Plant invasions, generalist herbivores, and novel defense weapons

One commonly accepted mechanism for biological invasions is that species, after introduction to a new region, leave behind their natural enemies and therefore increase in distribution and abundance. However, which enemies are escaped remains unclear. Escape from specialist invertebrate herbivores has been examined in detail, but despite the profound effects of generalist herbivores in natural communities their potential to control invasive species is poorly understood. We carried out parallel laboratory feeding bioassays with generalist invertebrate herbivores from the native (Europe) and from the introduced (North America) range using native and nonnative tetraploid populations of the invasive spotted knapweed, Centaurea stoebe. We found that the growth of North American generalist herbivores was far lower when feeding on C. stoebe than the growth of European generalists. In contrast, North American and European generalists grew equally well on European and North American tetraploid C. stoebe plants, lending no support for an evolutionary change in resistance of North American tetraploid C. stoebe populations against generalist herbivores. These results suggest that biogeographical differences in the response of generalist herbivores to novel plant species have the potential to affect plant invasions.     

Postdispersal seed predation limits the abundance of a long-lived perennial forb (Lithospermum ruderale)

Loss of seeds to consumers is common in plant communities, but the degree to which these losses influence plant abundance or population growth is often unclear. This is particularly the case for postdispersal seed predation by rodents, as most studies of rodent seed predation have focused on the sources of spatiotemporal variation in seed loss but not quantified the population consequences of this loss. In previous work we showed that seed predation by deer mice (Peromyscus maniculatus) substantially reduced seedling recruitment and establishment of Lithospermum ruderale (Boraginaceae), a long-lived perennial forb. To shed light on how rodent seed predation and the near-term effects on plant recruitment might influence longer-term patterns of L. ruderale population growth, we combined experimental results with demographic data in stage-based population models. Model outputs revealed that rodent seed predation had a significant impact on L. ruderale population growth rate (lambda). With the removal of postdispersal seed predation, the projected population growth rates increased between 0.06 and 0.12, depending on site (mean deltalambda across sites = 0.08). Seed predation shifted the projected stable stage distribution of populations from one with a high proportion of young plants to one in which larger adult size classes dominate. Elasticities of vital rates also changed, with germination and growth of seedlings and young plants becoming more important with the removal of seed predation. Simulations varying the magnitude of seed predation pressure while holding other vital rates constant showed that seed predation could lower lambda even if only 40% of available seeds were consumed. These results demonstrate that rodent granivory can be a potent force limiting the abundance of a long-lived perennial forb.     

Introduced Birds and the Fate of Hawaiian Rainforests

Abstract:  The Hawaiian Islands have lost nearly all their native seed dispersers, but have gained many frugivorous birds and fleshy-fruited plants through introductions. Introduced birds may not only aid invasions of exotic plants but also may be the sole dispersers of native plants. We assessed seed dispersal at the ecotone between native- and exotic-dominated forests and quantified bird diets, seed rain from defecated seeds, and plant distributions. Introduced birds were the primary dispersers of native seeds into exotic-dominated forests, which may have enabled six native understory plant species to become reestablished. Some native plant species are now as common in exotic forest understory as they are in native forest. Introduced birds also dispersed seeds of two exotic plants into native forest, but dispersal was localized or establishment minimal. Seed rain of bird-dispersed seeds was extensive in both forests, totaling 724 seeds of 9 native species and 2 exotics with over 85% of the seeds coming from native plants. Without suitable native dispersers, most common understory plants in Hawaiian rainforests now depend on introduced birds for dispersal, and these introduced species may actually facilitate perpetuation, and perhaps in some cases restoration, of native forests. We emphasize, however, that restoration of native forests by seed dispersal from introduced birds, as seen in this study, depends on the existence of native forests to provide a source of seeds and protection from the effects of ungulates. Our results further suggest that aggressive control of patches of non-native plants within otherwise native-dominated forests may be an important and effective conservation strategy.     

