Plant and insect cuticular lipids serve as behavioral cues for insects

The roles of plant and insect cuticular lipids in insect and plant interactions are reviewed. Emphasis is given to the influence that the host plant and the surface lipids of the host plant have upon insect herbivores and the predators and parasitoids of these herbivores. Variations in cuticular lipids of herbivorous insects are dependent upon the host plant, and these variations may affect the behavior of predators and parasitoids. The cuticular lipids of species which interact on multiple trophic levels are compared. Similarities were found between the hydrocarbons of herbivorous insects, their host plants, and their predators or parasitoids.     

Impacts of exotic invertebrates on New Zealand?s indigenous species and ecosystems

Biological invasions have significantly affected New Zealand?s native species and ecosystems. Most prominent are the effects of exotic mammals and plants, whereas few invertebrate invasions are known to have major effects on native ecosystems. Exceptions are the well-known cases of Vespula wasps in Nothofagus forest ecosystems and Eriococcus scale insects in Leptospermum shrublands. This limited impact is surprising because over 2000 exotic invertebrates have become established in New Zealand, among them many pests of exotic crop plants. The low impact of exotic invertebrates that invaded forests and other native ecosystems in New Zealand is in contrast to the situation in other parts of the world where many invertebrates have become important pests. We provide an overview of known invasions by exotic invertebrates in New Zealand, and explore in more detail several examples of invasive species, including herbivores, predators, parasitoids, decomposers and other groups in forests, grasslands, and other terrestrial ecosystems. Several hypotheses have been proposed to explain the comparative scarcity of such invasions that affect New Zealand?s indigenous ecosystems. There is a common view that New Zealand?s native species and ecosystems are inherently resistant to exotic invertebrate invaders, and there is some evidence to support this view. As a result of the high level of endemism in New Zealand?s flora, many native plants are phylogenetically distant from the host plants of many plant-feeding invaders. This provides some protection. Less host-specific plant-feeding insects, generalist predators, parasitoids and decomposers are less affected by such constraints, and these groups are perhaps more represented among the successful invaders of natural ecosystems. However, the shortage of studies on invader impacts on native species and ecosystems, compared with studies on economically important crops and production ecosystems, means that an unbiased comparison is not possible at this time. Furthermore, many invaders go through extended lag phases where their impacts are not easily noticed until they become more abundant and create more damage. Likewise, indirect effects of invaders, through more complex interactions in food webs, as well as impacts on ecosystem functions such as decomposition and pollination, are more subtle and difficult to detect without careful study. There is clearly a need for more research to determine more accurately which exotic invertebrates are already present, what their direct and indirect impacts are, and what generalisations and predictions about threats to native species and ecosystems are possible.     

Meta-analysis of tree diversity effects on the abundance,diversity and activity of herbivores' enemies

The natural enemies hypothesis predicts that the abundance and diversity of antagonists such as predators and parasitoids of herbivores increases with the diversity of plants, which can lead to more effective top-down control of insect herbivores. However, although the hypothesis has received large support in agricultural systems, fewer studies have been conducted in forest ecosystems and a comprehensive synthesis of previous research is still lacking.We conducted a meta-analysis of 65 publications comparing the diversity, abundance or activity of various groups of natural enemies (including birds, bats, spiders and insect parasitoids) in pure vs. mixed forest stands. We tested the effects of forest biome, natural enemy taxon and type of study (managed vs experimental forest).We found a significant positive effect of forest tree diversity on natural enemy abundance and diversity but not on their activity. The effect of tree diversity on natural enemies was stronger towards lower latitudes but was not contingent on the natural enemy taxon level.Overall, our study contributes toward a better understanding of the “natural enemies hypothesis” in forest systems and provides new insights about the mechanisms involved. Furthermore, we outline potential avenues for strengthening forest resistance to the growing threat of herbivorous insects.     

