Long-term dynamics of biotic and abiotic resistance to exotic species invasion in restored vernal pool plant communities

Invasion of native ecosystems by exotic species can seriously threaten native biodiversity, alter ecosystem function, and inhibit conservation. Moreover, restoration of native plant communities is often impeded by competition from exotic species. Exotic species invasion may be limited by unfavorable abiotic conditions and by competition with native species, but the relative importance of biotic and abiotic factors remains controversial and may vary during the invasion process. We used a long-term experiment involving restored vernal pool plant communities to characterize the temporal dynamics of exotic species invasion, and to evaluate the relative support for biotic and abiotic factors affecting invasion resistance. Experimental pools (n=256) were divided among controls and several seeding treatments. In most treatments, native vernal pool species were initially more abundant than exotic species, and pools that initially received more native seeds exhibited lower frequencies of exotic species over time. However, even densely seeded pools were eventually dominated by exotic species, following extreme climatic events that reduced both native and exotic plant densities across the study site. By the sixth year of the experiment, most pools supported more exotics than native vernal pool species, regardless of seeding treatment or pool depth. Although deeper pools were less invaded by exotic species, two exotics (Hordeum marinum and Lolium multiflorum) were able to colonize deeper pools as soon as the cover of native species was reduced by climatic extremes. Based on an information-theoretic analysis, the best model of invasion resistance included a nonlinear effect of seeding treatment and both linear and nonlinear effects of pool depth. Pool depth received more support as a predictor of invasion resistance, but seeding intensity was also strongly supported in multivariate models of invasion, and was the best predictor of resistance to invasion by H. marinum and L. multilorum. We conclude that extreme climatic events can facilitate exotic species invasions by both reducing abiotic constraints and weakening biotic resistance to invasion.     

A Two-Part Measure of Degree of Invasion for Cross-Community Comparisons

Abstract:  Invasibility is a critical feature of ecological communities, especially for management decisions. To date, invasibility has been measured in numerous ways. Although most researchers have used the richness (or number) of exotic species as a direct or indirect measure of community invasibility, others have used alternative measures such as the survival, density, or biomass of either a single or all exotic species. These different measures, even when obtained from the same communities, have produced inconsistent results and have made comparisons among communities difficult. Here, we propose a measure of the degree of invasion (DI) of a community as a surrogate for community invasibility. The measure is expressed as 2 independent components: exotic proportion of total species richness and exotic proportion of total species abundance (biomass or cover). By including richness and abundance, the measure reflects that the factors that control invasibility affect both of these components. Expressing exotic richness and abundance relative to the richness and abundance of all species in a community makes comparisons across communities of different sizes and resource availability possible and illustrates the importance of dominance of exotic species relative to natives, which is a primary management concern associated with exotic species.     

Inversion of plant dominance-diversity relationships along a latitudinal stress gradient

Species interactions affect plant diversity through the net effects of competition and facilitation, with the latter more prevalent in physically stressful environments when plant cover ameliorates abiotic stress. One explanation for species loss in invader-dominated systems is a shift in the competition-facilitation balance, with competition intensifying in areas formerly structured by facilitation. We test this possibility with a 10-site prairie meta-experiment along a 500-km latitudinal stress gradient, quantifying the relationships among abiotic stress, exotic dominance, and native plant recruitment over five years. The latitudinal gradient is inversely correlated with abiotic stress, with lower latitudes more moisture- and nutrient-limited. We observed strong negative effects by invasive dominant grasses on plant establishment, but only in northern sites with lower-stress environments. At these locations, disturbance was critical for recruitment by reducing the suppressive dominant (invasive) canopy. In more stressful environments to the south, the impacts of the dominant invaders on plant establishment became facilitative, and diversity was more limited by seed availability. Disturbance prevented recruitment because seedling survival depended on a protective plant canopy, presumably because the canopy reduced temperature or moisture stress. Seed limitation was similarly prevalent in all sites. Our work confirms the importance of facilitation as an organizing process for plants in higher-stress environments, even with transformations of species composition and dominance. It also demonstrates that the mechanisms regulating diversity, including invader impacts, can vary within the same plant community depending on environmental context. Because limits on native plant recruitment are environmentally contingent, management strategies that seek to increase diversity, including invader eradication, must account for site-level variations in the balance between biotic and abiotic constraints.     