Weed-biocontrol insects reduce native-plant recruitment through second-order apparent competition

Small-mammal seed predation is an important force structuring native-plant communities that may also influence exotic-plant invasions. In the intermountain West, deer mice (Peromyscus maniculatus) are prominent predators of native-plant seeds, but they avoid consuming seeds of certain widespread invasives like spotted knapweed (Centaurea maculosa). These mice also consume the biological-control insects Urophora spp. introduced to control C. maculosa, and this food resource substantially increases deer mouse populations. Thus, mice may play an important role in the invasion and management of C. maculosa through food-web interactions. We examined deer mouse seed predation and its effects on seedling emergence and establishment of a dominant native grass, Pseudoroegneria spicata, and forb, Balsamorhiza sagittata, in C. maculosa-invaded grasslands that were treated with herbicide to suppress C. maculosa or left untreated as controls. Deer mice readily took seeds of both native plants but removed 2-20 times more of the larger B. sagittata seeds than the smaller P. spicata seeds. Seed predation reduced emergence and establishment of both species but had greater impacts on B. sagittata. The intensity of seed predation corresponded with annual and seasonal changes in deer mouse abundance, suggesting that abundance largely determined mouse impacts on native-plant seeds. Accordingly, herbicide treatments that reduced mouse abundance by suppressing C. maculosa and its associated biocontrol food subsidies to mice also reduced seed predation and decreased the impact of deer mice on B. sagittata establishment. These results provide evidence that Urophora biocontrol agents may exacerbate the negative effects of C. maculosa on native plants through a form of second-order apparent competition-a biocontrol indirect effect that has not been previously documented. Herbicide suppressed C. maculosa and Urophora, reducing mouse populations and moderating seed predation on native plants, but the herbicide's direct negative effects on native forb seedlings overwhelmed the indirect positive effect of reducing deer mouse seed predation. By manipulating this four-level food chain, we illustrate that host-specific biological control agents may impact nontarget plant species through food-web interactions, and herbicides may influence management outcomes through indirect trophic interactions in addition to their direct effects on plants.     

Population-level compensation impedes biological control of an invasive forb and indirect release of a native grass

The intentional introduction of specialist insect herbivores for biological control of exotic weeds provides ideal but understudied systems for evaluating important ecological concepts related to top-down control, plant compensatory responses, indirect effects, and the influence of environmental context on these processes. Centaurea stoebe (spotted knapweed) is a notorious rangeland weed that exhibited regional declines in the early 2000s, attributed to drought by some and to successful biocontrol by others. We initiated an experiment to quantify the effects of the biocontrol agent, Cyphocleonus achates, on Ce. stoebe and its interaction with a dominant native grass competitor, Pseudoroegneria spicata, under contrasting precipitation conditions. Plots containing monocultures of each plant species or equal mixtures of the two received factorial combinations of Cy. achates herbivory (exclusion or addition) and precipitation (May-June drought or "normal," defined by the 50-year average) for three years. Cy. achates herbivory reduced survival of adult Ce. stoebe plants by 9% overall, but this effect was stronger under normal precipitation compared to drought conditions, and stronger in mixed-species plots compared to monocultures. Herbivory had no effect on Ce. stoebe per capita seed production or on recruitment of seedlings or juveniles. In normal-precipitation plots of mixed composition, greater adult mortality due to Cy. achates herbivory resulted in increased recruitment of new adult Ce. stoebe. Due to this compensatory response to adult mortality, final Ce. stoebe densities did not differ between herbivory treatments regardless of context. Experimental drought reduced adult Ce. stoebe survival in mixed-species plots but did not impede recruitment of new adults or reduce final Ce. stoebe densities, perhaps due to the limited duration of the treatment. Ce. stoebe strongly depressed P. spicata reproduction and recruitment, but these impacts were not substantively alleviated by herbivory on Ce. stoebe. Population-level compensation by dominant plants may be an important factor inhibiting top-down effects in herbivore-driven and predator-driven cascades.     

Mitigating exotic impacts: restoring deer mouse populations elevated by an exotic food subsidy.

