Importance of greater interdisciplinarity and geographic scope when tackling the driving forces behind biological invasions

Invasive non-native species are important drivers of ecosystem change, yet the driving forces of biological invasions themselves are poorly understood. Such information is essential to ensure policies focus on the most relevant drivers, and that future scenarios capture the full range of potential outcomes for invasive non-native species. I carried out a bibliometric analysis of articles published from 2000 to 2020 that address either invasive non-native species or biodiversity and ecosystem services and that also mention 1 or more drivers of ecosystem change. I examined 5 indirect drivers (demographic, economic, governance, sociocultural, and technological) and 6 direct drivers (climate change, invasive non-native species, land-use or sea-use change, natural hazards, pollution, and resource extraction). Using the Web of Science core collection of citation indexes, I undertook searches of article titles and keywords and retrieved 27,462 articles addressing invasive non-native species and 110,087 articles dealing with biodiversity or ecosystem services. Most research to date on biological invasions as well as on biodiversity and ecosystem services has focused on anthropogenic direct drivers of ecosystem change rather than indirect drivers. Yet currently, less than 18% of articles addressing biological invasions examined drivers of ecosystem change, a similar level to that found over 20 years ago for biodiversity or ecosystem services. Knowledge of the drivers of biological invasions is limited, emphasizes tractable drivers over those that require an interdisciplinary approach, and is biased toward developed economies. Drivers generally deemed important for biological invasions, such as governance and resource extraction, accounted for less than 2% of research effort. The absence of a systematic understanding of the forces that drive invasive non-native species and how they interact means that attempts to mitigate or forecast biological invasions are likely to fail. To address biological invasions requires a much better orientation of national and international research on drivers in relation to both their actual importance as well as their policy relevance.     

An Assessment of Invasion Risk from Assisted Migration

Abstract:  To reduce the risk of extinction due to climate change, some ecologists have suggested human-aided translocation of species, or assisted migration (AM), to areas where climate is projected to become suitable. Such intentional movement, however, may create new invasive species if successful introductions grow out of control and cause ecologic or economic damage. We assessed this risk by surveying invasive species in the United States and categorizing invaders based on origin. Because AM will involve moving species on a regional scale within continents (i.e., range shifts), we used invasive species with an intracontinental origin as a proxy for species that would be moved through AM. We then determined whether intracontinental invasions were more prevalent or harmful than intercontinental invasions. Intracontinental invasions occurred far less frequently than invasions from other continents, but they were just as likely to have had severe effects. Fish and crustaceans pose a particularly high threat of intracontinental invasion. We conclude that the risk of AM to create novel invasive species is small, but assisted species that do become invasive could have large effects. Past experience with species reintroductions may help inform policy regarding AM.     

Niche differentiation and fine-scale projections for Argentine ants based on remotely sensed data.

Modeling ecological niches of species is a promising approach for predicting the geographic potential of invasive species in new environments. Argentine ants (Linepithema humile) rank among the most successful invasive species: native to South America, they have invaded broad areas worldwide. Despite their widespread success, little is known about what makes an area susceptible--or not--to invasion. Here, we use a genetic algorithm approach to ecological niche modeling based on high-resolution remote-sensing data to examine the roles of niche similarity and difference in predicting invasions by this species. Our comparisons support a picture of general conservatism of the species' ecological characteristics, in spite of distinct geographic and community contexts.     

外来入侵植物在中国的分布及入侵能力研究进展

以近十几年来备受关注的外来植物入侵为背景,综述关于入侵植物在我国分布和入侵能力及其相关因素的研究进展,并探讨了未来研究需要加强的几个方面。不同起源地的入侵植物在我国分布区域不一样,在控制了起源地的作用后,入侵植物主要分布在经济较发达、人为活动较多的省份或地区（大尺度）,以及干扰较严重的生境（小尺度）。入侵能力强的多年生植物常有强大的无性繁殖能力、高光合速率和资源利用效率、强化感作用、以及适应异质生境的能力;起源于美洲的入侵能力强的1年生植物常能产生大量的易传播的种子、可自交亲和、强化感作用等;起源于欧亚大陆的入侵能力强的1年生植物常与作物非常相似,在农田中危害严重。未来需要研究干扰促进入侵的机制、化感作用与野外实际情况相结合研究、化感物质通过改变土壤微生态环境间接促进入侵的机制研究、1年生和多年生外来植物的入侵能力与其物种特性之间的关系、以及加强对弱入侵性外来植物的预警研究。这些研究不仅能使我们深入理解外来植物发生入侵和危害的规律,也可为人侵植物的防控和管理提供科学依据。     

