Potential establishment of alien-invasive forest insect species in the United States: where and how many?

International trade is widely acknowledged as a conduit for movement of invasive species, but few studies have directly quantified the invasion risk confronting individual locations of interest. This study presents estimates of the likelihood of successful entry for alien forest insect species at more than 3,000 urban areas in the contiguous United States (US). To develop these location-specific estimates, we first utilized historical merchandise imports and insect incursions data to estimate an annual US rate of alien insect species establishment. Next, we used historical pest interception data to calculate the proportion of all insects arriving at US ports of entry that are associated with forest hosts. We then combined these results to estimate a nationwide establishment rate specifically for alien forest insects. Finally, we employed international and domestic commodity flow networks to allocate this nationwide rate to individual US urban areas. For 2010, we estimated the nationwide rate as 1.89 new alien forest insect species per year. While the establishment rates observed at most urban areas were low (<0.005 new species/year), for a few select areas the rates predict new alien forest insect species establishments every 5–15 years. This national-scale assessment provides a realistic depiction of human-assisted establishment potential in the US as well as functional inputs for quantitative models of invasion. Overall, these analyses support broad-scale biosecurity and management strategies.     

Impacts of Alien Invasive Species on Freshwater Fauna at Risk in Canada

Freshwater aquatic organisms in North America are disproportionately imperilled when compared to their terrestrial counterparts due to widespread habitat alteration, pollution, overexploitation and the introduction of alien species. In this review, we examine the threat factors contributing to the endangerment of freshwater fishes and molluscs in Canada and further examine the nature of alien invasive species introductions affecting aquatic species at risk. Habitat loss and degradation is the predominant threat factor for Canadian freshwater fishes and molluscs that are listed as Extinct, Extirpated, Endangered and Threatened. Alien invasive species are the second most prevalent threat for fishes, affecting 26 of 41 listed species. Alien invasive species are a threat in most parts of Canada where listed fishes are found. Most (65%) of the alien invasive species affecting listed fishes are the result of intentional introductions related to sport fishing, and the majority of these introductions are unauthorized. Fifteen fishes and two plant species are cited as alien invasive species that impact listed fishes with brown bullhead (Ameiurus nebulosus) and pumpkinseed (Lepomis gibbosus) being the most prevalent. Alien species are a threat to 6 of 11 listed mollusc species. All six species are threatened by the alien zebra mussel (Dreissena polymorpha) in the Great Lakes basin. An erratum to this article is available at .     

Using a Network Model to Assess Risk of Forest Pest Spread via Recreational Travel

Long-distance dispersal pathways, which frequently relate to human activities, facilitate the spread of alien species. One pathway of concern in North America is the possible spread of forest pests in firewood carried by visitors to campgrounds or recreational facilities. We present a network model depicting the movement of campers and, by extension, potentially infested firewood. We constructed the model from US National Recreation Reservation Service data documenting more than seven million visitor reservations (including visitors from Canada) at campgrounds nationwide. This bi-directional model can be used to identify likely origin and destination locations for a camper-transported pest. To support broad-scale decision making, we used the model to generate summary maps for 48 US states and seven Canadian provinces that depict the most likely origins of campers traveling from outside the target state or province. The maps generally showed one of two basic spatial patterns of out-of-state (or out-of-province) origin risk. In the eastern United States, the riskiest out-of-state origin locations were usually found in a localized region restricted to portions of adjacent states. In the western United States, the riskiest out-of-state origin locations were typically associated with major urban areas located far from the state of interest. A few states and the Canadian provinces showed characteristics of both patterns. These model outputs can guide deployment of resources for surveillance, firewood inspections, or other activities. Significantly, the contrasting map patterns indicate that no single response strategy is appropriate for all states and provinces. If most out-of-state campers are traveling from distant areas, it may be effective to deploy resources at key points along major roads (e.g., interstate highways), since these locations could effectively represent bottlenecks of camper movement. If most campers are from nearby areas, they may have many feasible travel routes, so a more widely distributed deployment may be necessary.     

