Interactions between an invasive and a native bryozoan (<Emphasis Type="Italic">Membranipora membranacea</Emphasis> and <Emphasis Type="Italic">Electra pilosa</Emphasis>) species on kelp and <Emphasis Type="Italic">Fucus</Emphasis> substrates in Nova Scotia,Canada

Most research on biological invasions to date has focused on the population dynamics of very successful and disruptive introduced species; however, additional knowledge of the biology of the native species is essential for understanding interactions between the two and may reveal factors that limit invasion success. The invasive bryozoan Membranipora membranacea interacts with native Electra pilosa on two substrates in northwest Atlantic subtidal habitats: highly dynamic and fast-growing kelps; and smaller, more stable, and slow-growing fucoid algae. We quantified the relative abundance and evaluated encounter outcomes in different seasons of these two bryozoans on kelp and Fucus at four sites in Nova Scotia. We also examined the effects of substrate (kelp, Fucus), temperature (7, 10, 13°C), and food (limited, unlimited) on growth rates of E. pilosa in laboratory experiments and using field manipulations. We compared our findings on factors affecting the growth of E. pilosa to those on M. membranacea obtained in similar and thus directly comparable experiments from a previous study. The proportional abundance of M. membranacea was greater than that of E. pilosa on kelps, but the opposite was observed on Fucus. Competitive standoffs between the two bryozoans were more frequent than expected, with no differences recorded between substrates; most encounters were won by M. membranacea. Growth of E. pilosa was faster on Fucus than kelp, decreased with increasing temperature only on Fucus, and was not affected by food. Growth rate of E. pilosa in all treatments was slower than that previously measured for M. membranacea. Faster growth and strong overgrowth abilities likely interact on kelps to ensure success of the invasive bryozoan. Success can be limited by low space availability, which in turn restricts growth rate, and consequently, colony size, such as on fucoid substrates. The incorporation of alternative contexts into invasion research can reveal factors involved in the resilience of native communities.     

Geographic variation in resource dominance-discovery in Brazilian ant communities

A predictive framework for the ecology of species invasions requires that we learn what limits successful invaders in their native range. The red imported fire ant (Solenopsis invicta) is invasive in the United States, Puerto Rico, Australia, New Zealand, and China. Solenopsis invicta appears to be a superior competitor in its introduced range, where it can cause the local extirpation of native species, but little is known about its competitive ability in its native range in South America. Here we examine the competitive ability of S. invicta for food resources in three widely separated Brazilian ant communities. Each of these communities contains 20-40 ant species, 8-10 of which were common and frequently interacted with S. invicta. S. invicta at all three sites was attacked by several species-specific phorid parasitoids, and at one site, two other species were attacked by their own specialized parasitoids. We examined interactions in these local communities for evidence that trade-offs among ant species between resource dominance and resource discovery, and between resource dominance and parasitoid vulnerability facilitate local coexistence. The trade-off between resource dominance and resource discovery was strong and significant only at Santa Genebra, where parasitoids had no effect on the outcome of confrontations at resources. At Bonito, parasitoids significantly reduced the ability of S. invicta, which was the top-ranked behavioral dominant, from defending and usurping food resources from subordinate species. In the Pantanal, S. invicta ranked behind three other ant species in a linear hierarchy of behavioral dominance, and lost the majority of its interactions with a fourth more subordinate species, Paratrechina fulva, another invasive species. Parasitoids of S. invicta were uncommon in the Pantanal, and did not affect its low position in the hierarchy relative to the other two sites. Parasitoids, however, did affect the ability of Linepithema angulatum, the top-ranked behavioral dominant in this community, from defending and usurping resources from behavioral subordinates. These results indicate that both interspecific competition and trait-mediated indirect effects of phorid parasitoids affect the ecological success of the red imported fire ant in its native range, but that the relative importance of these factors varies geographically.     

