on the density of M. crenata. We found that M. crenata was more abundant at forest edges and invasion into forest interior was facilitated by pigs, paths, and canopy gaps, but that these effects were all additive, not synergistic (i.e., not multiplicative). These results highlight how modern invasions are driven by multiple disturbances as well as propagule pressure (e.g., urban birds dispersing seeds at forest edges where they establish in pig soil disturbances). Singapore's extensive native forest restoration efforts may have provided plentiful edge and secondary forests that are well suited to pigs and M. crenata, which in turn undermine the aims of fostering later-successional native plant communities. To prevent negative externalities, we suggest that plant restoration and rewilding projects consider the potential role of wildlife in facilitating non-native plants, and couple these actions with preliminary screening of unintended consequences and continued monitoring, as well as limiting human-mediated weed invasion to minimize propagule sources.     

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.

     

Relationship between native and exotic plant species at multiple savannah sites

We tested whether the recently proposed two‐part measure of degree of invasion (DI) of a community relating exotic proportion of cover to exotic proportion of richness can characterize patterns of plant invasions at multiple savannah sites in Southern Africa. Regression analysis was performed on transformed data to assess how this two‐part measure of DI compares to other metrics of community invasibility. The results indicate that at the plot level, the absolute cover of exotics was not significantly related to native cover for three sites out of four assessed (R² ≤ 0.17; P > 0.05). Also, at all four sites, no significant relationships were detected between native and exotic plant richness at both the 1‐m² and 400‐m² plot scales. By contrast, significant (P < 0.05) positive linear relationships were observed between exotic proportion of richness and exotic proportion of cover at all sites (R² was as high as 0.67 and 0.97 for two sites). Our results indicate that the new two‐part measure of DI is able to characterize patterns of plant invasions across plant communities in African savannahs.     

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.     

Recruitment limitation of native species in invaded coastal dune communities

Recruitment limitation may limit the ability of sites to regenerate after disturbances such as weed invasion and weed management. We investigated seed bank constraints and dispersal limitation in coastal dune communities on the east coast of Australia. The ability of sites to regenerate naturally following weed removal was assessed in coastal dune communities invaded by the invasive alien, bitou bush (Chrysanthemoides monilifera subsp. rotundata). To investigate recruitment limitation, seed banks and vegetation of invaded, native, intensively managed (selective application of herbicide and some re-vegetation) and extensively managed (large-scale, non-selective herbicide application) sites were compared. We investigated the dispersal mechanisms of species in the seed bank and vegetation to determine if communities might be dispersal-limited, i.e. contain significant numbers of species with only short-distance dispersal capabilities. Species richness and composition of soil seed banks differed from the vegetation in foredunes and hinddunes. Invasion depleted seed banks further. About half of the species had short-distance dispersal mechanisms indicating the potential for dispersal limitation. Secondary weed invasion following management was evident although alien species occurred in both seed banks and vegetation. Our results indicated that coastal dune communities suffer recruitment limitation. Native, managed and invaded dune communities appear to be both seed bank and dispersal-limited although management and invasion exacerbates recruitment. Regeneration of coastal dune communities will require active reintroduction of species, particularly those with short-distance dispersal mechanisms.     

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.     

Contrasting responses of native and alien plant species to soil properties shed new light on the invasion of dune systems

在陆地生态系统中,沿海沙丘特别容易受到外来植物的入侵。许多研究将沙丘生境的入侵归因于人为因素,但对土壤性质和植物性状在植物 入侵中的作用却知之甚少。本研究考察了沙丘系统土壤特征与外来植物入侵的关系,重点研究了土壤养分、土壤盐分和植物功能特征之间的相互作 用。研究地点是马拉诺和格拉多泻湖(北亚得里亚海)的沙质堰洲岛。根据沙丘系统生态梯度上的主要生境(前沙丘、后沙丘和盐沼),在10 个区域内选择100个地块(4 m × 4 m)。在每个地块中记录所有植物物种的发生和丰度,并收集一个土壤核。对每个土壤样品进行土壤质地、电导率(代表土壤盐 分)、有机碳和氮含量分析,并与本地和外来植物的种类数和覆盖度相关联。分析外来和本地物种主要的生殖功能和营养功能性状在生境中的变化。研究结果表明,土壤性质对外来物种库和本地物种库的影响不同,与植物整体多样性密切相关。在后沙丘(最易入侵)的生境中,高的土壤导电性限制了外来物种的数量,但土壤有机碳含量随着外来植物丰度的增加而增加,这也表明植物入侵与土壤之间存在潜在的反馈过程。只有在后沙丘生境中,随着环境条件的改善和植物竞争的加剧,本地植物和外来植物的功能性状谱有显著的趋同。本研究结果表明,在恶劣条件下,只有本地特有植物才能茁壮成长,而在中等条件下,土壤特性梯度与植物特性协同作用,抑制/促进外来植物丰富度。     

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.     

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.