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1.
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.  相似文献   

2.
The process of nonindigenous species (NIS) arrival has received limited theoretical consideration despite importance in predicting and preventing the establishment of NIS. We formulate a mechanistically based hierarchical model of NIS arrival and demonstrate simplifications leading to a marginal distribution of the number of surviving introduced individuals from parameters of survival probability and propagule pressure. The marginal distribution is extended as a stochastic process from which establishment emerges with a waiting time distribution. This provides a probability of NIS establishment within a specified period and may be useful for identifying patterns of successful invaders. However, estimates of both the propagule pressure and the individual survival probability are rarely available for NIS, making estimates of the probability of establishment difficult. Alternatively, researchers are able to measure proportional estimates of propagule pressure through models of NIS transport, such as gravity models, or of survival probability through habitat-matching indexes measuring the similarity between potentially occupied and native NIS ranges. Therefore, we formulate the relative waiting time between two locations and the probability of one location being invaded before the other.  相似文献   

3.
Determinants of vertebrate invasion success in Europe and North America   总被引:4,自引:0,他引:4  
Species that are frequently introduced to an exotic range have a high potential of becoming invasive. Besides propagule pressure, however, no other generally strong determinant of invasion success is known. Although evidence has accumulated that human affiliates (domesticates, pets, human commensals) also have high invasion success, existing studies do not distinguish whether this success can be completely explained by or is partly independent of propagule pressure. Here, we analyze both factors independently, propagule pressure and human affiliation. We also consider a third factor directly related to humans, hunting, and 17 traits on each species' population size and extent, diet, body size, and life history. Our dataset includes all 2362 freshwater fish, mammals, and birds native to Europe or North America. In contrast to most previous studies, we look at the complete invasion process consisting of (1) introduction, (2) establishment, and (3) spread. In this way, we not only consider which of the introduced species became invasive but also which species were introduced. Of the 20 factors tested, propagule pressure and human affiliation were the two strongest determinants of invasion success across all taxa and steps. This was true for multivariate analyses that account for intercorrelations among variables as well as univariate analyses, suggesting that human affiliation influenced invasion success independently of propagule pressure. Some factors affected the different steps of the invasion process antagonistically. For example, game species were much more likely to be introduced to an exotic continent than nonhunted species but tended to be less likely to establish themselves and spread. Such antagonistic effects show the importance of considering the complete invasion process.  相似文献   

4.
Propagule pressure is commonly considered a primary driver of invasive plant establishment and spread. However, the physical size or condition (i.e., quality) of propagules may also affect establishment, particularly under unfavorable habitat conditions such as low light environments. We used an outdoor mesocosm experiment to test the relative contribution of propagule size (number of individuals introduced) and quality (number of rhizome nodes) to the establishment and performance of the highly invasive cogongrass (Imperata cylindrica) under experimental sun and shade treatments. We found that the introduction of higher quality propagules (rhizome segments ≥3 nodes in length) significantly enhanced establishment across both light treatments, and increased final tiller count in the sun treatment. The sun treatment also enhanced rhizome growth, an effect that could increase spread rates and invasion success. Thus, while cogongrass is likely to establish in both sun and shade, introductions of large propagule sizes or large rhizomes in high light environments likely poses the greatest threat to native habitats. Our results demonstrate that propagule quality promoted both establishment and performance of a highly invasive grass species and suggest that propagule quality may play an important but underappreciated role in the invasion process.  相似文献   

