Urban trees: bridge-heads for forest pest invasions and sentinels for early detection

Urban trees have been increasingly appreciated for the many benefits they provide. As concentrated hubs of human-mediated movement, the urban landscape is, however, often the first point of contact for exotic pests including insects and plant pathogens. Consequently, urban trees can be important for accidentally introduced forest pests to become established and potentially invasive. Reductions in biodiversity and the potential for stressful conditions arising from anthropogenic disturbances can predispose these trees to pest attack, further increasing the likelihood of exotic forest pests becoming established and increasing in density. Once established in urban environments, dispersal of introduced pests can proceed to natural forest landscapes or planted forests. In addition to permanent long-term damage to natural ecosystems, the consequences of these invasions include costly attempts at eradication and post establishment management strategies. We discuss a range of ecological, economic and social impacts arising from these incursions and the importance of global biosecurity is highlighted as a crucially important barrier to pest invasions. Finally, we suggest that urban trees may be viewed as ‘sentinel plantings’. In particular, botanical gardens and arboreta frequently house large collections of exotic plantings, providing a unique opportunity to help predict and prevent the invasion of new pests, and where introduced pests with the capacity to cause serious impacts in forest environments could potentially be detected during the initial stages of establishment. Such early detection offers the only realistic prospect of eradication, thereby reducing damaging ecological impacts and long term management costs.     

Exploiting Allee effects for managing biological invasions

Biological invasions are a global and increasing threat to the function and diversity of ecosystems. Allee effects (positive density dependence) have been shown to play an important role in the establishment and spread of non-native species. Although Allee effects can be considered a bane in conservation efforts, they can be a benefit in attempts to manage non-native species. Many biological invaders are subject to some form of an Allee effect, whether due to a need to locate mates, cooperatively feed or reproduce or avoid becoming a meal, yet attempts to highlight the specific exploitation of Allee effects in biological invasions are surprisingly unprecedented. In this review, we highlight current strategies that effectively exploit an Allee effect, and propose novel means by which Allee effects can be manipulated to the detriment of biological invaders. We also illustrate how the concept of Allee effects can be integral in risk assessments and in the prioritization of resources allocated to manage non-native species, as some species beset by strong Allee effects could be less successful as invaders. We describe how tactics that strengthen an existing Allee effect or create new ones could be used to manage biological invasions more effectively.     

Establishment patterns of non-native insects in New Zealand

Insects comprise the majority of non-native animal species established around the world. However, geographic biases in knowledge hamper an overall understanding of biological invasions globally. A dataset of accidentally introduced non-native insect species established in New Zealand was compiled from databases, entomological literature, and examination of specimens in the New Zealand Arthropod Collection. For each non-native species, the first recorded location and first recorded date of detection was obtained. Excluding intentionally introduced species, there are 1477 non-native insect species successfully established in New Zealand across 16 orders, 234 families and 1017 genera. Four orders (Coleoptera, Hemiptera, Hymenoptera and Diptera) contributed 77.5% of all established insect species. Herbivores represented the largest feeding guild (47.7%), comprised of polyphagous (48.3%) or oligophagous (39.7%) species. The majority of these species originated in the Australasian (36.7%) and Palearctic regions (24.8%). Regression trees, using a binary recursive partitioning approach, found the number of international tourist arrivals, exotic vegetation cover, and regional gross domestic product were the main factors explaining spatial patterns of recently established species. Gross domestic product best explained temporal patterns of establishment over the last century. Our findings demonstrate that broad-scale analyses of non-native species have important applications for border biosecurity by providing insight into the extent of invasions. In New Zealand, the current trajectory indicates fewer non-native species are establishing annually, suggesting biosecurity efforts are being effective at reducing rates of establishment.     

Grain injury models for Prostephanus truncatus (Coleoptera: Bostrichidae) and Sitophilus zeamais (Coleoptera: Curculionidae) in rural maize stores in West Africa

Prostephanus truncatus (Horn) and Sitophilus zeamais Motschulsky have been reported as the two most serious pests of stored maize in sub-Saharan Africa and smallholder farmers are in urgent need of guidelines for their proper management. In this article we investigate the injury rates attributable to these two species in terms of percentage weight loss and percentage grain damage, and we derive functional response models for the two species on maize. The models successfully described the progression of grain injury in an extensive data set compiled from previously published studies, comprising 46 time series of data relating maize injury and insect pest density. The grain injury models can be used in conjunction with predictive models of pest population dynamics to guide the development of integrated management strategies for postharvest maize pests in West Africa and comparable regions elsewhere.     

