Predicting novel herbivore–plant interactions

As human‐aided range expansions and climate change alter the distributions of plants and their herbivores, predicting and addressing novel species interactions will become increasingly pressing for community ecologists. In this context, a key, surprisingly understudied question is: when an exotic plant is introduced, which herbivores will adopt this new potential host? Whether the plant is a weed, an ornamental, or a crop, the development versus non‐development of a novel plant–insect interaction can have profound effects for both economic and conservation applications. In this paper, we sketch mechanistic and statistical frameworks for predicting these interactions, based on how plant and herbivore traits as well as shared evolutionary history can influence detection, recognition, and digestion of novel plants. By emphasizing mechanisms at each of these steps, we hope to clarify different aspects of novel interactions and why they may or may not occur. We also emphasize prediction and forecasting, as a major goal is to know in advance which interactions will develop from the many plant or insect introductions that occur in natural and man‐made systems.     

The constraints of selecting for insect resistance in plantation trees

• 1 High productivity in plantations of exotic tree species is achieved by management for fast growth in the absence of the full complex of co‐evolved insect herbivores. In the case of Eucalyptus, silvicultural selection for desirable wood traits is concomitant with a trade‐off against defence and a reduction of chemical and genetic diversity. These factors, combined with accidental introductions, rapid insect evolution and the emergence of new pests, increase the likelihood that future plantations will need insect pest management to maintain productivity.
• 2 Forestry researchers have suggested that selecting for resistant genotypes may be beneficial in insect control. There are, however, significant differences between long‐lived trees and annual crops that make this approach unlikely to be successful. This is illustrated using several examples of research into resistance to insect herbivores in trees.
• 3 Selection for resistance to insects in trees requires an assessment of trial plantations for heritable variation in insect damage and then a determination of the effect of variation in resistance on insect population parameters. Identifying rare resistant genotypes using markers is difficult because many factors interact to produce a resistant phenotype, and phytophagous insects have less intimate relationships with their host than pathogens, resulting in weak associations with genetic loci.
• 4 If resistant genotypes are identified, their widespread deployment in plantations might not provide satisfactory management of insect pests when the use of extensive monocultures is continued. In this paper, experiments are suggested that would explore the effectiveness of polycultures or chemotype mixtures with respect to ameliorating the damage of insects on plantation productivity. In addition, mitigating the effects of some insects on plantation productivity by maintaining vigour of fast‐growing eucalypts should be considered.

     

Potential risks of systemic imidacloprid to parasitoid natural enemies of a cerambycid attacking Eucalyptus

A community of insect herbivores has established on eucalyptus species in California following their introduction from Australia. A number of the species are under complete or partial biological control. A response to introduction of additional pest species into the complex has been the application of systemic insecticides to infested trees. Natural enemies that have been introduced to control the various pest species feed on the nectar of treated trees, and thus may be affected by these pesticides. In this study, Eucalyptus rudis trees were treated at label rates with the neonicotinoid systemic insecticide imidacloprid and nectar sampled at 5 months post-treatment during the spring bloom. The concentration of imidacloprid and its toxic metabolites in nectar was measured by ELISA at 660 ppb. Adults of the encyrtid egg parasitoid Avetianella longoi that were fed floral nectar collected from treated trees had significantly lower survival and reproductive fitness than adults fed nectar from untreated trees. In feeding bioassays, in which the adults were fed a range of concentrations of imidacloprid in sugar water, the LC₅₀ for A. longoi was 212 ppb imidacloprid. Bioassays were also conducted with the braconid larval parasitoid, Syngaster lepidus. The LC₅₀ for S. lepidus was 288 ppb imidacloprid. The insecticide appears to be concentrating in the nectar at higher levels than reported from other plant species and at concentrations exceeding the LC₅₀ for two important parasitoids. If tree treatments become widespread as a result of continual introductions of new eucalypt herbivores, established biological control programs could be at significant risk.     

