Patterns and Pathways in the Post-Establishment Spread of Non-Indigenous Aquatic Species: The Slowing Invasion of North American Inland Lakes by the Zebra Mussel

The zebra mussel, Dreissena polymorpha, has spread through eastern North American aquatic ecosystems during the past 15 years. Whereas spread among navigable waterways was rapid, the invasion of isolated watersheds has progressed more slowly and less predictably. We examined the patterns of overland spread over multiple spatial and temporal extents including individual lake districts, states, and multi-state regions in the USA and found that only a small proportion (<8%) of suitable inland lakes have been invaded, with the rate of invasion appearing to be slowing. Of the 293 lakes known to be invaded, 97% are located in states adjacent to the Laurentian Great Lakes with over half located in Michigan. Only six states have more than 10 invaded lakes and only in Michigan and Indiana have more than 10% of suitable lakes become invaded. At smaller spatial extents, invaded lakes are often clustered within a lake-rich region across southern Michigan and northern Indiana. This clustering appears primarily due to multiple overland invasions originating from the Great Lakes followed to a lesser extent by subsequent secondary overland and downstream dispersal. Downstream spread appears responsible for only one third of the inland invasions. Temporally, invasions peaked in the late 1990s, with only 13 new invasions (0.4% of suitable lakes) reported in 2003 in the four-state region surrounding Lake Michigan. Peak rates of invasion occurred 4–6 years earlier in Michigan relative to Indiana and Wisconsin, but this time lag is likely due to differences in the establishment of Great Lake source populations rather than ‘stepping stone’ dispersal across the landscape.     

Deciphering the worldwide invasion of the Asian long‐horned beetle: A recurrent invasion process from the native area together with a bridgehead effect

Retracing introduction routes is crucial for understanding the evolutionary processes involved in an invasion, as well as for highlighting the invasion history of a species at the global scale. The Asian long‐horned beetle (ALB) Anoplophora glabripennis is a xylophagous pest native to Asia and invasive in North America and Europe. It is responsible for severe losses of urban trees, in both its native and invaded ranges. Based on historical and genetic data, several hypotheses have been formulated concerning its invasion history, including the possibility of multiple introductions from the native zone and secondary dispersal within the invaded areas, but none have been formally tested. In this study, we characterized the genetic structure of ALB in both its native and invaded ranges using microsatellites. In order to test different invasion scenarios, we used an approximate Bayesian “random forest” algorithm together with traditional population genetics approaches. The strong population differentiation observed in the native area was not geographically structured, suggesting complex migration events that were probably human‐mediated. Both native and invasive populations had low genetic diversity, but this characteristic did not prevent the success of the ALB invasions. Our results highlight the complexity of invasion pathways for insect pests. Specifically, our findings indicate that invasive species might be repeatedly introduced from their native range, and they emphasize the importance of multiple, human‐mediated introductions in successful invasions. Finally, our results demonstrate that invasive species can spread across continents following a bridgehead path, in which an invasive population may have acted as a source for another invasion.     

Modelling the introduction and spread of non‐native species: international trade and climate change drive ragweed invasion

Biological invasions are a major driver of global change, for which models can attribute causes, assess impacts and guide management. However, invasion models typically focus on spread from known introduction points or non‐native distributions and ignore the transport processes by which species arrive. Here, we developed a simulation model to understand and describe plant invasion at a continental scale, integrating repeated transport through trade pathways, unintentional release events and the population dynamics and local anthropogenic dispersal that drive subsequent spread. We used the model to simulate the invasion of Europe by common ragweed (Ambrosia artemisiifolia), a globally invasive plant that causes serious harm as an aeroallergen and crop weed. Simulations starting in 1950 accurately reproduced ragweed's current distribution, including the presence of records in climatically unsuitable areas as a result of repeated introduction. Furthermore, the model outputs were strongly correlated with spatial and temporal patterns of ragweed pollen concentrations, which are fully independent of the calibration data. The model suggests that recent trends for warmer summers and increased volumes of international trade have accelerated the ragweed invasion. For the latter, long distance dispersal because of trade within the invaded continent is highlighted as a key invasion process, in addition to import from the native range. Biosecurity simulations, whereby transport through trade pathways is halted, showed that effective control is only achieved by early action targeting all relevant pathways. We conclude that invasion models would benefit from integrating introduction processes (transport and release) with spread dynamics, to better represent propagule pressure from native sources as well as mechanisms for long‐distance dispersal within invaded continents. Ultimately, such integration may facilitate better prediction of spatial and temporal variation in invasion risk and provide useful guidance for management strategies to reduce the impacts of invasion.     