5.
Invasion ecology has been criticised for its lack of general principles. To explore this criticism, we conducted a meta-analysis that examined characteristics of invasiveness (i.e. the ability of species to establish in, spread to, or become abundant in novel communities) and invasibility (i.e. the susceptibility of habitats to the establishment or proliferation of invaders). There were few consistencies among invasiveness characteristics (3 of 13): established and abundant invaders generally occupy similar habitats as native species, while abundant species tend to be less affected by enemies; germination success and reproductive output were significantly positively associated with invasiveness when results from both stages (establishment/spread and abundance/impact) were combined. Two of six invasibility characteristics were also significant: communities experiencing more disturbance and with higher resource availability sustained greater establishment and proliferation of invaders. We also found that even though ‘propagule pressure’ was considered in only ~29% of studies, it was a significant predictor of both invasiveness and invasibility (55 of 64 total cases). Given that nonindigenous species are likely introduced non-randomly, we contend that ‘propagule biases’ may confound current paradigms in invasion ecology. Examples of patterns that could be confounded by propagule biases include characteristics of good invaders and susceptible habitats, release from enemies, evolution of ‘invasiveness’, and invasional meltdown. We conclude that propagule pressure should serve as the basis of a null model for studies of biological invasions when inferring process from patterns of invasion. An erratum to this article can be found at  相似文献   

6.
Mechanistic insights from invasion biology indicate that propagule pressure of exotic species and native community structure can independently influence establishment success. The role of native community connectivity via species dispersal and its potential interaction with propagule pressure on invasion success in metacommunities, however, remains unknown. Native community connectivity may increase biotic resistance to invasion by enhancing species richness and evenness, but the effects could depend upon the level of propagule pressure. In this study, a mesocosm experiment was used to evaluate the independent and combined effects of exotic propagule pressure and native community connectivity on invasion success. The effects of three levels of exotic Daphnia lumholtzi propagule pressure on establishment success, community structure and ecosystem attributes were evaluated in native zooplankton communities connected by species dispersal versus unconnected communities, and relative to a control without native species. Establishment of the exotic species exhibited a propagule dose‐dependent relationship with high levels of propagule pressure resulting in the greatest establishment success. Native community connectivity, however, effectively reduced establishment at the low level of propagule pressure and further augmented native species richness across propagule pressure treatments. Propagule pressure largely determined the negative impacts of the exotic species on native species richness, native biomass and edible producer biomass. The results highlight that native community connectivity can reduce invasion success at a low propagule dose and decrease extinction risk of native competitors, but high propagule pressure can overcome connectivity‐mediated biotic resistance to influence establishment and impact of the exotic species. Together, the results emphasize the importance of the interaction of propagule pressure and community connectivity as a regulator of invasion success, and argue for the maintenance of metacommunity connectivity to confer invasion resistance.  相似文献   

7.
One of the strongest generalities in invasion biology is the positive relationship between probability of establishment and the numbers of individuals introduced. Nevertheless, a number of significant questions remain regarding: (1) the relative importance of different processes during introduction (e.g., demographic, environmental, and genetic stochasticity, and Allee effects); (2) the relative effects of propagule pressure (e.g., number of introductions, size of introductions, and lag between introductions); and (3) different life history characteristics of the species themselves. Here, we adopt an individual-based simulation modeling approach to explore a range of such details in the relationship between establishment success and numbers of individuals introduced. Our models are developed for typical exotic bird introductions, for which the relationship between probability of establishment and the numbers of individuals introduced has been particularly well documented. For both short-lived and long-lived species, probability of establishment decreased across multiple introductions (compared with a single introduction of the same total size), and this decrease was greater when inbreeding depression was included. Sensitivity analyses revealed four predictors that together accounted for >95 % of model performance. Of these, R 0 (the average number of daughters produced per female over her lifetime) and propagule pressure were of primary importance, while random environmental effects and inbreeding depression exerted lesser influence. Initial founder size is undoubtedly going to be important for ensuring the persistence of introduced populations. However, we found the demographic traits, which influence how introduced individuals behave, to have the greatest effect on establishment success.  相似文献   