Effects of Cattle Grazing on Diversity in Ephemeral Wetlands

Abstract:  Cattle are usually thought of as a threat to biodiversity. In regions threatened by exotic species invasion and lacking native wild grazers, however, cattle may produce the type of disturbance that helps maintain diverse communities. Across 72 vernal pools, I examined the effect of different grazing treatments (ungrazed, continuously grazed, wet-season grazed and dry-season grazed) on vernal-pool plant and aquatic faunal diversity in the Central Valley of California. After 3 years of treatment, ungrazed pools had 88% higher cover of exotic annual grasses and 47% lower relative cover of native species than pools grazed at historical levels (continuously grazed). Species richness of native plants declined by 25% and aquatic invertebrate richness was 28% lower in the ungrazed compared with the continuously grazed treatments. Release from grazing reduced pool inundation period by 50 to 80%, making it difficult for some vernal-pool endemic species to complete their life cycle. My results show that one should not assume livestock and ranching operations are necessarily damaging to native communities. In my central California study site, grazing helped maintain native plant and aquatic diversity in vernal pools.     

Dominant species identity, not community evenness, regulates invasion in experimental grassland plant communities

While there has been extensive interest in understanding the relationship between diversity and invasibility of communities, most studies have only focused on one component of diversity: species richness. Although the number of species can affect community invasibility, other aspects of diversity, including species identity and community evenness, may be equally important. While several field studies have examined how invasibility varies with diversity by manipulating species identity or evenness, the results are often confounded by resource heterogeneity, site history, or disturbance. We designed a mesocosm experiment to examine explicitly the role of dominant species identity and evenness on the invasibility of grassland plant communities. We found that the identity of the dominant plant species, but not community evenness, significantly impacted invasibility. Using path analysis, we found that community composition (dominant species identity) reduced invasion by reducing early-season light availability and increasing late-season plant community biomass. Nitrogen availability was an important factor for the survival of invaders in the second year of the experiment. We also found significant direct effects of certain dominant species on invasion, although the mechanisms driving these effects remain unclear. The magnitude of dominant species effects on invasibility we observed are comparable to species richness effects observed in other studies, showing that species composition and dominant species can have strong effects on the invasibility of a community.     

The importance of quantifying propagule pressure to understand invasion: an examination of riparian forest invasibility

The widely held belief that riparian communities are highly invasible to exotic plants is based primarily on comparisons of the extent of invasion in riparian and upland communities. However, because differences in the extent of invasion may simply result from variation in propagule supply among recipient environments, true comparisons of invasibility require that both invasion success and propagule pressure are quantified. In this study, we quantified propagule pressure in order to compare the invasibility of riparian and upland forests and assess the accuracy of using a community's level of invasion as a surrogate for its invasibility. We found the extent of invasion to be a poor proxy for invasibility. The higher level of invasion in the studied riparian forests resulted from greater propagule availability rather than higher invasibility. Furthermore, failure to account for propagule pressure may confound our understanding of general invasion theories. Ecological theory suggests that species-rich communities should be less invasible. However, we found significant relationships between species diversity and invasion extent, but no diversity-invasibility relationship was detected for any species. Our results demonstrate that using a community's level of invasion as a surrogate for its invasibility can confound our understanding of invasibility and its determinants.     

Composite forces shape population dynamics of copepod crustaceans

Understanding the processes that control species abundance and distribution is a major challenge in ecology, yet for a large number of potentially important organisms, we know little about the biotic and abiotic factors that influence population size. One group of aquatic organisms that defies traditional demographic analyses is the Crustacea, particularly those with complex life cycles. We used likelihood techniques and information theoretics to evaluate a suite of models representing alternative hypotheses on factors controlling the abundance of two copepod crustaceans in a small, tropical floodplain lake. Quantitative zooplankton samples were collected at three stations in a Venezuelan floodplain lake from June through December 1984; the average sampling interval was two days. We constructed a series of models with stage structure that incorporated six biotic and abiotic covariates in various combinations to account for temporal changes in abundance of these target species and in their population growth rates. Our analysis produced several novel insights into copepod population dynamics. We found that multiple forces affected the abundance of particular stages, that these factors differed between species as well as among stages within each species, and that biotic processes had the largest effects on copepod population dynamics. Density dependence had a large effect on the survival of Oithona amazonica copepodites and on population growth rate of Diaptomus negrensis.     