A history of the biological and integrated control of red scale,Aonidiella aurantii on citrus in the lower Murray valley of Australia

In the lower Murray valley of Australia, the major insect pest of citrus, California red scaleAonidiella aurantii (Maskell), is controlled by a number of introduced hymenopterous parasites. Parasite introductions began in 1943, and continued until 1979. Eight species, especially in the Encyrtid genusAphytis, were introduced and mass released. The only species to become permanently established wereAphytis chrysomphali Mercet,Comperiella bifasciata (Howard),A. melinus DeBach, andProspaltella perniciosi (Tower).A. melinus, introduced in 1961, has become widely established throughout the region, is the dominant and most important parasite, and now appears to have completely displacedA. chrysomphali (1954) in all areas.C. bifasciata (1943) is also an important parasite and is distributed throughout the area.P. perniciosi (1970) is established on a few orchards near Mildura. Most of the work on the biological and integrated control of red scale began with the establishment of laboratory and insectary facilities at Mildura in 1967 and Loxton in 1968. A commercial insectary was also established at Loxton in 1971. Biological methods of insect control were first used commercially during 1944 (war period) by a few citrus growers at Merbein in Sunraysia. However, the adoption of biological and integrated methods for the control of red scale and other citrus pests, began on a large scale in the late 1960's. The use of organophosphate insecticides on citrus has declined since that time and has been close to zero since about 1977. Parasites and petroleum oil sprays have been used together in an integrated approach to the control of California red scale on some orchards. The biological control of this key pest has led to a decline in the incidence of secondary pests to the point where all insect pests of citrus are now regarded as being under effective biological control. The incidence of red scale, and other pests, has been less under biological and integrated control than it ever was when chemical control measures were widely used in the 1960's. Factors which have contributed to the success of the programme are discussed.     

Do trees in urban or ornamental plantings receive more damage by insects than trees in natural forests?

Abstract.

• 1 To determine if trees in urban or ornamental plantings are more susceptible to attack and receive more damage to foliage by herbivores than trees in natural forests, we compared the amount of leaf damage caused by several guilds of insects feeding on seven species of native, broadleaf trees in two geographic locations.
• 2 Total leaf damage did not differ significantly between urban or ornamental and natural forests, although trees in natural forests tended to have slightly higher levels of leaf damage.
• 3 Damage caused by chewing insects was consistently higher on trees in natural forests than in urban or ornamental plantings. All other feeding guilds showed no consistent pattern in levels of damage between the two habitats.
• 4 Total damage levels were highest on canopy trees and lowest on understorey trees.
• 5 These results are inconsistent with the view that trees in urban or ornamental settings are more susceptible to insect attack than trees in natural forests.
• 6 The lower level of foliar damage caused by chewing insects on trees in urban or ornamental plantings may arise because of low rates of dispersal by insects into urban environments, higher levels of plant resistance to insect attack in urban or ornamental plantings, or lower survival rates of herbivorous insects in urban environments.

     

Insect pests of eucalypts in California: implications of managing invasive species

For the first 150 years following their introduction, eucalypts planted in the California landscape were free of both insect pests and diseases. In the last 15 years, numerous herbivorous insect species have been introduced accidentally into the State and have caused significant damage to the trees. Several of these species, e.g. Phoracantha semipunctata (Fabricius), Phoracantha recurva Newman (Coleoptera: Cerambycidae) and Gonipterus scutellatus Gyllenhal (Coleoptera: Curculionidae), have also been introduced into other parts of the world where eucalypts are grown, whereas others, e.g. Glycaspsis brimblecombei Moore (Hemiptera: Spondyliaspidae) and Eucalyptolyma maideni Froggatt (Hemiptera: Spondyliaspidae), are currently restricted to California and Australia. Research programmes have provided management solutions to individual pest problems, but as more pest species are introduced, these solutions must be integrated across broad geographic, horticultural, and economic scales, in a systems approach.     

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

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

Diversity and impact of herbivorous insects on Brazilian peppertree in Florida prior to release of exotic biological control agents

Brazilian peppertree, Schinus terebinthifolia Raddi (Anacardiaceae), is a South American plant that is highly invasive in Florida. The impact of insect herbivores on the performance of Brazilian peppertree was evaluated at two locations in Florida using an insecticide exclusion method. Although 38 species of insect herbivores were collected on the invasive tree, there were no differences in growth or reproductive output of insecticide protected and unprotected trees, providing evidence that insect feeding had no measurable impact on tree performance. The majority of insects collected on Brazilian peppertree were generalists, and several were serious agricultural pests.     

