Global flows of insect transport and establishment: The role of biogeography,trade and regulations

Aim

Non-native species are part of almost every biological community worldwide, yet numbers of species establishments have an uneven global distribution. Asymmetrical exchanges of species between regions are likely influenced by a range of mechanisms, including propagule pressure, native species pools, environmental conditions and biosecurity. While the importance of different mechanisms is likely to vary among invasion stages, those occurring prior to establishment (transport and introduction) are difficult to account for. We used records of unintentional insect introductions to test (1) whether insects from some biogeographic regions are more likely to be successful invaders, (2) whether the intensity of trade flows between regions determines how many species are intercepted and how many successfully establish, and (3) whether the variables driving successful transport and successful establishment differ.

Location

Canada, mainland USA, Hawaii, Japan, Australia, New Zealand, Great Britain, South Korea, South Africa.

Methods

To disentangle processes occurring during the transport and establishment stages, we analysed border interceptions of 8199 insect species as a proxy for transported species flows, and lists of 2076 established non-native insect species in eight areas. We investigated the influence of biogeographic variables, socio-economic variables and biosecurity regulations on the size of species flows between regions.

Results

During transport, the largest species flows generally originated from the Nearctic, Panamanian and Neotropical regions. Insects native to 8 of 12 biogeographic regions were able to establish, with the largest flows of established species on average coming from the Western Palearctic, Neotropical and Australasian/Oceanian regions. Both the biogeographic region of origin and trade intensity significantly influenced the size of species flows between regions during transport and establishment. The transported species richness increased with Gross National Income in the source country, and decreased with geographic distance. More species were able to establish when introduced within their native biogeographic region.

Main Conclusions

Our results suggest that accounting for processes occurring prior to establishment is crucial for understanding invasion asymmetry in insects, and for quantifying regional biosecurity risks.     

Global compositional variation among native and non-native regional insect assemblages emphasizes the importance of pathways

Insects are among the world’s most ecologically and economically important invasive species. Here we assemble inventories of native and non-native species from 20 world regions and contrast relative numbers among these species assemblages. Multivariate ordination indicates that the distribution of species among insect orders is completely different between native and non-native assemblages. Some orders, such as the Psocoptera, Dictyoptera, Siphonaptera, Thysanoptera, and Hemiptera, are always over-represented in the non-native compared to native assemblages. Other orders, such as the Plecoptera, Trichoptera, Ephemeroptera, Odonata, Mecoptera and Microcoryphila, are consistently under-represented in non-native assemblages. These patterns most likely arise both as a result of variation among taxa in their association with invasion pathways responsible for transporting species among world regions, as well as variation in life-history traits that affect establishment potential. However, our results indicate that species compositions associated with invasiveness are fundamentally different from compositions related to insularity, indicating that colonization of islands selects for a different group of insect taxa than does selection for successful invaders. Native and non-native assemblage compositions were also related, to a lesser extent, to latitude of the region sampled. Together, these results illustrate the dominant role of invasion pathways in shaping the composition of non-native insect assemblages. They also emphasize the difference between natural background colonization of islands and anthropogenic colonization events, and imply that biological invasions are not a simple subset of a long-standing ecological process.     

Ecology of invasive mosquitoes: effects on resident species and on human health

Investigations of biological invasions focus on patterns and processes that are related to introduction, establishment, spread and impacts of introduced species. This review focuses on the ecological interactions operating during invasions by the most prominent group of insect vectors of disease, mosquitoes. First, we review characteristics of non-native mosquito species that have established viable populations, and those invasive species that have spread widely and had major impacts, testing whether biotic characteristics are associated with the transition from established non-native to invasive. Second, we review the roles of interspecific competition, apparent competition, predation, intraguild predation and climatic limitation as causes of impacts on residents or as barriers to invasion. We concentrate on the best-studied invasive mosquito, Aedes albopictus, evaluating the application of basic ecological theory to invasions by Aedes albopictus. We develop a model based on observations of Aedes albopictus for effects of resource competition and predation as barriers to invasion, evaluating which community and ecosystem characteristics favour invasion. Third, we evaluate the ways in which invasive mosquitoes have contributed to outbreaks of human and animal disease, considering specifically whether invasive mosquitoes create novel health threats, or modify disease transmission for existing pathogen-host systems.     

