Response of the fruit fly parasitoid Diachasmimorpha longicaudata to host and host‐habitat volatile cues

Chemical information is crucial to insect parasitoids for successful host location. Here, we evaluated the innate response of Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae), a fruit fly larval parasitoid, to cues from host and host habitat (i.e., fruit infested with host larvae). We first assessed the preference of female parasitoids between oranges infested with Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) and non‐infested fruit. Females were highly attracted towards infested oranges on the basis of volatile chemical cues. After this initial experiment, we aimed at revealing the potential sources of volatile cues present in an infested fruit. To this end, we considered five potential sources: (1) punctured fruit; (2) fly feeding, frass, or host‐marking pheromone deposited on the orange surface; (3) larval activity inside the fruit; (4) the larvae themselves; and (5) fungi associated with infestation of oranges. Habitat cues associated with host activity and those produced by rotten oranges or oranges colonized by fungi were highly attractive for female wasps, whereas odours associated with the activity of the adults on the surface of the fruit, and those released by the fruit after being damaged (as happens during fruit fly egg‐laying) were not used as cues by female parasitoids. Once the female had landed on the fruit, direct cues associated with larval activity became important although some indirect signals (e.g., products derived from larval activity inside the fruit) also increased host searching activity. Our findings indicate that naïve D. longicaudata uses chemical cues during host habitat searching and that these cues are produced both by the habitat and by the host larvae.     

Influence of presence and spatial arrangement of belowground insects on host‐plant selection of aboveground insects: a field study

Abstract 1. Several studies have shown that above‐ and belowground insects can interact by influencing each others growth, development, and survival when they feed on the same host‐plant. In natural systems, however, insects can make choices on which plants to oviposit and feed. A field experiment was carried out to determine if root‐feeding insects can influence feeding and oviposition preferences and decisions of naturally colonising foliar‐feeding insects. 2. Using the wild cruciferous plant Brassica nigra and larvae of the cabbage root fly Delia radicum as the belowground root‐feeding insect, naturally colonising populations of foliar‐feeding insects were monitored over the course of a summer season. 3. Groups of root‐infested and root‐uninfested B. nigra plants were placed in a meadow during June, July, and August of 2006 for periods of 3 days. The root‐infested and the root‐uninfested plants were either dispersed evenly or placed in clusters. Once daily, all leaves of each plant were carefully inspected and insects were removed and collected for identification. 4. The flea beetles Phyllotreta spp. and the aphid Brevicoryne brassicae were significantly more abundant on root‐uninfested (control) than on root‐infested plants. However, for B. brassicae this was only apparent when the plants were placed in clusters. Host‐plant selection by the generalist aphid M. persicae and oviposition preference by the specialist butterfly P. rapae, however, were not significantly influenced by root herbivory. 5. The results of this study show that the presence of root‐feeding insects can affect feeding and oviposition preferences of foliar‐feeding insects, even under natural conditions where many other interactions occur simultaneously. The results suggest that root‐feeding insects play a role in the structuring of aboveground communities of insects, but these effects depend on the insect species as well as on the spatial distribution of the root‐feeding insects.     

Behavioral evidence for contextual olfactory‐mediated avoidance of the ubiquitous phytopathogen Botrytis cinerea by Drosophila suzukii

Herbivorous insects may benefit from avoiding the smell produced by phytopathogens infecting plant host tissue if the infected tissue reduces insect fitness. However, in many cases the same species of phytopathogen can also infect host plant tissues that do not directly affect herbivore fitness. Thus, insects may benefit from differentiating between pathogen odors emanating from food and nonfood tissues. This is based on the hypothesis that unnecessarily staying attentive to pathogen odor from nonfood tissue may incur opportunity costs associated with not responding to other important survival functions. In this study adults of Drosophila suzukii Matsumura, an invasive larval frugivore, showed reduced attraction to the odor of raspberry fruit, a food tissue, when infected with Botrytis cinerea Pers., a ubiquitous phytopathogen, in favor of odors of uninfected raspberry fruit. Moreover, D. suzukii oviposited fewer eggs on infected raspberry fruit relative to uninfected raspberry fruit. Larval survival and adult size after eclosion were significantly reduced when reared on B. cinerea‐infected raspberry relative to uninfected fruit. Interestingly, when the behavioral choice experiment was repeated using Botrytis‐infected vs. ‐uninfected strawberry leaves, a nonfood tissue, in combination with fresh raspberry fruit, odor from B. cinerea‐infected leaves did not reduce D. suzukii attraction to raspberries relative to raspberries with uninfected leaves. These behavioral results illustrate the important role context can play in odor‐mediated interactions between insects, plants and microbes. We discuss implications of our findings for developing a repellent that can be useful for the management of D. suzukii.     