Temperature explains variable spread rates of the invasive woodwasp Sirex noctilio in the Southern Hemisphere

The frequency of introductions of non-indigenous forest insects into new habitats is increasing worldwide, often with profoundly adverse consequences on natural and plantation forest ecosystems. Understanding rates and patterns of spread of invasive forest insects is important for predicting when and where these species will expand their geographical range, with the potential to improve mitigation strategies. The woodwasp Sirex noctilio is a damaging invasive forest insect that kills numerous species of Pinus. Despite encountering highly variable eco-climatic conditions, S. noctilio has arrived and established in exotic pine forest production areas throughout the Southern Hemisphere. In this study, we compiled historical records of S. noctilio invasion to compare spread rates among eight contrasting eco-climatic regions in the Southern Hemisphere and to explore how spread rate is predicted by landscape variation in climate, habitat characteristics and anthropogenic effects. Spread rates for S. noctilio varied considerably among the invaded regions, ranging from 12 to 82 km per year. Among regions, spread rates of S. noctilio increased with increasing mean annual temperature and isothermality. We hypothesize that temperature may directly or indirectly influence S. noctilio population growth and dispersal, thereby influencing spread rates.     

Congener diversity,topographic heterogeneity and human‐assisted dispersal predict spread rates of alien herpetofauna at a global scale

Understanding the factors that determine rates of range expansion is not only crucial for developing risk assessment schemes and management strategies for invasive species, but also provides important insight into the ability of species to disperse in response to climate change. However, there is little knowledge on why some invasions spread faster than others at large spatiotemporal scales. Here, we examine the effects of human activities, species traits and characteristics of the invaded range on spread rates using a global sample of alien reptile and amphibian introductions. We show that spread rates vary remarkably among invaded locations within a species, and differ across biogeographical realms. Spread rates are positively related to the richness of native congeneric species and human‐assisted dispersal in the invaded range but are negatively correlated with topographic heterogeneity. Our findings highlight the importance of environmental characteristics and human‐assisted dispersal in developing robust frameworks for predicting species' range shifts.     

Exotic vascular plant invasiveness and forest invasibility in urban boreal forest types

The riverine forests of the northern city of Edmonton, Alberta, Canada display strong resilience to disturbance and are similar in species composition to southern boreal mixedwood forest types. This study addressed questions such as, how easily do exotic species become established in urban boreal forests (species invasiveness) and do urban boreal forest structural characteristics such as, native species richness, abundance, and vertical vegetation layers, confer resistance to exotic species establishment and spread (community invasibility)? Eighty-four forest stands were sampled and species composition and mean percent cover analyzed using ordination methods. Results showed that exotic tree/shrub types were of the most concern for invasion to urban boreal forests and that exotic species type, native habitat and propagule supply may be good indicators of invasive potential. Native forest structure appeared to confer a level of resistance to exotic species and medium to high disturbance intensity was associated with exotic species growth and spread without a corresponding loss in native species richness. Results provided large-scale evidence that diverse communities are less vulnerable to exotic species invasion, and that intermediate disturbance intensity supports species coexistence. From a management perspective, the retention of native species and native forest structure in urban forests is favored to minimize the impact of exotic species introductions, protect natural succession patterns, and minimize the spread of exotic species.     

Applying a spread model to identify the entry points from which the pine wood nematode, the vector of pine wilt disease, would spread most rapidly across Europe

Pine wilt disease, which can rapidly kill pines, is caused by the pine wood nematode, Bursaphelenchus xylophilus. It is expanding its range in many countries in Asia and measures are being taken at the EU level to prevent its spread from Portugal. Due to the threat to European forests, it is important to prevent additional introductions and target surveillance to the points of entry that pose the greatest risk. In this study, we present a model to identify the European ports from which the nematode can spread most rapidly across Europe. This model describes: (1) the potential spread of the pine wood nematode based on short-distance spread (the active flight of the vector beetles) and long-distance spread (primarily due to human-mediated transportation), and (2) the development of pine wilt disease based on climate suitability and the potential spread of the nematode. Separate introductions at 200 European ports were simulated under various climate change scenarios. We found that the pine wood nematode could invade 19–60% of the study area (30°00 N–72°00 N, 25°00 W–40°00 E) by 2030, with the highest spread from ports located in Eastern and Northern Europe. Based on climate change scenarios, the disease could affect 8–34% of the study area by 2030, with the highest spread from ports located in South-Eastern Europe. This study illustrates how a spread model can be used to determine the critical points of entry for invasive species, so that surveillance can be targeted more accurately and control measures prioritised.     