Effects of an Exotic Plant Invasion on Native Understory Plants in a Tropical Dry Forest

Abstract: The dry forests of southern India, which are endangered tropical ecosystems and among the world's most important tiger (Panthera tigris) habitats, are extensively invaded by exotic plants. Yet, experimental studies exploring the impacts of these invasions on native plants in these forests are scarce. Consequently, little is known about associated implications for the long‐term conservation of tigers and other biodiversity in these habitats. I studied the impacts of the exotic plant Lantana camara on understory vegetation in a dry‐forest tiger habitat in southern India. I compared the richness, composition, and abundance of tree seedlings, herbs, and shrubs and the abundance of grass among plots in which Lantana was cleared or left standing. These plots were distributed across two blocks—livestock free and livestock grazed. Removal of Lantana had an immediate positive effect on herb–shrub richness in the livestock‐free block, but had no effect on that of tree seedlings in either livestock block. Tree‐seedling and herb–shrub composition differed significantly between Lantana treatment and livestock block, and Lantana removal significantly decreased survival of tree seedlings. Nevertheless, the absence of trees, in any stage between seedling and adult, indicates that Lantana may stall tree regeneration. Lantana removal decreased the abundance of all understory strata, probably because forage plants beneath Lantana are less accessible to herbivores, and plants in Lantana‐free open plots experienced greater herbivory. Reduced access to forage in invaded habitats could negatively affect ungulate populations and ultimately compromise the ability of these forests to sustain prey‐dependent large carnivores. Additional research focused on understanding and mitigating threats posed by exotic plants may be crucial to the long‐term protection of these forests as viable tiger habitats.     

Seed and establishment limitation contribute to long-term native forb declines in California grasslands

The effects of exotic species invasions on biodiversity vary with spatial scale, and documentation of local-scale changes in biodiversity following invasion is generally lacking. Coupling long-term observations of local community dynamics with experiments to determine the role played by exotic species in recruitment limitation of native species would inform both our understanding of exotic impacts on natives at local scales and regional-scale management efforts to promote native persistence. We used field experimentation to quantify propagule and establishment limitation in a suite of native annual forbs in a California reserve, and compared these findings to species abundance trends within the same sites over the past 48 years. Observations at 11 paired sites (inside and outside the reserve) indicated that exotic annual plants have continued to increase in abundance over the past 48 years. This trend suggests the system has not reached equilibrium > 250 years after exotic species began to spread, and 70 years after livestock grazing ceased within the reserve. Long-term monitoring observations also indicated that six native annual forb species went extinct from more local populations than were colonized. To determine the potential role of exotic species in these native plant declines, we added seed of these species into plots adjacent to monitoring sites where plant litter and live grass competition were removed. Experimental results suggest both propagule and establishment limitation have contributed to local declines observed for these native forbs. Recruitment was highest at sites that had current or historical occurrences of the seeded species, and in plots where litter was removed. Grazing history (i.e., location within or outside the reserve) interacted with exotic competition removal, such that removal of live grass competition increased recruitment in more recently grazed sites. Abundance of forbs was positively related to recruitment, while abundance of exotic forbs was negatively related. Thus, exotic competition is likely only one factor contributing to local declines of native species in invaded ecosystems, with a combination of propagule limitation, site quality, and land use history also playing important and interactive roles in native plant recruitment.     

Invasive insect abundance varies across the biogeographic distribution of a native host plant.