Plant odour plumes as mediators of plant–insect interactions

Insect olfactory orientation along odour plumes has been studied intensively with respect to pheromonal communication, whereas little knowledge is available on how plant odour plumes (POPs) affect olfactory searching by an insect for its host plants. The primary objective of this review is to examine the role of POPs in the attraction of insects. First, we consider parameters of an odour source and the environment which determine the size, shape and structure of an odour plume, and we apply that knowledge to POPs. Second, we compare characteristics of insect pheromonal plumes and POPs. We propose a ‘POP concept’ for the olfactory orientation of insects to plants. We suggest that: (i) an insect recognises a POP by means of plant volatile components that are encountered in concentrations higher than a threshold detection limit and that occur in a qualitative and quantitative blend indicating a resource; (ii) perception of the fine structure of a POP enables an insect to distinguish a POP from an unspecific odorous background and other interfering plumes; and (iii) an insect can follow several POPs to their sources, and may leave the track of one POP and switch to another one if this conveys a signal with higher reliability or indicates a more suitable resource. The POP concept proposed here may be a useful tool for research in olfactory‐mediated plant–insect interactions.     

Primary attraction and random landing in host‐selection by wood‐feeding insects: a matter of scale?

1 Most plant‐feeding insects show some degree of specialization and use a variety of cues to locate their host. Two main mechanisms of host location, primary attraction and random landing, have been investigated for such insects. 2 Research has led to contradictory conclusions about those hypotheses, especially for wood‐feeding insects; however, recent studies suggest that both mechanisms may take place in a single taxon but at different scales. 3 We developed a field experiment to test the hypothesis that primary attraction occurs at larger scale and random landing at finer scale in wood‐feeding insects. Landing rates, measured using sticky traps, were compared first between patches and then between individual trees according to their distance to a baited central tree. 4 Polynomial functions describing landing rate to distance relationships were compared with a function produced by a null model describing what should occur under the random landing hypothesis. Scolytidae and Cerambycidae (Coleoptera) responded to volatiles at the patch scale, supporting the primary attraction hypothesis, but the landing patterns of some groups at finer scale matched closely the predictions of our null model, giving support to the random landing hypothesis. 5 Our results show that the primary attraction and random landing hypotheses are not mutually exclusive and that prelanding use of host‐produced volatile is scale‐dependant. Scale considerations should thus be included in the study of prelanding host‐selection of wood‐feeding insects.     

Cues used in host‐seeking behavior by frog‐biting midges (Corethrella spp. Coquillet)

We investigated the role of carbon dioxide and host temperature in host attraction in frog‐biting midges (Corethrella spp). In these midges, females are known to use frog calls to localize their host, but the role of other host‐emitted cues has yet not been investigated. We hypothesized that carbon dioxide acts as a supplemental cue to frog calls. To test this hypothesis, we determined the responses of the midges to carbon dioxide, frog calls, and both cues. A significantly lower number of midges are attracted to carbon dioxide and silent traps than to traps broadcasting frog calls. Adding carbon dioxide to the calls does not increase the attractiveness to the midges. Instead, carbon dioxide can have deterrent effects on frog‐biting midges. Temperature of calling frogs is not a cue potentially available to the midges. Contrary to our hypothesis, there was no supplemental effect of carbon dioxide when presented in conjunction to calls. Midge host‐seeking behavior strongly depends on the mating calls emitted by their anuran host. Overall, non‐acoustic cues such as host body temperature and carbon dioxide are not important in long‐distance host location by frog‐biting midges.     