Arrival rate of nonindigenous insect species into the United States through foreign trade

Introductions of invasive nonindigenous species, and the ensuing negative ecological and economic consequences, have increased with expanding global trade. Quantifying the influx of nonindigenous plant pest species through foreign trade is required for national and international risk assessments, monitoring and conservation efforts, and evaluation of ecological factors that affect invasion success. Here we use statistically robust data collected at US ports of entry and border crossings to estimate arrival rates of nonindigenous insect species via four cargo pathways and to evaluate the effectiveness of current efforts to monitor arrival of nonindigenous insect species. Interception rates were highest in refrigerated maritime cargo where a new insect species was intercepted on average every 54 inspections. Projected estimates of insect species richness stabilized only for non-refrigerated maritime cargo and US–Mexico border cargo, where inspectors likely detected 19–2% and 30–50% of the species being transported through these respective pathways. Conservative estimates of establishment suggest that 42 insect species may have become established through these four pathways between 1997 and 2001.     

The cobblers stick to their lasts: pollinators prefer native over alien plant species in a multi-species experiment

The majority of plant species rely, at least partly, on animals for pollination. Our knowledge on whether pollinator visitation differs between native and alien plant species, and between invasive and non-invasive alien species is still limited. Additionally, because numerous invasive plant species are escapees from horticulture, the transition from human-assisted occurrence in urbanized habitats to unassisted persistence and spread in (semi-)natural habitats requires study. To address whether pollinator visitation differs between native, invasive alien and non-invasive alien species, we did pollinator observations for a total of 17 plant species representing five plant families. To test whether pollinator visitation to the three groups of species during the initial stage of invasion depends on habitat type, we did the study in three urbanized habitats and three semi-natural grasslands, using single potted plants. Native plants had more but smaller flower units than alien plants, and invasive alien plants had more but smaller flowers than non-invasive alien plants. After accounting for these differences in floral display, pollinator visitation was higher for native than for alien plant species, but did not differ between invasive and non-invasive alien plant species. Pollinator visitation was on average higher in semi-natural than in urbanized habitats, irrespective of origin or status of the plant species. This might suggest that once an alien species has managed to escape from urbanized into more natural habitats, pollinator limitation will not be a major barrier to establishment and invasion.     

Tracking the invasive history of the green alga Codium fragile ssp. tomentosoides

The spread of nonindigenous species into new habitats is having a drastic effect on natural ecosystems and represents an increasing threat to global biodiversity. In the marine environment, where data on the movement of invasive species is scarce, the spread of alien seaweeds represents a particular problem. We have employed a combination of plastid microsatellite markers and DNA sequence data from three regions of the plastid genome to trace the invasive history of the green alga Codium fragile ssp. tomentosoides. Extremely low levels of genetic variation were detected, with only four haplotypes present in the species' native range in Japan and only two of these found in introduced populations. These invasive populations displayed a high level of geographical structuring of haplotypes, with one haplotype localized in the Mediterranean and the other found in Northwest Atlantic, northern European and South Pacific populations. Consequently, we postulate that there have been at least two separate introductions of C. fragile ssp. tomentosoides from its native range in the North Pacific.     

