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Although invasive alien species (IAS) are a major threat to biodiversity, human health, and economy, our understanding of the factors controlling their distribution and abundance is limited. Here, we determine how environmental factors, land use, life-history traits of the invaders, residence time, origin, and human usage interact to shape the spatial pattern of invasive alien plant species in South Africa. Relationships between the environmental factors and the extrinsic and intrinsic attributes of species were investigated using RLQ analysis, a multivariate method for relating a species-attribute table to an environmental table by way of a species presence/absence table. We then clustered species according to their position on the RLQ axes, and tested these groups for phylogenetic independence. The first three axes of the RLQ explained 99% of the variation and were strongly related to the species attributes. The clustering showed that, after accounting for environmental factors, the spatial pattern of IAS in South Africa was driven by human uses, life forms, and reproductive traits. The seven clusters of species strongly reflected geographical distribution, but also intrinsic species attributes and patterns of human use. Two of the clusters, centered on the genera Acacia and Opuntia, were phylogenetically non-independent. The remaining clusters comprised species of diverse taxonomic affinities, but sharing traits facilitating invasion in particular habitats. This information is useful for assessing the extent to which the potential spread of recent introductions can be predicted by considering the interaction of their biological attributes, region of origin, and human use. Key words: biological invasions; co-inertia analysis; environment; exotic species; human uses; invasive alien plants; life-history traits; plant invasions; RLQ analysis; South Africa.

African giant pouched rats (Cricetomys) are native to tropical Africa, where they range from Senegal and Gambia east across West Africa and the Congo Basin to the Indian Ocean coast of East Africa. Ecological niche models show that Cricetomys species differ in their invasive potential. Although neither of the presently recognized Cricetomys species appears to have genuinely broad distributional potential in North America, models predict that C. emini would have extremely restricted distributional potential, whereas C. gambianus would have a broader potential across the southeastern United States. Key words: African giant pouched rat, Cricetomys, ecological niche modeling, Genetic Algorithm for Rule-set Prediction, invasive species, rodent ecology

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. Key words: diversity: invasibility; multiple equilibria; niche differentiation; neutral theory; richness; species pool; vegetation.

The introduction of chronic, infectious diseases by colonizing populations (invasive or reintroduced) is a serious hazard in conservation biology, threatening the original host and other spillover species. Most research on spatial invasion of diseases has pertained to established host populations, either at steady state or fluctuating through time. Within a colonizing population, however, the spread of disease may be influenced by the expansion process of the population itself. Here we explore the simultaneous expansion of a colonizing population and a chronic, nonlethal disease introduced with it, describing basic patterns in homogeneous and structured landscapes and discussing implications for disease management. We describe expected outcomes of such introductions for three qualitatively distinct cases, depending on the relative velocities at which the population and epidemic expand. (1) If transmissibility is low the disease cannot be sustained, although it may first expand its range somewhat around the point of introduction. (2) If transmissibility is moderate but the wave-front velocity for the population, [v.sub.p], is higher than that for the disease, [v.sub.d], the disease wave front lags behind that of the population. (3) A highly transmissible disease, with [v.sub.d] > [v.sub.p], will invade sufficiently rapidly to track the spread of the host. To test these elementary theoretical predictions, we simulated disease outbreaks in a spatially structured host population occupying a real landscape. We used a spatially explicit, individual-based model of Persian fallow deer (Dama mesopotamica) reintroduced in northern Israel, considering a hypothetical introduction of bovine tuberculosis. Basic patterns of disease expansion in this realistic setting were similar to our conceptual predictions for homogeneous landscapes. Landscape heterogeneity, however, induced the establishment of population activity centers and disease foci within them, leading to jagged wave fronts and causing local variation in the relative velocities at which the population and epidemic expanded. Based on predictions from simple theory and simulations of managed outbreaks, we suggest that the relative velocities at which the population and epidemic expand have important implications for the impact of different management strategies. Recognizing which of our three general cases best describes a particular outbreak will aid in planning an efficient strategy to contain the disease. Key words: bovine tuberculosis; chronic disease; colonizing populations; Dama mesopotamica; infectious disease; invasive species; Israel; managed disease outbreak; reintroduction; Persian fallow deer; population expansion; spatially explicit model.

Abstract: The invasive potential of the nonnative Austrian black pine (Pinus nigra subsp. nigra Arn.) was analyzed in a 100-year-old Mediterranean mixed deciduous forest in the Massane Nature Reserve, eastern [...]

Abstract: While the spread of the exotic Lythrum salicaria L. across wetlands in North America is thought to Negatively affect the performance and diversity of native species, the actual impacts [...]

Abstract: Invasions by nonindigenous forest insects can have spectacular effects on the biodiversity, ecology, and economy of affected areas. This introduction explores several critical issues that are generally relevant to [...]

Abstract: Species-rich plant communities may be more resistant to invasive herbivores because of reduced host-plant accessibility and increased natural enemy diversity and abundance. We tested these hypotheses in Corsica, a [...]

Human activity is central in promoting invasion of arthropod generalist predators (AGP) and they have widespread and varied ecological impacts, many of which stem from displacement of similar native predators. Invasive AGP weakens or strengthens top-down control and they contribute to either invasional meltdown or to prevention of further damage from other invaders.

