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Impact of an Invasive Pest in Local Semiochemical Networks

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Impact of an Invasive Pest in Local Semiochemical Networks Dottorato di ricerca in Scienze Agrarie, Forestali e Ambientali Dipartimento di Scienze Agrarie, Alimentari e Forestali AGR/11 Entomologia generale e Applicata Impact of an invasive pest in local semiochemical networks DOTTORE COORDINATORE Dott. LETIZIA MARTORANA Prof. VINCENZO BAGARELLO TUTOR CO TUTOR Prof. STEFANO COLAZZA Prof. EZIO PERI CICLO XXXI 2019 Dottorato di ricerca in Scienze Agrarie, Forestali e Ambientali Dipartimento di Scienze Agrarie, Alimentari e Forestali AGR/11 Entomologia generale e Applicata Impact of an invasive pest in local semiochemical networks DOTTORE COORDINATORE Dott. LETIZIA MARTORANA Prof. VINCENZO BAGARELLO TUTOR CO TUTOR Prof. STEFANO COLAZZA Prof. EZIO PERI CICLO XXXI 2019 Table of contents Table of contents…………………………………………………………………pag. 1 Abstract…………………………………………………………………………..pag. 4 Riassunto…………………………………………………………………………pag. 6 Chapter 1 1.1 Introduction………………………………………………………….pag. 8 1.2 Study system…………………………………………………………pag. 12 1.2.1 Plant species……………………………………………...pag. 12 1.2.2 Phytophagous insect……………………………………..pag. 12 1.2.3 Zoophytophaugus insect…………………………………pag. 14 1.2.4 Parasitoid insect……………………………………….....pag. 15 1.3 Outline of this thesis…………………………………………………pag. 16 1.4 References……………………………………………………………pag. 18 Chapter 2 An invasive insect herbivore disrupts plant volatile-mediated tritrophic signalling………………………………………………………………………….pag. 27 2.1 Abstract……………………………………………………………… pag. 28 2.2 Introduction…………………………………………………………. pag. 29 2.3 Materials and methods………………………………………………pag. 31 2.3.1 Plants……………………………………………………...pag. 31 2.3.2 Insect rearing……………………………………………..pag. 31 2.3.3 Plant treatments………………………………………….pag. 32 2.3.4 Y-tube olfactometer bioassays…………………………..pag. 33 2.3.5 Statistical analysis………………………………………..pag. 34 2.4 Results………………………………………………………………...pag. 35 2.5 Discussion…………………………………………………………….pag. 36 2.6 References……………………………………………………………pag. 40 1 | P a g . Chapter 3 Attraction of the egg parasitoid, Telenomus podisi, to oviposition plant volatiles induced by the zoophytophagous predator, Podisus maculiventris, under biotic interactions with the invasive phytophagous pest, Halyomorpha halys……………..………………………………………………………………...pag. 45 3.1 Abstract……………………………………………………………….pag. 46 3.2 Introduction…………………………………………………………..pag. 47 3.3 Materials and methods……………………………………………….pag. 51 3.3.1 Plants………………………………………………………pag. 51 3.3.2 Insect rearing……………………………………………...pag. 51 3.3.3 Y-tube olfactometer bioassays …………………………...pag. 52 3.3.4 Plant treatments …………………………………………..pag. 52 3.3.5 Statistical analysis………………………………………….pag. 53 3.4 Results………………………………………………………………….pag. 54 3.5 Discussion………………………………………………………………pag. 56 3.6 References……………………………………………………………...pag. 60 Chapter 4 Role of contact semiochemicals in mediating response of local egg parasitoids to an invasive not associated host……………………………………………………….pag. 67 4.1 Abstract…………………………………………………………………pag. 68 4.2 Introduction…………………………………………………………….pag. 69 4.3 Materials and methods…………………………………………………pag. 72 4.3.1 Insect colonies……………………………………………….pag. 72 4.3.2 Parasitoid response to host chemical footprints…………..pag. 72 4.3.3 Host Preference……………………………………………...pag. 74 4.3.4 Host suitability……………………………………………....pag. 75 4.3.5 Statistical analysis…………………………………………..pag. 76 2 | P a g . 4.4 Results…………………………………………………………………...pag. 76 4.5 Discussion………………………………………………………………..pag. 78 4.6 References……………………………………………………………….pag. 82 Concluding remarks………………………………………………………………...pag. 87 Further prospective………………………………………………………………....pag. 89 Acknowledgment……………………………………………………………………pag. 90 3 | P a g . Abstract During the host location process, egg parasitoids rely on a series of chemical stimuli from the host/plant complex. In particular, in tritrophic systems consisting of plants, herbivorous stink bugs and their egg parasitoids, previous studies demonstrated that wasps are attracted by oviposition-induced plant volatiles (OIPVs). These compounds act as indirect plant defence mechanism allowing plants to recruit egg parasitoids, which are in general the most important natural enemies of stink bugs. Moreover, when wasps land on a plant, they can exploit chemical footprints left by male and female stink bugs walking on the leaves as indirect host-related cues, since chemicals lead them in an area where newly laid host eggs are likely to be found. Finally, since host female chemical footprints are the most promising signals of host eggs, wasps evolved the ability to distinguish between footprints left by females and males of their associated hosts. These foraging behaviours are adopted by egg parasitoids searching for native and associated hosts, but invasive insect