Overlooking the smallest matter: viruses impact biological invasions

Parasites and pathogens have recently received considerable attention for their ability to affect biological invasions, however, researchers have largely overlooked the distinct role of viruses afforded by their unique ability to rapidly mutate and adapt to new hosts. With high mutation and genomic substitution rates, RNA and single‐stranded DNA (ssDNA) viruses may be important constituents of invaded ecosystems, and could potentially behave quite differently from other pathogens. We review evidence suggesting that rapidly evolving viruses impact invasion dynamics in three key ways: (1) Rapidly evolving viruses may prevent exotic species from establishing self‐sustaining populations. (2) Viruses can cause population collapses of exotic species in the introduced range. (3) Viruses can alter the consequences of biological invasions by causing population collapses and extinctions of native species. The ubiquity and frequent host shifting of viruses make their ability to influence invasion events likely. Eludicating the viral ecology of biological invasions will lead to an improved understanding of the causes and consequences of invasions, particularly as regards establishment success and changes to community structure that cannot be explained by direct interspecific interactions among native and exotic species.     

Allee effects, aggregation, and invasion success

Understanding the factors that influence successful colonization can help inform ecological theory and aid in the management of invasive species. When founder populations are small, individual fitness may be negatively impacted by component Allee effects through positive density dependence (e.g., mate limitation). Reproductive and survival mechanisms that suffer due to a shortage of conspecifics may scale up to be manifest in a decreased per-capita population growth rate (i.e., a demographic Allee effect). Mean-field population level models are limited in representing how component Allee effects scale up to demographic Allee effects when heterogeneous spatial structure influences conspecific availability. Thus, such models may not adequately characterize the probability of establishment. In order to better assess how individual level processes influence population establishment and spread, we developed a spatially explicit individual-based stochastic simulation of a small founder population. We found that increased aggregation can affect individual fitness and subsequently impact population growth; however, relatively slow dispersal—in addition to initial spatial structure—is required for establishment, ultimately creating a tradeoff between probability of initial establishment and rate of subsequent spread. Since this result is sensitive to the scaling up of component Allee effects, details of individual dispersal and interaction kernels are key factors influencing population level processes. Overall, we demonstrate the importance of considering both spatial structure and individual level traits in assessing the consequences of Allee effects in biological invasions.     

Successful establishment of the invasive fire ant Solenopsis invicta in Taiwan: insights into interactions of alternate social forms

Aim Understanding the factors underlying the successful establishment of invasive ant species is critical for developing quarantine strategies to prevent additional invasions as well as for determining how such species overcome the selective pressures in invaded areas. Although several studies have revealed differences in the social organization and population genetics of invasive ants in their native and introduced ranges, few studies have considered the potential interactions between alternate social forms within newly‐invaded areas simply because many invasive ants are characterized as polygyne or unicolonial. Location Taoyuan and Chiayi County, Taiwan. Methods Both social forms (polygyne and monogyne) of the red imported fire ant, Solenopsis invicta, occur in two separately invaded areas in Taiwan (Taoyuan and Chiayi). We employed intensive sampling methods and diagnostic polymerase chain reaction assays to determine the distribution of these two social forms in both infested areas in Taiwan. Results The distribution of social forms differs dramatically between the two infested areas, consistent with different invasion histories regarding the original make‐up of individuals comprising the initial founder group. The Taoyuan population likely was colonized initially by ants of both social forms, with the subsequent spread characterized by continuous outward movement of the two forms, particularly the monogyne form. In contrast, the initial founders of the Chiayi population likely were of the polygyne social form only, suggesting the monogyne social form in this population appeared only recently and likely arose directly from polygyne queens converting to the alternate social form. Main conclusions Our results provide detailed insights into the invasion history of S. invicta in Taiwan and suggest that the distinct reproductive biology of the two social forms may have shaped the current distribution of S. invicta in these infested areas and that the dynamics of two forms may affect the long‐term persistence and potential for spread of this pest ant species.     