Historical introductions of species into new habitats can create rare opportunities to test evolutionary hypotheses, such as the role of natural selection in maintaining traits. This study examines two independent introductions of the African village weaverbird (Ploceus cucullatus) to islands where selection on egg appearance traits is expected to differ markedly from that of the source populations. The color and spotting of village weaver eggs in Africa are highly consistent within clutches, but highly variable between individuals. These two features may be an evolutionary response to brood parasitism. In Africa, weavers are parasitized by each other and by the diederik cuckoo (Chrysococcyx caprius), an egg mimic. African village weavers were introduced one century ago to Mauritius, and over two centuries ago to Hispaniola. Both islands are devoid of egg-mimicking brood parasites. In these two populations, between-individual variation and within-clutch consistency in egg appearance have both decreased, as has the incidence of spotting, relative to the source populations in Africa. These reductions are more pronounced on Hispaniola, the earlier introduction. Such changes support the hypothesis that egg appearance in the African village weaver has been maintained by natural selection as a counteradaptation to cuckoo brood parasitism. These results illustrate that the removal of an agent of selection can sometimes bring about rapid evolutionary consequences. brood parasitism | introduced species | Ploceidae | rapid evolution | trait loss

Hypothesized factors determining alien dominance were tested with structural equation modeling. Fire history is an important indirect factor as it affects both prefire stand structure and postfire alien seed banks.

Condition-specific competition, wherein competitive superiority varies with the abiotic environment, can determine species' distributions in a spatially heterogeneous environment. We investigated this phenomenon with two competing container-dwelling mosquitoes. We tested the hypothesis that habitat drying alters the outcome of interspecific competition, predicting that the competitive impact of Aedes albopictus on Aedes aegypti would be severe in wetter environments, but greatly reduced in drier environments. We tested these predictions in a laboratory experiment within cages, with aquatic larvae residing in water-filled cups within the cage, and adults emerging within the cage and ovipositing on the cups. We raised each species alone or with the competitor. Environmental treatments were fluctuating (water in cups evaporated to 50% of the original volume and then cups were refilled), and drying (water in cups evaporated completely and cups remained dry for two weeks before refilling). There was a significant interaction between treatment and species combination for adult populations of both species. Interspecific competition was highly asymmetrical. In the wetter fluctuating environment, interspecific competition had a large negative effect on A. aegypti, but in the drying environment, interspecific competition had a large negative effect on A. albopictus, and relatively little impact on A. aegypti. The main cause of the shift in competitive advantage appeared to be a greater increase in egg mortality for A. albopictus under dry conditions, compared to A. aegypti. Thus, mortality impinging on noncompeting life cycle stages can alter the population level impact of interspecific competition. The hypothesis that dry conditions shift competitive advantage away from A. albopictus is supported in this experiment. Key words: abiotic factors; Aedes aegypti; Aedes albopictus; environmental tolerances; invasive species.

The ability of weeds to proliferate into nonindigenous habitats has been attributed to escape from their native natural enemies, allowing reallocation of resources from chemical defense into growth and reproduction. Many invasive weeds, however, eventually encounter their native, coevolved enemies in areas of introduction. Examination of herbarium specimens of an invasive phototoxic European weed, Pastinaca sativa, through 152 years reveals phytochemical shifts coincident in time with the accidental introduction of a major herbivore, the parsnip webworm, Depressaria pastinacella. Plants collected before the introduction of webworms in North America and during the earliest stages of establishment (1850-1889) are lower in toxic furanocoumarins than all plants subsequently collected in North America and lower than European plant samples collected before 1889. Thus, introduction of a major specialist herbivore can increase noxiousness of a species in its area of introduction, illuminating a potential consequence of classical biocontrol programs involving insect herbivores and poisonous weeds. enemy release hypothesis | furanocoumarin | herbivory | invasive species | Lepidoptera

Introductions of non-indigenous species in ballast water are one of the greatest threats to freshwater and marine ecosystems worldwide. New approaches to reducing the release of organisms from ballast water are under consideration nationally and internationally. Unfortunately, the development of scientifically defensible policy for controlling introductions from ballast water has been retarded by the lack of relevant ecological theory and a shortage of information about the identity and numbers of organisms in ballast. Here, we present a novel quantitative approach to estimating the risk of species establishment by combining a model for population spread with known allometric correlations between body size and population growth rate for broad taxonomic categories. Our approach is applicable to sexually reproducing, planktonic taxonomic groups including ctenophores, cnidaria, arthropods, annelids, mollusks and (as an approximation) echinoderms and fishes. As expected, the allowable volume of ballast discharge depends strongly on the acceptable level of risk (which is a societal decision), the taxonomic group of concern and the characteristics of the receiving environment. For example, for a risk tolerance equivalent to establishment of one in one hundred introduced species, independent releases of untreated ballast water should not exceed around 50,000 metric tons. Because of differences in horizontal mixing in different environments, releases in harbors are more risky than releases in open water. These results provide quantitative guidelines that could immediately lower the risk of species invasion while other more permanent technological solutions are developed.

Despite recent advances in the study of invasive, nonnative species, we still understand very little about the variability of their population dynamics over space and time, and the processes that contribute to their demographic success. The goal of this study was to quantify the population dynamics of the invasive, subtropical shrub, Ardisia elliptica (Myrsinaceae), growing in a disturbed area of Everglades National Park, Florida, USA. We used bootstrap sampling and randomization tests to test hypotheses about spatial and temporal variation in population dynamics. Five populations spanning a range of habitats were studied for three years. Significant spatial and temporal variation in fruit production, relative growth, fates of vegetative individuals, and population growth rates were detected; however, spatial variation among populations had an overall stronger effect than temporal variation. Population growth rates were greater than 1 or not different from 1, implying that populations are on average increasing in size. Similar to most other shrubs, prospective analysis showed that stasis of large stage classes had the highest elasticities. However, a retrospective analysis indicated that it was variation in growth to larger stage classes that contributed the most to observed variation in population growth rates. Simulations suggest that historically higher population growth rates of A. elliptica were necessary to produce current population sizes in Everglades National Park. Key words: Ardisia elliptica; biological invasions; elasticity structure; invasive nonnative species; life table response experiment; matrix models; population dynamics; randomization tests.