Increasing numbers and intercontinental spread of invasive insects on eucalypts

Native to Australasia, Eucalyptus (sensu lato) is one of the most planted genera of trees in the world. However, the sustainability of Eucalyptus species as plantation trees in non-native areas is increasingly threatened by the introduction and spread of Eucalyptus-feeding insects from Australia. We examine patterns and potential trends with respect to the global spread of Eucalyptus-feeding insects. Likely pathways of introduction and drivers of the rapid distribution of these insects, as well as management options are considered. The rate of introductions is shown to have increased nearly fivefold since the 1980s. As a result, the number of non-native pests of eucalypts outside of Australia has doubled in less than three decades. Furthermore, the rate of secondary spread among continents has also increased. Surprisingly, we found no association between area planted and the number of pests or new introductions. Only a small number of countries have been the points of first detection outside the native range; these countries have acted as bridgeheads to other regions. Quarantine regulations aimed at reducing the spread of invasive organisms appear to be ineffective at a global scale, and pathways allowing these invasions to occur are poorly understood or unknown. An expanded suite of management options are needed to provide resilience against the rapid accrual and homogenization of eucalypt pests, thereby ensuring the sustainability of eucalypt forestry worldwide.     

Cross-scale management strategies for optimal control of trees invading from source plantations

Biological invasion by non-native tree species can transform landscapes, and as a consequence, has received growing attention from researchers and managers alike. This problem is driven primarily by the naturalisation and invasion of tree species escaping from cultivation or forestry plantations. Furthermore, these invasions can be strongly influenced by the land-use matrix of the surrounding region, specific management of the source populations, and environmental conditions that influence seed dispersal or habitat quality for the invader. A major unresolved challenge for managing tree invasions in landscapes is how management should be deployed to contain or slow the spread of invading populations from one or more sources (e.g. plantations). We develop a spatial simulation model to test: (1) how to best prioritise the control of invasive tree populations spatially to slow or contain the biological invader when habitat quality varies in the landscape, and (2) how to allocate control effort among different management units when trees spread from many source populations. We first show that to slow down spread effectively, management strategy is less important than management effort. We then identify the conditions affecting the relative performance of different management strategies. At the landscape scale, targeting peripheral stands consistently yielded the best results whereas at the regional scale, management strategies needed to account for both habitat quality and tree life-history. Overall, our findings demonstrate that knowledge of how habitat affects tree life-history stages can improve management to contain or slow tree invasions by improving the spatial match between management effort and efficacy.     

Economic impacts of non-native forest insects in the continental United States

Reliable estimates of the impacts and costs of biological invasions are critical to developing credible management, trade and regulatory policies. Worldwide, forests and urban trees provide important ecosystem services as well as economic and social benefits, but are threatened by non-native insects. More than 450 non-native forest insects are established in the United States but estimates of broad-scale economic impacts associated with these species are largely unavailable. We developed a novel modeling approach that maximizes the use of available data, accounts for multiple sources of uncertainty, and provides cost estimates for three major feeding guilds of non-native forest insects. For each guild, we calculated the economic damages for five cost categories and we estimated the probability of future introductions of damaging pests. We found that costs are largely borne by homeowners and municipal governments. Wood- and phloem-boring insects are anticipated to cause the largest economic impacts by annually inducing nearly $1.7 billion in local government expenditures and approximately $830 million in lost residential property values. Given observations of new species, there is a 32% chance that another highly destructive borer species will invade the U.S. in the next 10 years. Our damage estimates provide a crucial but previously missing component of cost-benefit analyses to evaluate policies and management options intended to reduce species introductions. The modeling approach we developed is highly flexible and could be similarly employed to estimate damages in other countries or natural resource sectors.     