动物入侵的行为机制

行为特征可在外来动物建立种群和扩张过程中发挥重要作用,因此,要正确理解动物入侵,常常需要仔细研究其行为机制。20世纪80年代以来,随着动物入侵规模在世界各地的迅速加剧,有关其行为机制的研究也受到了广泛关注。最近一些研究表明,一些入侵动物种内攻击和觅食等行为具有可塑性,因此它们能够灵活应对多变的环境条件,这对于种群的建立和维持至关重要;入侵动物与土著物种发生行为互作时,往往占据优势,从而取代土著物种,并有助于其地域扩张;入侵动物长距离扩散可以提高其地域扩张速度,许多行为可与扩散行为结合进一步促进扩张。今后需要加强对入侵动物的行为分析,使之全面地融合到生物入侵的研究之中。这不仅可以提高对外来物种入侵的预警和治理能力,而且为探索动物行为的奥秘以及动物间行为互作在物种进化中的意义提供了独特的机会。     

Testing population-level mechanisms of invasion by a mobile vertebrate: a simple conceptual framework for salmonids in streams

Invasion biology suffers from a lack of the ability to predict the outcome of particular invasions because of reliance on verbal models and lack of rigorous experimental data at the appropriate scale. More progress is likely to be made by considering invasions as population-level phenomena and initially focusing on specific taxa or particular categories of invasions. To this end, we propose a simple conceptual framework to motivate studies of invasion by salmonids (salmon, trout, grayling, and whitefish) in streams that emphasizes population-level mechanisms affecting native species and promoting spread by the invader. Specifically, the only direct mechanisms by which the abundance of the native species can decline are through biotic interactions which cause decreased reproductive rates or survival at specific life stages, net emigration, debilitating or fatal diseases introduced by the invader, or a combination of these factors. Conversely, abundance of the invader must increase by local reproduction, high survival, net immigration, or a combination of these factors. Review of existing salmonid invasion literature suggests that future studies could be improved by using manipulative field experiments at a spatial and temporal scale appropriate to address population-level processes, characterizing how movement affects the establishment and spread of an invader, and including abiotic context in experimental designs. Using the example of brook trout (Salvelinus fontinalis) invasion into streams containing native Colorado River cutthroat trout (Oncorhynchus clarki pleuriticus) in the central Rocky Mountains (USA), we demonstrate how the framework can be used to design a manipulative field experiment to test for population-level mechanisms causing ecological effects and promoting invasion success. Experiments of this type will give invasion ecologists a useful example of how a taxon-specific invasion framework can improve the ability to predict ecological effects, and provide fishery biologists with the quantitative foundation necessary to better manage stream salmonid invasions.     