Invasion biologists typically regard susceptibility of an invasive species to native predators as a fortuitous condition that increases biotic resistance to the invasion. The line of reasoning is that predation weakens the net impact of the invader and reduces its ability to displace native competitors. However, predation on invasives is a coupled interaction; every invader consumed also enhances the predator population. If these predators also consume native species, then the invader's indirect effect via predators (i.e., apparent competition) could be more harmful to natives than the effect of resource competition from the exotic. We apply general community ecology theory to determine the conditions under which the net effect of predation on the exotic species is to extirpate the native competitor. An approximation to these conditions provides a simple metric to estimate the threat of native species extinction due to apparent competition posed by an invader. Key words: apparent competition; biocontrol agent; biological control; biological invasions; biological resistance; competitive exclusion; exotic species; exploitative competition; nonindigenous species.

Sericea lespedeza [Lespedeza cuneata (Dumont) G. Don] is a non-native invasive plant that threatens the health of prairie and rangeland. A survey of rangeland in Cowley County, Kansas, was conducted [...]

Introduced American bullfrogs (Rana catesbeiana) come in contact with native amphibians on four continents and are well established in lowlands of western North America. To date, research on the effects of introduced bullfrogs on native frogs has focused on competition and predation, and is based largely on larval interactions. We present observations of interspecific amplexus between bullfrogs and two native ranid frogs (R. aurora and R. pretiosa) from six sites across the Pacific Northwest that imply that this interaction is more widespread than currently recognized. Our observations indicate that R. catesbeiana juveniles and subadults in this region are of appropriate size to elicit marked amplectic responses from males of both native species. Our literature review suggests that greater opportunity may exist for pairings between R. catesbeiana and native R. aurora or R. pretiosa than among syntopic native ranids in western North America. We hypothesize that interspecific amplexus with introduced R. catesbeiana could result in reproductive interference with negative demographic consequences in native ranid populations that have been reduced or altered by other stressors.

The relative importance of plasticity vs. adaptation for the spread of invasive species has rarely been studied. We examined this question in a clonal population of invasive freshwater snails (Potamopyrgus antipodarum) from the western United States by testing whether observed plasticity in life history traits conferred higher fitness across a range of temperatures. We raised isofemale lines from three populations from different climate regimes (high- and low-elevation rivers and an estuary) in a split-brood, common-garden design in three temperatures. We measured life history and growth traits and calculated population growth rate (as a measure of fitness) using an age-structured projection matrix model. We found a strong effect of temperature on all traits, but no evidence for divergence in the average level of traits among populations. Levels of genetic variation and significant reaction norm divergence for life history traits suggested some role for adaptation. Plasticity varied among traits and was lowest for size and reproductive traits compared to age-related traits and fitness. Plasticity in fitness was intermediate, suggesting that invasive populations are not general-purpose genotypes with respect to the range of temperatures studied. Thus, by considering plasticity in fitness and its component traits, we have shown that trait plasticity alone does not yield the same fitness across a relevant set of temperature conditions. Key words: adaptation; clonal reproduction; general-purpose genotypes; invasiveness; phenotypic plasticity; Potamopyrgus antipodarum; reaction norm.

Species become invasive if they (i) are introduced to a new range, (ii) establish themselves, and (iii) spread. To address the global problems caused by invasive species, several studies investigated steps ii and iii of this invasion process. However, only one previous study looked at step i and examined the proportion of species that have been introduced beyond their native range. We extend this research by investigating all three steps for all freshwater fish, mammals, and birds native to Europe or North America. A higher proportion of European species entered North America than vice versa. However, the introduction rate from Europe to North America peaked in the late 19th century, whereas it is still rising in the other direction. There is no clear difference in invasion success between the two directions, so neither the imperialism dogma (that Eurasian species are exceptionally successful invaders) is supported, nor is the contradictory hypothesis that North America offers more biotic resistance to invaders than Europe because of its less disturbed and richer biota. Our results do not support the tens rule either: that [approximately equal to] 10% of all introduced species establish themselves and that [approximately equal to] 10% of established species spread. We find a success of [approximately equal to] 50% at each step. In comparison, only [approximately equal to] 5% of native vertebrates were introduced in either direction. These figures show that, once a vertebrate is introduced, it has a high potential to become invasive. Thus, it is crucial to minimize the number of species introductions to effectively control invasive vertebrates. biotic resistance | ecological imperialism | invasive species | tens rule | time lags

The invasion of natural and anthropogenic habitats by alien plants is a global problem. To understand the factors that regulate the success of alien plant species we sampled alien plant biomass and species composition in 66 sites representing a wide variety of habitat types within Bruce Peninsula National Park, Ontario, Canada. We quantified the response of existing alien biomass to a suite of habitat variables including habitat productivity, standing biomass, species richness, soil nutrients and disturbance factors. Standing biomass in the sites varied from 0 to nearly 60 000 g/[m.sup.2] and annual productivity ranged from 0 to nearly 800 g*[m.sup.-2]-[yr.sup.-1]. The response of both absolute and relative alien biomass (dominance) to the independent variables was assessed using univariate plots and multivariate canonical correspondence analysis. The results show that total standing alien biomass and site disturbance history can predict alien dominance, but other factors including primary productivity, nutrient levels, and site species richness cannot. In the multivariate analysis, species composition of both aliens and natives was best explained by nutrient levels, disturbance type, biomass, and, to a lesser extent, productivity. The study illustrates the value of examining both absolute abundance and dominance of aliens across a broad array of habitat types, and in the context of a large suite of possible driving functions. Key words: alien plants; Bruce Peninsula National Park, Ontario, Canada; dominance; invasion biology; invasive species, success determinants; productivity; standing biomass.