E‐commerce trade in invasive plants

Biological invasions are a major concern in conservation, especially because global transport of species is still increasing rapidly. Conservationists hope to anticipate and thus prevent future invasions by identifying and regulating potentially invasive species through species risk assessments and international trade regulations. Among many introduction pathways of non‐native species, horticulture is a particularly important driver of plant invasions. In recent decades, the horticultural industry expanded globally and changed structurally through the emergence of new distribution channels, including internet trade (e‐commerce). Using an automated search algorithm, we surveyed, on a daily basis, e‐commerce trade on 10 major online auction sites (including eBay) of approximately three‐fifths of the world's spermatophyte flora. Many recognized invasive plant species (>500 species) (i.e., species associated with ecological or socio‐economic problems) were traded daily worldwide on the internet. A markedly higher proportion of invasive than non‐invasive species were available online. Typically, for a particular plant family, 30–80% of recognized invasive species were detected on an auction site, but only a few percentages of all species in the plant family were detected on a site. Families that were more traded had a higher proportion of invasive species than families that were less traded. For woody species, there was a significant positive relationship between the number of regions where a species was sold and the number of regions where it was invasive. Our results indicate that biosecurity is not effectively regulating online plant trade. In the future, automated monitoring of e‐commerce may help prevent the spread of invasive species, provide information on emerging trade connectivity across national borders, and be used in horizon scanning exercises for early detection of new species and their geographic source areas in international trade.     

Relatedness and environment affect traits associated with invasive and noninvasive introduced Commelinaceae.

Understanding the traits of invasive species may improve the ability to predict, prevent, and manage invasions. I compared morphological and performance traits of five congeneric pairs of invasive and noninvasive Commelinaceae across a factorial experiment using a range of water and nutrient availabilities. Invasive species had greater fecundity and vegetative reproduction than their noninvasive relatives. The invasive species also had higher relative growth rates, greater specific leaf area, and more plastic root-to-shoot ratios than noninvasive species. However, whether a trait was associated with invasiveness often depended on both environment and relatedness. Invasives had greater sexual and vegetative reproduction, higher specific leaf area, and greater relative growth rates than noninvasive congeners, but only in some environments. Differences between invasive and noninvasive taxa were greatest at high nutrient availabilities. These results suggest that studies of invasive species' traits must incorporate information on conditions under which the trait was measured. In addition, incorporating information on relatedness improved our ability to detect associations between species traits, such as specific leaf area and relative growth rate, and invasiveness, suggesting that such information may be required for a complete understanding of what makes a species invasive.     

Law and Large Carnivore Conservation in the Rocky Mountains of the U.S. and Canada

The law governing large carnivores in the western U.S. and western Canada abounds in jurisdictional complexity. In the U.S., different federal and state laws govern large carnivore conservation efforts; species listed under the Endangered Species Act are generally protected, whereas those subject to state regulation can be hunted, trapped, or otherwise taken. Neither federal nor state environmental or land management laws specifically protect large carnivores, though these laws can be used to protect habitat. A similar situation prevails in Canada. Canadian federal law does not address large carnivore conservation, although the national parks provide some secure habitat. Provincial laws vary widely; none of these laws specifically protect large carnivores, but some provisions can be invoked to protect habitat. Although the two nations have not entered any bilateral treaties to protect large carnivores, several species receive limited protection under multilateral treaty obligations. Despite these jurisdictional complexities, the existing legal framework can be built upon to promote large carnivore conservation efforts, primarily through a legally protected reserve system. Whether the political will exists to utilize fully the available legal authorities remains to be seen.     

