Neighbourhood of host plants influences oviposition decisions of a stem-boring weevil

The spatial arrangement of suitable host plants in the field may significantly constrain insects to find optimal hosts. Plant neighbours around a focal host plant can either lead to lower (associational resistance) or higher (associational susceptibility) herbivore loads. We tested whether the spatial arrangement of hosts of different suitability for the larval development of the shoot-base boring weevil Apion onopordi affects oviposition decisions in the field. Host plants in our study were healthy creeping thistles (Cirsium arvense; suboptimal hosts) and thistles infected by a rust pathogen (Puccinia punctiformis; optimal hosts). For analysis, we used nearest neighbour methods that disentangle the spatial distribution of organisms that are dependent on the position of other species (e.g. phytophagous insects and their host plants). Although theory predicts that the small-scale spatial infestation pattern can have major consequences for the population dynamics in insect–plant systems, field studies quantifying spatial pattern of phytophagous insects are rare.

The spatial arrangement of host plants clearly influenced oviposition pattern in A. onopordi. In contrast to previous studies, we demonstrated that not the rust infection itself determined if a plant was infested by weevils, but rather the density of rusted shoots within a certain neighbourhood. We found strong indications for associational susceptibility of healthy thistle shoots to weevil oviposition when growing in the neighbourhood of rusted thistles. Weevil-infested plants were spatially aggregated, indicating that A. onopordi is limited in its dispersal ability within patches. Other stem-boring insects on creeping thistle were affected in their oviposition decisions by other factors than A. onopordi. Thus, it may be difficult to find general rules for oviposition choice in phytophagous insects.

Our study showed that the spatial arrangement of host plants in the field critically determines oviposition choice and should thus be included as constraint in theories of optimal host selection.     

Establishing systemic rust infections in Cirsium arvense in the field

An existing method to induce systemic rust infections in creeping thistles (Cirsium arvense) using an insect vector, the weevil Ceratapion onopordi, was successfully applied in the field. Urediniospores of Puccinia punctiformis were experimentally transmitted to naturally grown thistles, to our knowledge for the first time resulting in systemically infected shoots in an established thistle population.     

Direct and Indirect Effects of a Shoot-Base Boring Weevil and Plant Competition on the Performance of Creeping Thistle, Cirsium arvense

Creeping thistle or Canada thistle, Cirsium arvense (L.) Scop., is considered one of the world's worst weeds and the third most important weed in Europe. Biological control of this indigenous weed in Europe by use of native agents may provide a low-cost alternative to use of chemical or mechanical control measures and contribute to a more sustainable weed management. We investigated the potential of a shoot-base boring weevil, Apion onopordi Kirby (Coleoptera: Apionidae), for biological weed control, in the presence or absence of plant competition by three grass species. Infestation of thistle shoots by A. onopordi at natural infestation levels reduced above- and belowground plant performance after 2 years. Plant competition at natural levels had an overall greater effect than that of herbivory, significantly reducing both above- and belowground thistle performance in both years, thereby slowing the propagation of the weed. Weevil infestation and grass competition had a synergistic effect on C. arvense growth; the combined effects of the two factors was greater than the sum of both single-factor effects. The experiment revealed that A. onopordi promotes systemic infections of the rust fungus Puccinia punctiformis (Str.) Röhl in the year following weevil infestation. Systemically infected thistle shoots died before the end of the growing season. Although the direct effect of A. onopordi may not be sufficient to control creeping thistle, the synergistic interaction with plant competition and the indirect effect via promotion of systemic rust infections makes A. onopordi a promising agent for the biological control of this weed.     

Different bioassays for investigating orientation responses of the banana weevil, Cosmopolites sordidus, show additive effects of host plant volatiles and a synthetic male-produced aggregation pheromone

Abstract Three different bioassay methods to investigate the orientation behaviour of the banana weevil, Cosmopolites sordidus (Germar) (Coleoptera: Curculionidae), to host plant volatiles and a synthetic pheromone (cosmolure+) were compared. A locomotion compensator was used to separately record walking tracks in response to three odour sources. The data show that C. sordidus uses odour‐conditioned anemotaxis in its orientation to the odour sources tested. Of the two olfactometers tested, a dual port olfactometer using a continuous airflow showed stronger discrimination by C. sordidus to the different odours compared with a double pitfall olfactometer. The results of all three bioassays indicate that C. sordidus responds in an additive way to the combination of fermentation plant volatiles and the synthetic pheromone.     