Worldwide marine invaders, such as the brown alga Undaria pinnatifida, offer challenging models for unraveling the apparent paradox of sustainable settlement of exotic species over a large spectrum of environments. Two intergenic noncoding mitochondrial loci were found to be highly informative at the within-species level. Twenty-five haplotypes were found over the whole dataset (333 base pairs, 524 individuals, and 24 populations). The native range showed striking population genetic structure stemming from low diversity within and high differentiation among populations, a pattern not observed in the introduced range of this seaweed. Contrary to classical expectations of founding effects associated with accidental introduction of exotic species, most of the introduced populations showed high genetic diversity. At the regional scale, genetic diversity and sequence divergence showed contrasting patterns in the two main areas of introduction (Europe and Australasia), suggesting different processes of introduction in the two regions. Gene genealogy analyses point to aquaculture as a major vector of introduction and spread in Europe but implicate maritime traffic in promoting recurrent migration events from the native range to Australasia. The multiplicity of processes and genetic signatures associated with the successful invasion confirms that multiple facets of global change, e.g., aquaculture practices, alteration of habitats, and increased traffic, act in synergy at the worldwide level, facilitating successful pandemic introductions. coastal habitats | gene genealogy | introduced species | kelp | mitochondrial DNA

We used airborne imaging spectroscopy and photon transport modeling to determine how biological invasion altered the chemistry of forest canopies across a Hawaiian montane rain forest landscape. The nitrogen-fixing tree Myrica faya doubled canopy nitrogen concentrations and water content as it replaced native forest, whereas the understory herb Hedychium gardnerianum reduced nitrogen concentrations in the forest overstory and substantially increased aboveground water content. This remote sensing approach indicates the geographic extent, intensity, and biogeochemical impacts of two distinct invaders; its wider application could enhance the role of remote sensing in ecosystem analysis and management. Hawaii | remote sensing | tropical forest | invasive species | imaging spectroscopy

Invasive species have the potential to alter trade-offs leading to selection in the populations they invade. Here we quantify the demographic and selective effects of herbivory by native insects and the introduced floral feeder Rhinocyllus conicus on Platte thistle (Cirsium canescens), a sparse monocarpic thistle endemic to North America. Rhinocyllus first invaded the Platte thistle population in 1993. Since then, its numbers have increased exponentially, while the Platte thistle population size has decreased. Data from 11 years were analyzed to determine how demographic rates varied with plant size and damage by native insects and Rhinocyllus. Individual growth, survival, flowering probability, and seed set were all size dependent. Damage to vegetative structures did not influence demographic rates; damage to flower heads did because Platte thistle is seed limited. These analyses were used to parameterize a series of integral projection models (IPMs) that investigated the effects of floral herbivory on the population growth rate [lambda], equilibrium population size, and the evolutionary stable (ES) flowering strategy. The IPMs showed that native insects have significant impact on the equilibrium population size and X, but not the ES flowering strategy, because they use the flowers of different-sized plants indiscriminately. In contrast, Rhinocyllus has the potential to drive Platte thistle extinct. Rhinocyllus preferentially fed and oviposited on the flowers of larger plants and therefore selected for a reduction in flowering size. However, as the thistle population went into decline, this pattern reversed. Thus, selection imposed by an invader may be complex and will reflect behavioral interactions between herbivore and host, as well as demographic changes in the host population. Key words: biological control; Cirsium canescens; evolutionary stable strategy; insect herbivore; integral projection model; invasive species; life history evolution; monocarpic perennial; Platte thistle; Rhinocyllus conicus; selection.