How Can the Operating Environment for Nutrition Research Be Improved in Sub-Saharan Africa? The Views of African Researchers

Optimal nutrition is critical for human development and economic growth. Sub-Saharan Africa is facing high levels of food insecurity and only few sub-Saharan African countries are on track to eradicate extreme poverty and hunger by 2015. Effective research capacity is crucial for addressing emerging challenges and designing appropriate mitigation strategies in sub-Saharan Africa. A clear understanding of the operating environment for nutrition research in sub-Saharan Africa is a much needed prerequisite. We collected data on the barriers and requirements for conducting nutrition research in sub-Saharan Africa through semi-structured interviews with 144 participants involved in nutrition research in 35 countries in sub-Saharan Africa. A total of 133 interviews were retained for coding. The main barriers identified for effective nutrition research were the lack of funding due to poor recognition by policymakers of the importance of nutrition research and under-utilisation of research findings for developing policy, as well as an absence of research priority setting from within Africa. Current research topics were perceived to be mainly determined by funding bodies from outside Africa. Nutrition researchers argued for more commitment from policymakers at national level. The low capacity for nutrition research was mainly seen as a consequence of insufficient numbers of nutrition researchers, limited skills and a poor research infrastructure. In conclusion, African nutrition researchers argued how research priorities need to be identified by African stakeholders, accompanied by consensus building to enable creating a problem-driven national research agenda. In addition, it was considered necessary to promote interactions among researchers, and between researchers and policymakers. Multidisciplinary research and international and cross-African collaboration were seen as crucial to build capacity in sub-Saharan nutrition research.     

Rearing beneficial insects for biological control purposes in resource poor areas of africa

Biological control of insect pest and weeds using beneficial insects in resource poor areas is not very well supported. Poor funding has affected in particular the importation of classical biological control agents, quarantine, rearing, research facilities, and the research programmes. Donor agencies, commercial and semi-commercial enterprises in a number of African countries have, however, been able to contribute to biological control efforts using beneficial insects by providing some of the resources needed. This has led to biological control becoming a real possibility to control insect pests and weeds in several resource poor countries. Examples are provided of the “spin-offs” of such programmes for resource poor areas in South Africa, Zambia, Kenya, Malawi, Benin and Nigeria. Conventional insect rearing, insect conservation and habitat management as an aid to biological control programmes are discussed.     

Issues Concerning the Eradication or Establishment and Biological Control of the Mediterranean Fruit Fly,Ceratitis capitata(Wiedemann) (Diptera: Tephritidae), in California

Classical biological control is suggested as a tool worth developing now for possible future use in the integrated pest management of the Mediterranean fruit fly (Medfly),Ceratitis capitata(Wiedemann), in California. Three factors that impact broadly on developing and implementing such a biological control program are: (1) the question of Medfly establishment, (2) quarantine considerations, and (3) agricultural and urban concerns. Each of these factors and their combined effects must be considered when discussing biological control of Medfly in California as shaped by historical perspectives on Medfly invasions, methods of Medfly eradication, and past biological control efforts against Medfly. We believe that biological control research should play a foundational role in any future Medfly management programs in California. Development of biological control should involve life history studies of Medfly and its natural enemies in their area of endemicity in sub-Saharan, southeast Africa. Medfly has been studied and should continue to be studied in areas it has invaded, because information derived from such studies provides insights into the potential distribution, abundance, and impact of Medfly populations in California. A plan for a biological research program on Medfly and its relatives and a biological control strategy are presented.     

Strain improvement of fungal insecticides for controlling insect pests and vector-borne diseases

Insect pathogenic fungi play an important natural role in controlling insect pests. However, few have been successfully commercialized due to low virulence and sensitivity to abiotic stresses that produce inconsistent results in field applications. These limitations are inherent in most naturally occurring biological control agents but development of recombinant DNA techniques has made it possible to significantly improve the insecticidal efficacy of fungi and their tolerance to adverse conditions, including UV. These advances have been achieved by combining new knowledge derived from basic studies of the molecular biology of these pathogens, technical developments that enable very precise regulation of gene expression, and genes encoding insecticidal proteins from other organisms, particularly spiders and scorpions. Recent coverage of genomes is helping determine the identity, origin, and evolution of traits needed for diverse lifestyles and host switching. In future, such knowledge combined with the precision and malleability of molecular techniques will allow design of multiple pathogens with different strategies and host ranges to be used for different ecosystems, and that will avoid the possibility of the host developing resistance. With increasing public concern over the continued use of synthetic chemical insecticides, these new types of biological insecticides offer a range of environmental-friendly options for cost-effective control of insect pests.     