The threat posed by exotic organisms to native systems has led to extensive research on exotic invaders, yet management of invasives has progressed relatively slowly. This is partly due to poor understanding of how exotic species management influences native organisms. To address this shortfall, we experimentally evaluated the efficacy of an invasives management tool for restoring native deer mouse (Peromyscus maniculatus) populations elevated by exotic species. The exotic insects, Urophora spp., were introduced in North America for biological control of the Eurasian invader, spotted knapweed (Centaurea maculosa), but instead of controlling C. maculosa, Urophora have become an important food resource that doubles P. maniculatus populations, with substantial indirect effects on other organisms. We hypothesized that herbicide suppression of Urophora's host plant would reduce the Urophora food resource and restore P. maniculatus populations to natural levels. Prior to treatment, mouse populations did not differ between controls and treatments, but following treatment, P. maniculatus were half as abundant where treatment reduced Urophora. Peromyscus maniculatus is insensitive to direct herbicide effects, and herbicide-induced habitat changes could not explain the P. maniculatus response. Treatment-induced reductions of the Urophora food resource offered the most parsimonious explanation for the mouse response: Multistate mark-recapture models indicated that P. maniculatus survival declined where Urophora were removed, and survival rates were more correlated with variation in population size than movement rates. Other demographic and reproductive parameters (sex ratios, reproductive status, pregnancy rates, and juvenile recruitment) were unaffected by treatment. These results suggest the Urophora biocontrol elevated P. maniculatus survival, and the herbicide treatment restored mouse populations by removing the exotic food and reducing survival. This work illustrates the importance of mechanistic understandings of community and population ecology for improving invasive species management.     

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.     

Vertebrate herbivory impacts seedling recruitment more than niche partitioning or density-dependent mortality

In tropical forests, resource-based niches and density-dependent mortality are mutually compatible mechanisms that can act simultaneously to limit seedling populations. Differences in the strengths of these mechanisms will determine their roles in maintaining species coexistence. In the first assessment of these mechanisms in a Congo Basin forest, we quantified their relative strengths and tested the extent to which density-dependent mortality is driven by the distance-dependent behavior of seed and seedling predators predicted by the Janzen-Connell hypothesis. We conducted a large-scale seed addition experiment for five randomly selected tropical tree species, caging a subset of seed addition quadrats against vertebrate predators. We then developed models to assess the mechanisms that determine seedling emergence (three months after seed addition) and survival (two years after seed addition). As predicted, both niche differentiation and density-dependent mortality limited seedling recruitment, but predation had the strongest effects on seedling emergence and survival. Seedling species responded differently to naturally occurring environmental variation among sites, including variation in light levels and soil characteristics, supporting predictions of niche-based theories of tropical tree species coexistence. The addition of higher densities of seeds into quadrats initially led to greater seedling emergence, but survival to two years decreased with seed density. Seed and seedling predation reduced recruitment below levels maintained by density-dependent mortality, an indication that predators largely determine the population size of tree seedlings. Seedling recruitment was unrelated to the distance to or density of conspecific adult trees, suggesting that recruitment patterns are generated by generalist vertebrate herbivores rather than the specialized predators predicted by the Janzen-Connell hypothesis. If the role of seed and seedling predation in limiting seedling recruitment is a general phenomenon, then the relative abundances of tree species might largely depend on species-specific adaptations to avoid, survive, and recover from damage induced by vertebrate herbivores. Likewise, population declines of herbivorous vertebrate species (many of which are large and hunted) may trigger shifts in species composition of tropical forests.     