Humans are altering the composition of biological communities through a variety of activities that increase rates of species invasions and species extinctions, at all scales, from local to global. These changes in components of the Earth's biodiversity cause concern for ethical and aesthetic reasons, but they also have a strong potential to alter ecosystem properties and the goods and services they provide to humanity. Ecological experiments, observations, and theoretical developments show that ecosystem properties depend greatly on biodiversity in terms of the functional characteristics of organisms present in the ecosystem and the distribution and abundance of those organisms over space and time. Species effects act in concert with the effects of climate, resource availability, and disturbance regimes in influencing ecosystem properties. Human activities can modify all of the above factors; here we focus on modification of these biotic controls. The scientific community has come to a broad consensus on many aspects of the relationship between biodiversity and ecosystem functioning, including many points relevant to management of ecosystems. Further progress will require integration of knowledge about biotic and abiotic controls on ecosystem properties, how ecological communities are structured, and the forces driving species extinctions and invasions. To strengthen links to policy and management, we also need to integrate our ecological knowledge with understanding of the social and economic constraints of potential management practices. Understanding this complexity, while taking strong steps to minimize current losses of species, is necessary for responsible management of Earth's ecosystems and the diverse biota they contain. Based on our review of the scientific literature, we are certain of the following conclusions: 1) Species' functional characteristics strongly influence ecosystem properties. Functional characteristics operate in a variety of contexts, including effects of dominant species, keystone species, ecological engineers, and interactions among species (e.g., competition, facilitation, mutualism, disease, and predation). Relative abundance alone is not always a good predictor of the ecosystem-level importance of a species, as even relatively rare species (e.g., a keystone predator) can strongly influence pathways of energy and material flows. 2) Alteration of biota in ecosystems via species invasions and extinctions caused by human activities has altered ecosystem goods and services in many well-documented cases. Many of these changes are difficult, expensive, or impossible to reverse or fix with technological solutions. 3) The effects of species loss of changes in composition, and the mechanisms by which the effects manifest themselves, can differ among ecosystem properties, ecosystem types, and pathways of potential community change. 4) Some ecosystem properties are initially insensitive to species loss because (a) ecosystems may have multiple species that carry out similar functional roles, (b) some species may contribute relatively little to ecosystem properties, or (c) properties may be primarily controlled by abiotic environmental conditions. 5) More species are needed to insure a stable supply of ecosystem goods and services as spatial and temporal variability increases, which typically occurs as longer time periods and larger areas are considered. We have high confidence in the following conclusions: 1) Certain combinations of species are complementary in their patterns of resource use and can increase average rates of productivity and nutrient retention. At the same time, environmental conditions can influence the importance of complementarity in structuring communities. Identification of which and how many species act in a complementary way in complex communities is just beginning. 2) Susceptibility to invasion by exotic species is strongly influenced by species composition and, under similar environmental conditions, generally decreases with increasing species richness. However, several other factors, such as propagule pressure, disturbance regime, and resource availability also strongly influence invasion success and often override effects of species richness in comparisons across different sites or ecosystems. 3) Having a range of species that respond differently to different environmental perturbations can stabilize ecosystem process rates in response to disturbances and variation in abiotic conditions. Using practices that maintain a diversity of organisms of different functional effect and functional response types will help preserve a range of management options. Uncertainties remain and further research is necessary in the following areas: 1) Further resolution of the relationships among taxonomic diversity, functional diversity, and community structure is important for identifying mechanisms of biodiversity effects. 2) Multiple trophic levels are common to ecosystems but have been understudied in biodiversity/ecosystem functioning research. The response of ecosystem properties to varying composition and diversity of consumer organisms is much more complex than responses seen in experiments that vary only the diversity of primary producers. 3) Theoretical work on stability has outpaced experimental work, especially field research. We need long-term experiments to be able to assess temporal stability, as well as experimental perturbations to assess response to and recovery from a variety of disturbances. Design and analysis of such experiments must account for several factors that covary with species diversity. 4) Because biodiversity both responds to and influences ecosystem properties, understanding the feedbacks involved is necessary to integrate results from experimental communities with patterns seen at broader scales. Likely patterns of extinction and invasion need to be linked to different drivers of global change, the forces that structure communities, and controls on ecosystem properties for the development of effective management and conservation strategies. 5) This paper focuses primarily on terrestrial systems, with some coverage of freshwater systems, because that is where most empirical and theoretical study has focused. While the fundamental principles described here should apply to marine systems, further study of that realm is necessary. Despite some uncertainties about the mechanisms and circumstances under which diversity influences ecosystem properties, incorporating diversity effects into policy and management is essential, especially in making decisions involving large temporal and spatial scales. Sacrificing those aspects of ecosystems that are difficult or impossible to reconstruct, such as diversity, simply because we are not yet certain about the extent and mechanisms by which they affect ecosystem properties, will restrict future management options even further. It is incumbent upon ecologists to communicate this need, and the values that can derive from such a perspective, to those charged with economic and policy decision-making. Key words: biodiversity; complementary resource use; ecosystem goods and services; ecosystem processes; ecosystem properties; functional characteristics; functional diversity; net primary production; sampling effect; species extinction; species invasions; species richness; stability.

Few invaded ecosystems are free from habitat loss and disturbance, leading to uncertainty whether dominant invasive species are driving community change or are passengers along for the environmental ride. The ''driver' model predicts that invaded communities are highly interactive, with subordinate native species being limited or excluded by competition from the exotic dominants. The 'passenger' model predicts that invaded communities are primarily structured by noninteractive factors (environmental change, dispersal limitation) that are less constraining on the exotics, which thus dominate. We tested these alternative hypotheses in an invaded, fragmented, and fire-suppressed oak savanna. We examined the impact of two invasive dominant perennial grasses on community structure using a reduction (mowing of aboveground biomass) and removal (weeding of above- and belowground biomass) experiment conducted at different seasons and soil depths. We examined the relative importance of competition vs. dispersal limitation with experimental seed additions. Competition by the dominants limits the abundance and reproduction of many native and exotic species based on their increased performance with removals and mowing. The treatments resulted in increased light availability and bare soil: soil moisture and N were unaffected. Although competition was limiting for some, 36 of 79 species did not respond to the treatments or declined in the absence of grass cover. Seed additions revealed that some subordinates are dispersal limited: competition alone was insufficient to explain their rarity even though it does exacerbate dispersal inefficiencies by lowering reproduction. While the net effects of the dominants were negative, their presence restricted woody plants, facilitated seedling survival with moderate disturbance (i.e., treatments applied in the fall), or was not the primary limiting factor for the occurrence of some species. Finally, the species most functionally distinct from the dominants (forbs, woody plants) responded most significantly to the treatments. This suggests that relative abundance is determined more by trade-offs relating to environmental conditions (long-term fire suppression) than to traits relating to resource capture (which should most impact functionally similar species). This points toward the passenger model as the underlying cause of exotic dominance, although their combined effects (suppressive and facilitative) on community structure are substantial. Key words: British Columbia, Canada, dominance; facilitation; functional redundancy; oak savanna; plant competition; plant invasion; recruitment limitation; removal experiment; repeated-measures ANOVA.