紫茎泽兰入侵过程中生物群落的交互作用

生物入侵在全球范围内影响了生物群落的结构与功能,打破了群落内物种共存的生态格局,继而反馈性影响全球环境。该文就外来杂草紫茎泽兰入侵对生物群落之间交互作用的影响进行了分析。1）紫茎泽兰通过竞争排斥降低了土著植物群落的多样性,造成依赖于土著植物的节肢动物群落减少或丧失适宜的栖息环境。2）打破了土著植物与节肢动物之间相互依存的状态,并通过单优群落优势和强烈化感作用制约天敌昆虫的自然控制作用。3）通过改变地表生境和枯落物种类影响土壤动物群落。4）引起土壤微生物群落组成和功能的变化,改变土壤中可利用资源的形式和数量,影响并重塑了生物种间互作模式,并动态反馈于地面植物群落新格局的形成。分析指出：1）入侵过程中群落之间的交互作用通过多层次生态过程对群落结构与功能的生态改变发挥影响。2）入侵对生物群落的改变所产生的生态驱动反馈性作用于群落互作模式的重塑、群落和生态系统新格局的重建。同时,指出了生物入侵对群落影响的复杂性以及后续研究的方向。     

Examining the colonization process of exotic species varying in competitive abilities using a cellular automaton model

Current threats of invasive species have significant implications for ecological systems. Given their potential impacts, invasive species have been the subject of extensive empirical and theoretical studies. However, these studies have tended to focus on species that produce highly visible ecological and economic impacts. In our study, we take a step back from focusing on these high-impact invasive species, and examine the general colonization (invasion) process of exotic species that have various “competitive abilities” against the native species. Using a two-species cellular automaton model, we demonstrate that: (1) a threshold level of competitive ability is required for the exotic species to successfully establish in a new landscape and (2) an exotic species with superior competitive ability does not necessarily become dominant in a landscape (alternatively, a species that has inferior competitive ability may successfully colonize a new system). Our findings have significant implications for the study of species invasions and also provide clues to how species assemble in ecological communities.     

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

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

异质生境空心莲子草植物群落组成及物种多样性研究

研究草海湿地空心莲子草（Alternanthera philoxeroides）入侵对异质生境植物群落生活型、物种组成、重要值及物种多样性的影响,结果表明,①异质生境中群落生活型组成比例有差异,水生生境地面芽植物（H）∶隐芽植物（Cr）∶一年生植物（Th）为56∶38∶6;湿生生境为43∶36∶21。②异质生境中物种组成差异明显,水生生境试验样方包含9科13属16种,对照样方（无空心莲子草入侵）包含10科19属23种;湿生生境试验样方包含8科14属14种,对照样方有10科19属22种。空心莲子草入侵不同生境后群落内物种重要值变化及差异均较大,部分乡土物种在试验样方中减少甚至消失,水生生境和湿生生境群落中分别有30.44%、36.36%物种消失。③异质生境中空心莲子草重要值与物种多样性指数、生态优势度指数、均匀度指数均呈二项式函数关系;两种生境的多样性指数和均匀度指数均表现为随空心莲子草重要值先增加后减少,生态优势度指数均为先减少后增加,空心莲子草入侵对乡土植物群落物种多样性产生正向影响和负向影响,在群落中其重要值达25.8（水生生境）、18.9（湿生生境）时,是乡土植物多样性下降的临界点。     

Alternative strategies in avian scavengers: how subordinate species foil the despotic distribution

Trade-offs in species’ traits can mediate competition and enable coexistence. A key challenge in ecology is understanding the role of species’ trade-offs in maintaining diversity, and evolutionary trade-offs between the abilities of competing species are best understood by considering how competitive advantages change along an environmental gradient. Previous studies of such trade-offs are generally limited to two-species systems and a single trade-off. In this study, I consider the effect of trade-offs in search efficiency and competitive abilities on habitat use patterns among a diverse avian scavenger guild. I hypothesize that species’ dominance status and search efficiency will both be correlated with patch quality. Using counts of searching birds in areas that vary in habitat quality in terms of both wildlife and human settlement density and observations at experimental carcasses, I assess the competitive ability, search efficiency, and habitat use of seven avian scavenger species in Masai Mara National Reserve, Kenya. Findings support the hypothesis with Bateleurs, a species with high search efficiency, and Ruppell’s, Lappet-faced, and White-backed vultures, species with high individual or social dominance, preferentially exploiting habitats of high quality, while Tawny eagles and Hooded vultures, species with low search efficiency and competitive ability, prefer habitats of low quality. This paper demonstrates the importance of considering multiple strategies for assessing the effect of competition on habitat use within complex communities.     