Five Potential Consequences of Climate Change for Invasive Species

Abstract:  Scientific and societal unknowns make it difficult to predict how global environmental changes such as climate change and biological invasions will affect ecological systems. In the long term, these changes may have interacting effects and compound the uncertainty associated with each individual driver. Nonetheless, invasive species are likely to respond in ways that should be qualitatively predictable, and some of these responses will be distinct from those of native counterparts. We used the stages of invasion known as the "invasion pathway" to identify 5 nonexclusive consequences of climate change for invasive species: (1) altered transport and introduction mechanisms, (2) establishment of new invasive species, (3) altered impact of existing invasive species, (4) altered distribution of existing invasive species, and (5) altered effectiveness of control strategies. We then used these consequences to identify testable hypotheses about the responses of invasive species to climate change and provide suggestions for invasive-species management plans. The 5 consequences also emphasize the need for enhanced environmental monitoring and expanded coordination among entities involved in invasive-species management.     

Plant invasions, generalist herbivores, and novel defense weapons

One commonly accepted mechanism for biological invasions is that species, after introduction to a new region, leave behind their natural enemies and therefore increase in distribution and abundance. However, which enemies are escaped remains unclear. Escape from specialist invertebrate herbivores has been examined in detail, but despite the profound effects of generalist herbivores in natural communities their potential to control invasive species is poorly understood. We carried out parallel laboratory feeding bioassays with generalist invertebrate herbivores from the native (Europe) and from the introduced (North America) range using native and nonnative tetraploid populations of the invasive spotted knapweed, Centaurea stoebe. We found that the growth of North American generalist herbivores was far lower when feeding on C. stoebe than the growth of European generalists. In contrast, North American and European generalists grew equally well on European and North American tetraploid C. stoebe plants, lending no support for an evolutionary change in resistance of North American tetraploid C. stoebe populations against generalist herbivores. These results suggest that biogeographical differences in the response of generalist herbivores to novel plant species have the potential to affect plant invasions.     

Implementing CITES regulations for timber.

Foresters are currently confronted with a new challenge. For the first time a commonly traded timber species has been listed on the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). At the 12th Conference of the Parties in November 2002, countries voted 68 to 30 to place the premier timber species of Latin America, big-leaf mahogany (Swietenia macrophylla King [Meliaceae]), on CITES Appendix II. Under Appendix II regulations, trade in mahogany requires that exporting countries verify that each shipment was legally obtained and that its harvest was non-detrimental to the survival of the species. Unfortunately, implementation has been weak, in part because countries have yet to develop a common, pragmatic, cost-effective system to make the legal and non-detriment findings. This paper recommends what such a system might include.     

Insects mediate the effects of propagule supply and resource availability on a plant invasion

Invasive species are a global threat to biodiversity and the functioning of natural ecosystems. Here, we report on a two-year experiment aimed at elucidating the combined and relative effects of three key controls on plant invasions: propagule supply, soil nitrogen (N) availability, and herbivory by native insects. We focus on the exotic species Lespedeza cuneata, a Rank 1 invasive species. Propagule supply and soil N-availability interacted to control the density and foliar cover of L. cuneata. In low N plots, density and foliar cover of L. cuneata were higher in the propagule addition plots than in the plots to which propagules were not added. Surprisingly, this interaction was significant only when the abundance of herbivores was experimentally reduced. This experiment provides evidence that native insect herbivores mediate the interactive effects of propagule supply and resources on invasion by a widespread invasive plant species.     