Invasive competitor and native seed predators contribute to rarity of the narrow endemic Astragalus sinuatus Piper

The conservation of rare plant species hinges on our ability to identify the underlying mechanisms that limit rare plant populations. Theory on rarity suggests that both predispersal seed predation and competition can be important mechanisms influencing abundance and/or distribution of rare plant populations. Yet few studies have tested these interactions, and those that have evaluated each mechanism independently. Astragalus sinuatus Piper (Whited's milkvetch) is a narrow endemic plant species restricted to eight populations within a 10-km2 area in eastern Washington. We used experimental and observational methods to test the effects of native insect predispersal seed predators and an invasive grass (Bromus tectorum L. [cheatgrass]) on seed set and population density of A. sinuatus. We quantified per capita seed production and pod predation rates across four sites and among four years. Seed predation rates were high across four sites (66-82%) and all years (65-82%). Experimental reduction of predispersal seed predators significantly increased per capita seed set of A. sinuatus (164-345%) at two experimental sites. Concurrently, two seed addition experiments demonstrated the effect of seed loss and presence of B. tectorum on seedling recruitment and establishment of A. sinuatus over four growing seasons. In the first seed addition experiment, we found no difference in recruitment and establishment between low (40) and high (120) seed addition levels. In the second addition experiment (one level of addition; 40 seeds), we found that recruitment and survivorship increased 200% in plots where B. tectorum was removed compared to plots where B. tectorum was present. Thus, seed addition had no impact in the presence of B. tectorum; conversely, in the absence of B. tectorum, seed addition was highly effective at increasing population numbers. Results suggest that, in areas where B. tectorum is present, recruitment is site limited, and it is seed limited when B. tectorum is absent. We recommend that managers reduce B. tectorum in an effort to increase population growth of A. sinuatus; in areas where B. tectorum is absent, short-term reduction of insect predators should be considered as a strategy to increase population growth of this rare species.     

Seed predation by Neotropical rain forest mammals increases diversity in seedling recruitment

Seed dispersal and seedling recruitment (the transition of seeds to seedlings) set the spatiotemporal distribution of new individuals in plant communities. Many terrestrial rain forest mammals consume post-dispersal seeds and seedlings, often inflicting density-dependent mortality. In part because of density-dependent mortality, diversity often increases during seedling recruitment, making it a critical stage for species coexistence. We determined how mammalian predators, adult tree abundance, and seed mass interact to affect seedling recruitment in a western Amazonian rain forest. We used exclosures that were selectively permeable to three size classes of mammals: mice and spiny rats (weighing <1 kg), medium-sized rodents (1-12 kg), and large mammals (20-200 kg). Into each exclosure, we placed seeds of 13 tree species and one canopy liana, which varied by an order of magnitude in adult abundance and seed mass. We followed the fates of the seeds and resulting seedlings for at least 17 months. We assessed the effect of each mammalian size class on seed survival, seedling survival and growth, and the density and diversity of the seedlings that survived to the end of the experiment. Surprisingly, large mammals had no detectable effect at any stage of seedling recruitment. In contrast, small- and medium-sized mammals significantly reduced seed survival, seedling survival, and seedling density. Furthermore, predation by small mammals increased species richness on a per-stem basis. This increase in diversity resulted from their disproportionately intense predation on common species and large-seeded species. Small mammals thereby generated a rare-species advantage in seedling recruitment, the critical ingredient for frequency dependence. Predation by small (and to a lesser extent, medium-sized) mammals on seeds and seedlings significantly increases tree species diversity in tropical forests. This is the first long-term study to dissect the effects of various mammalian predators on the recruitment of a diverse set of tree species.     

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.     

Beyond the ecological: biological invasions alter natural selection on a native plant species

Biological invasions can have strong ecological effects on native communities by altering ecosystem functions, species interactions, and community composition. Even though these ecological effects frequently impact the population dynamics and fitness of native species, the evolutionary consequences of biological invasions have received relatively little attention. Here, I show that invasions impose novel selective pressures on a native plant species. By experimentally manipulating community composition, I found that the exotic plant Medicago polymorpha and the exotic herbivore Hypera brunneipennis alter the strength and, in some instances, the direction of natural selection on the competitive ability and anti-herbivore defenses of the native plant Lotus wrangelianus. Furthermore, the community composition of exotics influenced which traits were favored. For example, high densities of the exotic herbivore Hypera selected for increased resistance to herbivores in the native Lotus; however, when Medicago also was present, selection on this defense was eliminated. In contrast, selection on tolerance, another plant defense trait, was highest when both Hypera and Medicago were present at high densities. Thus, multiple exotic species may interact to influence the evolutionary trajectories of native plant populations, and patterns of selection may change as additional exotic species invade the community.     

