Electrophysiological activity of the Sirex noctilio ovipositor: You know the drill?

Sirex woodwasp (Sirex noctilio F: Hymenoptera: Siricidae) is a major worldwide pest of pine (Pinus) species. The female woodwasp undertakes exploratory drills with the ovipositor prior to egg-laying to assess tree suitability. Previous work has shown that this behaviour is associated with assessing the osmotic pressure of the tree. Here we show that, in addition, the ovipositor is electrophysiologically active and capable of detecting ethanol and chemical components of solvent extracts of pine needles and bark. Scanning electron micrographs of the ovipositor show the presence of structures which may have a chemoreceptive function. Our research expands our knowledge of the role that the Sirex ovipositor plays in egg-laying site selection.

Nature's Swiss Army Knives: Ovipositor Structure Mirrors Ecology in a Multitrophic Fig Wasp Community

Background: Resource partitioning is facilitated by adaptations along niche dimensions that range from morphology to behaviour. The exploitation of hidden resources may require specially adapted morphological or sensory tools for resource location and utilisation. Differences in tool diversity and complexity can determine not only how many species can utilize these hidden resources but also how they do so. Methodology and Principal Findings: The sclerotisation, gross morphology and ultrastructure of the ovipositors of a seven-member community of parasitic wasps comprising of gallers and parasitoids developing within the globular syconia (closed inflorescences) of Ficus racemosa (Moraceae) was investigated. These wasps also differ in their parasitism mode (external versus internal oviposition) and their timing of oviposition into the expanding syconium during its development. The number and diversity of sensilla, as well as ovipositor teeth, increased from internally ovipositing to externally ovipositing species and from gallers to parasitoids. The extent of sclerotisation of the ovipositor tip matched the force required to penetrate the syconium at the time of oviposition of each species. The internally ovipositing pollinator had only one type of sensillum and a single notch on the ovipositor tip. Externally ovipositing species had multiple sensilla types and teeth on their ovipositors. Chemosensilla were most concentrated at ovipositor tips while mechanoreceptors were more widely distributed, facilitating the precise location of hidden hosts in these wasps which lack larval host-seeking behaviour. Ovipositor traits of one parasitoid differed from those of its syntopic galler congeners and clustered with those of parasitoids within a different wasp subfamily. Thus ovipositor tools can show lability based on adaptive necessity, and are not constrained by phylogeny. Conclusions/Significance: Ovipositor structure mirrored the increasingly complex trophic ecology and requirements for host accessibility in this parasite community. Ovipositor structure could be a useful surrogate for predicting the biology of parasites in other communities.

Host location by the fruit fly parasitoid Diachasmimorpha krausii : role of fruit fly species, life stage and host plant

1 Diachasmimorpha krausii is a braconid parasitoid of larval tephritid fruit flies, which feed cryptically within host fruit. At the ovipositor probing stage, the wasp cannot discriminate between hosts that are physiologically suitable or unsuitable for offspring development and must use other cues to locate suitable hosts. 2 To identify the cues used by the parasitoid to find suitable hosts, we offered, to free flying wasps, different combinations of three fruit fly species (Bactrocera tryoni, Bactrocera cacuminata, Bactrocera cucumis), different life stages of those flies (adults and larvae) and different host plants (Solanum lycopersicon, Solanum mauritianum, Cucurbita pepo). In the laboratory, the wasp will readily oviposit into larvae of all three flies but successfully develops only in B. tryoni. Bactrocera tryoni commonly infests S. lycopersicon (tomato), rarely S. mauritianum (wild tobacco) but never C. pepo (zucchini). The latter two plant species are common hosts for B. cacuminata and B. cucumis, respectively. 3 The parasitoid showed little or no response to uninfested plants of any of the test species. The presence of adult B. tryoni, however, increased parasitoid residency time on uninfested tomato. 4 When the three fruit types were all infested with larvae, parasitoid response was strongest to tomato, regardless of whether the larvae were physiologically suitable or unsuitable for offspring development. By contrast, zucchini was rarely visited by the wasp, even when infested with B. tryoni larvae. 5 Wild tobacco was infrequently visited when infested with B. cacuminata larvae but was more frequently visited, with greater parasitoid residency time and probing, when adult flies (either B. cacuminata or B. tryoni) were also present. 6 We conclude that herbivore-induced, nonspecific host fruit wound volatiles were the major cue used by foraging D. krausii. Although positive orientation to infested host plants is well known from previous studies on opiine braconids, the failure of the wasp to orientate to some plants even when infested with physiologically suitable larvae, and the secondary role played by adult fruit flies in wasp host searching, are newly-identified mechanisms that may aid parasitoid host location in environments where both physiologically suitable and unsuitable hosts occur.