Native vegetation structure,landscape features and climate shape non-native plant richness and cover in New Zealand native shrublands

Aim

Studies investigating the determinants of plant invasions rarely examine multiple factors and often only focus on the role played by native plant species richness. By contrast, we explored how vegetation structure, landscape features and climate shape non-native plant invasions across New Zealand in mānuka and kānuka shrublands.

Location

New Zealand.

Method

We based our analysis on 247 permanent 20 × 20-m plots distributed across New Zealand surveyed between 2009 and 2014. We calculated native plant species richness and cumulative cover at ground, understorey and canopy tiers. We examined non-native species richness and mean species ground cover in relation to vegetation structure (native richness and cumulative cover), landscape features (proportion of adjacent anthropogenic land cover, distance to nearest road or river) and climate. We used generalized additive models (GAM) to assess which variables had greatest importance in determining non-native richness and mean ground cover and whether these variables had a similar effect on native species in the ground tier.

Results

A positive relationship between native and non-native plant species richness was not due to their similar responses to the variables examined in this study. Higher native canopy richness resulted in lower non-native richness and mean ground cover, whereas higher native ground richness was associated with higher native canopy richness. Non-native richness and mean ground cover increased with the proportion of adjacent anthropogenic land cover, whereas for native richness and mean ground cover, this relationship was negative. Non-native richness increased in drier areas, while native richness was more influenced by temperature.

Main Conclusions

Adjacent anthropogenic land cover seems to not only facilitate non-native species arrival by being a source of propagules but also aids their establishment as a result of fragmentation. Our results highlight the importance of examining both cover and richness in different vegetation tiers to better understand non-native plant invasions.     

Determinants for the successful establishment of exotic ants in New Zealand

Biological invasions can dramatically alter ecosystems. An ability to predict the establishment success for exotic species is important for biosecurity and conservation purposes. I examine the exotic New Zealand ant fauna for characteristics that predict or determine an exotic species’ ability to establish. Quarantine records show interceptions of 66 ant species: 17 of which have established, 43 have failed to establish, whereas nests of another six are periodically observed but have failed to establish permanently (called ‘ephemeral’ establishment). Mean temperature at the highest latitude and interception variables were the only factors significantly different between established, failed or ephemeral groups. Aspects of life history, such as competitive behaviour and morphology, were not different between groups. However, in a stepwise discriminant analysis, small size was a key factor influencing establishment success. Interception rate and climate were also secondarily important. The resulting classification table predicted establishment success with 71% accuracy. Because not all exotic species are represented in quarantine records, a further discriminant model is described without interception data. Though with less accuracy (65%) than the full model, it still correctly predicted the success or failure of four species not used in the previous analysis. Techniques for improving the prediction accuracy are discussed. Predicting which species will establish in a new area appears an achievable goal, which will be a valuable tool for conservation biology.     

Patterns of bird invasion are consistent with environmental filtering

Predicting invasion potential has global significance for managing ecosystems as well as important theoretical implications for understanding community assembly. Phylogenetic relationships of introduced species to the extant community may be predictive of establishment success because of the opposing forces of competition/shared enemies (which should limit invasions by close relatives) versus environmental filtering (which should allow invasions by close relatives). We examine here the association between establishment success of introduced birds and their phylogenetic relatedness to the extant avifauna within three highly invaded regions (Florida, New Zealand, and Hawaii). Published information on both successful and failed introductions, as well as native species, was compiled for all three regions. We created a phylogeny for each avifauna including all native and introduced bird species. From the estimated branch lengths on these phylogenies, we calculated multiple measurements of relatedness between each introduced species and the extant avifauna. We used generalized linear models to test for an association between relatedness and establishment success. We found that close relatedness to the extant avifauna was significantly associated with increased establishment success for exotic birds both at the regional (Florida, Hawaii, New Zealand) and sub‐regional (islands within Hawaii) levels. Our results suggest that habitat filtering may be more important than interspecific competition in avian communities assembled under high rates of anthropogenic species introductions. This work also supports the utility of community phylogenetic methods in the study of vertebrate invasions.     