The arrival of invasive species can devastate natural ecosystems but the long-term effects of invasion are less clear. If native organisms can adapt to the presence of the invader, the severity of impact will decline with time. In Australia, invasive cane toads (Bufo marinus) are highly toxic to most snakes that attempt to eat them. Because snakes are gape-limited predators with strong negative allometry for head size, maximum relative prey mass (and thus, the probability of eating a toad large enough to be fatal) decreases with an increase in snake body size. Thus, the arrival of toads should exert selection on snake morphology, favoring an increase in mean body size and a decrease in relative head size. We tested these predictions with data from specimens of four species of Australian snakes, collected over >80 years. Geographic information system layers provided data on the duration of toad exposure for each snake population, as well as environmental variables (latitude, precipitation, and temperature). As predicted, two toad-vulnerable species (Pseudechis porphyriacus and Dendrelaphis punctulatus) showed a steady reduction in gape size and a steady increase in body length with time since exposure to toads. In contrast, two species at low risk from toads (Hemiaspis signata and Tropidonophis mairii) showed no consistent change in these morphological traits as a function of the duration of toad exposure. These results provide strong evidence of adaptive changes in native predators as a result of the invasion of toxic prey. adaptation | conservation | contemporary evolution | coevolution

Invasive species impacts on native species and communities have been widely recognised for decades and may involve important economical losses. In this study, we examined two marine muricid gastropods: an invader, Ocinebrellus inornatus, and a resident, Ocenebra erinacea. Both species co-occur on French Atlantic coasts and probably have economical impacts on oyster farming areas of the Charente-Maritime region of France. In previous studies, we investigated the introduction source and the expansion patterns of O. inornatus, using molecular markers. However, these studies are not sufficient to fully understand the expansion dynamics of the exotics. The present framework is devoted to comparing life-history traits between the introduced and resident species. Our results first showed that O. inornatus has more favourable traits, such as a better mean growth rate and a higher reproductive effort, in comparison with O. erinacea. These traits may explain the invader establishment and, partly, its spread along the coast of France. Secondly, the resident species drilled a higher rate of oysters than the invader. Finally, the establishment of O. inornatus in France does not seem to be at the expense of O. erinacea because: (1) resources are not limiting in oyster farming areas and (2) there does not appear to be competition by interference between the species.

A widespread but controversial idea in ecology states that the number of invaders of a species assemblage depends on its species richness. Both negative and positive relationships have been reported. We examined whether a simple neutral model where assemblages are generated by drawing individuals from two pools of identical species (native and alien) can predict this relationship. We performed a meta-analysis of published data on this relationship. The neutral model showed that in communities with low and fixed numbers of individuals, the relationship between the number of aliens and native species is strong and negative. This becomes weaker as the proportion of species from both pools already present in the community increases. The relationship between alien and native species richness becomes positive when the number of individuals is allowed to vary, because the richness depends on number of individuals and/or area sampled. The meta-analysis showed that scale-dependence of the relationship between alien and native species richness is universal and compatible with the neutral model. Unless more evidence is available to refute it, the relationship between native and alien species richness should be considered a result of a neutral processes due to constraints on the number of individuals in the community. Key words: alien species; disturbance rate; meta-analysis; neutral process; scale; species--area relationship; species richness; transient dynamics.

Surveys of natural habitats often indicate that native and exotic species richness are positively correlated at large scales and negatively correlated at small scales. The small-scale relationship is often presented as evidence that native richness can repel invasion or conversely that exotic invasions can reduce native diversity. The larger scale pattern has been interpreted as evidence of the importance of facilitation, variable habitat quality, propagule supply, and other ecological phenomena. However, these explanations fail to consider expected native-exotic richness relationships under a null model assuming no species interactions. We show via simulation that the null expectation for a randomly assembled community is a negative relationship between native and exotic species richness at the smallest scales and, when plots vary in total richness, a positive relationship at larger scales. We outline a procedure to compare observational data to this null expectation using a permutation test of labels of species origin (native or exotic). Our use of this technique on plant community data indicates that patterns of native and exotic richness are remarkably similar to those generated by a null model. We argue that a null model approach is needed to evaluate whether observed native-exotic richness data deviate significantly from expected patterns generated by sampling and statistical artifacts. Key words: assembly rules; ecosystem functioning; invasion biology; invasion frequency; permutation test; plant diversity; riparian plant communities; scale dependence; strandline plant communities.

The damage caused by biological invasions has traditionally been thought to result from alien species taking advantage of ecological differences between the native and introduced ranges. In contrast, the role of evolutionary forces has received relatively little attention. Our results show that evolutionary change in Silene latifolia, a North American weed that was introduced from Europe about 200 years ago, can help explain the plant's successful North American invasion. By growing plants from seed collected in 40 populations from Europe and North America under common garden greenhouse and field conditions, we found significant genetic differences in life history, reproductive, and defensive characters. In general, morphological traits and competitive ability remained unchanged, while North American plants germinated earlier, grew faster, produced more flowers, had greater survival, and invested less into defensive traits (trichomes, fruit capsule) than their European conspecifics. We suggest that as S. latifolia escaped a suite of specialist enemies, natural selection favored individuals that invest more in growth and reproduction and less in defense. Key words: alien plant: biological invasion: common garden; EICA hypothesis; escape from enemies; Silene latifolia; trichome.