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.     

Park Management of Exotic Plant Species: Problems and Issues

Abstract: Vegetation management policies in public parks in the United States call for the removal of exotic species to the extent feasible. The underlying goal is to preserve samples of wilderness by restoring plant communities to the "natural state" that existed prior to extensive human influence. With limited budgets, park managers are necessarily selective in targeting exotic species for control. If the focus is on the more readily controlled species, however, park landscapes may gradually become populated by more resistant exotics Further, because plants exhibit some redundancy in ecosystem function, exotic plant species can substitute in part for natives in performing a range of ecosystem functions, including wildlife support and soil binding. Consequently the removal of exotics can result in significant perturbations to certain ecosystem functions during the period of transition to native cover. The individualistic paradigm of plant distribution implies that the impact of exotic plant species on invaded communities will vary. Choosing which species to remove requires careful evaluation of the impact of the removal on ecosystem structure and function. The effective balancing of park management goals for wilderness maintenance and recreational use requires clearer recognition of the adaptive response of ecosystems to invasion and a rethinking of the bases for prioritizing which species are to be removed.     

Indirect effects of prey swamping: differential seed predation during a bamboo masting event

Resource pulses often involve extraordinary increases in prey availability that "swamp" consumers and reverberate through indirect interactions affecting other community members. We developed a model that predicts predator-mediated indirect effects induced by an epidemic prey on co-occurring prey types differing in relative profitability/preference and validated our model by examining current-season and delayed effects of a bamboo mass seeding event on seed survival of canopy tree species in mixed Patagonian forests. The model shows that predator foraging behavior, prey profitability, and the scale of prey swamping influence the character and strength of short-term indirect effects on various alternative prey. When in large prey-swamped patches, nonselective predators decrease predation on all prey types. Selective predators, instead, only benefit prey of similar quality to the swamping species, while very low or high preference prey remain unaffected. Negative indirect effects (apparent competition) may override such positive effects (apparent mutualism), especially for highly preferred prey, when prey-swamped patches are small enough to allow predator aggregation and/or predators show a reproductive numerical response to elevated food supply. Seed predation patterns during bamboo (Chusquea culeou) masting were consistent with predicted short-term indirect effects mediated by a selective predator foraging in large prey-swamped patches. Bamboo seeds and similarly-sized Austrocedrus chilensis (ciprés) and Nothofagus obliqua (roble) seeds suffered lower predation in bamboo flowered than nonflowered patches. Predation rates on the small-seeded Nothofagus dombeyi (coihue) and the large-seeded Nothofagus alpina (rauli) were independent of bamboo flowering. Indirect positive effects were transient; three months after bamboo seeding, granivores preyed heavily upon all seed types, irrespective of patch flowering condition. Moreover, one year after bamboo seeding, predation rates on the most preferred seed (rauli) was higher in flowered than in nonflowered patches. Despite rapid predator numerical responses, short-term positive effects can still influence community recruitment dynamics because surviving seeds may find refuge beneath the litter produced by bamboo dieback. Together, our theoretical analysis and experiments indicate that indirect effects experienced by alternative prey during and after prey-swamping episodes need not be universal but can change across a prey quality spectrum, and they critically depend on predator-foraging rules and the spatial scale of swamping.     

Human impacts, plant invasion, and imperiled plant species in California.