Utility of the CLIMEX ‘match climates regional’ algorithm for pest risk analysis: an evaluation with non-native ants in New Zealand

Pest risk analysts frequently ask if the climate of a pest risk analysis area could be suitable for the establishment of an organism of concern. Species distribution models can help to answer this question, but constructing them is technically complex, time consuming, and uninformative for additional non-modelled species. A quicker more broadly applicable approach involves using environmental distance metrics, including climate matching algorithms such as the ‘match climates regional’ function of CLIMEX (CLIMEX-MCR), to generate indices of climatic similarity between different locations without reference to particular species. Several studies have shown that various environmental distance metrics can provide biologically meaningful results. However, the veracity of the CLIMEX-MCR algorithm remains unevaluated, despite its application in numerous published studies. We used CLIMEX-MCR and high resolution New Zealand climate data to measure climatic similarities between New Zealand and the rest of the world. We then tested the veracity of the climatic match estimates by evaluating if their predictions regarding the suitability of New Zealand’s climate for 43 non-native ant species corresponded with empirical observations of those species in New Zealand. Non-native ants that are, or were once, established outdoors in New Zealand had overseas distributions that were climatically well matched with New Zealand. In contrast, species that either are established only indoors in New Zealand, or were observed to temporarily nest outdoors then die in New Zealand, had overseas distributions that were poorly matched. Species that are frequently intercepted at New Zealand’s border, but are not established there, generally also had overseas distributions with low climatic similarities to New Zealand. We also measured climatic similarities between New Zealand’s 13 national parks and the rest of the world. The overseas distributions of the non-native ants showed poor climatic matches with New Zealand’s national parks, which was consistent with the absence of persistent outdoor non-native ant populations in those parks. Our results support the utility of CLIMEX-MCR algorithm for pest risk analysis.     

The importance of globalisation in driving the introduction and establishment of alien species in Europe

Understanding the role of globalisation in promoting introduction and establishment of alien species is an important step towards successful management of biological invasions. We aimed to quantify the taxon‐dependent association of globalisation with the introduction and establishment of alien species in Europe. The availability of the KOF index of globalisation that measures all economic, social and political dimensions of global connectivity enables a study of this factor. Based on an extensive database of alien species, we used model selections based on the Akaike information criterion and hierarchical partitioning to identify the importance of globalisation in predicting the number of all introduced species and established species of ten mainly terrestrial taxa in countries across Europe. The association of globalisation with alien species establishment varied depending on taxon type. While the gross domestic product (GDP) of countries was a strong predictor for all but one taxon, globalisation was also found to be an important predictor for three taxa including those of high (e.g. insects) and low mobility (e.g. magnoliophyta). Globalisation explained 3.1 to 22% independently, and 5.5 to 35% jointly with other variables, of among‐country variations in the number of established alien species. The effect of globalisation on the distribution of all introduced species is not substantially different from that on the established alien species. This study highlights how globalisation among habitat availability and environmental conditions can determine the patterns of alien species introduction and establishment across Europe. The results also emphasise the varying degree of importance between different taxa. Knowledge of the relative significance of various pathways with regard to different taxa is important for correctly focusing efforts to reduce the spread of these species.     

First record of the Australian guava moth Coscinoptycha improbana Meyrick (Lepidoptera: Carposinidae) in New Caledonia: Implication for quarantine and biosecurity surveys in insular territories

We recorded the Australian guava moth for the first time in New Caledonia. Given its biology and recent spread into New Zealand, this moth may be a pest risk for many fruit crops and native plant species if it is proved to have been introduced in New Caledonia. Indeed, this record challenges our capabilities to identify insect interceptions in the context of a high gap of taxonomic knowledge in New Caledonia. It also urges high endemism islands to implement early detection protocols to prevent establishment and spread of new invasive species.     

Non-native grass invasion alters native plant composition in experimental communities

Invasions of non-native species are considered to have significant impacts on native species, but few studies have quantified the direct effects of invasions on native community structure and composition. Many studies on the effects of invasions fail to distinguish between (1) differential responses of native and non-native species to environmental conditions, and (2) direct impacts of invasions on native communities. In particular, invasions may alter community assembly following disturbance and prevent recolonization of native species. To determine if invasions directly impact native communities, we established 32 experimental plots (27.5 m²) and seeded them with 12 native species. Then, we added seed of a non-native invasive grass (Microstegium vimineum) to half of the plots and compared native plant community responses between control and invaded plots. Invasion reduced native biomass by 46, 64, and 58%, respectively, over three growing seasons. After the second year of the experiment, invaded plots had 43% lower species richness and 38% lower diversity as calculated from the Shannon index. Nonmetric multidimensional scaling ordination showed a significant divergence in composition between invaded and control plots. Further, there was a strong negative relationship between invader and native plant biomass, signifying that native plants are more strongly suppressed in densely invaded areas. Our results show that a non-native invasive plant inhibits native species establishment and growth following disturbance and that native species do not gain competitive dominance after multiple growing seasons. Thus, plant invaders can alter the structure of native plant communities and reduce the success of restoration efforts.     