城市绿地外来物种风险分析体系构建及其在上海世博会管理中的应用

外来物种风险分析是防止生物入侵的有效手段之一。本文按照风险识别、风险评估和风险管理3个阶段,构建了城市绿地外来物种风险分析体系。文中提出了4个层次、26个指标构成风险评估指标体系,此体系囊括了城市绿地外来物种的传入、定殖、扩散、危害等入侵风险形成的基本要素,并规范了风险指数的计算方法。以2010年上海世博会引进日本景观苗木可能携带的外来物种为对象,对该体系在生产实践中进行了应用。结果表明:高风险物种共7种,涉及害虫4种、植物病原微生物2种、植物线虫1种;中风险物种共10种,涉及害虫3种、植物病原微生物4种、植物线虫3种;低、极低风险物种各1种。根据风险分析结果,对以上物种提出了有针对性的风险管理措施。实践表明,该风险分析体系实用性强,在上海世博会植物引种过程中为防止外来物种入侵起到了较好的预警效果,为管理者提供了有价值的决策参考,有力保障了上海世博会期间的生态安全。     

The exotic mammals of Argentina

Exotic mammals in South America represent about 20% of world mammal introductions. The aim of our paper is to provide a global assessment of the exotic mammals of Argentina, their pathways, impacts, and a synthesis of their attributes as potential invasive species. We reviewed and compiled data from a diversity of sources and databases on alien mammals occurring in feral state exclusively. We recorded 18 species of exotic mammals for Argentina. The majority of introductions occurred between the 18th and 19th centuries and their ports of entry were located in temperate ecosystems, between 34° and 55° SL. Most of their entry pathways were associated with human activities (e.g. sport hunting, food and fur industry). The exotic mammals occupy ecoregions similar to their original distributions, but most of them have experienced a range expansion to novel habitats. The fauna of exotic mammals of Argentina represents a good opportunity to understand the dynamics of the invasion process as they represent a diversity of ecological groups and environmental contexts.     

The status quo of invasive alien insect species and plant quarantine in Korea

The recent increase in agricultural commodities from abroad and travelers due to the economic growth and globalization in Korea has resulted in an increase in invasive alien insect species establishing in Korea. When “alien” is defined as species introduced after the beginning of the Greater Korean Empire Era (1897), 171 insect species are considered invasive alien species. On average, 0.85 alien insect species have become established each year for the last 40 years, and 76.6% of the alien insect species are economic agricultural pests. The annual invasion rate and the pest interceptions from imported cut flowers, planting material, and vegetables have been increasing rapidly. Traveler's baggage is an important pathway for fruit flies. Most of the alien pests that were first found in Japan were confirmed in Korea at least 3 years later until the 1990s, but the pattern has been reversed since the 2000s. Thus, continuous information sharing with neighboring countries such as Japan and China is needed to develop a system for early detection and prompt action against invading insect species. Thus, we introduce and summarize the present status of invasive alien insect species in Korea.     

Optimization and validation of a cost‐effective protocol for biosurveillance of invasive alien species

Environmental DNA (eDNA) metabarcoding has revolutionized biodiversity monitoring and invasive pest biosurveillance programs. The introduction of insect pests considered invasive alien species (IAS) into a non‐native range poses a threat to native plant health. The early detection of IAS can allow for prompt actions by regulating authorities, thereby mitigating their impacts. In the present study, we optimized and validated a fast and cost‐effective eDNA metabarcoding protocol for biosurveillance of IAS and characterization of insect and microorganism diversity. Forty‐eight traps were placed, following the CFIA''s annual forest insect trapping survey, at four locations in southern Ontario that are high risk for forest IAS. We collected insects and eDNA samples using Lindgren funnel traps that contained a saturated salt (NaCl) solution in the collection jar. Using cytochrome c oxidase I (COI) as a molecular marker, a modified Illumina protocol effectively identified 2,535 Barcode Index Numbers (BINs). BINs were distributed among 57 Orders and 304 Families, with the vast majority being arthropods. Two IAS (Agrilus planipennis and Lymantria dispar) are regulated by the Canadian Food Inspection Agency (CFIA) as plant health pests, are known to occur in the study area, and were identified through eDNA in collected traps. Similarly, using 16S ribosomal RNA and nuclear ribosomal internal transcribed spacer (ITS), five bacterial and three fungal genera, which contain species of regulatory concern across several Canadian jurisdictions, were recovered from all sampling locations. Our study results reaffirm the effectiveness and importance of integrating eDNA metabarcoding as part of identification protocols in biosurveillance programs.     