Recent non‐native invertebrate plant pest establishments in Great Britain: origins,pathways, and trends

1 An appraisal of non‐native invertebrate plant pest establishments in Great Britain, between 1970 and 2004, was carried out to improve our understanding of current invasion processes by non‐native plant pests, and to assist national strategies in managing the risks they pose. 2 A total of 164 establishments, comprising 50 natural colonists and 114 human‐assisted introductions, were recorded across 13 major taxonomic groups. 3 The mean rate of establishment was 22.1 species per 5‐year period: 19.1 and 3.0 species outside and inside protected cultivation, respectively. Despite the continuing rapid growth in international trade and a general perception that rates of pest invasions are accelerating, no significant temporal trends in the rate of establishments in Great Britain were detected, either for natural colonists or human‐assisted introductions, or for pests of plants grown indoors or outside. 4 The plant trade, particularly in ornamental plants, accounted for nearly 90% of human‐assisted introductions; apiculture, biological control, timber imports, transport stowaways and intentional releases each contributed less than 5%. Only eight (4.9%) of the establishments could be considered as having no direct potential economic impact because all other species have been recorded as feeding on cultivated plants. A greater proportion of establishments by both natural colonists and human‐assisted introductions occurred on non‐native, woody plants. 5 The present study confirms previous work in other European countries that highlight the predominant role of the ornamental plant trade in introducing new plant pests to the European continent, mainly from Asia and North America.     

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.     

Lepidopteran pests in the Upper Volga area (Lepidoptera)

Data on 154 species of lepidopteran pests in the Upper Volga region, of which 9 are considered as principal, 29 as minor, and 116 as potential pests are summarized. Most of harmful species belong to the families Tortricidae (27), Noctuidae (24), Gelechiidae (11), Gracillariidae (9), Geometridae (8), and Lymantriidae (6). The pests of forest trees and shrubs are most numerous, followed by the pests of fruit trees and berry shrubs, pests of ornamental trees and shrubs, etc. There is no lepidopteran pest of field and forage crops in the region. On the whole, the economic importance of Lepidoptera in the Upper Volga region is lower than that of the other insect orders.     

Lianas as a food resource for herbivorous insects: a comparison with trees

Woody climbers or, ‘lianas’, are one of the features that characterise rainforests. They contribute substantially to plant diversity and leaf biomass which makes them a potentially important food source for herbivores. Here, we focus on insect herbivores, folivores in particular, to show how disparities in the quantitative and qualitative availability of leaves between lianas and trees may differentially influence insect folivory and the herbivore communities themselves. We develop a conceptual model and show that lianas in general have lower structural and chemical defences, a greater nutritional profile and a preferable phenology in comparison with trees, which, contrary to our expectations, has led to assemblages of more‐specialised insects. The impacts this has on higher trophic levels and broader ecological networks, however, are poorly known. We show through a study of four tropical floras from different biogeographic realms that lianas are likely to be a target for a wide range of insect herbivore taxa as they are a phylogenetically diverse group and increase diversity of higher taxa at local scales. This, in combination with their highly palatable leaves, may also make them a suitable temporary food source for insects during times when preferred host plants are scarce. This phenomenon has been observed in mammalian herbivores but awaits investigation in insects as does the effects this may have on survival and fitness. Apparent recent increases in liana abundances in some forests, likely due to climate change, makes understanding their role in supporting and maintaining biodiversity an increasingly important and necessary challenge. Since trees or saplings have usually been the subject of studies on insect herbivory, major knowledge gaps remain about the ways in which lianas contribute to, support and maintain the ecosystems in which they exist. We use our conceptual model to guide future research directions and express the necessity for caution when extrapolating explanations of herbivory derived from data on trees to growth forms with fundamentally different ecologies.     

The effects of gibberellin A3 on white clover during floral development

Hokkanen & Pimentel (1984) proposed a novel approach for the selection of biocontrol agents. They advocated the selection of agents from a relative of the weed plant rather than from the target weed species itself. The new relationship that such agents would have with the weed would be characterised by a relative lack of homeostasis compared with the old herbivore-plant associations traditionally used in weed biocontrol, and would consequently be more effective. There are few examples to support these contentions because of the traditional use of old herbivore-plant associations in weed biocontrol. In the present study, herbivore-plant associations in agricultural situations, which are analogous to agent-weed associations, are examined to assess the potential of new associations for weed biocontrol. The herbivores on 14 introduced crop plants which have salient similarities to the major weeds in the south-western Cape were surveyed: (a) 68% of the 188 insect and mite herbivores are indigenous species in new associations with these host plants, and (b) of the five most damaging pests on each of 13 of the crop plants, 53% are in new associations with the plants. Of the 40 most important agricultural pests in South Africa, 58% are in new associations, confirming these results. About 50% of the insect and mite herbivores in new associations with their host plants are oligophagous, indicating that new associations are not necessarily characterised by polyphagy and hence unsafe for use in weed biocontrol. We conclude that new associations between herbivore species and host plants have strong potential in weed biocontrol because (a) their frequency in agriculture indicates that they can easily be established in weed biocontrol situations, (b) they are as damaging as old associations and (c) they are not necessarily unsafe as regards host specificity. We therefore advocate the use of both the classical approach and that of Hokkanen & Pimentel (1984). Our support for Hokkanen & Pimentel (1984) is, however, based on evidence and rationale clearly different to theirs, and we provide novel guidelines, which can be routinely and practically applied in the selection of agents for weed biocontrol.     