Studies of biogeographic variation in species abundances are fundamental to understanding and predicting the impacts of invasive exotic species. We quantified the abundance of the introduced and now invasive biocontrol weevil, Rhinocyllus conicus, on a newly adopted native host plant, Cirsium canescens (Platte thistle), across the plant's distributional range. We used regression and structural equation analyses to examine variation in weevil abundance at 92-108 sites over three years in relation to variation in abiotic and biotic parameters hypothesized to be important in insect or plant dynamics and distribution. We found that R. conicus now occurs throughout the majority of the range of C. canescens, even in the center of the native plant's distribution where its coevolved, targeted weed host (Carduus nutans, musk thistle) is absent. In fact, weevil densities were greater in the center of the native plant's distribution in the Sand Hills formation than in peripheral sand patches closer to areas where the targeted 'exotic thistle is common. None of the macroclimatic variables examined were consistent predictors of the large-scale variation in weevil abundance on C. canescens. In addition to biogeographic position, the only other consistent predictor of weevil densities across sites was the number of flower heads per C. canescens plant. These results exclude the "spillover" hypothesis to explain nontarget feeding on this newly adopted native host species. Instead, the results are consistent with the alternative hypothesis that exotic weevil abundance on C. canescens is related to the local availability of native floral resources. Because C. canescens densities have declined with increases in R. conicus at sites central in the plant's distribution, these results suggest that isolated, peripheral populations of C. canescens are likely to be critical for persistence of Platte thistle. More generally, this study suggests that the persistence of a native species that is impacted by an exotic natural enemy may require preservation of populations in habitats outside the optimal portion of the native species' distribution.     

Acceleration of Exotic Plant Invasion in a Forested Ecosystem by a Generalist Herbivore

Abstract: The successful invasion of exotic plants is often attributed to the absence of coevolved enemies in the introduced range (i.e., the enemy release hypothesis). Nevertheless, several components of this hypothesis, including the role of generalist herbivores, remain relatively unexplored. We used repeated censuses of exclosures and paired controls to investigate the role of a generalist herbivore, white‐tailed deer (Odocoileus virginianus), in the invasion of 3 exotic plant species (Microstegium vimineum, Alliaria petiolata, and Berberis thunbergii) in eastern hemlock (Tsuga canadensis) forests in New Jersey and Pennsylvania (U.S.A.). This work was conducted in 10 eastern hemlock (T. canadensis) forests that spanned gradients in deer density and in the severity of canopy disturbance caused by an introduced insect pest, the hemlock woolly adelgid (Adelges tsugae). We used maximum likelihood estimation and information theoretics to quantify the strength of evidence for alternative models of the influence of deer density and its interaction with the severity of canopy disturbance on exotic plant abundance. Our results were consistent with the enemy release hypothesis in that exotic plants gained a competitive advantage in the presence of generalist herbivores in the introduced range. The abundance of all 3 exotic plants increased significantly more in the control plots than in the paired exclosures. For all species, the inclusion of canopy disturbance parameters resulted in models with substantially greater support than the deer density only models. Our results suggest that white‐tailed deer herbivory can accelerate the invasion of exotic plants and that canopy disturbance can interact with herbivory to magnify the impact. In addition, our results provide compelling evidence of nonlinear relationships between deer density and the impact of herbivory on exotic species abundance. These findings highlight the important role of herbivore density in determining impacts on plant abundance and provide evidence of the operation of multiple mechanisms in exotic plant invasion.     

A new method for broad-scale modeling and projection of plant assemblages under climatic,biotic, and environmental cofiltering

There is increasing interestin broad-scale analysis, modeling, and prediction of the distribution and composition of plant species assemblages under climatic, environmental, and biotic change, particularly for conservation purposes. We devised a method to reliably predict the impact of climate change on large assemblages of plant communities, while also considering competing biotic and environmental factors. To this purpose, we first used multilabel algorithms in order to convert the task of explaining a large assemblage of plant communities into a classification framework able to capture with high cross-validated accuracy the pattern of species distributions under a composite set of biotic and abiotic factors. We applied our model to a large set of plant communities in the Swiss Alps. Our model explained presences and absences of 175 plant species in 608 plots with >87% cross-validated accuracy, predicted decreases in α, β, and γ diversity by 2040 under both moderate and extreme climate scenarios, and identified likely advantaged and disadvantaged plant species under climate change. Multilabel variable selection revealed the overriding importance of topography, soils, and temperature extremes (rather than averages) in determining the distribution of plant species in the study area and their response to climate change. Our method addressed a number of challenging research problems, such as scaling to large numbers of species, considering species relationships and rarity, and addressing an overwhelming proportion of absences in presence–absence matrices. By handling hundreds to thousands of plants and plots simultaneously over large areas, our method can inform broad-scale conservation of plant species under climate change because it allows species that require urgent conservation action (assisted migration, seed conservation, and ex situ conservation) to be detected and prioritized. Our method also increases the practicality of assisted colonization of plant species by helping to prevent ill-advised introduction of plant species with limited future survival probability.     