Chinese tallow (Sapium sebiferum) is an invasive tree in the southeastern United States that has been a management concern in Florida. We used observational and experimental approaches to explore factors influencing Chinese tallow invasion around the perimeter of Lake Jackson, Leon County, Florida. Twenty-one transect sites were randomly chosen, 12 with, and 9 without Chinese tallow. We used abundances of co-occurring species and abiotic environmental data to study factors associated with Chinese tallow abundance among and within transects. Canonical correspondence analysis indicated that soil moisture availability and other soil characteristics correlated with Chinese tallow invasion, but that Chinese tallow was not associated with other species distributions. A seeding experiment demonstrated that higher elevations with lower soil moisture have higher germination and seedling survival. This experiment predicts specific elevations around the lake that are most vulnerable to Chinese tallow invasion. Presence or absence of Chinese tallow among transects was not a significant predictor of germination success, suggesting that not all sites suitable for invasion have been invaded. In addition, controlled burning decreased germination probability and may therefore be useful for invasion control.

With field, laboratory, and modeling approaches, we examined the interplay among habitat structure, intraguild predation (IGP), and parasitism in an ongoing species invasion. Native Gammarus duebeni celticus (Crustacea: Amphipoda) are often, but not always, replaced by the invader Gammarus pulex through differential IGP. The muscle-wasting microsporidian parasite Pleistophora mulleri infects the native but not the invader. We found a highly variable prevalence of P. mulleri in uninvaded rivers, with 0-91% of hosts parasitized per sample. In addition, unparasitized natives dominated fast-flowing riffle patches of river, whereas parasitized individuals dominated slower-flowing, pooled patches. We examined the survivorship of invader and native in single and mixed-species microcosms with high, intermediate, and zero parasite prevalence. G. pulex survivorship was high in all treatments, whereas G. duebeni subsp. celticus survivorship was significantly lower in the presence of the invader. Further, parasitized G. duebeni subsp. celticus experienced near-total elimination. Models of the species replacement process implied that parasite-enhanced IGP would make invasion by G. pulex more likely, regardless of habitat and parasite spatial structure. However, where heterogeneity in parasite prevalence creates a landscape of patches with different susceptibilities to invasion, G. pulex may succeed in cases where invasion would not be possible if patches were equivalent. The different responses of parasitized and unparasitized G. duebeni subsp. celticus to environmental heterogeneity potentially link landscape patterns to the success or failure of the invasion process.

The demography of the fresh water fish species vendace during its development over a decade, from first occurrence to an established population, in the Pasvik watercourse is documented. The invasive ability of vendace appears to result from a combination of high fecundity and strong competitive ability in a disturbed environment.

The discovery record of introduced species reflects a combination of the introduction process and the discovery process. For this reason, the discovery record does not provide a direct proxy for the record of introductions. We describe a general method for estimating the rate of introductions from the discovery record. The method is based on a statistical model of the discovery record that includes both the introduction and discovery processes. The method is illustrated using the discovery record of introduced species in the San Francisco estuary (California, USA). The estimated mean rate of introductions increases from 0.3 introductions in 1850 to 2.3 introductions in 1995. Key words: maximum likelihood estimation; Poisson distribution; species introductions, estimating rate.

The conceptualization of alien invasive species conflates two axes of variability that have become unhelpfully blurred. The nativeness/alienness axis refers to the presumed belonging of a species in ecological or social space. Invasiveness refers to the behavior of the species in question, particularly in relation to other species. The overlay of nation introduces further variability. Teasing these axes apart is important for more effective environmental management. We examine these concepts using two influential forms of ecological knowledge: the biogeographical and ecological literature and the vernacular experiences of suburban backyarders. Three case studies, the invasive native Pittosporum undulatum and two invasive exotics, Lantana camara and Cinnamomum camphora, illustrate the complex and contingent nature of human interactions with such species and the potential for human interactions to increase and/or reduce the propagation of plant species. Keywords: Australia, backyards, Cinnamomum camphora, Lantana camara, Pittosporum undulatum, weeds., Species have been moving around the globe throughout the ecological history of the Earth. These movements have accelerated in the last few hundred years as a consequence of European colonialism, [...]

Environmentalists and ecologists typically regard invasive species as ecologically detrimental and undesirable. Although the ecological impacts of invasive species are well documented, less attention has been devoted to the sociocultural contexts guiding responses to species invasion. In this study the roles of invasive species are reconsidered through three prominent green spaces in Toronto, Canada: the Don Valley Brick Works, High Park, and the Leslie Street Spit. The case studies challenge popular negative assumptions about invasive species and suggest that they can serve important functions both for local ecosystems and for human communities. The case studies also provide lessons on the tension between and within different environmental imaginaries informing invasive species management. Invasives are often compatible with recreational interests, whereas naturalization efforts are ecologically sensitive and costly. Invasives can help restore human-made wastelands, and naturalization efforts often benefit wealthy rather than poor neighborhoods. Keywords: Canada, invasive species, public spaces, Toronto., In the recently published Nature by Design, Eric Higgs wonders about future views of weedy and exotic species. He asks, 'Will we be so overwhelmed by invasions on native flora [...]