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.     

Biological invasions in developing and developed countries: does one model fit all?

There is a strong bias concerning the regions of the globe where research on biological invasions is conducted, with notably lower representation of developing countries. However, in developing countries, effective management strategies to control invasions could be more beneficial in conserving global biodiversity since these countries tend to have larger, highly diverse natural habitats. Lower levels of development are seen as an obstacle to tackling biological invasions, but little thought is given to the advantages of developing countries in dealing with invasive species. We analyzed differences between developed and developing countries regarding the problem of invasive species and their historical and current patterns of international trade, disturbance levels and land use, research and monitoring, control and mitigation, and social awareness. Developed nations have some advantages, especially in levels of social awareness and means for controlling and studying exotics, but developing nations also enjoy important advantages given their lower levels of international trade and the availability of low-cost labor. Also, there is evidence that the process of economic development, which results in more efficient ways to transform landscapes and increases international trade, is strongly associated with increasing rates of biological invasion. Differences in data quality and availability between developed and developing countries make comparative analyses of biological invasions a difficult task. Thus, these differences creates a challenge in forming global strategies to deal with invasions. There have been calls for creating international plans to deal with invasive species, but we believe that it is important first to acknowledge the challenges and understand both the advantages and disadvantages of developing countries.     

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.     

Planting Sentinel European Trees in Eastern Asia as a Novel Method to Identify Potential Insect Pest Invaders

Quarantine measures to prevent insect invasions tend to focus on well-known pests but a large proportion of the recent invaders were not known to cause significant damage in their native range, or were not even known to science before their introduction. A novel method is proposed to detect new potential pests of woody plants in their region of origin before they are introduced to a new continent. Since Asia is currently considered to be the main supplier of insect invaders to Europe, sentinel trees were planted in China during 2007-2011 as an early warning tool to identify the potential for additional Asian insect species to colonize European trees. Seedlings (1-1.5 m tall) of five broadleaved (Quercus petraea, Q. suber, Q. ilex, Fagus sylvatica, and Carpinus betulus) and two conifer species (Abies alba and Cupressus sempervirens) were planted in blocks of 100 seedlings at two widely separated sites (one in a nursery near Beijing and the other in a forest environment near Fuyang in eastern China), and then regularly surveyed for colonization by insects. A total of 104 insect species, mostly defoliators, were observed on these new hosts, and at least six species were capable of larval development. Although a number of the insects observed were probably incidental feeders, 38 species had more than five colonization events, mostly infesting Q. petraea, and could be considered as being capable of switching to European trees if introduced to Europe. Three years was shown to be an appropriate duration for the experiment, since the rate of colonization then tended to plateau. A majority of the identified species appeared to have switched from agricultural crops and fruit trees rather than from forest trees. Although these results are promising, the method is not appropriate for xylophagous pests and other groups developing on larger trees. Apart from the logistical problems, the identification to species level of the specimens collected was a major difficulty. This situation could be improved by the development of molecular databases.     

Plant establishment and invasions: an increase in a seed disperser combined with land abandonment causes an invasion of the non-native walnut in Europe

Successful invasive species often are established for a long time period before increasing exponentially in abundance. This lag phase is one of the least understood phenomena of biological invasions. Plant invasions depend on three factors: a seed source, suitable habitat and a seed disperser. The non-native walnut, Juglans regia, has been planted for centuries in Central Europe but, until recently, has not spread beyond planted areas. However, in the past 20 years, we have observed a rapid increase in walnut abundance, specifically in abandoned agricultural fields. The dominant walnut disperser is the rook, Corvus frugilegus. During the past 50 years, rooks have increased in abundance and now commonly inhabit human settlements, where walnut trees are planted. Central Europe has, in the past few decades, experienced large-scale land abandonment. Walnut seeds dispersed into ploughed fields do not survive, but when cached into ploughed and then abandoned fields, they successfully establish. Rooks preferentially cache seeds in ploughed fields. Thus, land-use change combined with disperser changes can cause rapid increase of a non-native species, allowing it to become invasive. This may have cascading effects on the entire ecosystem. Thus, species that are non-native and not invasive can become invasive as habitats and dispersers change.     