Earthworm Invasion as the Driving Force Behind Plant Invasion and Community Change in Northeastern North American Forests

Abstract:  Identification of factors that drive changes in plant community structure and contribute to decline and endangerment of native plant species is essential to the development of appropriate management strategies. Introduced species are assumed to be driving causes of shifts in native plant communities, but unequivocal evidence supporting this view is frequently lacking. We measured native vegetation, non-native earthworm biomass, and leaf-litter volume in 15 forests in the presence and absence of 3 non-native plant species ( Microstegium vimineum, Alliaria petiolata, Berberis thunbergii ) to assess the general impact of non-native plant and earthworm invasions on native plant communities in northeastern United States. Non-native plant cover was positively correlated with total native plant cover and non-native earthworm biomass. Earthworm biomass was negatively associated with cover of native woody and most herbaceous plants and with litter volume. Graminoid cover was positively associated with non-native earthworm biomass and non-native plant cover. These earthworm-associated responses were detected at all sites despite differences in earthworm species and abundance, composition of the native plant community, identity of invasive plant species, and geographic region. These patterns suggest earthworm invasion, rather than non-native plant invasion, is the driving force behind changes in forest plant communities in northeastern North America, including declines in native plant species, and earthworm invasions appear to facilitate plant invasions in these forests. Thus, a focus on management of invasive plant species may be insufficient to protect northeastern forest understory species.     

Level of environmental threat posed by horticultural trade in Cactaceae

Ornamental horticulture has been identified as an important threat to plant biodiversity and is a major pathway for plant invasions worldwide. In this context, the family Cactaceae is particularly challenging because it is considered the fifth most threatened large taxonomic group in the world; several species are among the most widespread and damaging invasive species; and Cactaceae is one of the most popular horticultural plant groups. Based on the Convention on International Trade in Endangered Species of Wild Flora and Fauna and the 11 largest online auction sites selling cacti, we documented the international cactus trade. To provide an in‐depth look at the dynamics of the industry, we surveyed the businesses involved in the cactus trade in South Africa (a hotspot of cactus trade and invasions). We purchased seeds of every available species and used DNA barcoding to identify species to the genus level. Although <20% of this trade involved threatened species and <3% involved known invasive species, many species were identified by a common name. However, only 0.02% of the globally traded cacti were collected from wild populations. Despite a large commercial network, all South African imports (of which 15% and 1.5% were of species listed as threatened and invasive, respectively) came from the same source. With DNA barcoding, we identified 24% of the species to genus level. Based on our results, we believe that if trade restrictions are placed on the small proportion of cacti that are invasive and there is no major increase in harvesting of native populations, then the commercial trade in cactus poses a negligible environmental threat. However, there are currently no effective methods for easily identifying which cacti are traded, and both the illicit harvesting of cacti from the wild and the informal trade in invasive taxa pose on‐going conservation challenges.     

Declines in Woodland Salamander Abundance Associated with Non-Native Earthworm and Plant Invasions

Abstract:  Factors that negatively affect the quality of wildlife habitat are a major concern for conservation. Non-native species invasions, in particular, are perceived as a global threat to the quality of wildlife habitat. Recent evidence indicates that some changes to understory plant communities in northern temperate forests of North America, including invasions by 3 non-native plant species, are facilitated by non-native earthworm invasion. Furthermore, non-native earthworm invasions cause a reduction in leaf litter on the forest floor, and the loss of forest leaf litter is commonly associated with declines in forest fauna, including amphibians. We conducted a mark-recapture study of woodland salamander abundance across plant invasion fronts at 10 sites to determine whether earthworm or plant invasions were associated with reduced salamander abundance. Salamander abundance declined exponentially with decreasing leaf litter volume. There was no significant relationship between invasive plant cover and salamander abundance, independent of the effects of leaf litter loss due to earthworm invasion. An analysis of selected salamander prey abundance (excluding earthworms) at 4 sites showed that prey abundance declined with declining leaf litter. The loss of leaf litter layers due to non-native earthworm invasions appears to be negatively affecting woodland salamander abundance, in part, because of declines in the abundance of small arthropods that are a stable resource for salamanders. Our results demonstrate that earthworm invasions pose a significant threat to woodland amphibian fauna in the northeastern United States, and that plant invasions are symptomatic of degraded amphibian habitat but are not necessarily drivers of habitat degradation.     

The Quagga Mussel Crisis at Lake Mead National Recreation Area,Nevada (U.S.A.)