Garlic mustard (Alliaria petiolata, Brassicaceae) is an invasive, nonindigenous species currently invading the understory of North American woodlands where it is a serious threat to the native flora. Part of this success might be due to allelopathic interference by garlic mustard. Two congeneric species, the European Geum urbanum and the North American Geum laciniatum, were tested for allelopathic inhibition of germination by garlic mustard. Seeds were germinated either on substrate contaminated by garlic mustard or on substrate with contamination neutralized by activated carbon. Allelopathic effects of native European and invasive North American garlic mustard populations were also compared. Activated carbon increased germination by 14%, indicating that garlic mustard contaminated the substrate through root exudates. Activated carbon in turn counteracted this effect. The two test species differed in their sensitivity to allelopathic interference. North American G. laciniatum had a much stronger increase in germination when activated carbon was added to the substrate, independent of the origin of garlic mustard. In contrast, the European G. urbanum germinated better in substrate precultivated with North American garlic mustard, whereas activated carbon increased its germination only in substrate precultivated with European garlic mustard. Key words: allelopathy; biological invasion; competition; garlic mustard; Geum laciniatum; Geum urbanum.

The dynamics of invasive species may depend on their abilities to compete for resources and exploit disturbances relative to the abilities of native species. We test this hypothesis and explore its implications for the restoration of native ecosystems in one of the most dramatic ecological invasions worldwide, the replacement of native perennial grasses by exotic annual grasses and forbs in 9.2 million hectares of California grasslands. The long-term persistence of these exotic annuals has been thought to imply that the exotics are superior competitors. However, seed-addition experiments in a southern California grassland revealed that native perennial species, which had lower requirements for deep soil water, soil nitrate, and light, were strong competitors, and they markedly depressed the abundance and fecundity of exotic annuals after overcoming recruitment limitations. Native species reinvaded exotic grasslands across experimentally imposed nitrogen, water, and disturbance gradients. Thus, exotic annuals are not superior competitors but rather may dominate because of prior disturbance and the low dispersal abilities and extreme current rarity of native perennials. If our results prove to be general, it may be feasible to restore native California grassland flora to at least parts of its former range.

Invasive species that penetrate habitat boundaries are likely to experience strong selection and rapid evolution. This study documents evolutionary shifts in tolerance and performance following the invasion of fresh water by the predominantly estuarine and salt marsh copepod Eurytemora affinis. Commongarden experiments were performed on freshwater-invading (Lake Michigan) and ancestral saline (St. Lawrence marsh) populations to measure shills in adult survival (at 0, 5, and 25 PSU), and survival during development and development time (both using full-sib clutches split across 0, 5, 15, and 25 PSU). Results showed clear evidence of heritable shifts in tolerance and performance associated with freshwater invasions. The freshwater population exhibited a gain in low-salinity tolerance and a reduction in high-salinity tolerance relative to the saline population, suggesting tradeoffs. These tradeoffs were supported by negative genetic correlations between survival at fresh (0 PSU) versus higher salinities. Mortality in response to salinity occurred primarily before metamorphosis, suggesting that selection in response to salinity had acted primarily on the early life-history stages. The freshwater population exhibited curious patterns of life-history evolution across salinities, relative to the saline population, of retarded development to metamorphosis but accelerated development from metamorphosis to adulthood. This pattern might reflect tradeoffs between development rate and survival in fresh water at the early life-history stages, but some other selective force acting on later life-history stages. Significant effects of clutch (genotype) and clutch-by-salinity interaction (G x E) on survival and development time in both populations indicated ample genetic variation as substrate for natural selection. Variation for high-salinity tolerance was present in the freshwater population despite negative genetic correlations between high- and low-salinity tolerance. Results implicate the importance of natural selection and document the evolution of reaction norms during freshwater invasions.

Populus deltoides subsp, wislizinii (Salicaceae), a cottonwood native to the Middle Rio Grande of New Mexico, must potentially compete against exotic Tamarix ramosissima (Tamaricaceae) during establishment after flooding. We investigated competitive interactions between seedlings of Tamarix and Populus in two substrates representing field textures and declining (i.e., draw-down) or stagnant water tables. The experiment was performed using a full-additive series design and interpreted with response surface models for each species. As reflected in both aboveground mass and height, Populus suppressed aboveground growth of Tamarix across all treatments, whereas competitive effects of Tamarix against Populus could only be seen at low Populus densities. Clay substrates with draw-down stimulated the greatest growth and created the most intense competitive environment for both species. Tamarix was competitively suppressed in every substrate tested, with the weakest response in sand with no draw-down, where growth of Populus was poorest. These results suggest that stream flow management that promotes Populus establishment could also aid in controlling Tamarix invasion across a range of substrates. Key words: competition; flooding disturbance; invasion; Populus deltoides subsp, wislizinii; riparian management; Salicaceae; Tamaricaceae; Tamarix ramosissima.

Invasions of nonindigenous species have caused ecological devastation to natural communities worldwide, yet the biological bases for invasiveness remain poorly understood. Our studies of invasive watermilfoil (Myriophyllum) populations revealed widespread polymorphisms in biparentally inherited nuclear ribosomal DNA sequences, which were not detected in populations of native North American species. Subclones of the polymorphic regions revealed the occurrence of distinct sequences matching those acquired from both nonindigenous and native North American species. Molecular data demonstrate clearly that invasive watermilfoil populations in North America have resulted from hybridization between nonindigenous and native species. These observations suggest that invasiveness in these aggressive aquatic weeds may be linked to heterosis maintained by vegetative propagation.