Initial conditions and their effect on invasion velocity across heterogeneous landscapes

Accurate, time dependent control options are required to halt biological invasions prior to equilibrium establishment, beyond which control efforts are often impractical. Although invasions have been successfully modeled using diffusion theory, diffusion models are typically confined to providing simple range expansion estimates. In this work, we use a Susceptible/Infected cellular automaton (CA) to simulate diffusion. The CA model is coupled with a network model to track the speed and direction of simulated invasions across heterogeneous landscapes, allowing for identification of locations for targeted control in both time and space. We evaluated the role of the location of initial establishment insofar as it affected the pattern and rate of spread and how these are influenced by patch attributes such as size. Our results show that the location of initial establishment can significantly affect the temporal dynamics of an invasion. Traditional network metrics such as degree and measures of topological distance were insufficient for predicting the direction and speed of the invasion. Our coupled models allow the dynamic tracking of invasions across fragmented landscapes for both theoretical and practical applications.     

Mechanisms of invasive population establishment and spread of pinewood nematodes in China

This paper summarizes the results of our study of the pinewood nematode (Bursaphelenchus xylophilus). By population genetic analysis, it was determined that there was no genetic bottle caused by the founder effect and genetic drift in the Chinese invasive population. Multiple invasions with large amounts of nematodes from different sources led to rich genetic diversity in the invasive population. Keeping high genetic diversity in the invasive process may be one of the genetic mechanisms in its successful invasion. By testing interspecies competition, it was shown that, with high fecundity and a strong competitive ability, B. xylophilus outcompeted the native species B. mucronatus in the natural ecosystem during the invasion process. Competitive displacement may be one of the ecological mechanisms of B. xylophilus’s invasion. In addition, an unequal interspecific hybridization with introgress was in favor of the invasive species which also accelerated the replacement of B. mucronatus by B. xylophilus. The structures, functions and evolutions of a few important genes that are closely related to the ecological adaptation of pinewood nematodes were studied to explore the molecular mechanism of its ecological adaptations. Further, the resistance and resilience mechanism of the pine ecosystem invaded by pinewood nematodes was also investigated. The results of these studies uncovered a portion of the genetic and ecological mechanisms of PWN’s successful invasion and laid a foundation for further study to obtain a comprehensive interpretation of the mechanisms of the nematode invasion. The results also provided a scientific basis for effectively controlling the occurrence and spread of pine wilt disease which is caused by nematodes. Various aspects requiring further investigation are considered. Supported by the National Key Basic Research and Development Program of China (Grant Nos. 2009CB119200 and 2002CB111400).     

Dynamic expansion in recently introduced populations of fire ant parasitoids (Diptera: Phoridae)

Combating invasive species requires a detailed, mechanistic understanding of the manner and speed with which organisms expand their ranges. Biological control efforts provide an opportunity to study the process of species invasions and range expansions under known initial conditions. This study examines the rate, pattern and mechanisms of spread for two populations of the biological control agent Pseudacteon tricuspis, phorid-fly parasitoids of imported fire ants. We employ a trap-based survey method that detects phorid flies in low-density populations, and provides data on abundance. This technique allows us to differentiate between continuous population spread and effective long-distance dispersal and to examine density gradients of phorid flies across the expanding population front. We find that occupied sites in front of the leading edge of continuous populations were common; forming small populations we refer to as satellite populations. Satellite populations are tens of kilometers from the nearest possible source. Wind governs the dynamics of spread in these two central Texas populations. Population edges expanding with the wind exhibited a higher frequency of effective long-distance dispersal than did populations expanding into the wind. This enhanced effective long-distance dispersal rate translated into a five times faster rate of spread for population edges traveling with the wind. This planned invasion shares many characteristics in common with unplanned species invasions including: protracted establishment phase during which densities were below detection thresholds, and slow initial spread immediately after establishment followed by rapid, accelerating spread rates as population sizes grew.     