Life histories, dispersal, invasions, and global change: progress and prospects in New Zealand ecology, 1989?2029

We highlight three areas of significant progress in ecology since 1989 which are particularly relevant to New Zealand, and three major challenges for the next two decades. Progress: (1) The unusual life histories of New Zealand organisms, including extreme longevity and low reproductive rates, are now seen as efficient responses to the low-disturbance environment present before the arrival of large mammals, including humans. (2) Recent data show that long distance dispersal has been far more common than previously supposed, changing our image of New Zealand from a Gondwanan ark to the ?flypaper of the Pacific?. (3) Greatly improved techniques for pest control, and innovative species management, have stabilised numbers of many of the most charismatic of New Zealand?s threatened species. Problems: (1) Native species continue to decline, including many previously thought to be stable, and improved phylogenetics and new discoveries have added threatened species. (2) Despite increased emphasis on biosecurity, biological invasions are continuing, driven by increased trade and lags in naturalisation. (3) Conservation efforts risk being overwhelmed by the direct effects of increasing human population, resource use, invasions, and global climate change at a time when human food supplies and economies are coming under increasing pressure from environmental constraints. Conclusions: (1) We need improved ecological understanding and more management tools for invasive and threatened species, especially for species other than birds. (2) In these decades of rapid climate change and habitat conversion, there is an urgent need for more widespread and sustainable integration of native species into New Zealand?s rural and urban lowland landscapes.     

Fish Invasions in the World's River Systems: When Natural Processes Are Blurred by Human Activities

Because species invasions are a principal driver of the human-induced biodiversity crisis, the identification of the major determinants of global invasions is a prerequisite for adopting sound conservation policies. Three major hypotheses, which are not necessarily mutually exclusive, have been proposed to explain the establishment of non-native species: the “human activity” hypothesis, which argues that human activities facilitate the establishment of non-native species by disturbing natural landscapes and by increasing propagule pressure; the “biotic resistance” hypothesis, predicting that species-rich communities will readily impede the establishment of non-native species; and the “biotic acceptance” hypothesis, predicting that environmentally suitable habitats for native species are also suitable for non-native species. We tested these hypotheses and report here a global map of fish invasions (i.e., the number of non-native fish species established per river basin) using an original worldwide dataset of freshwater fish occurrences, environmental variables, and human activity indicators for 1,055 river basins covering more than 80% of Earth's surface. First, we identified six major invasion hotspots where non-native species represent more than a quarter of the total number of species. According to the World Conservation Union, these areas are also characterised by the highest proportion of threatened fish species. Second, we show that the human activity indicators account for most of the global variation in non-native species richness, which is highly consistent with the “human activity” hypothesis. In contrast, our results do not provide support for either the “biotic acceptance” or the “biotic resistance” hypothesis. We show that the biogeography of fish invasions matches the geography of human impact at the global scale, which means that natural processes are blurred by human activities in driving fish invasions in the world's river systems. In view of our findings, we fear massive invasions in developing countries with a growing economy as already experienced in developed countries. Anticipating such potential biodiversity threats should therefore be a priority.     

新西兰鸟类入侵成功的有关因素

了解外来入侵物种（alien invasive species,AIS)的生物学涉及纯生物学及应用生物学的问题,但是靠预先设计的实验来加深人们的了解显然是不可能的,然而我们可以研究现有的入侵,这也是一个很好的途径,本文利用新西兰历史上的一些记录,探讨了物种在新环境中成功定居的因素,令人惊奇的是,物种间的生物学差异对成功的定居几乎没有什么作用,相反,真正起作用的是一个物种被引入新环境的频率及数量。     

Fish Invasions in the World's River Systems: When Natural Processes Are Blurred by Human Activities