Differential acceptance of and survivorship on a non‐host plant between two geographically separate populations of the phytophagous ladybird beetle Henosepilachna yasutomii

Phytophagous specialists among insects occasionally accept plants they do not or only rarely utilize under natural conditions. This non‐host acceptance could represent an initial stage in host‐range evolution. Here, we examined adult acceptance of and larval survivorship on a non‐host plant – wild thistle, Cirsium kamtschaticum Ledeb. (Asteraceae) – for two geographically separate populations of the phytophagous ladybird beetle Henosepilachna yasutomii Katakura (Coleoptera: Coccinellidae), which normally utilizes blue cohosh, Caulophyllum robustum Maxim. (Berberidaceae), in the wild. The two beetle populations showed lower acceptance of and survivorship on the non‐host plant (thistle) than on the normal host plant (blue cohosh). Furthermore, they differed significantly in their responses to thistle but not to blue cohosh. Even for the beetle population that more readily accepted thistle as adults, acceptance was moderate and the eclosion rate was low, indicating reduced performance on the non‐host plant. Although studies have infrequently focused on non‐host acceptance, this is likely common in phytophagous specialists and thus is a potentially important evolutionary factor, as it may determine the future direction of food‐range evolution.     

Orientation of the pine weevil Hylobius abietis to underground sources of host volatiles

Adults of Hylobius abietis (L.) (Coleoptera: Curculionidae) were found to locate conifer roots suitable for oviposition by utilizing host volatiles diffusing through the soil. Underground sources of host volatiles were presented to weevils in a laboratory bioassay. A cold-trapping condensate of Scots pine, Pinus sylvestris L., and fractions of it were tested. Various fractions containing host terpenes attracted weevils in the bioassay, but the complete pine condensate caused the highest response. Ethanol was also found to be attractive. Weevils caged underground in the absence of host material did not attract weevils on the surface.

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Zusammenfassung Rüsselkäfer (Hylobius abietis) können auf der Erdoberfläche die Lage im Boden verborgenen Brutmaterials feststellen und Wurzeln senkrecht grabend auffinden. Mit einer neu entwickelten Methode für Laborversuche, die sich auf die spezifische Reaktion des Eingrabens zum Brutmaterial gründet, wurden das Orientierungsverhalten der Käfer und die zum Brutmaterial führenden Geruchsstoffe des Wirtes studiert.Beide Geschlechter von H. abietis reagierten gleichartig mit Eingraben auf die Geruchsstoffe des Wirtes. In Wahlversuchen zwischen gleichwertigen Anlockungspunkten kam häufig Aggregation der Käfer an einem Punkt vor. Die stärkere Ansammlung an einzelnen Stellen stand im Zusammenhang mit dem Vorhandensein einer Erdröhre zur Quelle der Geruchsstoffe. In den folgenden Versuchen wurde deshalb den Tieren jeweils nur eine Geruchsstoffquelle angeboten. Weder Weibchen noch Männchen im Boden übten eine Anziehung auf Käfer an der Oberfläche aus.Kiefernstücke und Kondensate flüchtiger Kieferinhaltsstoffe sowie Kondensat nach Passage durch die Kolonne des Gaschromatographen waren stark attraktiv. Alle durch präparative Gaschromatographie hergestellten Fraktionen des Kondensats waren ebenfalls, aber schwächer attraktiv. Zehnfache Verdünnung des Kondensats und der Fraktionen verminderte ihre Attraktivität mit etwa 40–70%. Auch Äthanol, Methanol und Pentan wurden geprüft; nur Äthanol hatte eine mässig anlockende Wirkung.