The functional significance of fruit exocarp on host selection and oviposition by Queensland fruit fly, Bactrocera tryoni (Froggatt) (Tephritidae: Diptera)

Queensland fruit fly is Australia's most serious insect pest of horticulture. The fly lays its eggs into fruit, where they hatch into maggots which destroy the fruit. Understanding egg laying behaviour, known as oviposition, is a critical but under-researched aspect of fruit fly biology. This thesis focused on three aspects of oviposition: the role of fruit peel as a physical barrier to oviposition; the quality of fruit for maggot development; and the structure and wear of the egg laying organ – the ovipositor. Results showed that flies selected fruit based on their suitability for offspring survival, not because of the softness or hardness of fruit peel. Previously reported use of holes or wounds in fruit peel by ovipositing females was determined to be a mechanism which saved the female time, not a mechanism to reduce ovipositor wear. The results offer insights into the evolution of host use by fruit flies and their sustainable management.

Biomechanics of Substrate Boring by Fig Wasps: Role of Zinc in Insect Cuticle

There are many biomechanical challenges that a female insect must meet to successfully oviposit and ensure her evolutionary success. These begin with selection of a suitable substrate through which the ovipositor must penetrate without itself buckling or fracturing. The second phase corresponds to steering and manipulating the ovipositor to deliver eggs at desired locations. Finally, the insect must retract her ovipositor fast to avoid possible predation and repeat this process multiple times during her lifetime. From a materials perspective, insect oviposition is a fascinating problem and poses many questions. Specifically, are there diverse mechanisms that insects use to drill through hard substrates without itself buckling or fracturing? What are the structure-property relationships in the ovipositor material? These are some of the questions we address with a model system consisting of a parasitoid fig wasp - fig substrate system. To characterize the structure of ovipositors, we use scanning electron microscopy with a detector to quantify the presence of transition elements. Our results show that parasitoid ovipositors have teeth like structures on their tips and contain high amounts of zinc as compared to remote regions. Sensillae are present along the ovipositor to aid detection of chemical species and mechanical deformations. To quantify the material properties of parasitoid ovipositors, we use an atomic force microscope and show that tip regions have higher modulus as compared to remote regions. Finally, we use videography to show that ovipositors buckle during oviposition and estimate the forces needed to cause substrate boring based on Euler buckling analysis. Such methods may be useful for the design of functionally graded surgical tools.

Biomechanics of substrate boring by fig wasps

Female insects of diverse orders bore into substrates to deposit their eggs. Such insects must overcome several biomechanical challenges to successfully oviposit, which include the selection of suitable substrates through which the ovipositor can penetrate without itself fracturing. In many cases, the insect may also need to steer and manipulate the ovipositor within the substrate to deliver eggs at desired locations before rapidly retracting her ovipositor to avoid predation. In the case of female parasitoid ichneumonid wasps, this process is repeated multiple times during her lifetime, thus testing the ability of the ovipositioning apparatus to endure fracture and fatigue. What specific adaptations does the ovipositioning apparatus of a female ichneumonoid wasp possess to withstand these challenges? We addressed this question using a model system composed of parasitoid and pollinator fig wasps. First, we show that parasitoid ovipositor tips have teeth-like structures, preferentially enriched with zinc, unlike the smooth morphology of pollinator ovipositors. We describe sensillae present on the parasitoid ovipositor tip that are likely to aid in the detection of chemical species and mechanical deformations and sample microenvironments within the substrate. Second, using atomic force microscopy, we show that parasitoid tip regions have a higher modulus compared with regions proximal to the abdomen in parasitoid and pollinator ovipositors. Finally, we use videography to film wasps during substrate boring and analyse buckling of the ovipositor to estimate the forces required for substrate boring. Together, these results allow us to describe the biomechanical principles underlying substrate boring in parasitoid ichneumonid wasps. Such studies may be useful for the biomimetic design of surgical tools and in the use of novel mechanisms to bore through hard substrates.