Reevaluating the conceptual framework for applied research on host‐plant resistance

Applied research on host‐plant resistance to arthropod pests has been guided over the past 60 years by a framework originally developed by Reginald Painter in his 1951 book, Insect Resistance in Crop Plants. Painter divided the “phenomena” of resistance into three “mechanisms,” nonpreference (later renamed antixenosis), antibiosis, and tolerance. The weaknesses of this framework are discussed. In particular, this trichotomous framework does not encompass all known mechanisms of resistance, and the antixenosis and antibiosis categories are ambiguous and inseparable in practice. These features have perhaps led to a simplistic approach to understanding arthropod resistance in crop plants. A dichotomous scheme is proposed as a replacement, with a major division between resistance (plant traits that limit injury to the plant) and tolerance (plant traits that reduce amount of yield loss per unit injury), and the resistance category subdivided into constitutive/inducible and direct/indirect subcategories. The most important benefits of adopting this dichotomous scheme are to more closely align the basic and applied literatures on plant resistance and to encourage a more mechanistic approach to studying plant resistance in crop plants. A more mechanistic approach will be needed to develop novel approaches for integrating plant resistance into pest management programs.     

Intraspecific variation in herbivore‐induced plant volatiles influences the spatial range of plant–parasitoid interactions

Chemical information influences the behaviour of many animals, thus affecting species interactions. Many animals forage for resources that are heterogeneously distributed in space and time, and have evolved foraging behaviour that utilizes information related to these resources. Herbivore‐induced plant volatiles (HIPVs), emitted by plants upon herbivore attack, provide information on herbivory to various animal species, including parasitoids. Little is known about the spatial scale at which plants attract parasitoids via HIPVs under field conditions and how intraspecific variation in HIPV emission affects this spatial scale. Here, we investigated the spatial scale of parasitoid attraction to two cabbage accessions that differ in relative preference of the parasitoid Cotesia glomerata when plants were damaged by Pieris brassicae caterpillars. Parasitoids were released in a field experiment with plants at distances of up to 60 m from the release site using intervals between plants of 10 or 20 m to assess parasitism rates over time and distance. Additionally, we observed host‐location behaviour of parasitoids in detail in a semi‐field tent experiment with plant spacing up to 8 m. Plant accession strongly affected successful host location in field set‐ups with 10 or 20 m intervals between plants. In the semi‐field set‐up, plant finding success by parasitoids decreased with increasing plant spacing, differed between plant accessions, and was higher for host‐infested plants than for uninfested plants. We demonstrate that parasitoids can be attracted to herbivore‐infested plants over large distances (10 m or 20 m) in the field, and that stronger plant attractiveness via HIPVs increases this distance (up to at least 20 m). Our study indicates that variation in plant traits can affect attraction distance, movement patterns of parasitoids, and ultimately spatial patterns of plant–insect interactions. It is therefore important to consider plant‐trait variation in HIPVs when studying animal foraging behaviour and multi‐trophic interactions in a spatial context.     