herbivores can interfere with local multitrophic interactions by affecting the chemical cues emitted by plants and disrupting the attraction of egg parasitoids. In this thesis the potential impact of the brown marmorated stink bug, Halyomorpha halys (Stål) (Heteroptera: Pentatomidae) on local infochemical networks at long and short range was evaluated. Halyomorpha halys is a species native to Asia, which was recently introduced in the Europe and North America, and to evaluate the effect of its establishment, laboratory experiments were conducted using two local tritriphoc systems, in Europe, the one consisting of Vicia faba L. - Nezara viridula (L.) (Heteroptera: Pentatomidae) - Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae), and in North America, the one consisting of V. faba - Podisus maculiventris (Say) (Heteroptera: Pentatomidae) - Telenomus podisi (Ashmead) (Hymenoptera: Scelionidae). The results showed that both egg parasitoid species, T. basalis and T. podisi, were attracted by the OIPVs of their associated hosts, N. viridula and P. maculiventris, respectively, but not by the OIPVs of non-associated host H. halys. However, in both systems, the parasitoid attraction toward OIPVs emitted following the attack of the associated host was disrupted when host and non-associated host were concurrently present on the same plant. Concerning the substrate-born cues, T. basalis females (naïve or rewarded by an oviposition experience) ability to discriminate between female and male of H. halys were tested. Naïve T. basalis wasps did not discriminate between areas contaminated by chemical footprints left by the non-associated host female or male on the contrary experienced female of T. basalis, re-encountering after 1 hour, discriminated between male 4 | P a g . and female chemical footprints but they were not able to discriminate H. halys gender after 24 hours. This indicates that invasion by an alien herbivore can interfere with local established semiochemical webs. Overall the outcomes are discussed by evaluating the possible consequences of alien insect spread on local parasitoid foraging behaviour. 5 | P a g . Riassunto Durante il processo di localizzazione dell’ospite i parassitoidi oofagi si affidano a una serie di segnali chimici provenienti dal complesso pianta/ospite. In particolare studi precedenti hanno dimostrato che nei sistemi tri-trofici costituiti da piante coltivate, insetti fitofagi e i loro parassitoidi oofagi associati, questi ultimi sono attratti dagli “oviposition induced plant volatile” (OIPVs). Questi composti agiscono come meccanismo indiretto di difesa delle piante per attrarre i parassitoidi oofagi, che sono in generale tra i più importanti antagonisti degli insetti fitofagi tra cui i Pentatomidi. Inoltre quando i parassitoidi riescono ad arrivare sulla pianta sfruttano le tracce chimiche lasciate sulle foglie dagli ospiti di entrambi i sessi per essere guidati nelle zone dove è più probabile che trovino le uova dell’ospite. Infine poiché le tracce appartenenti alle femmine dell’ospite sono i segnali più promettenti per localizzare l’ovatura i parassitoidi oofagi hanno evoluto la capacità di distinguere tra le tracce lasciate dai maschi e delle femmine dei loro ospiti. Questo comportamento di foraggiamento è adottato dai parassitoidi per la ricerca degli ospiti associati, ma cosa accadrebbe se un insetto alieno erbivoro entrasse a fare parte di questo network infochimico? Per valutare il possibile impatto su di un sistema tri-trofico locale sia a breve che a lunga distanza, della cimice asiatica Haliomorpha halys (Stål) (Heteroptera: Pentatomidae), una specie nativa dell’Asia che recentemente è stata introdotta nel vecchio e nel nuovo mondo sono stati condotti esperimenti in condizione di laboratorio su due sistemi, uno Europeo costituito da Vicia faba L. - Nezara viridula (L.) (Heteroptera: Pentatomidae) - Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae), e l’altro tipico del Nord America costituito da V. faba - Podisus maculiventris (Say) (Heteroptera: Pentatomidae) - Telenomus podisi (Ashmead) (Hymenoptera: Scelionidae). I risultati hanno evidenziato che entrambi i parassitoidi T. basalis e T. podisi sono stati attratti dagli OIPVs indotti dall’azione di ovideposizione dei loro ospiti associati, rispettivamente N. viridula e P. maculiventris, ma non dagli OIPVs indotti dall’ospite non associato H. halys. In entrambi i sistemi tritrofici l’attrazione verso gli OIPVs indotti dai rispettivi ospiti associati N. viridula e P. maculiventris è risultata essere interrotta quando l’ospite associato e l’ospite non associato infestavano contemporaneamente la stessa pianta. Questo indica che l’invasione da parte di un insetto fitofago alieno interferisce con la ben stabilita rete semiochimica locale. Per quanto riguarda le tracce chimiche lasciate sul substrato dall’insetto
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