Because species invasions are a principal driver of the human-induced biodiversity crisis, the identification of the major determinants of global invasions is a prerequisite for adopting sound conservation policies. Three major hypotheses, which are not necessarily mutually exclusive, have been proposed to explain the establishment of non-native species: the “human activity” hypothesis, which argues that human activities facilitate the establishment of non-native species by disturbing natural landscapes and by increasing propagule pressure; the “biotic resistance” hypothesis, predicting that species-rich communities will readily impede the establishment of non-native species; and the “biotic acceptance” hypothesis, predicting that environmentally suitable habitats for native species are also suitable for non-native species. We tested these hypotheses and report here a global map of fish invasions (i.e., the number of non-native fish species established per river basin) using an original worldwide dataset of freshwater fish occurrences, environmental variables, and human activity indicators for 1,055 river basins covering more than 80% of Earth's surface. First, we identified six major invasion hotspots where non-native species represent more than a quarter of the total number of species. According to the World Conservation Union, these areas are also characterised by the highest proportion of threatened fish species. Second, we show that the human activity indicators account for most of the global variation in non-native species richness, which is highly consistent with the “human activity” hypothesis. In contrast, our results do not provide support for either the “biotic acceptance” or the “biotic resistance” hypothesis. We show that the biogeography of fish invasions matches the geography of human impact at the global scale, which means that natural processes are blurred by human activities in driving fish invasions in the world's river systems. In view of our findings, we fear massive invasions in developing countries with a growing economy as already experienced in developed countries. Anticipating such potential biodiversity threats should therefore be a priority.     

The Role of Tourism and Recreation in the Spread of Non-Native Species: A Systematic Review and Meta-Analysis

Managing the pathways by which non-native species are introduced and spread is considered the most effective way of preventing species invasions. Tourism and outdoor recreation involve the frequent congregation of people, vehicles and vessels from geographically diverse areas. They are therefore perceived to be major pathways for the movement of non-native species, and ones that will become increasingly important with the continued growth of these sectors. However, a global assessment of the relationship between tourism activities and the introduction of non-native species–particularly in freshwater and marine environments–is lacking. We conducted a systematic review and meta-analysis to determine the impact of tourism and outdoor recreation on non-native species in terrestrial, marine and freshwater environments. Our results provide quantitative evidence that the abundance and richness of non-native species are significantly higher in sites where tourist activities take place than in control sites. The pattern was consistent across terrestrial, freshwater and marine environments; across a variety of vectors (e.g. horses, hikers, yachts); and across a range of taxonomic groups. These results highlight the need for widespread biosecurity interventions to prevent the inadvertent introduction of invasive non-native species (INNS) as the tourism and outdoor recreation sectors grow.     

Ecology of forest insect invasions

Forests in virtually all regions of the world are being affected by invasions of non-native insects. We conducted an in-depth review of the traits of successful invasive forest insects and the ecological processes involved in insect invasions across the universal invasion phases (transport and arrival, establishment, spread and impacts). Most forest insect invasions are accidental consequences of international trade. The dominant invasion ‘pathways’ are live plant imports, shipment of solid wood packaging material, “hitchhiking” on inanimate objects, and intentional introductions of biological control agents. Invading insects exhibit a variety of life histories and include herbivores, detritivores, predators and parasitoids. Herbivores are considered the most damaging and include wood-borers, sap-feeders, foliage-feeders and seed eaters. Most non-native herbivorous forest insects apparently cause little noticeable damage but some species have profoundly altered the composition and ecological functioning of forests. In some cases, non-native herbivorous insects have virtually eliminated their hosts, resulting in major changes in forest composition and ecosystem processes. Invasive predators (e.g., wasps and ants) can have major effects on forest communities. Some parasitoids have caused the decline of native hosts. Key ecological factors during the successive invasion phases are illustrated. Escape from natural enemies explains some of the extreme impacts of forest herbivores but in other cases, severe impacts result from a lack of host defenses due to a lack of evolutionary exposure. Many aspects of forest insect invasions remain poorly understood including indirect impacts via apparent competition and facilitation of other invaders, which are often cryptic and not well studied.     

Introduction history of non-native freshwater fish in Okinawa-jima Island: ornamental aquarium fish pose the greatest risk for future invasions

We examined the introduction history of non-native freshwater fish in Okinawa-jima Island, Japan, to identify the pathway posing the greater risk for future invasions and to measure the strength of ornamental fish introductions as a vector of new invasions using all available records of freshwater fish introductions to date. At least 45 non-native freshwater fish have been introduced to Okinawa-jima Island. Of those, 22 species have successfully established in natural waters and, consequently, non-native fish species represent 76 % of the total freshwater fish fauna. The most important pathway was via the dumping of unwanted ornamental aquarium fish, which accounted for 71 % of the total introductions. The introduction of ornamental fish has occurred extensively in recent decades, and all fish introduced since the 1980s have been ornamental aquarium fish. The overall rate of establishment success was estimated at 51 %, although this is undoubtedly an overestimate because of unreported failed introductions. Establishment rates varied between stocked fish (100 %) and dumped ornamental fish (39 %), which may reflect the low propagule pressure of ornamental fish. From the observed trends in freshwater fish introductions, the dumping of ornamental fish into natural waters is expected to continue if no action is taken to reduce this most important pathway. We recommend regulations against the release of unwanted pet fish into natural waters, better programs for returning unwanted pet fish to stores or local zoos, educational programs for aquarists and retailers, and regulations against selling and trading ornamental fish that are judged to pose a high risk of causing economic or ecological harm.     