8.
High propagule pressure is arguably the only consistent predictor of colonization success. More individuals enhance colonization success because they aid in overcoming demographic consequences of small population size (e.g. stochasticity and Allee effects). The number of founders can also have direct genetic effects: with fewer individuals, more inbreeding and thus inbreeding depression will occur, whereas more individuals typically harbour greater genetic variation. Thus, the demographic and genetic components of propagule pressure are interrelated, making it difficult to understand which mechanisms are most important in determining colonization success. We experimentally disentangled the demographic and genetic components of propagule pressure by manipulating the number of founders (fewer or more), and genetic background (inbred or outbred) of individuals released in a series of three complementary experiments. We used Bemisia whiteflies and released them onto either their natal host (benign) or a novel host (challenging). Our experiments revealed that having more founding individuals and those individuals being outbred both increased the number of adults produced, but that only genetic background consistently shaped net reproductive rate of experimental populations. Environment was also important and interacted with propagule size to determine the number of adults produced. Quality of the environment interacted also with genetic background to determine establishment success, with a more pronounced effect of inbreeding depression in harsh environments. This interaction did not hold for the net reproductive rate. These data show that the positive effect of propagule pressure on founding success can be driven as much by underlying genetic processes as by demographics. Genetic effects can be immediate and have sizable effects on fitness.  相似文献   

9.
We studied the effect of propagule pressure on the establishment and subsequent spread of the invasive little fire ant Wasmannia auropunctata in a Gabonese oilfield in lowland rain forest. Oil well drilling, the major anthropogenic disturbance over the past 21 years in the area, was used as an indirect measure of propagule pressure. An analysis of 82 potential introductions at oil production platforms revealed that the probability of successful establishment significantly increased with the number of drilling events. Specifically, the shape of the dose–response establishment curve could be closely approximated by a Poisson process with a 34% chance of infestation per well drilled. Consistent with our knowledge of largely clonal reproduction by W. auropunctata , the shape of the establishment curve suggested that the ants were not substantially affected by Allee effects, probably greatly contributing to this species' success as an invader. By contrast, the extent to which W. auropunctata spread beyond the point of initial introduction, and thus the extent of its damage to diversity of other ant species, was independent of propagule pressure. These results suggest that while establishment success depends on propagule pressure, other ecological or genetic factors may limit the extent of further spread. Knowledge of the shape of the dose–response establishment curve should prove useful in modelling the future spread of W. auropunctata and perhaps the spread of other clonal organisms.  相似文献   

10.
Invasive species are spreading at high rates, yet fundamental processes allowing them to progress through the stages of invasion are unclear. The establishment stage is a critical point because this is when exotic species can survive, reproduce, and begin to spread. Unfortunately, inference of population dynamics during this stage may be impossible if historical and observational data are incomplete. Nonetheless, critical inferences on population dynamics during the establishment stage can be acquired indirectly by characterizing demographic history via the population genetics of recently introduced populations. Geckos have been introduced at a global scale and are one of the most successfully establishing families of alien reptile known. Here we conduct a series of population genetic analyses among five close subpopulations of the introduced Mediterranean gecko Hemidactylus turcicus. We tested for non-equilibrium genetic signatures, a pattern expected during early stages of invasion if there were few founders or repeated introductions led to population turnover. Genetic analyses showed no evidence of non-equilibrium dynamics such as genetic bottlenecks. Moreover, we found strong support for population genetic equilibrium dynamics. The observed results may have been generated via an introduction that involved high propagule pressure. However, given the life history of H. turcicus including generation time and dispersal potential, we favor the hypothesis that the invasive metapopulation has rapidly reached the establishment stage as indicated by relatively constant effective sizes and migration rates among introduced subpopulations. The ability to rapidly pass through the establishment stage may in part explain the invasion success of these geckos.  相似文献   

11.
Disturbances create fluctuations in resource availability that alter abiotic and biotic constraints. Exotic invader response may be due to multiple factors related to disturbance regimes and complex interactions between other small- and large-scale abiotic and biotic processes that may vary across invasion stages. We explore how cheatgrass responds to both frequency and season of prescribed burning for a 10-year period in ponderosa pine forested stands. To understand interactions of fire disturbance, other abiotic factors, biotic resistance, and propagule pressure, we use long-term data from different spatial scales representing different invasion stages (local establishment or spread and broader scale extent/impact) to model cheatgrass dynamics. We found that after 10 years, cheatgrass cover increased with fall burning regardless of burn frequency (1 burn vs. 3 burns). There was no evidence that cheatgrass invasion is decreasing through time even in areas burned only once. Factors important for explaining local fine-scale cheatgrass establishment and spread, and broader scale extent/impact varied. The spatial extent of the first burns facilitated fine-scale cheatgrass establishment while bare soil cover constrained establishment. Biotic resistance, in the form of native annual forb cover, constrained fine-scale cheatgrass spread. Initial cheatgrass abundance in 2002, a factor related to propagule pressure, was key for explaining the broader scale extent/impact of cheatgrass by 2012. Biotic resistance, in the form of native perennial grass cover, constrained extent/impact but only when initial cheatgrass abundance was low. Our findings regarding factors affecting invasion dynamics may be useful to consider for future restoration and conservation efforts in burned ponderosa pine forests.  相似文献   