Prediction and error in multi‐stage models for spread of aquatic non‐indigenous species

Aim Predictions of spread of non‐indigenous species allow for greater efficiency in managing invasions by targeting areas for preventative measures. The invasion sequence is a useful concept in predictions of spread, as it allows us to test hypotheses about the transport and establishment of propagules in novel habitats. Our aims are twofold: (1) to develop and validate multi‐stage invasion models for the introduced fishhook waterflea, Cercopagis pengoi, and (2) to assess how variability in the transport patterns of the propagules influences the accuracy and spatial extent for predictions of spread. Location New York State, USA. Methods We developed a two‐stage model for the spread of C. pengoi. First, we developed a stochastic gravity model for dispersal based on surveys of recreational boat traffic in New York State as a proxy for propagule pressure. We then modelled the probability of establishment based on predicted levels of propagule pressure and measures of lakes’ physicochemistry. In addition, we used Monte Carlo simulations based on the gravity model to propagate variability in boater traffic through the establishment model to assess how uncertainty in dispersal influenced predictions of spread. Results The amount recreationalists were willing to spend, lake area and population size of the city nearest to the destination lake were significant factors affecting boater traffic. In turn, boater traffic, lake area, specific conductance and turbidity were significant predictors of establishment. The inclusion of stochastic dispersal reduced the rate of false positives (i.e. incorrect prediction of an invasion) in detecting invasions at the upper 95% prediction interval for the probability of establishment. Main conclusions Combinations of measures of propagule pressure, habitat suitability and stochastic dispersal allow for the most accurate predictions of spread. Further, multi‐stage spread models may overestimate the extent of spread if stochasticity in early stages of the models is not considered.     

Global networks for invasion science: benefits,challenges and guidelines

Much has been done to address the challenges of biological invasions, but fundamental questions (e.g., which species invade? Which habitats are invaded? How can invasions be effectively managed?) still need to be answered before the spread and impact of alien taxa can be effectively managed. Questions on the role of biogeography (e.g., how does biogeography influence ecosystem susceptibility, resistance and resilience against invasion?) have the greatest potential to address this goal by increasing our capacity to understand and accurately predict invasions at local, continental and global scales. This paper proposes a framework for the development of ‘Global Networks for Invasion Science’ to help generate approaches to address these critical and fundamentally biogeographic questions. We define global networks on the basis of their focus on research questions at the global scale, collection of primary data, use of standardized protocols and metrics, and commitment to long-term global data. Global networks are critical for the future of invasion science because of their potential to extend beyond the capacity of individual partners to identify global priorities for research agendas and coordinate data collection over space and time, assess risks and emerging trends, understand the complex influences of biogeography on mechanisms of invasion, predict the future of invasion dynamics, and use these new insights to improve the efficiency and effectiveness of evidence-based management techniques. While the pace and scale of global change continues to escalate, strategic and collaborative global networks offer a powerful approach to inform responses to the threats posed by biological invasions.     

Green Frogs Show Reduced Foraging Success in Habitats Invaded by Japanese knotweed

Habitat loss is causing amphibian population declines worldwide, so there is increased attention to forces that degrade remaining habitats. Terrestrial habitats surrounding wetlands are critical foraging areas for temperate anurans. We investigated plant community changes in two old fields invaded by Japanese knotweed (Fallopia japonica) and the foraging success of Green frogs (Rana clamitans) in invaded and non-invaded portions of those fields. Within each field, vegetation data were recorded in quadrats located along two transects bisecting the invasion fronts. We placed frogs in ‘foraging buckets’ along transects and measured their change in mass over a 38 h period. There were significant changes in vegetation structure and composition associated with Japanese knotweed invasion. Diverse assemblages of native plants that covered non-invaded plots were absent from areas invaded by Japanese knotweed. There was also a significant change in vegetation architecture between invaded and non-invaded habitats. Change in frog mass declined significantly along transects, with most frogs in non-invaded plots gaining mass and no frogs in invaded plots gaining mass. Most frogs from non-invaded plots but only two from invaded plots defecated shortly after removal from foraging buckets (verification of recent feeding). We hypothesize that Japanese knotweed invasions degrade terrestrial habitat quality for frogs by indirectly reducing arthropod abundance. Nonnative plant invasions may be another factor contributing to amphibian population declines.     