Resposta de Gryon gallardoi (Brèthes) (Hymenoptera: Scelionidae) à qualidade dos ovos de Spartocera dentiventris (Berg) (Hemiptera: Coreidae) e evidências dos mecanismos de seleção envolvidos

Foi investigada, em condições de laboratório, a resposta de Gryon gallardoi a grupos de ovos de Spartocera dentiventris de diferentes idades (2, 3, 4, 5, 6, 7, 8 e 12 dias), registrando-se o efeito destas nos processos de seleção do parasitóide e na prole produzida. Os estímulos envolvidos em tais processos e a descrição morfológica das sensilas antenais também foram estudados. O aumento da idade do hospedeiro modificou o comportamento de oviposição, acarretando em um maior tempo de tamborilamento e de inserção do ovipositor, não influenciando o percentual de parasitismo, auto-parasitismo e a proporção de machos, porém acarretou em um maior tempo de desenvolvimento, redução no percentual de emergência, diminuição na aceitação das fêmeas e redução do tamanho corporal da prole do parasitóide. Testes em olfatômetro demonstraram que folhas de fumo provocaram uma maior resposta nas fêmeas recém-emergidas do parasitóide do que nas experientes. (Continua Através do contato, as fêmeas não diferenciaram extratos de ovos jovens e velhos, quando aplicados em ovos lavados com solvente. Ovos não parasitados, impregnados com extrato de parasitados, foram evitados. Cilindros de acrílico de maior tamanho, usados como hospedeiro artificial, resultou em um maior tempo de tamborilamento em relação aos menores. A análise cromatográfica detectou três substâncias diferentes em extrato de ovos de um dia de idade em relação a extrato de ovos de 5 e 12 dias. Nas antenas das fêmeas do parasitóide encontrou-se diferentes tipos de sensilas porosas e aporosas. Os resultados deste trabalho sugerem que a condição do hospedeiro acarreta em modificações no comportamento bem como no sucesso da prole, sendo a aceitação e a discriminação dos mesmos possivelmente influenciada por uma complexa mistura de estímulos.

Female reproductive system of the decapitating fly Pseudacteon wasmanni Schmitz (Diptera : Phoridae)

Pseudacteon wasmanni is a South American decapitating fly that parasitizes workers of Solenopsis fire ants. We used light microscopy (historesin serial-sectioning stained with Haematoxylin/Eosin) and scanning electron microscopy to show and analyze internal and whole external views of the female reproductive system. All specimens analyzed (n = 9) by light microscopy showed post-vitellogenic oocytes inside the ovaries. The lack of typical follicles (oocyte-nurse cell complexes) in all specimens suggests that oogenesis occurs during the pupal stage. The total number of eggs found ranged from 31 to 280 (X = 142 +/- 73, SD). The egg has a slugform or torpedo shape (about 130 by 20 mum) with a pointed apex at the posterior pole as defined by the fly; the micropyle appears to be in a depression or invagination at the anterior pole. An acute hypodermic-like ovipositor is evaginated from the hard sclerotized external genitalia during egg laying. The existence of a muscular bulb associated with the end of the common oviduct suggests that the egg is injected into the ant's body by a strong contraction of the bulb which probably is stimulated by bending of several ventral sensilla. During contraction, the abdomen extends out along a large fold between the sixth and seventh tergites in such a way that the sclerotized genitalia is rotated ventrally into a slightly anterior orientation in preparation for oviposition. (C) 2003 Elsevier B.V. Ltd. All rights reserved.