12.
Successful alien species invasion depends on many factors studied mostly in post invasion habitats, and subsequently summarized in frameworks tailored to describe the studied invasion. We used an existing expanded framework with three groups of contributing factors: habitat invisibility, system context and species invasiveness, to analyze the probability of alien species invasions in terrestrial communities of Maritime Antarctic in the future. We focused on the first two factor groups. We tested if the expanded framework could be used under a different scenario. We chose Point Thomas Oasis on King George Island to perform our analysis. Strong geographical barrier, low potential bioclimatic suitability and resource availability associated with habitat invasibility significantly reduce the likelihood of biological invasion in Antarctica. An almost full enemy release (low pressure of consumers), the high patchiness of the habitat, and the prevalence of open gaps also associated with habitat invasibility increase the possibility of invasion. The dynamics of functional connectivity, propagule pressure and spatio-temporal patterns of propagule arrival associated with human activity and climate change belonging to the system context contribute to an increase in the threat of invasions. Due to the still low land transport activity migration pathways are limited and will reduce the spread of alien terrestrial organisms by land. An effective way of preventing invasions in Antarctica seems to lie in reducing propagule pressure and eliminating alien populations as early as possible. The expanded conceptual framework opens up wider possibilities in analyzing invasions taking place in different systems and with multiple taxa.  相似文献   

13.
Colonization is of longstanding interest in theoretical ecology and biogeography, and in the management of weeds and other invasive species, including insect pests and emerging infectious diseases. Due to accelerating invasion rates and widespread economic costs and environmental damages caused by invasive species, colonization theory has lately become a matter of considerable interest. Here we review the concept of propagule pressure to inquire if colonization theory might provide quantitative tools for risk assessment of biological invasions. By formalizing the concept of propagule pressure in terms of stochastic differential equation models of population growth, we seek a synthesis of invasion biology and theoretical population biology. We focus on two components of propagule pressure that affect the chance of invasion: (1) the number of individuals initially introduced, and (2) the rate of subsequent immigration. We also examine how Allee effects, which are expected to be common in newly introduced populations, may inhibit establishment of introduced propagules. We find that the establishment curve (i.e., the chance of invasion as a function of initial population size), can take a variety of shapes depending on immigration rate, carrying capacity, and the severity of Allee effects. Additionally, Allee effects can cause the stationary distribution of population sizes to be bimodal, which we suggest is a possible explanation for time lags commonly observed between the detection of an introduced population and widespread invasion of the landscape.  相似文献   

14.
The monk parakeet (Myiopsitta monachus) is a successful invasive species that does not exhibit life history traits typically associated with colonizing species (e.g., high reproductive rate or long‐distance dispersal capacity). To investigate this apparent paradox, we examined individual and population genetic patterns of microsatellite loci at one native and two invasive sites. More specifically, we aimed at evaluating the role of propagule pressure, sexual monogamy and long‐distance dispersal in monk parakeet invasion success. Our results indicate little loss of genetic variation at invasive sites relative to the native site. We also found strong evidence for sexual monogamy from patterns of relatedness within sites, and no definite cases of extra‐pair paternity in either the native site sample or the examined invasive site. Taken together, these patterns directly and indirectly suggest that high propagule pressure has contributed to monk parakeet invasion success. In addition, we found evidence for frequent long‐distance dispersal at an invasive site (~100 km) that sharply contrasted with previous estimates of smaller dispersal distance made in the native range (~2 km), suggesting long‐range dispersal also contributes to the species’ spread within the United States. Overall, these results add to a growing body of literature pointing to the important role of propagule pressure in determining, and thus predicting, invasion success, especially for species whose life history traits are not typically associated with invasiveness.  相似文献   