Abstract: Parks are cornerstones of conservation; and non‐native invasive species drive extensive changes to biological diversity in parks. Knowing this, national park staff at Lake Mead National Recreation Area in the southwestern United States had a program in place for early detection of the non‐native, invasive quagga mussel (Dreissena rostriformis bugensis). Upon finding the mussel in January 2007, managers moved quickly to access funding and the best available science to implement a response. Managers considered four options—doing nothing, closing the park, restricting movement on the lakes, and educating and enforcing park visitors—and decided to focus on education and enforcing existing laws. Nonetheless, quagga spread throughout the park and soon began to appear throughout the western United States. I examined why efforts to control the expansion failed and determined the general lessons to be learned from this case. Concentrating human visitation on the lakes through land‐use zoning opened a pathway for invasion, reduced management options, and led to the rapid spread of quagga. To reconcile competing mandates to protect nature and provide recreation, zoning in parks has become a common practice worldwide. It reduces stress on some areas of a park by restricting and thus concentrating human activity in particular areas. Concentrating the human activity in one area does three things: cements pathways that repeatedly import and export vectors of non‐native invasive species; creates the disturbed area necessary to enable non‐native invasive species to gain a foothold; and, establishes a source of invasions that, without appropriate controls, can quickly spread to a park's wilderness areas.     

Eradication via destratification: whole-lake mixing to selectively remove rainbow smelt, a cold-water invasive species

Invasive species can have severe effects on aquatic ecosystems. After invasions occur, eradication should be considered whenever the potential loss of ecosystem services outweighs the cost of the eradication method. Here we evaluate the possibility of destratifying Crystal Lake, Wisconsin, USA, to eradicate the invasive fish rainbow smelt (Osmerus mordax). We modeled the effects of three destratification scenarios (non-, low-, and high-mixing) using both physical and biological models. Field observations were used to calibrate the models. Water temperatures estimated from 18 unique DYRESM simulations were used in a bioenergetics model to estimate growth of five age classes of rainbow smelt under normal and destratified conditions. Our simulations indicate that destratification can eliminate optimal rainbow smelt thermal habitat resulting in mortality. Destratified lake temperatures also surpassed several physiological critical temperatures. Bioenergetics simulations predicted a weight loss of 45-55% in yearling and adult rainbow smelt. We found that destratification is potentially effective for eradicating cold-water species in temperate lakes.     

Influence of invasive plant Coreopsis grandiflora on functional diversity of soil microbial communities

Biological invasions are increasingly attracting the ecologists' attention. Invasive plants threaten the natural ecosystems not only by competing with the native plants, but also by altering the structure and function of soil microbial communities belowground. In this study, we studied the effects of the invasive plant Coreopsis grandiflora (C. grandiflora) on the functional diversity of soil microbial communities in Laoshan mountain in the province of Shandong, North of China. We sampled soil from plots that were invaded or not invaded by C. grandiflora. The functional diversity of microbial communities in the sampled soils was assessed by the Biolog procedure test. By the ANOVA analysis of average well color development (AWCD), Shannon index (H'), Shannon evenness (E), principle components analysis of the level physiological profiles (CLPP) and correlation analysis between the studied parameters, we found that the invasive species C. grandiflora enhanced the functional diversity of soil microbial communities where the habitat was invaded by the C. grandiflora. The study indicated thatthe successful invasive plants have profound effects on the function of soil microbial communities.     

Resolving whether botanic gardens are on the road to conservation or a pathway for plant invasions