沈阳城市绿化生态工程树种综合评价分级选择

开展城市绿化生态工程建设是保护城市生态环境、促进城市可持续发展的必由之路,对城市绿化树种生态适应性及生态功能性相结合的综合评价,是构建城市生态工程、合理应用选择城市绿化植物的科学依据,通过专家咨询评价的方法,从生态适应能力、绿化生态效益、美化效果、抗病虫害性、抗污染性、经济效益等方面全面系统地建立了沈阳市绿化树种生态功能与生态适应性评价应用的综合指标体系。并根据专家第一轮评价和沈阳市特定区域和范围,在对绿化树种的综合应用评价的基础上,选定耐寒性、耐旱性、耐瘠薄、抗病虫能力、抗污染能力、观赏性和生态效益7项评价指标为主要的评价指标因子。对沈阳常见的200种绿化树种进行了综合评判与分级,评价结果表明,综合效能为1级植物有58种;2级植物为93种;3级植物38种;4级植物11种。     

Population,community and ecosystem effects of exotic herbivores: A growing global concern

Exotic herbivores represent a serious threat to native biodiversity, producing large scale changes in native communities and altering ecosystem processes. In this special issue, we present a series of case studies and reviews from different areas of the world that highlight (1) the consequences of herbivore introductions are a global problem; (2) they can result in wholesale shifts in the distribution of dominant plants on the landscape and; (3) the effects of herbivore introductions extend from the population to the community and ecosystem level. These studies suggest that introduced herbivores often retard ecosystem recovery after disturbance, facilitate invasion of plant species and can act as selective agents on native plant communities. These studies also suggest that several topics, including facilitation between exotic herbivores and exotic plants and animals (i.e., invasional meltdown) and the effect of exotic herbivores on ecosystem processes, require more research attention. Overall the papers in this special feature suggest that introduced herbivores are a global problem with wide-ranging ecological and evolutionary effects.     

Non-protein amino acids in plant defense against insect herbivores: representative cases and opportunities for further functional analysis

Chemical defense against herbivores is of utmost importance for plants. Primary and secondary metabolites, including non-protein amino acids, have been implicated in plant defense against insect pests. High levels of non-protein amino acids have been identified in certain plant families, including legumes and grasses, where they have been associated with resistance to insect herbivory. Non-protein amino acids can have direct toxic effects via several mechanisms, including misincorporation into proteins, obstruction of primary metabolism, and mimicking and interfering with insect neurological processes. Additionally, certain non-protein amino acids allow nitrogen to be stored in a form that is metabolically inaccessible to herbivores and, in some cases, may act as signals for further plant defense responses. Specialized insect herbivores often possess specific mechanisms to avoid or detoxify non-protein amino acids from their host plants. Although hundreds of non-protein amino acids have been found in nature, biosynthetic pathways and defensive functions have been elucidated in only a few cases. Next-generation sequencing technologies and the development of additional plant and insect model species will facilitate further research on the production of non-protein amino acids, a widespread but relatively uninvestigated plant defense mechanism.     