     

Response of the parasitoidEupelmus vuilleti to the odours of the phytophagous host and its host plant in an olfactometer

Eupelmus vuilleti (CRW) is a solitary ectoparasitoid which, under our experimental conditions, developed at the expense of the larvae and pupae ofBruchidius atrolineatus (Pic) (Coleoptera, Bruchidae). The larvae of this beetle are found inside the seeds of cowpea,Vigna unguiculata (Walp). Using a tubular olfactometer, we determined the olfactory stimuli involved in the orientation of females of the hymenopterous speciesE. vuilleti towards one of its hosts;B. atrolineatus. Several odours were tested including those from host plants ofB. atrolineatus, from non-host plants, from the host itself and from the waste products of the host. The odours of pods or dry seeds ofV. unguiculata (host plant of the phytophagous insect) affected the locomotor behaviour of females ofE. vuilleti. The odours from the plant in the vegetative stage had no effect. Kairomones from larvae, pupae or faeces ofB. atrolineatus were attractive to the parasitoid females. Odours from adult male or femaleB. atrolineatus, as well as the sex pheromone emitted by females were not attractive toE. vuilleti females. This sex pheromone did however stimulate the locomotor activity of the parasitoid. The adaptive significance of these olfactory stimuli is discussed.

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Résumé Eupelmus vuilleti (CRW) est un ectoparasito?de solitaire qui se développe dans nos conditions expérimentales aux dépens des larves et des nymphes deBruchidius atrolineatus (Pic) (Coléoptère:Bruchidae). Les larves de ce Coléoptère se trouvent à l'intérieur des graines d'une LégumineuseVigna unguiculata (Walp). Grace à un olfactomètre tubulaire, nous avons déterminé les différents stimuli olfactifs intervenant dans l'orientation des femelles de cet hyménoptère, vers l;un de ses h?tes;Bruchidius atrolineatus. Plusieurs odeurs émanant soit de plantes-h?tes ou nonh?tes deB. atrolineatus, soit de l'h?te lui-même ou résultant de son métabolisme ont été testées. Les odeurs de gousses ou de graines sèches deVigna unguiculata Walp (plante-h?te du phytophage) influencent le comportement locomoteur des femelles. Par contre, les odeurs de plantes au stade végétatif n'ont pas d'effet. Les kairomones émanant de larves, nymphes ou fèces de larves deB. atrolineatus exercent un effect attractif sur les femelles parasito?des. Les odeurs émanant d'adultes males ou femelles deB. atrolineatus ainsi que la phéromone sexuelle émise par les femelles ne sont pas attractives pour les femelles deE. vuilleti. Cette phéromone sexuelle, par contre, stimule l'activité locomotrice des parasito?des. La signification adaptative de ces différents stimuli olfactifs est discutée.

     

Non‐host plant essential oil volatiles with potential for a ‘push‐pull’ strategy to control the tea green leafhopper,Empoasca vitis

The tea green leafhopper, Empoasca vitis Göthe (Hemiptera: Cicadellidae), is an economically important pest of tea crops, Camellia sinensis (L.) O. Kuntze (Theaceae), in China. The use of non‐host plant essential oils for manipulation of E. vitis was investigated for potential incorporation into a ‘push‐pull’ control strategy for this pest. The effectiveness of 14 plant essential oils in repelling E. vitis was investigated in laboratory assays. Rosemary oil, geranium oil, lavender oil, cinnamon oil, and basil oil repelled leafhoppers in a Y‐shaped olfactometer. We also compared the efficacy of these five plant essential oils to repel E. vitis in the presence of a host plant volatile‐based leafhopper attractant, (Z)‐3‐hexenyl acetate, in a tea plantation. In the treatment combination, four plates (north, south, east, and west) treated with an essential oil surrounded a central sticky plate treated with (Z)‐3‐hexenyl acetate. Fewer E. vitis were found on the plates treated with rosemary oil (12.5% reduction) than on the four water‐sprayed control treatment plates surrounding a central plate with (Z)‐3‐hexenyl acetate. We compared the distribution of E. vitis on the plates, and the relative numbers of E. vitis on each plate were compared with similar plates in the control treatment. When four plates treated with rosemary oil surrounded a central (Z)‐3‐hexenyl acetate‐treated plate, the distribution of E. vitis on the different plates changed significantly compared with that of the control. Relatively fewer E. vitis were found on the east (13.0% reduction) rosemary oil‐treated plates and more E. vitis (11.3% increase) were found on the central attractant‐treated plate. Our findings indicate that rosemary oil is a promising leafhopper repellent that should be tested further in a ‘push‐pull’ strategy for control of E. vitis.     