Biosecurity and Vector Behaviour: Evaluating the Potential Threat Posed by Anglers and Canoeists as Pathways for the Spread of Invasive Non-Native Species and Pathogens

Invasive non-native species (INNS) endanger native biodiversity and are a major economic problem. The management of pathways to prevent their introduction and establishment is a key target in the Convention on Biological Diversity''s Aichi biodiversity targets for 2020. Freshwater environments are particularly susceptible to invasions as they are exposed to multiple introduction pathways, including non-native fish stocking and the release of boat ballast water. Since many freshwater INNS and aquatic pathogens can survive for several days in damp environments, there is potential for transport between water catchments on the equipment used by recreational anglers and canoeists. To quantify this biosecurity risk, we conducted an online questionnaire with 960 anglers and 599 canoeists to investigate their locations of activity, equipment used, and how frequently equipment was cleaned and/or dried after use. Anglers were also asked about their use and disposal of live bait. Our results indicate that 64% of anglers and 78.5% of canoeists use their equipment/boat in more than one catchment within a fortnight, the survival time of many of the INNS and pathogens considered in this study and that 12% of anglers and 50% of canoeists do so without either cleaning or drying their kit between uses. Furthermore, 8% of anglers and 28% of canoeists had used their equipment overseas without cleaning or drying it after each use which could facilitate both the introduction and secondary spread of INNS in the UK. Our results provide a baseline against which to evaluate the effectiveness of future biosecurity awareness campaigns, and identify groups to target with biosecurity awareness information. Our results also indicate that the biosecurity practices of these groups must improve to reduce the likelihood of inadvertently spreading INNS and pathogens through these activities.     

Overestimation of establishment success of non-native birds in Hawaii and Britain

The tens rules states that 10 % of all introduced species establish and about 10 % of those species become invasive. Several studies have failed to support the tens rule. However, these studies are beset by a general weakness: many unsuccessful invasions are never reported, and without these data tests of the tens rules are inadequate. Here, using data on the establishment success of non-native birds in Hawaii and Britain and comparing these data with those from a previous study, we show that lack of information about failed species introductions, and the tendency to report species that have become invasive more than those that have not, result in an overestimate of the establishment success and invasion rates of non-native species.     

Diversity of non-native terrestrial arthropods on woody plants in Canada

A list of non-native phytophagous insects and mites on woody plants (trees, shrubs, vines) in Canada was compiled using information from literature and input from taxonomists. The 419 recorded species include Hemiptera (53% of species), Lepidoptera (22%), Coleoptera (13%) and Hymenoptera (9%). Almost all species originate from the Palearctic, especially Europe, reflecting historical trade patterns. About 41% of species were directly introduced to Canada from countries of origin, and the remainder spread from the United States of America (USA) after initial establishment there. Major ports on the east and west coasts, on Lake Erie and Lake Ontario are the main points of entry for exotic species directly introduced, and southern British Columbia (BC), Ontario (ON) and Quebec (QC) are the major points of entry for species spreading from the USA. Consequently, BC, ON, QC and Nova Scotia have the highest diversity of non-native species, and the prairie provinces and northern territories have the lowest. The extent of the distribution of individual species is related to length of time in Canada, number of introductions and dispersal abilities. Almost all native woody plant genera in Canada have been invaded by exotic phytophages. The large majority of phytophages occur on angiosperms. Woody plant genera with the largest distribution, highest species diversity and highest local abundances tend to host the greatest number of non-native species, including Picea, Pinus, Malus, Prunus, Salix, Betula, Quercus, Pyrus and Populus. The arrival rate of species in Canada increased from the late nineteenth century until about 1960, and declined rapidly thereafter. Quarantine legislation enacted in the USA in 1912 and in Canada in 1976 seems to have reduced the rate of insect invasion. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.