Are bottlenecks associated with colonization? Genetic diversity and diapause variation of native and introduced Rhagoletis completa populations

The success of invasive species appears to be a paradox: despite experiencing strong population bottlenecks, invasive species are able to successfully establish in new environments. We studied how the walnut husk fly, Rhagoletis completa, was able to successfully colonize California from the Midwestern United States, by examining genetic diversity and diapause variation of native and introduced fly populations. Climate plays an important role in the successful establishment of introduced insects, because insect diapause is highly dependent upon external climatic conditions. We examined if: (1) fly populations show signs of a population bottleneck, (2) native and introduced flies differ in diapause length when exposed to California and Midwestern climatic conditions, and (3) population genetic diversity is related to variation in diapause length. We assessed if fly diapause conformed more to a model of establishment by local adaptation or to a model of a highly plastic “general-purpose genotype”. Our results indicate that only two populations close to the original introduced location showed signs of a population bottleneck, and native and introduced populations did not differ in genetic diversity. Genetic diversity increased in the northern introduced populations, suggesting that multiple introductions have occurred. Flies emerged about 2 weeks earlier under the Midwestern treatment than the California treatment, and introduced flies emerged about a week earlier than native flies. All flies emerged when walnuts are typically available in California. Although variance in diapause length differed between populations, it did not vary between populations or regions. Furthermore, genetic diversity was not associated with diapause variation. Therefore, multiple introductions and a “general-purpose genotype” appear to have facilitated the fly’s invasion into California.     

林木营养与虫害——油松施氮肥的抗虫效应

本文探讨了油松施氮肥后的抗虫效应。1981—1983年试验油松每年以两种方法施尿素5次。根施尿素每次用量为0.1和0.2公斤/株或采用重量比是1:200的尿素溶液进行叶面喷雾。实验结果如下:(1)枯梢率低于对照树;(2)松大蚜、松梢螟和红蜘蛛的虫口数量减少;(3)树皮和树叶中的总氮量、粗纤维素、粗灰分含量增加,而可溶性糖含量下降。经统计分析,上述结果和对照树相比有显著差异。结果表明,施氮肥不仅会增强树势,还会增强树的抗虫性而减轻虫害。我们认为,其主要作用机理是树木营养成分的明显改变影响了昆虫的生长发育和繁殖。     

应用马尔可夫链理论对褐飞虱和白背飞虱发生程度的预测

褐飞虱Nilaparvata lugens (Stál)和白背飞虱Sogatella furcifera (Horváth) 是我国稻区的主要迁飞性害虫,给水稻生产带来严重危害.以重庆市秀山县1983～2007年田间褐飞虱和白背飞虱发生程度的时间序列资料,运用基于马尔可夫链理论的转移概率预测法,构建了1～5阶转移概率矩阵,根据每一阶概率矩阵单独进行回报的历史符合率计算各阶转移概率矩阵的权重,以预报年前5a褐飞虱和白背飞虱的连续发生状态预测第6年的发生级别.回报20a的结果历史符合率均达到90%,对2008年的预测也符合当年田间实际发生情况.该方法构建的5阶转移概率矩阵及其权重对于秀山县及其邻近地区褐飞虱和白背飞虱发生程度的长期预报具有重要指导意义,为害虫的长期可预测性提供了一种快速、有效的方法.     

Evidence for genetic erosion of a California native tree, <Emphasis Type="Italic">Platanus racemosa</Emphasis>, via recent,ongoing introgressive hybridization with an introduced ornamental species

When non-native, genetically diverse species are introduced, hybridization with native congeners may erode the genetic composition of local species, perhaps even resulting in extinction. While such events may lead to adverse consequences at the community and ecosystem level, few studies exist on ecologically important tree species. In the genus Platanus, introgressive hybridization is widespread, and one common ornamental species, introduced to California during the late 19th century, is itself a hybrid. Our microsatellite analysis of more than 400 Platanus trees from north-central California reveals a complex pattern of invasion and hybridization in an age-structured population. By using size as a proxy for age, we have demonstrated that the Platanus population of north-central California has recently gained genetic diversity and effective population size. Principal coordinate analysis (PCoA) and genetic admixture analysis (STRUCTURE) both reveal a strong differentiation of genotypes into two main genetic clusters, with a large number of admixed genotypes. One of the genetic clusters identified is heavily biased towards younger trees, including samples from locations with relatively recently planted ornamental trees likely to be P. × hispanica (formerly known as P. × acerifolia). We conclude that the two genetic clusters correspond to the native P. racemosa and the introduced invasive hybrid species P. × hispanica. Additional hybridization between the invasive ornamental and the native species has occurred in California, and recent hybrid trees are more likely to be younger than trees without admixture. Our findings suggest that the observed increase in genetic diversity among California Platanus is due to rampant ongoing introgression, which may be threatening the continued genetic distinctiveness of the native species. This is cause for concern from a conservation standpoint, due to a direct loss of genetic distinctiveness, and a potential reduction in habitat value of associated species.