Invasion dynamics of round goby (<Emphasis Type="Italic">Neogobius melanostomus</Emphasis>) in Hamilton Harbour,Lake Ontario

Most introduced non-native species fail to establish as a result of mortality or reproductive failure. An established population can increase the probability of survival and reproductive success of newly introduced individuals by reducing both Allee effects and demographic stochasticity. Previously, attention has been paid to the establishment phase of the invasion process and its probability modelled as a stochastic process, while the spread phase has received less attention. By analyzing data collected during the spread phase of an invasion of the round goby, Neogobius melanostomus, in Hamilton Harbour, Lake Ontario, we develop an analytical approach to backcalculate the time to establishment and to determine the time to habitat saturation. Our modelling shows that: (1) during the transition between arrival and establishment, propagule pressure in the form of new adults entering the area can be very low and still represent a significant probability of establishment; (2) much higher concentrations of juveniles would be needed to pose a significant risk of invasion; (3) the demographic contribution of propagule pressure during the spread phase is low and its total elimination will not halt population growth and spread; (4) a short elapsed time between arrival and establishment indicated that the transition between these two phases can be characterized as a deterministic process with high propagule pressure and low adult mortality rates; and, (5) very aggressive management actions would be needed to halt population growth after population establishment, suggesting that preventative measures are the most effective management options available to reduce risk of future invasions.     

Multiple introductions and population structure during the rapid expansion of the invasive Sahara mustard (Brassica tournefortii)

The specific mechanisms that result in the success of any species invasion case are difficult to document. Reproductive strategies are often cited as a primary driver of invasive success, with human activities further facilitating invasions by, for example, acting as seed vectors for dispersal via road, train, air, and marine traffic, and by producing efficient corridors for movement including canals, drainages, and roadways. Sahara mustard (Brassica tournefortii) is a facultative autogamous annual native to Eurasia that has rapidly invaded the southwestern United States within the past century, displacing natives, and altering water‐limited landscapes in the southwest. We used a genotyping‐by‐sequencing approach to study the population structure and spatial geography of Sahara mustard from 744 individuals from 52 sites across the range of the species’ invasion. We also used herbaria records to model range expansion since its initial introduction in the 1920s. We found that Sahara mustard occurs as three populations in the United States unstructured by geography, identified three introduction sites, and combined herbaria records with genomic analyses to map the spread of the species. Low genetic diversity and linkage disequilibrium are consistent with self‐fertilization, which likely promoted rapid invasive spread. Overall, we found that Sahara mustard experienced atypical expansion patterns, with a relatively constant rate of expansion and without the lag phase that is typical of many invasive species.     

Evaluating methods to quantify spatial variation in the velocity of biological invasions

Invading species rarely spread homogeneously through a landscape and invasion patterns typically display irregular frontal boundaries as the invasion progresses through space. Those irregular patterns are generally produced by local environmental factors that may slow or accelerate movement of the frontal boundary. While there is an abundant literature on species distribution modelling methods that quantify local suitability for species establishment, comparatively few studies have examined methods for measuring the local velocity of invasions that can then be statistically analysed in relation to spatially variable environmental factors. Previous studies have used simulations to compare different methods for estimating the overall rate of spread of an invasion. We adopted a similar approach of simulating invasions that resemble two real case‐studies, both in terms of their spatial resolution (i.e. considering the size of one cell as one km) and their spatial extent (> 600 000 km²). Simulations were sampled to compare how different methods used to measure local spread rate, namely the neighbouring, nearest distance and Delaunay methods, perform for spatio‐temporal comparisons. We varied the assessment using three levels of complexity of the spatio‐temporal pattern of invasion, three sample sizes (500, 1000 and 2000 points), three different spatial sampling patterns (stratified, random, aggregated), three interpolation methods (generalized linear model, kriging, thin plate spline regression) and two spatio‐temporal modelling structures (trend surface analysis and boundary displacement), resulting in a total of 486 different scenarios. The thin plate spline regression interpolation method, in combination with trend surface analysis, was found to provide the most robust local spread rate quantification as it was able to reliably accommodate different sampling conditions and invasion patterns. This best approach was successfully applied to two case‐studies, the invasion of France by the horse‐chestnut leafminer Cameraria ohridella and by the bluetongue virus, generally in agreement with previously published values of spread rates. Potential avenues for further research are discussed.     