Widespread density-dependent seedling mortality promotes species coexistence in a highly diverse Amazonian rain forest

Negative density-dependent mortality can promote species coexistence through a spacing mechanism that prevents species from becoming too locally abundant. Negative density-dependent seedling mortality can be caused by interactions among seedlings or between seedlings and neighboring adults if the density of neighbors affects the strength of competition or facilitates the attack of natural enemies. We investigated the effects of seedling and adult neighborhoods on the survival of newly recruited seedlings for multiple cohorts of known age from 163 species in Yasuni National Park, Ecuador, an ever-wet, hyper-diverse lowland Amazonian rain forest. At local scales, we found a strong negative impact on first-year survival of conspecific seedling densities and adult abundance in multiple neighborhood sizes and a beneficial effect of a local tree neighborhood that is distantly related to the focal seedling. Once seedlings have survived their first year, they also benefit from a more phylogenetically dispersed seedling neighborhood. Across species, we did not find evidence that rare species have an advantage relative to more common species, or a community compensatory trend. These results suggest that the local biotic neighborhood is a strong influence on early seedling survival for species that range widely in their abundance and life history. These patterns in seedling survival demonstrate the role of density-dependent seedling dynamics in promoting and maintaining diversity in understory seedling assemblages. The assemblage-wide impacts of species abundance distributions may multiply with repeated cycles of recruitment and density-dependent seedling mortality and impact forest diversity or the abundance of individual species over longer time scales.     

Community assembly in experimental grasslands: suitable environment or timely arrival?

It is hard to defend the view that biotic communities represent a simple and predictable response to the abiotic environment. Biota and the abiotic environment interact, and the environment of an individual certainly includes its neighbors and visitors in the community. The complexity of community assembly calls forth a quest for general principles, yet current results and theories on assembly rules differ widely. Using a grassland microcosm as a model system, we manipulated fertility, disturbance by defoliation, soil/microclimate, and arrival order of species belonging to two groups differing in functional attributes. We analyzed the outcome of community assembly dynamics in terms of species richness, invasibility, and species composition. The analyses revealed strong environmental control over species richness and invasibility. Species composition was mainly determined by the arrival order of species, indicating that historical contingency may change the outcome of community assembly. The probability for multiple equilibria appeared to increase with productivity and environmental stability. The importance of arrival order offers an explanation of the difficulties in predicting local occurrences of species in the field. In our experiment, variation in fertility and disturbance was controlling colonization with predictable effects on emergent community properties such as species richness. The key mechanism is suggested to be asymmetric competition, and our results show that this mechanism is relatively insensitive to the species through which it works. While our analyses indicate a positive and significant correlation between richness and invasibility, the significance disappears after accounting for the effect of the environment. The importance of arrival order (historical contingency) and environmental control supports the assumption of the unified neutral theory that different species within a trophic level can be considered functionally equivalent when it comes to community assembly. However, our results indicate that variation in asymmetric competition is the key factor determining the richness of the resulting communities, and this is far from neutral.     