Geographic variation in oviposition choice of a leaf beetle: the relationship between host plant ranking, specificity, and motivation

The degree of adaptation of herbivorous insects to their local flora is an important component of the evolutionary processes that lead to host plant specialization in insects. In this study we investigated geographic variations in the oviposition preference of the leaf beetle Oreina elongata Suffrian (Coleoptera: Chrysomelidae: Chrysolini) in relation to differences in host plant specialization, in the field. We focused on the mechanisms of host choice and asked whether potential differences among populations are due to variations in host plant ranking and/or host plant specificity. We performed a combination of simultaneous choice and sequential no‐choice experiments with two of the major host plants of the beetle [Cirsium spinosissimum (L.) and Adenostyles alliariae (Gouan) (Asteraceae)]. The results suggested that spatial variation in host plant specialization has resulted in differences between populations in some aspects of the oviposition choice of O. elongata, while other aspects seem unaffected. We found no variation in host plant ranking among populations, as estimated in simultaneous choice tests. In contrast, the sequential no‐choice test indicated that host plant specificity was lower in a population that never encountered the highest ranked plant in the field. This finding agreed with our expectations, and we discuss our results in relation to the commonly used hierarchical threshold model. The results suggested that the mechanism for the differences in specificity is the variation among populations in the general motivation to oviposit, rather than quantitative differences in relative preference for the two hosts. We stress that it is essential to establish which of the two mechanisms is most important, as it will affect the probability of evolutionary change in host plant ranking.     

Diapause incidence of progeny in relation to parental geographic origin,host plant and rearing density in the cabbage beetle,Colaphellus bowringi

The cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), which is widely distributed in China, undergoes an imaginal diapause in the soil. In this study, we investigated the influences of parental geographic origin, host plant, and rearing density on the diapause incidence of progeny in this beetle. In studies conducted at 25 °C and L12:D12 photoperiod, the proportions of adults entering diapause were significantly different among latitudes from which the parents had been collected. The incidence of diapause increased with increasing latitude. Reciprocal cross tests between post‐diapause adults from different latitudes showed that there were significant differences in diapause incidence between pure‐bred and hybrid adults, suggesting that diapause incidence among progeny was determined by both the female and male, although the female appeared to have a stronger effect than the male. These results revealed that parental origin has an important influence on progeny diapause. Parental host plant species not only affected diapause incidence of the current generation, but also affected the incidence of diapause in the progeny produced by the non‐diapause parents; yet, the parental host plant had no influence on the incidence of diapause in the progeny produced by the post‐diapause parents. Rearing density of the parental generation had no significant effect on the incidence of diapause in the progeny.     

Sex ratio in catches of American mink – How to catch the females

To protect seabirds breeding on small nearshore islands (“sites”) along the coast of west Scotland from American mink, continuous year-round conventional cage trapping was carried out at constant effort at ten sites for three years (2003–2005) and at five of these sites for 10 years (1997–2006). In both periods, trapped male mink outnumbered females for most of the year, usually by more than two to one, with peaks of males in April–May and in September–October. However, a striking change took place in August–September, when the sex ratio of catches became close to unity. Most of the trapped females were caught in these two months. The chance of catching a female by this method was seven or eight times greater in August–September than during the rest of the year. Relatively few mink of either sex were caught in November–February. These findings, particularly the greatly increased catch of females in August–September, may be useful to others working to control alien American mink.     

Molecular tracking of individual host use in the Shiny Cowbird – a generalist brood parasite

Generalist parasites exploit multiple host species at the population level, but the individual parasite's strategy may be either itself a generalist or a specialist pattern of host species use. Here, we studied the relationship between host availability and host use in the individual parasitism patterns of the Shiny Cowbird Molothrus bonariensis, a generalist avian obligate brood parasite that parasitizes an extreme range of hosts. Using five microsatellite markers and an 1120‐bp fragment of the mtDNA control region, we reconstructed full‐sibling groups from 359 cowbird eggs and chicks found in nests of the two most frequent hosts in our study area, the Chalk‐browed Mockingbird Mimus saturninus and the House Wren Troglodytes aedon. We were able to infer the laying behavior of 17 different females a posteriori and found that they were mostly faithful to a particular laying area and host species along the entire reproductive season and did not avoid using previously parasitized nests (multiple parasitism) even when other nests were available for parasitism. Moreover, we found females using the same host nest more than once (repeated parasitism), which had not been previously reported for this species. We also found few females parasitizing more than one host species. The use of an alternative host was not related to the main hosts' nest availability. Overall, female shiny cowbirds use a spatially structured and host species specific approach for parasitism, but they do so nonexclusively, resulting in both detectable levels of multiple parasitism and generalism at the level of individual parasites.     