Dufour glands in the hymenopterans (Apidae, Formicidae, Vespidae): a review

Anexas ao aparelho do ferrão dos himenópteros aculeados encontram-se as glândulas de veneno e as de Dufour. A glândula de veneno é originada das glândulas associadas ao ovopositor dos himenópteros ancestrais não aculeados, já a glândula de Dufour é menos derivada, homóloga das glândulas colateriais dos outros insetos, sendo encontrada em todas as fêmeas dos himenópteros. Nestes insetos sua função é, em grande parte, desconhecida, mas, em formigas, parece estar envolvida com a comunicação e a defesa e, nas abelhas não sociais, com a construção e a proteção do ninho. Nas vespas pode estar relacionada ao reconhecimento parental. Foram observadas diferenças morfológicas e na composição química da secreção da glândula de Dufour entre as espécies, bem como na mesma espécie, entre as castas dos himenópteros sociais e entre indivíduos da mesma casta desempenhando diferentes funções ou pertencentes a ninhos diferentes. Portanto, nos himenópteros, sua função original de produzir substâncias para proteger os ovos ou favorecer a ovoposição parece ter sido substituída ou complementada com a função de produzir semioquímicos com função na comunicação.

SPATIAL AUTOCORRELATION OF MORPHOMETRIC VARIATION IN LUTOSA-BRASILIENSIS (BRUNNER VONWATTENWYL, 1888) (ORTHOPTERA, GRYLLACRIDOIDEA, HENICIDAE)

Geographic differentiation and sexual dimorphism in eighteen morphometric characters of Lutosa brasiliensis (Orthoptera: Henicidae) collected in eight localities of the State of São Paulo (Brazil) were analysed. A two-way Multivariate Analysis of Variance (MONOVA) was used to assess simultaneously the effects of sex and geographic location (plus their interaction) on morphometric variability. The spatial patterns of variation were analysed by Factor and Spatial Autocorrelation Analyses (Moran's I coefficient in four distance classes). Both indicate that the main direction of variation is, for males and females, a north-south cline in overall body size. In females, however, ovipositor length is not correlated with overall body size and displays a different pattern of variation over geographic space, indicating that distinct evolutionary forces produced the geographic differentiation in the species.

Análise filogenética de Scotaenini (Hymenoptera, Vespoidea, Tiphiidae, Thynninae)

Pós-graduação em Biologia Animal - IBILCE

A new genus and two new species of Opilioacaridae (Acari: Parasitiformes) from Amazonia, Brazil with a key to world genera

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Parasitoid-host interaction: sensory structures involved in the parasitism behavior of Bracon vulgaris (Hymenoptera: Braconidae)

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Aspectos morfológicos dos aparelhos reprodutores, maturação de ovários e avaliação do desenvolvimento de ninfas de Quesada gigas (Olivier, 1790) (Hemíptera: Cicadidae) em relação a diferentes hospedeiros

Pós-graduação em Agronomia (Entomologia Agrícola) - FCAV

Diversification in the use of resources by Idarnes species: bypassing functional constraints in the fig-fig wasp interaction

Mutualisms such as the figfig wasp mutualism are generally exploited by parasites. We demonstrate that amongst nonpollinating fig wasps (NPFWs) parasitic on Ficus citrifolia, a species of Idarnes galls flowers and another species feeds on galls induced by other wasps killing their larvae. The galling wasp inserts its ovipositor through the fig wall into the fig cavity. The ovipositor then follows a sinuous path and is introduced through the stigma and style of the flower. The egg is deposited between the integument and nucellus, in the exact location where the pollinating mutualistic wasp would have laid its egg. Gall induction is a complex process. In contrast, the path followed by the ovipositor of the other species is straightforward: attacking a larva within a developed gall poses different constraints. Shifts in feeding regime have occurred repeatedly in NPFWs. Monitoring traits associated with such repeated evolutionary shifts may help understand underlying functional constraints. (c) 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106, 114122.