蚊虫搜寻吸血寄主和产卵行为的调节因子及相关嗅觉机理

嗅觉在蚊虫的吸血寄主搜寻、产卵和糖源搜寻行为中起决定作用,而在交配行为中的作用并不清楚。本文系统全面地综述了近20年来蚊虫化学生态学和嗅觉识别的分子机理的研究。蚊虫的触角、下颚须和口喙上的嗅觉感器感觉环境中释放的各种挥发性化合物。气味分子与嗅觉气味结合蛋白和气味受体的结合所启动的一系列生化反应产生神经动作电位。蚊虫嗅觉神经元编码气味中化合物的组成、浓度及其暂时瞬间的浓度变化和空间分布。吸血前后神经元的活性在数量和质量上有变化,反映了蚊虫在搜寻吸血寄主和产卵行为上的调节。在吸血寄主搜寻中,人体和动物释放的二氧化碳、乳酸以及其他气味协同引诱蚊虫向目标气味源定向飞行,最后找到吸血寄主。而成熟产卵雌蚊是利用产卵场所释放的腐烂气味寻找适宜的产卵场所,一些蚊虫卵、幼虫或蛹分泌的产卵信息素引诱和刺激雌蚊产卵,并与产卵生境气味起协同作用。植物气味尤其是花香味引诱蚊虫找到蜜源。驱避剂也是直接或间接通过嗅觉起作用,一些驱蚊剂由于阻断嗅觉反应而抑制蚊虫的定向飞行。从植物、动物或人体以及产卵场所释放的气味中有望找到有效的引诱和驱避化合物。对蚊虫嗅觉识别机理的认识将使我们开发出有效的蚊虫诱捕技术,进而应用于种群监测和控制。     

Olfactory cues used in host‐habitat location and host location by the parasitoid Cotesia urabae

There is a growing body of evidence that many hymenopteran parasitoids make use of olfaction as the primary mechanism to detect and locate hosts. In this study, a series of bioassays was conducted to investigate the orientation behaviour of the gum leaf skeletonizer larval parasitoid Cotesia urabae Austin & Allen (Hymenoptera: Braconidae) in both Y‐tube and four‐arm olfactometers. In a Y‐tube olfactometer, male C. urabae were attracted only to virgin conspecific females. Host‐plant leaves, damaged leaves, host larvae, and host larvae feeding on leaves were highly attractive to female C. urabae, whereas host frass and conspecific males were not. The multiple‐comparison bioassay conducted in a four‐arm olfactometer clearly indicates that C. urabae females were significantly more attracted to the host Uraba lugens Walker (Lepidoptera: Nolidae) larvae feeding on Eucalyptus fastigata H Deane & Maiden (Myrtaceae) leaves than to any other of the odour sources tested. The results of this study show that C. urabae individuals responded to chemical cues specific to the host plant and target host insect, and support hypotheses that unreliable cues are not utilized for host location by specific natural enemies.     

Chemical aspects of host‐acceptance behaviour in the bird cherry‐oat aphid Rhopalosiphum padi: host‐acceptance signals used by different morphs with the same genotype

Host acceptance by gynoparae and winged virginoparae of the bird cherry‐oat aphid Rhopalosiphum padi (L.) is investigated utilizing leaves and aqueous extracts of the primary and secondary hosts, as well as nonhost plants. Gynoparae are specialized to reproduce on bird cherry Prunus padus L., whereas virginoparae reproduce and feed on various grasses. Host acceptance is assessed using levels of reproduction and survival for adults, as well as survival for nymphs. Little is known of host acceptance by nymphs. The data show that gynoparae and winged virginoparae produce nymphs almost exclusively on their host plants, bird cherry and barley leaves, respectively, over 72 h. When tested with aqueous plant extracts, however, gynoparae produce nymphs almost exclusively on bird cherry extract and progeny numbers are found to be similar to those on plant leaves. Few nymphs are produced on artificial diet. By contrast, winged virginoparae produce nymphs on aqueous extracts of barley, bird cherry and bean, as well as on artificial diet. The numbers of nymphs deposited by gynoparae are similar on aqueous extracts of bird cherry leaves collected at different times during the growing season. When extracts from leaves of various Prunus species are tested, only leaves of P. padus and Prunus virginiana stimulate parturition. Oviparae, the sexual female nymphs of gynoparae, survive well for 96 h on both bird cherry and barley leaves but not on bean seedlings, whereas nymphs of winged virginoparae survive well only on barley leaves. They do not survive for 96 h on any plant‐leaf extracts, although they do survive on artificial diet.     