Reprint of: Biological control of rice insect pests in China

Rice is one of the most important food crops in the world. China has the second largest area of the rice growing in the world and the highest yield of rice produced. Infestation by insect pests, especially rice planthoppers, stem borers and leaf folders, is always a serious challenge to rice production in China. Current methods for controlling insect pests in China mainly include good farming practices, biological control, breeding and growing resistant varieties, and the use of chemical insecticides. However, for farmers, the favorite method for insect pest control is still the application of chemical insecticide, which not only causes severe environmental pollution and the resurgence of herbivores but also reduces populations of the natural enemies of herbivores. To control insect pests safely, effectively and sustainably, strategies encouraging biological control are currently demanded. Here we review the progress that has been made in the development and implementation of biological controls for rice in China since the 1970s. Such progress includes the species identification of the natural enemies of rice insect pests, the characterization of their biology, and the integration of biological controls in integrated pest management. To develop effective ecological engineering programs whose aim is to implement conservation biological controls, further research, including the evaluation of the roles of plants in non-crop habitats in conservation biological controls, volatiles in enhancing efficiency of natural enemies and natural enemies in manipulating insect pests, and education to increase farmers’ knowledge of biological controls, is proposed.     

Biosafety in Populus spp. and other forest trees: from non-native species to taxa derived from traditional breeding and genetic engineering

Forest trees are fundamental components of our environment, mainly due to their long lifetime and important role in forest ecology. In the past, some non-native tree species and taxa from traditional breeding have induced severe environmental impacts such as biological invasion, changes in the ‘gene pool’, and spread of diseases in forestry. Genetically modified trees obtained in different research groups worldwide are particularly confronted with increased concerns regarding biosafety issues. In the light of current biosafety research worldwide, various threats facing forests and natural tree populations are evaluated in this review: biological invasions, horizontal gene transfer, vertical gene transfer and effects on other organisms. Results available from groups working in biosafety research and risk avoidance using forest trees, with emphasis on transgenic trees, are reviewed. Independent biosafety research as well as the establishment of biosafety research programs for forest trees financed by national and international authorities is now more important than ever before. Biosafety problems detected in the past clearly show the importance of a prior case-by-case evaluation of non-native species, new taxa and also genetically modified trees according to the precautionary principle before their release to avoid risks to the environment and human health.     

Comparing the refuge strategy for managing the evolution of insect resistance under different reproductive strategies

Genetically modified (GM) crops are used extensively worldwide to control diploid agricultural insect pests that reproduce sexually. However, future GM crops will likely soon target haplodiploid and parthenogenetic insects. As rapid pest adaptation could compromise these novel crops, strategies to manage resistance in haplodiploid and parthenogenetic pests are urgently needed. Here, we developed models to characterize factors that could delay or prevent the evolution of resistance to GM crops in diploid, haplodiploid, and parthenogenetic insect pests. The standard strategy for managing resistance in diploid pests relies on refuges of non-GM host plants and GM crops that produce high toxin concentrations. Although the tenets of the standard refuge strategy apply to all pests, this strategy does not greatly delay the evolution of resistance in haplodiploid or parthenogenetic pests. Two additional factors are needed to effectively delay or prevent the evolution of resistance in such pests, large recessive or smaller non-recessive fitness costs must reduce the fitness of resistance individuals in refuges (and ideally also on GM crops), and resistant individuals must have lower fitness on GM compared to non-GM crops (incomplete resistance). Recent research indicates that the magnitude and dominance of fitness costs could be increased by using specific host–plants, natural enemies, or pathogens. Furthermore, incomplete resistance could be enhanced by engineering desirable traits into novel GM crops. Thus, the sustainability of GM crops that target haplodiploid or parthenogenetic pests will require careful consideration of the effects of reproductive mode, fitness costs, and incomplete resistance.     

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.