An experimental study of competition between fire ants and Argentine ants in their native range

An understanding of why introduced species achieve ecological success in novel environments often requires information about the factors that limit the abundance of these taxa in their native ranges. Although numerous recent studies have evaluated the importance of natural enemies in this context, relatively few have examined how ecological success may result from differences in the magnitude of interference competition between communities in the native and introduced ranges of nonnative species. Here we examine how native-range competitive environments may relate to invasion success for two important invasive species, the red imported fire ant (Solenopsis invicta) and the Argentine ant (Linepithema humile), in a region of native-range sympatry. At two study sites in northern Argentina, we used stable-isotope analysis, a variety of observational approaches, and two different reciprocal removal experiments to test (1) whether S. invicta competes asymmetrically with L. humile (as suggested by the 20th century pattern of replacement in the southeastern United States) and (2) the extent to which these two species achieve behavioral and numerical dominance. Stable-isotope analysis and activity surveys indicated that S. invicta and L. humile are both omnivores and forage during broadly overlapping portions of the diel cycle. Short-term removal experiments at baits revealed no competitive asymmetry between S. invicta and L. humile. Longer-term colony removal experiments illustrated that S. invicta and L. humile experience an approximately equal competitive release upon removal of the other. Our results indicate that neither S. invicta nor L. humile achieves the same degree of behavioral or ecological dominance where they co-occur in native populations as they do in areas where either is common in their introduced range. These results strongly suggest that interspecific competition is an important limiting factor for both S. invicta and L. humile in South America.     

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

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

Forecasting the Spread of Invasive Rainbow Smelt in the Laurentian Great Lakes Region of North America

Abstract:  Rainbow smelt ( Osmerus mordax ) have invaded many North American lakes, often resulting in the extirpation of native fish populations. Yet, their invasion is incipient and provides the rationale for identifying ecosystems likely to be invaded and where management and prevention efforts should be focused. To predict smelt presence and absence, we constructed a classification-tree model based on habitat data from 354 lakes in the native range for smelt in southern Maine. Maximum lake depth, lake area, and Secchi depth (surrogate measure of lake productivity) were the most important predictors. We then used our model to identify lakes vulnerable to invasion in three regions outside the smelt's native range: northern Maine (52 of 244 lakes in the non-native range), Ontario (4447 of 8110), and Wisconsin (553 of 5164). We further identified a subset of lakes with a strong potential for impact (potential–impact lakes) based on the presence of fish species that are affected by rainbow smelt. Ninety-four percent of vulnerable lakes in the non-native range in Maine are also potential–impact lakes, as are 94% and 58% of Ontario and Wisconsin's vulnerable lakes, respectively. Our modeling approach can be applied to other invaders and regions to identify invasion-prone ecosystems, thus aiding in the management of invasive species and the efficient allocation of invasive species mitigation and prevention resources.     

Twenty years of salt marsh succession on a Dutch coastal barrier island

After a formerly grazed salt marsh was released from cattle grazing, changes in plant species composition were monitored for 20 yr, using vegetation maps and permanent plots. Three areas, differing in age and nutrient status were compared. The number of plant species and plant communities decreased.Elymus athericus (Elytrigia pungens) became dominant in most plant communities after 5–20 yr on the oldest and most productive salt marsh. In younger areas it took more time forE. athericus to become dominant. At least 7 cm of clay seemed to be a prerequisite for this plant species to increase in dominance. The results from monitoring over decades are discussed in view of the knowledge on succession over centuries as derived from a chronosequence.     

Consequences of behavioral vs. numerical dominance on foraging activity of desert seed-eating ants

Dominance relationships among species play a major role in the structure of animal communities. Yet, dominant species with different trade-offs in resource exploitation and monopolization could affect community structure in variable ways. In ants, dominant species could be classified into either behavioral dominants that exhibit territorial aggression or numerical dominants that exhibit high biomass or frequency of occurrence. While each class of dominance has generally been found to negatively affect the foraging activity of species in ant communities, the concurrent effect of both classes of species has never been tested. Here, we examined the effects of two behaviorally dominant species, Crematogaster inermis and Monomorium salomonis, and a numerically dominant species, Messor arenarius, on the foraging behavior of seed-eating species in a desert ant assemblage. In a 1-year study, the foraging activity of the ant species was assessed using seed baits, which were sampled during night and day. While the numerically dominant species exhibited high foraging efficiency and negatively affected the ability of other seed-eating species to obtain seeds, significantly more seeds remained at baits that were occupied the previous night by each of the two behaviorally dominant species, possibly due to aggressive exclusion of M. arenarius foragers from the baits. This exclusion also facilitated greater foraging activity of the seed-eating species. Our results demonstrate how these two types of dominance could differently affect the foraging activity of ant species in 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.     