Effects of human–livestock–wildlife interactions on habitat in an eastern Kenya rangeland

Human–livestock–wildlife interactions have increased in Kenyan rangelands in recent years, but few attempts have been made to evaluate their impact on the rangeland habitat. This study identified drivers of increased human–livestock–wildlife interactions in the Meru Conservation Area between 1980 and 2000 and their effects on the vegetation community structure. The drivers were habitat fragmentation, decline in pastoral grazing range, loss of wildlife dispersal areas and increase in livestock population density. Agricultural encroachment increased by over 76% in the western zone adjoining Nyambene ranges and the southern Tharaka area, substantially reducing the pastoral grazing range and wildlife dispersal areas. Livestock population increased by 41%, subjecting areas left for pastoral grazing in the northern dispersal area to prolonged heavy grazing that gave woody plant species a competitive edge over herbaceous life‐forms. Consequently, open wooded grassland, which was the dominant vegetation community in 1980, decreased by c. 40% as bushland vegetation increased by 42%. A substantial proportion of agro pastoralists were encountered around Kinna and Rapsu, areas that were predominantly occupied by pastoralists three decades ago, indicating a possible shift in land use in order to spread risks associated with habitat alterations.     

Fertilisers and insect herbivores: a meta‐analysis

Despite the importance of a thorough understanding of the effect of synthetic fertiliser on insect population dynamics, existing literature is conflicting and an area of intense debate. Here, a categorical random‐effects meta‐analysis and a vote count meta‐analysis are employed to examine the effects of nitrogen (N), phosphorus (P), potassium (K) and NPK fertiliser on insect population dynamics. In agreement with the general consensus, insects were found to respond positively, overall, to fertilisers. Sucking insects showed a much stronger response to fertilisers than chewing insects. The environment in which a study is conducted can have a marked effect on insect responses to fertiliser, with natural environments showing the potential for buffering effects of nitrogen fertilisers in particular. As well as highlighting the potential shortfall in the amount of research investigating particularly the effects of potassium and phosphorus, this study provides an invaluable flag post in the ongoing research investigating fertiliser effects on ecosystems.     

Aspects of insect chemical ecology: exploitation of reception and detection as tools for deception of pests and beneficial insects

Empirical exploitation of insect reception and detection at the peripheral neurosensory level has been extremely valuable for identifying pheromones and other semiochemicals, mainly by electroantennogram or single cell preparations coupled with capillary gas chromatography. Differential sensitivity to semiochemicals at the single‐cell level has allowed the identification of some of the most active semiochemicals relating to host location and, more importantly, to the avoidance of nonhosts. However, in terms of molecular recognition, there is still a considerable gap in understanding the detection of particular molecules and their discrimination from closely‐related chemical structures. New approaches will be needed to understand the processes of molecular recognition more precisely. Nevertheless, from electrophysiological studies to the most advanced molecular techniques, it has been possible to identify semiochemicals for the deception of pests in their quest to find plant and animal hosts, as well as mates. Even the deception of insects antagonistic to pests, particularly parasitoids, can now be exploited for managing pests in more sustainable systems. Successes in exploiting insect semiochemicals in the interests of better agriculture and animal husbandry are exemplified, and potential new ways of learning more about reception and detection for deception are discussed. This takes the subject beyond the management of pest and beneficial insects to wider commercial and social opportunities.     

Inter‐oceanic variation in patterns of host‐associated divergence in a seabird ectoparasite

Aim Parasites with global distributions and wide host spectra provide excellent models for exploring the factors that drive parasite diversification. Here, we tested the relative force of host and geography in shaping population structure of a widely distributed and common ectoparasite of colonial seabirds, the tick Ixodes uriae. Location Two natural geographic replicates of the system: numerous seabird colonies of the North Pacific and North Atlantic Ocean basins. Methods Using eight microsatellite markers and tick samples from a suite of multi‐specific seabird colonies, we examined tick population structure in the North Pacific and compare patterns of diversity and structure to those in the Atlantic basin. Analyses included population genetic estimations of diversity and population differentiation, exploratory multivariate analyses, and Bayesian clustering approaches. These different analyses explicitly took into account both the geographic distance among colonies and host use by the tick. Results Overall, little geographic structure was observed among Pacific tick populations. However, host‐related genetic differentiation was evident, but was variable among host types and lower than in the North Atlantic. Main conclusions Tick population structure is concordant with the genetic structure observed in seabird host species within each ocean basin, where seabird populations tend to be less structured in the North Pacific than in the North Atlantic. Reduced tick genetic structure in the North Pacific suggests that host movement among colonies, and thus tick dispersal, is higher in this region. In addition to information on parasite diversity and gene flow, our findings raise interesting questions about the subtle ways that host behaviour, distribution and phylogeographic history shape the genetics of associated parasites across geographic landscapes.     