Root herbivores influence the behaviour of an aboveground parasitoid through changes in plant-volatile signals

It is widely reported that plants emit volatile compounds when they are attacked by herbivorous insects, which may be used by parasitoids and predators to locate their host or prey. The study of herbivore-induced plant volatiles and their role in mediating interactions between plants, herbivores and their natural enemies have been primarily based on aboveground systems, generally ignoring the potential interactions between above and belowground infochemical- and food webs. This study examines whether herbivory by Delia radicum feeding on roots of Brassica nigra (black mustard) affects the behaviour of Cotesia glomerata , a parasitoid of the leaf herbivore Pieris brassicae , mediated by changes in plant volatiles. In a semi-field experiment with root-damaged and root-undamaged plants C. glomerata prefers to oviposit in hosts feeding on root-undamaged plants. In addition, in a flight-cage experiment the parasitoid also prefers to search for hosts on plants without root herbivores. Plants exposed to root herbivory were shown to emit a volatile blend characterized by high levels of specific sulphur volatile compounds, which are reported to be highly toxic for insects, combined with low levels of several compounds, i.e. beta-farnesene, reported to act as attractants for herbivorous and carnivorous insects. Our results provide evidence that the foraging behaviour of a parasitoid of an aboveground herbivore can be influenced by belowground herbivores through changes in the plant volatile blend. Such indirect interactions may have profound consequences for the evolution of host selection behaviour in parasitoids, and may play an important role in the structuring and functioning of communities.     

Non-native plants rarely provide suitable habitat for native gall-inducing species

For insect herbivores, a critical niche requirement—possibly the critical niche requirement—is the presence of suitable host plants. Current research suggests that non-native plants are not as suitable as native plants for native herbivores, resulting in decreases in insect abundance and richness on non-native plants. Like herbivores, gall-forming insects engage in complex, species-specific interactions with host plants. Galls are plant tissue tumors (including bulbous or spindle-shaped protrusions on leaves, stems and other plant organs) that are induced by insects through physical or chemical damage (prompting plants to grow a protective tissue shell around the insect eggs and larvae). As such, we hypothesized that gall-inducing insect species richness would be higher on native than non-native plants. We also predicted higher gall-inducing insect species richness on woody than herbaceous plants. We used an extensive literature review in which we compiled gall host plant species by genus, and we assigned native or non-native (or mixed) status to each genus. We found that native plants host far more gall-inducing insect species than non-native plants; woody plants host more gall-inducing species than herbaceous plants; and native woody plants host the most gall-inducing species of all. Gall-inducing species generally are a very cryptic group, even for experts, and hence do not elicit the conservation efforts of more charismatic insects such as plant pollinators. Our results suggest that non-native plants, particularly non-native woody species, diminish suitable habitat for gall-inducing species in parallel with similar results found for other herbivores, such as Lepidopterans. Hence, the landscape-level replacement of native with non-native species, particularly woody ones, degrades taxonomically diverse gall-inducing species (and their inquilines and parasitoids), removing multiple layers of diversity from forest ecosystems.

     

Physiological function and ecological aspects of fatty acid-amino acid conjugates in insects†

In tritrophic interactions, plants recognize herbivore-produced elicitors and release a blend of volatile compounds (VOCs), which work as chemical cues for parasitoids or predators to locate their hosts. From detection of elicitors to VOC emissions, plants utilize sophisticated systems that resemble the plant–microbe interaction system. Fatty acid–amino acid conjugates (FACs), a class of insect elicitors, resemble compounds synthesized by microbes in nature. Recent evidence suggests that the recognition of insect elicitors by an ancestral microbe-associated defense system may be the origin of tritrophic interactions mediated by FACs. Here we discuss our findings in light of how plants have customized this defense to be effective against insect herbivores, and how some insects have successfully adapted to these defenses.     

Ecological effects of invasive alien insects

A literature survey identified 403 primary research publications that investigated the ecological effects of invasive alien insects and/or the mechanisms underlying these effects. The majority of these studies were published in the last 8 years and nearly two-thirds were carried out in North America. These publications concerned 72 invasive insect species, of which two ant species, Solenopsis invicta and Linepithema humile, accounted for 18% and 14% of the studies, respectively. Most publications investigated effects on native biodiversity at population or community level. Genetic effects and, to a lesser extent, effects on ecosystem services and processes were rarely explored. We review the effects caused by different insect invaders according to: their ecosystem roles, i.e. herbivores, predators, parasites, parasitoids and pollinators; the level of biological organisation at which they occur; and the direct and indirect mechanisms underlying these effects. The best documented effects occur in invasive ants, Eurasian forest herbivores invasive in North America, and honeybees. Impacts may occur through simple trophic interactions such as herbivory, predation or parasitism. Alien species may also affect native species and communities through more complex mechanisms such as competition for resources, disease transmission, apparent competition, or pollination disruption, among others. Finally, some invasive insects, particularly forest herbivores and ants, are known to affect ecosystem processes through cascading effects. We identify biases and gaps in our knowledge of ecological effects of invasive insects and suggest further opportunities for research. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Contrasting patterns of herbivore and predator pressure on invasive and native plants