15.
BACKGROUND AND AIMS: Success during the early stages of the life-history of alien plants is essential for invasion to occur. The reproductive components of plant invaders have mostly been studied in species reproducing sexually but little is known about invaders that depend exclusively on vegetative reproduction. In this paper, the importance of the different recruitment stages on population growth is quantified and, thus, the invasion potential of the South African annual geophyte Oxalis pes-caprae invading Mediterranean ecosystems is assessed. METHODS: Tests and experiments were conducted across Menorca (Balearic Islands) to analyse the spatial variability of Oxalis pes-caprae reproductive components (i.e. bulb production, bulb bank, bulb predation, bulb mortality, bulb dormancy, bulb germination, plant establishment and survival). KEY RESULTS: Oxalis pes-caprae has a transient bulb bank that remains dormant in the soil during summer. High levels of bulb predation after dispersal, followed by bulb mortality during summer or a failure to germinate in autumn were the most critical factors limiting plant establishment. Bulb germination was high. However, plant establishment and bulb production is constrained by intraspecific competition, but is not affected by soil disturbance. No symptoms of spatial discordance could be found between recruitment stages because the spatial variability of the life cycle was extremely low at all the scales examined (i.e. among populations, habitats and microsites). It was estimated that, on average, 4 % of bulbs can become plants the following year and the field rate of population increase (lambda) to be 0.08. CONCLUSIONS: The results suggest that invasion is constrained by post-dispersal bulb predation, loss of viability of the propagule bank due to summer drought and high intraspecific competition. However, a high spatial concordance between recruitment stages and probably a high propagule pressure due to human and livestock bulb dispersal determine the success of this invader across Menorca Island.  相似文献   

16.
Risk is defined with many minor variations in the biological literature. Common to most definitions are the following elements: the probability of a future event; and the consequences of the event, usually with respect to some predefined human value. Risk analysis includes elements of risk assessment (quantification of risk), uncertainty (of the event and its consequences), risk management (reducing risk to an acceptable level), and development of policy to balance finite resources with uncertainty and risk tolerance. When biological invasion and its risk are jointly examined, it is common that the consequences of invasion are not explicitly quantified, but understood to be sufficiently negative that it must be minimized to the extent possible. Risk analysis then becomes quantification of the probabilities of an introduction (event) and that the introduction leads to establishment, and the uncertainty of those probabilities. I describe a risk analysis framework for the Asian gypsy moth—a known invader—in its pathway. The framework uses the available information regarding the transportation route of the vector (ships), and a phenology model that estimates vector contamination (propagule size), the probability of introduction, and the probability of initial establishment given an introduction. Reducing propagule pressure is arguably the most important factor in reducing biological invasion; propagule pressure can be reduced by inspection and sanitation of the pathway vector (e.g., ships, trucks, humans) at the point(s) of departure and at the point of entry. I demonstrate how the risk analysis framework can be used to more efficiently target incoming ships for inspection and propagule pressure reduction.  相似文献   

17.
Wolbachia are very common, maternally transmitted endosymbionts of insects. They often spread by a mechanism termed cytoplasmic incompatibility (CI) that involves reduced egg hatch when Wolbachia-free ova are fertilized by sperm from Wolbachia-infected males. Because the progeny of Wolbachia-infected females generally do not suffer CI-induced mortality, infected females are often at a reproductive advantage in polymorphic populations. Deterministic models show that Wolbachia that impose no costs on their hosts and have perfect maternal transmission will spread from arbitrarily low frequencies (though initially very slowly); otherwise, there will be a threshold frequency below which Wolbachia frequencies decline to extinction and above which they increase to fixation or a high stable equilibrium. Stochastic theory was used to calculate the probability of fixation in populations of different size for arbitrary current frequencies of Wolbachia, with special attention paid to the case of spread after the arrival of a single infected female. Exact results are given based on a Moran process that assumes a specific demographic model, and approximate results are obtained using the more general Wright-Fisher theory. A new analytical approximation for the probability of fixation is derived, which performs well for small population sizes. The significance of stochastic effects in the natural spread of Wolbachia and their relevance to the use of Wolbachia as a drive mechanism in vector and pest management are discussed.  相似文献   