Response of the grasshopper <Emphasis Type="Italic">Myrmeleotettix maculatus</Emphasis> (Orthoptera: Acrididae) to invasion by the exotic moss <Emphasis Type="Italic">Campylopus introflexus</Emphasis> in acidic coastal dunes

In Europe, acidic coastal dunes are threatened by the invasion of the exotic moss Campylopus introflexus. While the effect of the moss encroachment on the vegetation is well analysed, knowledge of possible impact on arthropods is lacking. Thus, an experiment was conducted in acidic coastal dunes on the Baltic island of Hiddensee, Germany. Myrmeleotettix maculatus, a common Orthoptera species of open and dry habitats, was sampled by pitfall trapping in eleven plots invaded by C. introflexus and in eleven native, non-invaded plots rich in lichens. Overall, 826 individuals of M. maculatus were captured (266 nymphs, 560 adults). Mean number of adults was significantly higher in native plots. This maybe explained by a higher proportion of grasses (food supply), a higher availability of shelter (from predators, weather), or more favourable microclimate conditions in native plots and a higher mortality rate in invaded plots. However, mean number of both young and old nymphs did not differ significantly between both types. This could imply that invaded areas at least serve as favourable oviposition sites and larval habitats. The observed negative effect of the moss invasion on M. maculatus remains to be studied on other arthropods.     

Evolutionary cause of the vulnerability of insular communities

Communities on oceanic islands are considered to be vulnerable to biological invasion. However, because the detailed structures of such communities have not yet been revealed, the relationship between their vulnerability and structure is not clear. Because such communities evolved without biological invasion, they are expected to have structures different from those of mainland communities, and this difference is expected to affect their vulnerability to invasion. I conducted computer simulations based on a food web model and investigated the difference in structure between mainland and insular model communities, the former of which evolved with frequent invasion and the latter without invasion. In addition, by conducting computer simulations of invasion of these model communities, I investigated the relationship between community structure and vulnerability to biological invasion. The insular model community evolved to have an unstable structure, in that a small number of plant species supported a large number of animal species, and each species in the community had a small biomass. When a plant species invaded and disturbed the base of the insular model community, many animal species relying on the plants easily became extinct. In addition, when a carnivorous species invaded, animal species with small biomass tended to become extinct. Community collapses caused by biological invasion occurred more frequently in the insular model community than in the mainland model community. These results indicated that those communities that evolved without invasion were vulnerable to invasion. The available data on real insular communities suggest that some have reached the endangered state predicted by this model.     

Using vertebrate environmental DNA from seawater in biomonitoring of marine habitats

Conservation and management of marine biodiversity depends on biomonitoring of marine habitats, but current approaches are resource-intensive and require different approaches for different organisms. Environmental DNA (eDNA) extracted from water samples is an efficient and versatile approach to detecting aquatic animals. In the ocean, eDNA composition reflects local fauna at fine spatial scales, but little is known about the effectiveness of eDNA-based monitoring of marine communities at larger scales. We investigated the potential of eDNA to characterize and distinguish marine communities at large spatial scales by comparing vertebrate species composition among marine habitats in Qatar, the Arabian Gulf (also known as the Persian Gulf), based on eDNA metabarcoding of seawater samples. We conducted species accumulation analyses to estimate how much of the vertebrate diversity we detected. We obtained eDNA sequences from a diverse assemblage of marine vertebrates, spanning 191 taxa in 73 families. These included rare and endangered species and covered 36% of the bony fish genera previously recorded in the Gulf. Sites of similar habitat type were also similar in eDNA composition. The species accumulation analyses showed that the number of sample replicates was insufficient for some sampling sites but suggested that a few hundred eDNA samples could potentially capture >90% of the marine vertebrate diversity in the study area. Our results confirm that seawater samples contain habitat-characteristic molecular signatures and that eDNA monitoring can efficiently cover vertebrate diversity at scales relevant to national and regional conservation and management.     

Ideal dominance distributions: a test using red-winged blackbirds (Agelaius phoeniceus)

Summary The ideal dominance distribution model predicts that competition between individuals of a species for territories will result in socially dominant individuals acquiring territories in higher quality habitat than their subordinates. Although the dispersion and relative reproductive success of male red-winged blackbirds (Agelaius phoeniceus) across habitats in eastern Ontario appears to conform to the ideal dominance distribution model, data from a study of three captive groups comprised of males from both high (marsh) and low (upland field) quality habitats failed to support the prediction that males from marsh habitat are dominant to those from upland habitat. Contrary to the prediction males from uplands were generally dominant to males from marshes. We found a significant positive correlation between dominance and both increased epaulet size and increased body size. Controlling for these positive effects, upland males remained generally dominant to marsh males. Measurements of independent samples of males from both habitats indicated that the overall distribution of males does not conform to ideal dominance. We suggest that the strong between-year territory fidelity shown by male red-winged blackbirds and chance events when they initially acquire territories may contribute to this lack of conformity.     