Effect of Invasive Plant Species on Temperate Wetland Plant Diversity

Abstract:  Invasive species are a major threat to global biodiversity and an important cause of biotic homogenization of ecosystems. Exotic plants have been identified as a particular concern because of the widely held belief that they competitively exclude native plant species. We examined the correlation between native and invasive species richness in 58 Ontario inland wetlands. The relationship between exotic and native species richness was positive even when we controlled for important covarying factors. In addition, we examined the relationship between the abundance of four native species (  Typha latifolia, T. angustifolia, Salix petiolaris, Nuphar variegatum ) and four invasive species (  Lythrum salicaria, Hydrocharis morsus-ranae, Phalaris arundinacea, Rhamnus frangula ) that often dominate temperate wetlands and native and rare native species richness. Exotic species were no more likely to dominate a wetland than native species, and the proportion of dominant exotic species that had a significant negative effect on the native plant community was the same as the proportion of native species with a significant negative effect. We conclude that the key to conservation of inland wetland biodiversity is to discourage the spread of community dominants, regardless of geographical origin.     

Remote analysis of biological invasion and the impact of enemy release

Escape from natural enemies is a widely held generalization for the success of exotic plants. We conducted a large-scale experiment in Hawaii (USA) to quantify impacts of ungulate removal on plant growth and performance, and to test whether elimination of an exotic generalist herbivore facilitated exotic success. Assessment of impacted and control sites before and after ungulate exclusion using airborne imaging spectroscopy and LiDAR, time series satellite observations, and ground-based field studies over nine years indicated that removal of generalist herbivores facilitated exotic success, but the abundance of native species was unchanged. Vegetation cover <1 m in height increased in ungulate-free areas from 48.7% +/- 1.5% to 74.3% +/- 1.8% over 8.4 years, corresponding to an annualized growth rate of lambda = 1.05 +/- 0.01 yr(-1) (median +/- SD). Most of the change was attributable to exotic plant species, which increased from 24.4% +/- 1.4% to 49.1% +/- 2.0%, (lambda = 1.08 +/- 0.01 yr(-1)). Native plants experienced no significant change in cover (23.0% +/- 1.3% to 24.2% +/- 1.8%, lambda = 1.01 +/- 0.01 yr(-1)). Time series of satellite phenology were indistinguishable between the treatment and a 3.0-km2 control site for four years prior to ungulate removal, but they diverged immediately following exclusion of ungulates. Comparison of monthly EVI means before and after ungulate exclusion and between the managed and control areas indicates that EVI strongly increased in the managed area after ungulate exclusion. Field studies and airborne analyses show that the dominant invader was Senecio madagascariensis, an invasive annual forb that increased from < 0.01% to 14.7% fractional cover in ungulate-free areas (lambda = 1.89 +/- 0.34 yr(-1)), but which was nearly absent from the control site. A combination of canopy LAI, water, and fractional cover were expressed in satellite EVI time series and indicate that the invaded region maintained greenness during drought conditions. These findings demonstrate that enemy release from generalist herbivores can facilitate exotic success and suggest a plausible mechanism by which invasion occurred. They also show how novel remote-sensing technology can be integrated with conservation and management to help address exotic plant invasions.     

The invasibility of marine algal assemblages: role of functional diversity and identity

The emergence of the biodiversity-ecosystem functioning debate in the last decade has renewed interest in understanding why some communities are more easily invaded than others and how the impact of invasion on recipient communities and ecosystems varies. To date most of the research on invasibility has focused on taxonomic diversity, i.e., species richness. However, functional diversity of the communities should be more relevant for the resistance of the community to invasions, as the extent of functional differences among the species in an assemblage is a major determinant of ecosystem processes. Although coastal marine habitats are among the most heavily invaded ecosystems, studies on community invasibility and vulnerability in these habitats are scarce. We carried out a manipulative field experiment in tide pools of the rocky intertidal to test the hypothesis that increasing functional richness reduces the susceptibility of macroalgal communities to invasion. We selected a priori four functional groups on the basis of previous knowledge of local species characteristics: encrusting, turf, subcanopy, and canopy species. Synthetic assemblages containing one, two, three, or four different functional groups of seaweeds were created, and invasion by native species was monitored over an eight-month period. Cover and resource availability in the assemblages with only one functional group showed different patterns in the use of space and light, confirming true functional differences among our groups. Experimental results showed that the identity of functional groups was more important than functional richness in determining the ability of macroalgal communities to resist invasion and that resistance to invasion was resource-mediated.     