A global conservation goal is to understand the pathways through which invasive species are introduced into new regions. Botanic gardens are a pathway for the introduction of invasive non‐native plants, but a quantitative assessment of the risks they pose has not been performed. I analyzed data on the living collections of over 3000 botanic gardens worldwide to quantify the temporal trend in the representation of non‐native species; the relative composition of threatened, ornamental, or invasive non‐native plant species; and the frequency with which botanic gardens implement procedures to address invasive species. While almost all of the world's worst invasive non‐native plants occurred in one or more living collections (99%), less than one‐quarter of red‐listed threatened species were cultivated (23%). Even when cultivated, individual threatened species occurred in few living collections (7.3), while non‐native species were on average grown in 6 times as many botanic gardens (44.3). As a result, a botanic garden could, on average, cultivate four times as many invasive non‐native species (20) as red‐listed threatened species (5). Although the risk posed by a single living collection is small, the probability of invasion increases with the number of botanic gardens within a region. Thus, while both the size of living collections and the proportion of non‐native species cultivated have declined during the 20th century, this reduction in risk is offset by the 10‐fold increase in the number of botanic gardens established worldwide. Unfortunately, botanic gardens rarely implement regional codes of conduct to prevent plant invasions, few have an invasive species policy, and there is limited monitoring of garden escapes. This lack of preparedness is of particular concern given the rapid increase in living collections worldwide since 1950, particularly in South America and Asia, and highlights past patterns of introduction will be a poor guide to determining future invasion risks.     

Stakeholder participation in management of invasive vertebrates

Stakeholders are increasingly involved in species conservation. We sought to understand what features of a participatory conservation program are associated with its ecological and social outcomes. We conducted a case study of the management of invasive vertebrates in Australia. Invasive vertebrates are a substantial threat to Australia's native species, and stakeholder participation in their management is often necessary for their control. First, we identified potential influences on the ecological and social outcomes of species conservation programs from the literature. We used this information to devise an interview questionnaire, which we administered to managers of 34 participatory invasive-vertebrate programs. Effects of invasive species were related to program initiator (agency or citizen), reasons for use of a participatory approach, and stakeholder composition. Program initiator was also related to the participation methods used, level of governance (i.e., governed by an agency or citizens), changes in stakeholder interactions, and changes in abundance of invasive species. Ecological and social outcomes were related to changes in abundance of invasive species and stakeholder satisfaction. We identified relations between changes in the number of participants, stakeholder satisfaction, and occurrence of conflict. Potential ways to achieve ecological and social goals include provision of governmental support (e.g., funding) to stakeholders and minimization of gaps in representation of stakeholder groups or individuals to, for example, increase conflict mitigation. Our findings provide guidance for increasing the probability of achieving ecological and social objectives in management of invasive vertebrates and may be applicable to other participatory conservation programs.     

Invasive species distribution modeling (iSDM): Are absence data and dispersal constraints needed to predict actual distributions?

Species distribution models (SDMs) based on statistical relationships between occurrence data and underlying environmental conditions are increasingly used to predict spatial patterns of biological invasions and prioritize locations for early detection and control of invasion outbreaks. However, invasive species distribution models (iSDMs) face special challenges because (i) they typically violate SDM's assumption that the organism is in equilibrium with its environment, and (ii) species absence data are often unavailable or believed to be too difficult to interpret. This often leads researchers to generate pseudo-absences for model training or utilize presence-only methods, and to confuse the distinction between predictions of potential vs. actual distribution. We examined the hypothesis that true-absence data, when accompanied by dispersal constraints, improve prediction accuracy and ecological understanding of iSDMs that aim to predict the actual distribution of biological invasions. We evaluated the impact of presence-only, true-absence and pseudo-absence data on model accuracy using an extensive dataset on the distribution of the invasive forest pathogen Phytophthora ramorum in California. Two traditional presence/absence models (generalized linear model and classification trees) and two alternative presence-only models (ecological niche factor analysis and maximum entropy) were developed based on 890 field plots of pathogen occurrence and several climatic, topographic, host vegetation and dispersal variables. The effects of all three possible types of occurrence data on model performance were evaluated with receiver operating characteristic (ROC) and omission/commission error rates. Results show that prediction of actual distribution was less accurate when we ignored true-absences and dispersal constraints. Presence-only models and models without dispersal information tended to over-predict the actual range of invasions. Models based on pseudo-absence data exhibited similar accuracies as presence-only models but produced spatially less feasible predictions. We suggest that true-absence data are a critical ingredient not only for accurate calibration but also for ecologically meaningful assessment of iSDMs that focus on predictions of actual distributions.