Aquatic plant community invasibility and scale-dependent patterns in native and invasive species richness

Invasive species richness often is negatively correlated with native species richness at the small spatial scale of sampling plots, but positively correlated in larger areas. The pattern at small scales has been interpreted as evidence that native plants can competitively exclude invasive species. Large-scale patterns have been understood to result from environmental heterogeneity, among other causes. We investigated species richness patterns among submerged and floating-leaved aquatic plants (87 native species and eight invasives) in 103 temperate lakes in Connecticut (northeastern USA) and found neither a consistently negative relationship at small (3-m2) scales, nor a positive relationship at large scales. Native species richness at sampling locations was uncorrelated with invasive species richness in 37 of the 60 lakes where invasive plants occurred; richness was negatively correlated in 16 lakes and positively correlated in seven. No correlation between native and invasive species richness was found at larger spatial scales (whole lakes and counties). Increases in richness with area were uncorrelated with abiotic heterogeneity. Logistic regression showed that the probability of occurrence of five invasive species increased in sampling locations (3 m2, n = 2980 samples) where native plants occurred, indicating that native plant species richness provided no resistance against invasion. However, the probability of three invasive species' occurrence declined as native plant density increased, indicating that density, if not species richness, provided some resistance with these species. Density had no effect on occurrence of three other invasive species. Based on these results, native species may resist invasion at small spatial scales only in communities where density is high (i.e., in communities where competition among individuals contributes to community structure). Most hydrophyte communities, however, appear to be maintained in a nonequilibrial condition by stress and/or disturbance. Therefore, most aquatic plant communities in temperate lakes are likely to be vulnerable to invasion.     

Consequences of dominance: a review of evenness effects on local and regional ecosystem processes

The composition of communities is strongly altered by anthropogenic manipulations of biogeochemical cycles, abiotic conditions, and trophic structure in all major ecosystems. Whereas the effects of species loss on ecosystem processes have received broad attention, the consequences of altered species dominance for emergent properties of communities and ecosystems are poorly investigated. Here we propose a framework guiding our understanding of how dominance affects species interactions within communities, processes within ecosystems, and dynamics on regional scales. Dominance (or the complementary term, evenness) reflects the distribution of traits in a community, which in turn affects the strength and sign of both intraspecifc and interspecific interactions. Consequently, dominance also mediates the effect of such interactions on species coexistence. We review the evidence for the fact that dominance directly affects ecosystem functions such as process rates via species identity (the dominant trait) and evenness (the frequency distribution of traits), and indirectly alters the relationship between process rates and species richness. Dominance also influences the temporal and spatial variability of aggregate community properties and compositional stability (invasibility). Finally, we propose that dominance affects regional species coexistence by altering metacommunity dynamics. Local dominance leads to high beta diversity, and rare species can persist because of source-sink dynamics, but anthropogenically induced environmental changes result in regional dominance and low beta diversity, reducing regional coexistence. Given the rapid anthropogenic alterations of dominance in many ecosystems and the strong implications of these changes, dominance should be considered explicitly in the analysis of consequences of altered biodiversity.     

The invasion paradox: reconciling pattern and process in species invasions

The invasion paradox describes the co-occurrence of independent lines of support for both a negative and a positive relationship between native biodiversity and the invasions of exotic species. The paradox leaves the implications of native-exotic species richness relationships open to debate: Are rich native communities more or less susceptible to invasion by exotic species? We reviewed the considerable observational, experimental, and theoretical evidence describing the paradox and sought generalizations concerning where and why the paradox occurs, its implications for community ecology and assembly processes, and its relevance for restoration, management, and policy associated with species invasions. The crux of the paradox concerns positive associations between native and exotic species richness at broad spatial scales, and negative associations at fine scales, especially in experiments in which diversity was directly manipulated. We identified eight processes that can generate either negative or positive native-exotic richness relationships, but none can generate both. As all eight processes have been shown to be important in some systems, a simple general theory of the paradox, and thus of the relationship between diversity and invasibility, is probably unrealistic. Nonetheless, we outline several key issues that help resolve the paradox, discuss the difficult juxtaposition of experimental and observational data (which often ask subtly different questions), and identify important themes for additional study. We conclude that natively rich ecosystems are likely to be hotspots for exotic species, but that reduction of local species richness can further accelerate the invasion of these and other vulnerable habitats.