The adaptive value of remnant native plants in invaded communities: an example from the Great Basin

Changes in the species composition of biotic communities may alter patterns of natural selection occurring within them. Native perennial grass species in the Intermountain West are experiencing a shift in the composition of interspecific competitors from primarily perennial species to an exotic, annual grass. Thus traits that confer an advantage to perennial grasses in the presence of novel annual competitors may evolve in invaded communities. Here I show that such traits are apparent in populations of a native perennial grass, big squirreltail (Elymus multisetus M.E. Jones), exposed to cheatgrass (Bromus tectorum L.) competitors. Dormant big squirreltail plants were collected from cheatgrass-invaded and uninvaded sites near Bordertown, California, USA, a mid-elevation (1600 m) sagebrush community, and transplanted into pots in a greenhouse. Individual plants were split into equal halves. One half was grown with competition from cheatgrass, and the other half was grown without competition. Plants collected from invaded sites responded more quickly to watering, growing more leaves in the first 10 days after transplanting. In addition, big squirreltail plants collected from invaded areas experienced a smaller decrease in plant size when grown with competition than did plants collected from uninvaded areas. Accordingly, while there were fewer big squirreltail individuals in the invaded sites, they were more competitive with cheatgrass than were the more abundant conspecifics in nearby uninvaded areas. It is possible that annual grasses were the selective force that caused these population differences, which may contribute to the long-term persistence of the native populations. While it is tempting to restore degraded areas to higher densities of natives (usually done by bringing in outside seed material), such actions may impede long-term adaptation to new conditions by arresting or reversing the direction of ongoing natural selection in the resident population. If hot spots of rapid evolutionary change can be identified within invaded systems, these areas should be managed to promote desirable change and could serve as possible sources of restoration material or reveal traits that should be prioritized during the development of restoration seed material.     

Multi-trophic invasion resistance in Hawaii: bioacoustics, field surveys, and airborne remote sensing

We used airborne imaging spectroscopy and scanning light detection and ranging (LiDAR), along with bioacoustic recordings, to determine how a plant species invasion affects avian abundance and community composition across a range of Hawaiian submontane ecosystems. Total avian abundance and the ratio of native to exotic avifauna were highest in habitats with the highest canopy cover and height. Comparing biophysically equivalent sites, stands dominated by native Metrosideros polymorpha trees hosted larger native avian communities than did mixed stands of Metrosideros and the invasive tree Morella faya. A multi-trophic analysis strongly suggests that native avifauna provide biotic resistance against the invasion of Morella trees and exotic birds, thus slowing invasion "meltdowns" that disrupt the functioning of native Hawaiian ecosystems.     

Additive effects of aboveground polyphagous herbivores and soil feedback in native and range-expanding exotic plants

Plant biomass and plant abundance can be controlled by aboveground and belowground natural enemies. However, little is known about how the aboveground and belowground enemy effects may add up. We exposed 15 plant species to aboveground polyphagous insect herbivores and feedback effects from the soil community alone, as well as in combination. We envisaged three possibilities: additive, synergistic, or antagonistic effects of the aboveground and belowground enemies on plant biomass. In our analysis, we included native and phylogenetically related range-expanding exotic plant species, because exotic plants on average are less sensitive to aboveground herbivores and soil feedback than related natives. Thus, we examined if lower sensitivity of exotic plant species to enemies also alters aboveground-belowground interactions. In a greenhouse experiment, we exposed six exotic and nine native plant species to feedback from their own soil communities, aboveground herbivory by polyphagous insects, or a combination of soil feedback and aboveground insects and compared shoot and root biomass to control plants without aboveground and belowground enemies. We observed that for both native and range-expanding exotic plant species effects of insect herbivory aboveground and soil feedback added up linearly, instead of enforcing or counteracting each other. However, there was no correlation between the strength of aboveground herbivory and soil feedback. We conclude that effects of polyphagous aboveground herbivorous insects and soil feedback add up both in the case of native and related range-expanding exotic plant species, but that aboveground herbivory effects may not necessarily predict the strengths of soil feedback effects.