How to become a tree without wood – biomechanical analysis of the stem of Carica papaya L

Carica papaya L. does not contain wood, according to the botanical definition of wood as lignified secondary xylem. Despite its parenchymatous secondary xylem, these plants are able to grow up to 10‐m high. This is surprising, as wooden structural elements are the ubiquitous strategy for supporting height growth in plants. Proposed possible alternative principles to explain the compensation for lack of wood in C. papaya are turgor pressure of the parenchyma, lignified phloem fibres in the bark, or a combination of the two. Interestingly, lignified tissue comprises only 5–8% of the entire stem mass. Furthermore, the phloem fibres do not form a compact tube enclosing the xylem, but instead form a mesh tubular structure. To investigate the mechanism of papaya's unusually high mechanical strength, a set of mechanical measurements were undertaken on whole stems and tissue sections of secondary phloem and xylem. The structural Young's modulus of mature stems reached 2.5 GPa. Since this is low compared to woody plants, the flexural rigidity of papaya stem construction may mainly be based on a higher second moment of inertia. Additionally, stem turgor pressure was determined indirectly by immersing specimens in sucrose solutions of different osmolalities, followed by mechanical tests; turgor pressure was between 0.82 and 1.25 MPa, indicating that turgor is essential for flexural rigidity of the entire stem.     

The responses of the cabbage seed weevil Ceutorhynchus assimilis to volatile compounds from oilseed rape in a linear track olfactometer

The cabbage seed weevil, Ceutorhynchus assimilis Payk. [syn. Ceutorhynchus obstrictus (Marsham)] (Coleoptera: Curculionidae), a crucifer-feeding insect, is a pest of oilseed rape (Brassica napus L.). It is known to be attracted by isothiocyanates, crucifer-specific volatiles that are metabolites of the glucosinolates. The responses of this insect to other electrophysiologically-active volatiles from rape were tested in a linear track olfactometer. Attraction was demonstrated to nitriles (phenylacetonitrile, 4-pentenenitrile and 5-hexenenitrile), which are also glucosinolate metabolites, and to volatiles emitted by a wider spectrum of plant families ((Z)-3-hexen-1-ol and methyl salicylate). Combination of an isothiocyanate mixture with phenylacetonitrile increased attraction, but there was no such increase when the isothiocyanate mixture was combined with methyl salicylate. A mixture of 23 volatiles, emulating an attractive air-entrainment extract of oilseed rape, was not significantly attractive, although a high proportion of weevils (60%) turned towards it. The potential of these volatiles for inclusion into an isothiocyanate-based monitoring system is discussed.     

Response to host plant odors and aggregation pheromone by larvae of the Colorado potato beetle on a servosphere

Using a servosphere (locomotion compensator), locomotory behavior of Colorado potato beetle Leptinotarsa decemlineata Say (Coleoptera: Chrysomelidae) larvae was measured in detail in response to pulsed and non-pulsed odors of hostplant and conspecific pheromone. Second instars showed decreased straightness of movement, and all larvae showed decreased speed, in response to potato odor. Change in straightness by 2nd instars was also significantly affected by the interaction of pheromone and pulsing treatments. Fourth instars showed increased positive anemotaxis in response to the combined hostplant and pheromone odors. To our knowledge, this is the first demonstration of larval behavioral responses to adult pheromone in a holometabolous insect.     