Invasive species are one of the fastest growing conservation problems. These species homogenize the world's flora and fauna, threaten rare and endemic species, and impose large economic costs. Here, we examine the distribution of 834 of the more than 1000 exotic plant taxa that have become established in California, USA. Total species richness increases with net primary productivity; however, the exotic flora is richest in low-lying coastal sites that harbor large numbers of imperiled species, while native diversity is highest in areas with high mean elevation. Weedy and invasive exotics are more tightly linked to the distribution of imperiled species than the overall pool of exotic species. Structural equation modeling suggests that while human activities, such as urbanization and agriculture, facilitate the initial invasion by exotic plants, exotics spread ahead of the front of human development into areas with high numbers of threatened native plants. The range sizes of exotic taxa are an order of magnitude smaller than for comparable native taxa. The current small range size of exotic species implies that California has a significant "invasion debt" that will be paid as exotic plants expand their range and spread throughout the state.     

Fungal endophytes directly increase the competitive effects of an invasive forb

Competitive outcomes among plants can vary in different abiotic and biotic conditions. Here we tested the effects of two phylotypes of Alternaria endophytes on the growth, competitive effects, and competitive responses of the exotic invasive forb Centaurea stoebe. Centaurea stoebe was a better competitor against North American grass species than grasses from its European home range in the absence of endophytes. However, one endophyte both increased the biomass of C. stoebe and reduced the competitive effect of North American grasses on C. stoebe. The competitive effects of C. stoebe on grass species native to North America were enhanced by both fungal endophytes, but not for native European grasses. We do not know the mechanism by which endophytes increased C. stoebe's competitive ability, and particularly against biogeographically new neighbors, but one endophyte increased the competitive ability of C. stoebe without increasing its size, suggesting mechanisms unrelated to increased growth. We tested only a fraction of the different endophytic fungi that have been found in C. stoebe, only scratching the surface of understanding their indirect effects. However, our results are the first to demonstrate such effects of a fungal endophyte infecting an invasive forb, and one of the few to show that endophyte effects on competition do not have to be mediated through herbivory.     

Habitat-specific impacts of multiple consumers on plant population dynamics

Multiple consumers often attack seeds, seedlings, and adult plants, but their population-level consequences remain uncertain. We examined how insect and small mammal consumers influenced the demography and abundance of the perennial shrub, bush lupine (Lupinus arboreus). In grassland and dune habitats we established replicate experimental lupine populations in 81-m2 plots that were either protected from, or exposed to, herbivorous voles and granivorous mice (via fencing) and/or root feeding insects (via insecticide treatment). Populations were initiated with transplanted seedlings in 1999 and 2000. We followed the demography of these cohorts, subsequent generations, and the seed bank for 5.5 years. Voles and insects killed many seedlings in dune (1999 only) and grassland (1999 and 2000) habitats. After 2000, insects and voles had minimal effects on seedling or adult survival. Seed predation by granivorous mice, however, greatly depressed seedling recruitment, resulting in lower adult lupine abundance in control plots vs. those protected from rodents. In grasslands, initial effects of voles and insects on seedling survival produced large differences among treatments in adult plant density and the cumulative number of seeds produced throughout the experiment. Differences among grassland populations in seed rain, however, had little influence on the magnitude of seedling recruitment into this habitat. Instead, recruitment out of a preexisting seed bank compensated for the lack of seed production in populations exposed to consumers. Shading by dense adults in plots protected from consumers limited seedling establishment within these populations. Although differences among populations in cumulative seed rain did not influence adult establishment, populations protected from consumers accumulated substantially larger seed banks than controls. These results illustrate how density dependence, habitat-specific seed dynamics, and particular demographic impacts of consumers interact to shape plant population responses to consumers.     

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.     

The natives are restless, but not often and mostly when disturbed

The argument that the threat posed by introduced species is overblown is often buttressed by the observation that native species sometimes also become invasive. An examination of the literature on plant invasions in the United States shows that six times more nonnative species have been termed invasive than native species, and that a member of the naturalized nonnative pool is 40 times more likely than a native species to be perceived as invasive. In the great majority of instances in which a native plant species is seen as invasive, the invasion is associated with an anthropogenic disturbance, especially changed fire or hydrological regime, livestock grazing, and changes wrought by an introduced species. These results suggest that natives are significantly less likely than nonnatives to be problematic for local ecosystems.