Gypsy moth invasion in North America: A simulation study of the spatial pattern and the rate of spread

Gypsy moth is regarded as one of the top most harmful invasive species. Its invasion in the northeastern US has led to widespread forest defoliation, wildlife disruption and even a change in biogeochemical conditions over the area of 10⁶ km². Spread of gypsy moth has a few distinct features such as a patchy spatial distribution of the gypsy moth population, which is largely uncorrelated to the environmental heterogeneity, and a high variability (almost over an order of magnitude) in the spread rates. These features are usually explained by human-assisted dispersal, e.g. when masses of gypsy moth eggs are inadvertently transported by cars and vehicles. This theory, however, somewhat disagrees with the existence of the strong Allee effect that tends to wipe out small new colonies. In this paper, we suggest an alternative explanation that the patchy structure can result from the interplay between two natural factors such as wind dispersal and viral infection. In order to check this hypothesis, we describe the gypsy moth spread with a diffusive SI model and study its properties by means of extensive computer simulations. Interestingly, in a certain parameter range our model shows formation of spatial patterns that are qualitatively similar to those observed in the field. To find out the relevant parameter range, we make a careful review of available literature sources. For biologically meaningful parameter values, we then show that the rates of gypsy moth spread predicted by our model are in good agreement with the lower band of the rates observed in nature.     

Differences in endemic insect assemblages among vegetation types on a small island of the oceanic Ogasawara Islands

Natural vegetation is often replaced by invasive alien plants on isolated oceanic islands. To determine how invasive alien plants affect insect diversity, we compared flying insects captured using Malaise traps among different vegetation types on a small island (Nishijima; 0.49 km²) in the oceanic Ogasawara (Bonin) Islands in the north‐western Pacific. The numbers of individuals and species, and the species composition of pollinators (bees), predators (wasps) and wood borers (cerambycid, mordellid and elaterid beetles) were compared among three vegetation types: Casuarina equisetifolia (an invasive alien tree) forest, natural forest and natural grassland (forest edge), during two seasons (June and October–November 2005). In traps, 80.0, 66.7, 87.5, 85.7 and 100.0% of bee, wasp, cerambycid, mordellid and elaterid beetle species, respectively, were endemic to the Ogasawara Islands. Grassland had the highest wasp and bee species richness, whereas natural forest had the highest species richness of wood‐boring beetles. The C. equisetifolia forest had the poorest species richness for most insect groups (except mordellid beetles). More individuals of most insect groups (except bees) were captured in June than in October–November. More individual bees and wasps were captured in grassland than in forests, whereas more individual mordellid and elaterid beetles were captured in forests than in grassland. The number of cerambycid individuals did not differ among vegetation types. Redundancy analysis suggested that most insect species preferred natural forest or grassland to alien forest. Therefore, further invasion of natural grassland and forest by the alien tree C. equisetifolia may negatively affect the endemic insect fauna of Nishijima.     

2008-2016年间广州市外来入侵植物的变化分析

为了解2008-2016年广州市外来入侵植物的变化,采用文献查阅和实地考察相结合的方式,近期对广州市外来入侵植物进行了调查。结果表明,2016年广州市外来入侵植物共有38科112属144种,比2008年分别增加了40.7%、89.8%和97.3%。新增加的71种植物主要有豆科（Fabaceae） 14种、菊科（Asteraceae） 13种、旋花科（Convolvulaceae） 6种、禾本科（Poaceae） 6种和茜草科（Rubiaceae） 5种。外来入侵植物主要来源于美洲（共104种）,多为陆生草本植物。广州约一半以上的外来植物是通过人为有意引进,进而形成入侵的。恶意和严重入侵的植物种类约占入侵物种总数的40%,表明广州的城市森林生态系统已经面临着严重威胁。因此,需加强对广州市外来植物入侵动态的监测,提高对外来陆生草本植物引入危害风险的评估,并开展对外来入侵植物资源开发和利用的研究,以构建稳定的生态安全格局,保障绿色生态城市建设。     