非嗜食寄主气味蛹期处理对不同寄主来源美洲斑潜蝇雌成虫嗅觉定向的影响

测定用菜豆和丝瓜饲养的美洲斑潜蝇Liriomyza sativae Blanchard蛹暴露于用正己烷稀释的番茄或南瓜叶片挥发性抽提物(TLE或PLE)72h后,羽化雌成虫在Y形嗅觉仪中对TLE或PLE的定向率、平均反应时间以及对正己烷的定向率和平均反应时间,比较幼虫寄主经历和寄主气味对美洲斑潜蝇成虫嗅觉记忆的影响。结果表明,蛹期使用非嗜食寄主植物气味处理,能增加美洲斑潜蝇羽化雌成虫对非嗜食寄主气味的定向;幼虫期的寄主经历对处理后的成虫嗅觉定向也产生影响,饲喂丝瓜的美洲斑潜蝇蛹期处理后,羽化雌成虫显著偏向选择处理气味而非正己烷,饲喂菜豆的美洲斑潜蝇对南瓜的气味也表现出同样的特点,但对番茄的气味则相反,即使处理后对番茄气味的选择比率增加,仍然偏向选择正己烷。     

Trade‐off between attraction to aggregation pheromones and repellent effects of spike lavender essential oil and its main constituent linalool in the flour beetle Tribolium confusum

Essential oils of aromatic plants and their individual volatile components have been tested in pest management strategies for their toxic and often repellent effects on target insects. When evaluating their possible effects on crucial behaviours of the pest insects, the olfactory environment including intraspecific communication cues has to be considered. We used the flour beetle Tribolium confusum du Val (Coleoptera: Tenebrionidae), a common stored‐product pest, to investigate the influence of oil of spike lavender, Lavandula spica Medik. (Labiaceae), and its main component, linalool, at various doses on olfactory‐guided behaviour. Using four‐way olfactometers, a dose‐dependent repellent effect of L. spica oil and linalool alone was revealed. On the other hand, we confirmed that T. confusum is attracted by conspecifics, by means of an aggregation pheromone and by 10 ng of one of its components, 1‐tetradecene. Twenty‐four hour pre‐exposure to 10 μl of L. spica oil abolished subsequent attraction to 1‐tetradecene and reduced attraction to five conspecifics. Simultaneous exposure to L. spica oil or linalool and five conspecifics reduced the repellent effect of the volatiles in a dose‐dependent manner, whereas simultaneous exposure to 1‐tetradecene at 10 ng abolished the repellent effect of L. spica oil only at a dose of 0.01 mg. These results indicate a dose‐dependent trade‐off between attractive and plant‐derived repellent volatiles, which may influence the effectiveness of such volatiles in their potential use in alternative pest management strategies.     

Non‐random segregation of an autosomal gene in males of the flea beetle,Phyllotreta nemorum: implications for colonization of a novel host plant

The flea beetle, Phyllotreta nemorum (L.) (Coleoptera: Chrysomelidae: Alticinae), is currently expanding its host plant range in Europe. The ability to utilize a novel host plant, Barbarea vulgaris R. Br. (Brassicaceae), is controlled by major dominant genes named R‐genes. The present study used extensive crossing experiments to illustrate a peculiar mode of inheritance of the R‐gene in a population from Delemont (Switzerland). When resistant males from Delemont are mated with recessive females from a laboratory line, the female F₁ offspring contains the R‐allele and is able to utilize B. vulgaris, whereas the male offspring contains the r‐allele and is unable to utilize the plant. This outcome suggests X‐linkage of the R‐gene, but further crossing experiments demonstrated that this was not the case. When the R‐gene is present in offspring from males from a laboratory line that originates from Taastrup (Denmark), it is transmitted to female and male offspring in equal proportions as a normal autosomal gene. The results demonstrate a polymorphism in segregation patterns of an autosomal R‐gene in P. nemorum males. Males from Delemont contain a factor which causes non‐random segregation of the R‐gene (NRS‐factor). This factor is inherited patrilineally (from fathers to sons). Males with the NRS‐factor transmit the R‐gene to their female offspring, whereas males without the NRS‐factor transmit the R‐gene to female and male offspring in equal proportions. Various models for the non‐random segregation of autosomes in P. nemorum males are discussed – e.g., fusions between autosomes and sex chromosomes, and genomic imprinting. The implications of various modes of inheritance of R‐genes for the ability of P. nemorum populations to colonize novel patches of B. vulgaris are discussed.