18.
Arthropods make up the largest group of invasive alien species (IAS) worldwide. Although invasion research has been biased towards alien plants and vertebrates, it has suggested potential mechanisms for the success of IAS and provided a theoretical framework for further investigation. Here we address key concepts from invasion biology that are essential to our understanding of the success of invasive alien arthropod predators and parasitoids including human intervention, environmental characteristics, propagule pressure, biological traits, and biological interactions. To gain a greater understanding of the factors most likely to influence the different stages of invasion (arrival, establishment, and spread) for alien arthropod predators and parasitoids, we use a comparative approach to compare and contrast the differential success of invasions by alien phytophagous and carnivorous arthropods. Insights gained from this comparison suggest that future research will require a multitrophic approach in order to enhance our understanding of invasions at higher trophic levels.  相似文献   

19.
Real-world uncertainties and data limitations make it difficult to predict how, when and where non-indigenous species (NIS) will spread. Typically only a small fraction of sites are sampled during only a few time intervals, such that we know neither the full spatial extent nor the true temporal progress of invasion. Yet, these unsampled locations might affect the invasion dynamics. We extend propagule pressure models to incorporate both human-mediated and natural fluvial dispersal vectors, and develop techniques to incorporate missing spatial and temporal data on invasions. We apply our model to Bythotrephes longimanus, a high-risk aquatic NIS, using a regional-scale 311-lake survey in a popular watershed in Ontario and extending our analysis to 1,300 unsampled lakes. Of 100 model runs with different random subsets of 50 sampled lakes reserved for validation, we were able to obtain an average area under the curve value of 0.89. Human-mediated dispersal accounted for 99.75% of the contribution of propagules to probability of establishment. Although the discovery rate is accelerating, our results suggest the annual rate of lake invasions is decelerating over time. Management efforts controlling recreational boating traffic out of the largest lakes in the system will be the most effective way of slowing the spread of B. longimanus in lakes within this system.  相似文献   

20.
Aim We investigated watershed‐scale abiotic environmental factors associated with population establishment of one of the ‘world’s 100 worst alien invaders’ on a temperate Atlantic island. Within the context of the conservation implications, we aimed to quantify (1) the early history and demographics (numbers and origins) of human‐mediated brown trout (Salmo trutta) introductions, (2) the current distribution of established populations, and (3) the watershed‐scale environmental factors that may resist or facilitate trout establishment. Location Island of Newfoundland, Canada. Methods We combined field sampling with historical and contemporary records from literature to assemble a presence–absence and physical habitat database for 312 watersheds on Newfoundland. Probability of watershed establishment was modelled with general additive ANCOVA models to control for nonlinear effects of propagule pressure (i.e. the distance to and number of invasion foci within a biologically relevant range) and model performance based on AIC. Results Between 1883 and 1906, 16 watersheds were introduced with brown trout from the Howietoun Hatchery, near Stirling, Scotland. Since that time, populations have established in 51 additional watersheds at an estimated rate of spread of 4 km per year. We did not detect any obvious abiotic barriers to resist trout establishment, but showed that for a given amount of propagule pressure that relatively large and productive watersheds were most likely to be established. Main conclusions Brown trout have successfully invaded and established populations in watersheds of Newfoundland and are currently slowly expanding on the island. Populations are more likely to establish in relatively large and productive watersheds, thereby supporting predictions of island biogeography theory. However, we suggest that all watersheds in Newfoundland are potentially susceptible to successful brown trout invasion and that abiotic factors alone are unlikely to act sufficiently as barriers to population establishment.  相似文献   

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