Invasive non-native plant species often harbor fewer herbivorous insects than related native plant species. However, little is known about how herbivorous insects on non-native plants are exposed to carnivorous insects, and even less is known on plants that have recently expanded their ranges within continents due to climate warming. In this study we examine the herbivore load (herbivore biomass per plant biomass), predator load (predator biomass per plant biomass) and predator pressure (predator biomass per herbivore biomass) on an inter-continental non-native and an intra-continental range-expanding plant species and two congeneric native species. All four plant species co-occur in riparian habitat in north-western Europe. Insects were collected in early, mid and late summer from three populations of all four species. Before counting and weighing the insects were classified to trophic guild as carnivores (predators), herbivores, and transients. Herbivores were further subdivided into leaf-miners, sap-feeders, chewers and gallers. Total herbivore loads were smaller on inter-continental non-native and intra-continental range-expanding plants than on the congeneric natives. However, the differences depended on time within growing season, as well as on the feeding guild of the herbivore. Although the predator load on non-native plants was not larger than on natives, both non-native plant species had greater predator pressure on the herbivores than the natives. We conclude that both these non-native plant species have better bottom-up as well as top-down control of herbivores, but that effects depend on time within growing season and (for the herbivore load) on herbivore feeding guild. Therefore, when evaluating insects on non-native plants, variation within season and differences among feeding guilds need to be taken into account.     

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

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

Insect communities on experimental mugwort (Artemisia vulgaris L.) plots along an urban gradient

We studied the ability of insect herbivores and their natural enemies to colonize exposed, potted mugwort plants (Artemisia vulgaris L.) along a rural-urban gradient in 1994 in Hamburg (northern Germany). Ectophagous insects, leafmines and galls were monitored weekly from mid-May to mid-September. Endophagous insects were counted by harvesting and dissecting the stems at the end of the growing season. The rural-urban gradient was characterized by a gradient of vegetation-free areas and increasing proportion of ground covered in concrete, tarmac, paving and other impermeable surfaces surrounding the Artemisia plots, i.e. six different zones of increasing isolation. Numbers of insect species (herbivores, parasitoids and predators) decreased along the gradient from 43 to 12. Monophagous herbivores were not more affected than polyphagous herbivores, but parasitoids, especially rare species, were more strongly affected by isolation than predators. Some dominant herbivorous species were very successful colonizers and occurred in inner city sites devoid of all natural vegetation. Sometimes their abundance increased in the inner city to significantly higher densities than in the urban fringe. Isolation appeared to be the main reason for the observed patterns, since area and soil conditions were held constant in the experiment. Microclimate and pollution were considered to play a minor role. Received: 16 January 1997 / Accepted: 26 August 1997     

Nitrogen in insects: implications for trophic complexity and species diversification

Disparities in nutrient content (nitrogen and phosphorus) between herbivores and their plant resources have lately proven to have major consequences for herbivore success, consumer-driven nutrient cycling, and the fate of primary production in ecosystems. Here we extend these findings by examining patterns of nutrient content between animals at higher trophic levels, specifically between insect herbivores and predators. Using a recently compiled database on insect nutrient content, we found that predators exhibit on average 15% greater nitrogen content than herbivores. This difference persists after accounting for variation from phylogeny and allometry. Among herbivorous insects, we also found evidence that recently derived lineages (e.g., herbivorous Diptera and Lepidoptera) have, on a relative basis, 15%-25% less body nitrogen than more ancient herbivore lineages (e.g., herbivorous Orthoptera and Hemiptera). We elaborate several testable hypotheses for the origin of differences in nitrogen content between trophic levels and among phylogenetic lineages. For example, interspecific variation in insect nitrogen content may be directly traceable to differences in dietary nitrogen (including dilution by gut contents), selected for directly in response to the differential scarcity of dietary nitrogen, or an indirect consequence of adaptation to different feeding habits. From some functional perspectives, the magnitude rather than the source of the interspecific differences in nitrogen content may be most critical. We conclude by discussing the implications of the observed patterns for both the trophic complexity of food webs and the evolutionary radiation of herbivorous insects.     