Modelling the Arrival of Invasive Organisms via the International Marine Shipping Network: A Khapra Beetle Study

Species can sometimes spread significant distances beyond their natural dispersal ability by anthropogenic means. International shipping routes and the transport of shipping containers, in particular are a commonly recognised pathway for the introduction of invasive species. Species can gain access to a shipping container and remain inside, hidden and undetected for long periods. Currently, government biosecurity agencies charged with intercepting and removing these invasive species when they arrive to a county's border only assess the most immediate point of loading in evaluating a shipping container's risk profile. However, an invasive species could have infested a container previous to this point and travelled undetected before arriving at the border. To assess arrival risk for an invasive species requires analysing the international shipping network in order to identify the most likely source countries and the domestic ports of entry where the species is likely to arrive. We analysed an international shipping network and generated pathway simulations using a first-order Markov chain model to identify possible source ports and countries for the arrival of Khapra beetle (Trogoderma granarium) to Australia. We found Kaohsiung (Taiwan) and Busan (Republic of Korea) to be the most likely sources for Khapra beetle arrival, while the port of Melbourne was the most likely point of entry to Australia. Sensitivity analysis revealed significant stability in the rankings of foreign and Australian ports. This methodology provides a reliable modelling tool to identify and rank possible sources for an invasive species that could arrive at some time in the future. Such model outputs can be used by biosecurity agencies concerned with inspecting incoming shipping containers and wishing to optimise their inspection protocols.     

Climate change and biological invasions: evidence,expectations, and response options

A changing climate may directly or indirectly influence biological invasions by altering the likelihood of introduction or establishment, as well as modifying the geographic range, environmental impacts, economic costs or management of alien species. A comprehensive assessment of empirical and theoretical evidence identified how each of these processes is likely to be shaped by climate change for alien plants, animals and pathogens in terrestrial, freshwater and marine environments of Great Britain. The strongest contemporary evidence for the potential role of climate change in the establishment of new alien species is for terrestrial arthropods, as a result of their ectothermic physiology, often high dispersal rate and their strong association with trade as well as commensal relationships with human environments. By contrast, there is little empirical support for higher temperatures increasing the rate of alien plant establishment due to the stronger effects of residence time and propagule pressure. The magnitude of any direct climate effect on the number of new alien species will be small relative to human‐assisted introductions driven by socioeconomic factors. Casual alien species (sleepers) whose population persistence is limited by climate are expected to exhibit greater rates of establishment under climate change assuming that propagule pressure remains at least at current levels. Surveillance and management targeting sleeper pests and diseases may be the most cost‐effective option to reduce future impacts under climate change. Most established alien species will increase their distribution range in Great Britain over the next century. However, such range increases are very likely be the result of natural expansion of populations that have yet to reach equilibrium with their environment, rather than a direct consequence of climate change. To assess the potential realised range of alien species will require a spatially explicit approach that not only integrates bioclimatic suitability and population‐level demographic rates but also simulation of landscape‐level processes (e.g. dispersal, land‐use change, host/habitat distribution, non‐climatic edaphic constraints). In terms of invasive alien species that have known economic or biodiversity impacts, the taxa that are likely to be the most responsive are plant pathogens and insect pests of agricultural crops. However, the extent to which climate adaptation strategies lead to new crops, altered rotations, and different farming practices (e.g. irrigation, fertilization) will all shape the potential agricultural impacts of alien species. The greatest uncertainty in the effects of climate change on biological invasions exists with identifying the future character of new species introductions and predicting ecosystem impacts. Two complementary strategies may work under these conditions of high uncertainty: (i) prioritise ecosystems in terms of their perceived vulnerability to climate change and prevent ingress or expansion of alien species therein that may exacerbate problems; (ii) target those ecosystem already threatened by alien species and implement management to prevent the situation deteriorating under climate change.