The tri‐trophic niche concept and adaptive radiation of phytophagous insects

A conceptual divide exists between ecological and evolutionary approaches to understanding adaptive radiation, although the phenomenon is inherently both ecological and evolutionary. This divide is evident in studies of phytophagous insects, a highly diverse group that has been frequently investigated with the implicit or explicit goal of understanding its diversity. Whereas ecological studies of phytophagous insects increasingly recognize the importance of tri‐trophic interactions as determinants of niche dimensions such as host‐plant associations, evolutionary studies typically neglect the third trophic level. Here we attempt to reconcile ecological and evolutionary approaches through the concept of the ecological niche. We specifically present a tri‐trophic niche concept as a foil to the traditional bi‐trophic niche concept for phytophagous insects. We argue that these niche concepts have different implications for understanding herbivore community structure, population divergence, and evolutionary diversification. To this end, we offer contrasting empirical predictions of bi‐ and tri‐trophic niche concepts for patterns of community structure, the process of population divergence, and patterns of evolutionary diversification of phytophagous insects.     

Effects of cucumber mosaic virus‐infected chilli plants on non‐vector Bemisia tabaci (Hemiptera: Aleyrodidae)

Plant virus infections are known to alter host plant attractiveness and suitability for insect herbivores.This study was conducted to determine how cucumber mosaic virus (CMV)-infected chilli plants affect the fitness and settling preferences ofnonvector whitefly,Bemisia tabaci adults under dual-choice conditions with volatile organic compounds analyzed using solid phase microextraction coupled with gas chromatography-mass spectrometry (GC-MS).Results showed that the presence of CIVIV in chilli plants substantially affects the settling preferences of the B.tabaci,which preferred to settle on noninfected plants.Duration of the egg stage and the longevity and fecundity of adult B.tabaci on CMV-infected chilli plants were not markedly different from those on noninfected chilli plants.In contrast,the developmental time from egg to adult was significantly reduced in CMV-infected chilli plants compared to the noninfected plants.The results also showed that CMV-infected chilli plants released significantly more linalool and phenylacetaldehyde than noninfected plants.Overall,it was suggested that the behavioral response of B.tabaci might be modified by CMV-infected plants,which alter the release of specific headspace volatiles.Based on these results,the modification of plant volatile profiles may help in enhancing the effectiveness of biological control and the protection of crop plants against B.tabaci.     

Stimuli affecting selection of oviposition sites by female peach twig borer, Anarsia lineatella Zeller (Lepidoptera: Gelechiidae)

In laboratory and field experiments, stimuli were tested that might affect oviposition decisions by female peach twig borer moths, Anarsia lineatella Zeller (Lepidoptera: Gelechiidae). When given a choice between immature green peach fruits, green mature peach fruits and soft-ripe peach fruits, the latter received the fewest eggs. Fuzzy halves of peach fruits received ten times more eggs then shaved hairless halves. Volatiles from both almond and peach shoots induced more oviposition by females than by control stimuli. Similarly, volatiles from immature green peach fruits, mature green or mature hard-ripe peach fruits induced more oviposition than their respective control stimuli. In a choice experiment, volatiles from immature peach fruit stimulated three times more oviposition than those from soft-ripe peach fruit. Discrimination against mature soft-ripe peach fruits as potential oviposition sites may lie in the phenology of A. lineatella and host peach fruits. Larval development to the pupal stage takes 15–27 days. Therefore, any eggs laid on a ripe fruit 14 days before it falls from the tree will not likely develop into adult insects because developing larvae will only reach third or fourth instar before the fruit is decomposed, and only first and second instar larvae can overwinter.