Advances in research of induced resistance to insects in cotton

Any change in a plant that occurs following herbivory or environmental factors is an induced response. These changes include phytochemical induction, increases in physical defenses, emission of volatiles that attract predators and parasitoids of herbivores, and reduction in plant nutritional quality for herbivores, which is termed induced resistance. Induced resistance has been demonstrated ubiquitously in plants. It is one of our goals to review what is known about the induced resistance to herbivorous insects in cotton, including three resistance secondary metabolites (terpenoid, tannin, and flavonoids) that are contained at any significant levels of resistance to herbivorous insects in cotton cultivates. In many cases, the quantities or quality of secondary metabolites in plant are changed after attacked by insects. This review focuses on induced plant resistance as quantitative or qualitative enhancement of defense mechanism against insect pests, especially on the abiotic-elicitors-induced resistance in cotton plants. The abiotic-elicitor of cupric chloride, an exogenous inorganic compound, may induce the secondary metabolites accumulation and is referred to as a copperinducible elicitor (CIE). Finally, we discuss how copperinducible elicitor may be used in the Integrated Pest Management (IPM) system for cotton resistance control.     

Characterizing an invasion assemblage: first comparison of insect communities on native and introduced subspecies of Phragmites australis in Ontario,Canada

Biological Invasions - Introduced Phragmites (Phragmites australis ssp. australis) forms part of an invasion assemblage in North America that includes non-native insect herbivores and parasitoids,...     

Long-term effects of ungulates on phytophagous insects

Abstract.  1. Most plants interact with a diverse suite of herbivores, allowing the opportunity for the existence of positive and negative interactions between highly dissimilar organisms. However, most studies on herbivorous interactions have been performed under the assumption that they occur mainly between similar species. Consequently, ecologists are still far from a full understanding of the ecological factors that determine insect population dynamics.
2. In this study, a 7-year field experiment was conducted that manipulated the presence of ungulates to evaluate their effects on the abundance, attack rate, and survival of four guilds of co-occurring herbivorous insects living on the same host plant: seed predators, stem borers, gall makers and sap suckers. These four guilds differed in habits and behaviour, the first three being sessile and endophytic and the last being free-living.
3. This study shows that the abundance of all four guilds was negatively affected by ungulates. However, the effect on attack rate differed among guilds, as mammals do not affect the seed predator attack rate. Ungulates also differentially affected insect survival, ingesting only seed predators and gall makers.
4. In summary, this study suggests that diverse mechanisms may affect different insect guilds in different ways. Therefore, competition between disparate herbivores appears to be complex and can be provoked by multiple mechanisms.     

The impact of resource limitation and the phenology of parasitoid attack on the duration of insect herbivore outbreaks

Fluctuations in resource quality and quantity, and changes in mortality due to predators and parasites are thought to be of prime importance in the regular fluctuations of forest insects. We examine how food limitation and parasitoids with different phenologies of attack regulate the population cycles of insect hosts. Our analysis of the limit cycle of a model with a biologically realistic form of density dependence in the host yields two novel predictions. First, outbreaks will typically last for only 2 generations after parasitoids begin to reduce the host population below the maximum density. Second, host growth rate is important in determining cycle length only when parasitoids attack before the impacts of resource limitation affect the host. The robustness of these predictions are tested using a more general form of density dependence in the host, revealing that our predictions are valid as long as density dependence in the host is not too overcompensatory.     

转基因抗虫作物对非靶标昆虫的影响

转基因抗虫作物自 1996年被批准商业化种植以来 ,它的抗虫性和经济效益已得到了普遍肯定 ,同时 ,转基因抗虫作物对非靶标生物的影响 ,如转基因抗虫作物的长期种植 ,是否会导致次要害虫上升为主要害虫 ,是否会影响有益昆虫 ,包括重要经济昆虫、捕食性和寄生性天敌以及重要蝶类的种类及种群数量 ,已成为转基因抗虫作物生态风险评估的重要内容。一些研究结果表明 ,转基因抗虫作物在对靶标害虫有效控制的同时 ,一些对杀虫蛋白不敏感的非靶标害虫有加重危害的趋势 ,由于种植转基因抗虫作物 ,减少了化学农药的使用 ,客观上也使非靶标害虫种群数量上升 ,这对转基因抗虫作物害虫综合治理提出了新的要求。靶标害虫数量的减少直接影响了害虫天敌种群数量 ,靶标害虫取食转基因抗虫作物后发育迟缓 ,也间接影响了天敌昆虫的生长发育 ,转基因抗虫作物的花粉或花蜜是一些重要经济昆虫如蜜蜂、熊蜂和一些寄生蜂 ,甚至捕食性天敌的食物来源 ,或花粉飘落到一些鳞翅目昆虫如家蚕或重要蝶类昆虫的寄主植物上 ,直接或间接对这些昆虫造成一定影响。目前大多数研究表明转基因抗虫作物对非靶标昆虫 ,特别是对有益昆虫没有明显的不利影响 ,也有研究报道认为对某些有益昆虫有一定的不良影响。这为深入开展转基因抗虫作物的生态安全     

Can hyperparasitoids cause large‐scale outbreaks of insect herbivores?