Pine invasion decreases density and changes native tree communities in woodland Cerrado

ABSTRACT

Background: Invasive plants can negatively impact native communities, but the majority of the effects of these invasions have been demonstrated only for temperate ecosystems. Tropical ecosystems, including the Cerrado, a biodiversity hotspot, are known to be invaded by numerous non-native species, but studies of their impacts are largely lacking.

Aims: Our research aimed at quantifying how Pinus spp. presence and density affected Cerrado plant communities.

Methods: We sampled areas invaded and non-invaded by Pinus spp. to determine if pine invasion affected native tree richness, diversity, evenness, and density. We also evaluated if community composition differed between invaded and non-invaded sites.

Results: We found invaded plots had lower native tree densities than non-invaded plots and that Pinus spp. invasions changed native tree communities by reducing native species abundances.

Conclusion: Invasive pines had negative impacts on the native Cerrado tree community by reducing native plant density and changing species abundances. Reduced density and abundance at early invasion stages can result in reduction in biodiversity in the long term.     

Niche conservatism and spread of seaweed invasive lineages with different residence time in the Mediterranean Sea

Marine algae invasions attract a lot of interest as they are altering the structure of marine ecosystems. However, niche dynamics and risk predictions of marine invasions integrating phylogeographic structure in the analyses have not yet been investigated. In this study, we perform a comprehensive analysis of two invasive lineages of Caulerpa taxifolia with different residence time in the Mediterranean Sea for a better understanding of their invasive processes. We performed lineage-based and species-based niche models to assess the risk of invasion, the spatial overlap, and the variables delimiting the distribution of the two lineages. We also compared the effect of using different extents on niche overlap and niche shift analyses. Intraspecific models with pooled occurrences accurately found two separate regions susceptible of invasion for each invasive lineage in the Mediterranean, while species-based predictions underestimated invaded regions. The invasive lineages spread across colder coastal areas than the species. Altogether, we provide evidence that different invasive lineages of algae show dissimilar environmental responses and invasive ranges that are not detectable by species-based analyses. Moreover, niche overlap and niche shift analyses seem to depend greatly on the geographical extent used. According to the most appropriate extent (worldwide), the invaded range did not show niche shift, and thus, no evidence of a post-introduction adaptation scenario was found as both lineages invaded habitats similar to their Australian native locations. Actions to prevent further spreading of the most recent invasive lineage are needed.     

Occupation,body size and sex ratio of round goby (<Emphasis Type="Italic">Neogobius melanostomus</Emphasis>) in established and newly invaded areas of an Ontario river

Invasive species represent a challenge because the particular characteristics of a species’ invasion are often unknown before the invasion. To provide some clarity as to how invasive species demographic structure might change as a population advances its range, we compared the proportion of occupied sites, size structure and sex ratio of round gobies in the area where they first invaded with more recently invaded areas at the extent of their range in a river in south-eastern Ontario. We used a standardized angling technique to sample gobies larger than 45-mm total length in the summer and early autumn of 2007. Round goby at the upstream and downstream extent of their range occupied a lower proportion of randomly selected sites, and contained a wider distribution of sizes as well as significantly larger individuals. Sex ratios in all areas were male-biased and the male-to-female ratio was significantly higher in the upstream segment of the river (2.2:1) compared to the area of first introduction (1.4:1). The difference between the newly invaded and the established sites suggests that round goby population structure may be affected by density. The results of this study help us further describe the demographic characteristics of biological invasions whilst examples of population structure and behaviour in gobies and other species provide a basis for generating hypotheses for range expansion.