Synchronous population fluctuations occur in many species and have large economic impacts, but remain poorly understood. Dispersal, climate and natural enemies have been hypothesized to cause synchronous population fluctuations across large areas. For example, insect herbivores cause extensive forest defoliation and have many natural enemies, such as parasitoids, that may cause landscape‐scale changes in density. Between outbreaks, parasitoid‐caused mortality of hosts/herbivores is high, but it drops substantially during outbreak episodes. Because of their essential role in regulating herbivore populations, we need to include parasitoids in spatial modelling approaches to more effectively manage insect defoliation. However, classic host‐parasitoid population models predict parasitoid density, and parasitoid density is difficult to relate to host‐level observations of parasitoid‐caused mortality. We constructed a novel model to study how parasitoids affect insect outbreaks at the landscape scale. The model represents metacommunity dynamics, in which herbivore regulation, colonisation and extinction are driven by interactions with the forest, primary parasitoids and hyperparasitoids. The model suggests that parasitoid spatial dynamics can produce landscape‐scale outbreaks. Our results propose the testable prediction that hyperparasitoid prevalence should increase just before the onset of an outbreak because of hyperparasitoid overexploitation. If verified empirically, hyperparasitoid distribution could provide a biotic indicator that an outbreak will occur.     

Advances in research of induced resistance to insects in cotton

Any change in a plant that occurs following herbivory or environmental factors is an induced response. These changes include phytochemical induction, increases in physical defenses, emission of volatiles that attract predators and parasitoids of herbivores, and reduction in plant nutritional quality for herbivores, which is termed induced resistance. Induced resistance has been demon-strated ubiquitously in plants. It is one of our goals to review what is known about the induced resistance to herbivorous insects in cotton, including three resistance secondary metabolites (terpenoid, tannin, and flavonoids) that are contained at any significant levels of resistance to herbivorous insects in cotton cultivates. In many cases, the quantities or quality of secondary metabolites in plant are changed after attacked by insects. This review focuses on induced plant resistance as quantitative or qualitative enhancement of defense mechanism against insect pests, especially on the abiotic-elicitors-induced resistance in cotton plants. The abiotic-elicitor of cupric chloride, an exogenous inorganic compound, may induce the second-ary metabolites accumulation and is referred to as a copper-inducible elicitor (CIE). Finally, we discuss how copper-inducible elicitor may be used in the Integrated Pest Management (IPM) system for cotton resistance control.     

The ecology and management of mammal invasions in forests

Populations of invasive wild mammals have contributed significantly to the total unwanted impacts of biological invasions. They are known to impact forest ecosystems globally, but reviews summarizing this information are currently lacking. Here we (1) review the ecological characteristics of mammal invasions in forests; (2) characterize the range of ecological impacts on forest communities and the economic consequences of those impacts; (3) review what is known about interactions between the impacts of invasive mammals and other drivers of global change; and (4) consider the complex ecological and socio-economic challenges of simultaneously managing multiple invasive mammals and native biota affected by them. The unwanted impacts of invasive herbivores and predators are intensifying in many parts of the world and the need to manage their impacts to prevent further loss of indigenous biodiversity and damage to productive assets is greater than ever. However, management needs to be conducted within appropriate social, cultural, ethical, and animal welfare frameworks. Achieving effective management of populations of mammals invasive in forest ecosystems will require the filling of many knowledge gaps, including: better understanding their impacts; strategic options and tactical solutions for managing them; and achieving social licence to operate.     

Insect herbivores as potential causes of mortality and adaptation in gallforming insects

Summary Recent studies have suggested that plant galls benefit only the insects living in them and not the host plants, and that galls are induced by insects primarily to improve the plant as a microenvironment or a food source. The potential advantage to insects of protection from their predators and parasitoids has been considered unclear and perhaps minor in importance. However, the potential threat to gallforming insects from other insect herbivores has usually been relatively neglected. This paper notes literature and observations which suggest that herbivores may either consume or be deterred by galls. Even soft leaf galls produced by Hormaphis and Phylloxera aphids appeared to deter some herbivores in the field. The threat of herbivory to galls might help explain general patterns of gall ecology and morphology, and deserves closer attention.