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A Review of Natural Parasitism Levels in Oak Galls in Great Britain

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A Review of Natural Parasitism Levels in Oak Galls in Great Britain A Defra Network partnership delivering interdisciplinary plant health FUTURE PROOFING research to improve biosecurity and build capability Plant Health Task 5.4.4 Collect baseline data on native oak gall parasitoids in England A review of natural parasitism levels in oak galls in Great Britain Rachel Down, Damian De Marzo, Chris Malumphy and Neil Audsley (Fera) 31st March 2019 Work Package 5 Control Contents Abstract …………………………………………………………………………………………………………………………………………. 3 Chapter 1. Introduction …………………………………………………………………………………………………………………. 4 Chapter 2. Cynipid oak galls of Great Britain …………………………………………………………………………….…… 6 Chapter 3. Interactions between oak galls, their associated communities and Torymus sinensis ….. 87 Summary ……………………………………………………………………………………………………………………………………. 104 References ……………………………………………………………………………………………………………………………….… 105 Parasitism levels in oak galls 2 Abstract Oak gall wasps (Hymenoptera: Cynipidae) are typically characterised by their complex life-cycles, alternating between sexual and asexual generations. The galls that they induce are both species and generation specific and show an astonishing degree of diversity and complexity. Oak galls also support a very rich and diverse, well-structured, community of parasitoids and cynipid inquilines (Hymenoptera: Cynipidae, tribe Synergini). The majority of parasitoids associated with cynipid oak galls belong to the hymenopteran super-family Chalcidoidea, and are distributed across six families: Pteromalidae, Eulophidae, Torymidae, Eurytomidae, Eupelmidae and Ormyridae. Inquiline species live within the cynipid galls but do not themselves induce primary gall formation. They are classified as lethal and non-lethal according to whether or not their chambers encroach upon those of the gall inducer thus killing the gall wasp larva inside. Relationships within the galls are complex: parasitoids can attack the gall wasp larvae, inquiline species, and other parasitoid species (known as hyperparasitism) present within the gall. Parasitoid species are rarely host specific and will usually attack many different species of gall wasp. However, the sexual and asexual generation galls usually support different species of parasitoids although it is often the case that the same generation of closely related gall wasps will support similar communities of parasitoids and inquilines. There are approximately 200 species of cynipid gall wasps, and 100 species of chalcid parasitoids and 38 species of inquilines recorded as associated with them across the western Palaearctic region. This review aims to provide life-cycle details, and levels of attack for associated parasitoids and inquilines for oak gall wasp species known to be present in Great Britain. This information is being gathered prior to the potential release of a non-native biocontrol agent, the chalcid parasitoid Torymus sinensis Kamijo (Hymenoptera: Torymidae), for control of the invasive Oriental chestnut gall wasp Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae), a species of gall wasp that attacks sweet chestnut trees (Castanea spp.). Concern has been raised that this potential biocontrol agent may be able to utilise native and naturalised oak gall wasps in Great Britain as an alternative host. Indeed, this has been observed in other areas of Europe (Italy) where T. sinensis has already been released. Obtaining baseline data on natural parasitism levels in oak galls prior to release will allow for effective monitoring of competition between T. sinensis and native parastoid species should T. sinensis be released into the environment. Parasitism levels in oak galls 3 Chapter 1. Introduction The Oriental or Asian chestnut gall wasp, Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae), is a pest species that attacks sweet chestnut trees (Castanea spp.). It is native to China but has been accidentally introduced into other geographical regions including Japan (Moriya et al., 2003), the USA (Payne et al., 1975), and western Europe in 2002 when it was discovered in Italy (Brussino et al., 2002). Since its discovery in Italy it has spread into numerous western European countries (Down and Audsely, 2016), and was first discovered in the UK in Farningham Wood, Kent in 2015 (Everatt, 2015; Malumphy, 2015). Since 2015 it has spread and is now found at multiple locations in London and the south east of England (Forestry Commission, 2018). Control and management options for Oriental chestnut gall wasp have previously been reviewed (Everatt, 2015; Down and Audsley, 2016), the authors concluding that the only feasible control option for managing D. kuriphilus is a classical biological control programme involving the release of a non- native parasitoid wasp. There are 30 species of chalcid parasitoid wasps known to attack D. kuriphilus in China, but one in particular, Torymus sinensis Kamijo (Hymenoptera: Torymidae), was found to be a very common parasitoid of D. kuriphilus, and the strain native to China is phenologically synchronised with the life cycle of D. kuriphilus thus providing good control of the gall wasp populations (Moriya et al., 2003). Torymus sinensis is univoltine. Adults emerge from D. kuriphilus galls in the early spring and oviposit inside fresh developing galls, on the surface of the gall wasp larvae. The T. sinensis larvae feed on the gall former and remain in the galls until the following spring. Torymus sinensis has been released in Japan, North America and Italy to control Oriental chestnut gall wasp and has been very successful (detailed in Down and Audsley, 2016); up to 95% parasitisation rates have been reported in parts of Italy (Ferracini and Alma, 2015). More recently other European countries such as France (2011; Borowiec et al., 2014), Croatia (2014; Matošević et al., 2014) and Hungary (2014; Matošević et al., 2014), have followed suit. Torymus sinensis has also been introduced into areas of Slovenia although surveys have since indicated that the parasitoid may have arrived from other areas of Europe by natural spread prior to the first deliberate introduction in 2015 (Jurc et al., 2017). Test releases have been reported in Spain and Portugal (Paparella et al., 2016), and preparations are currently underway in the UK to gain approval for the use of T. sinensis to control Oriental chestnut gall wasp. Prior to release of the exotic parasitoid wasp it is important to understand the native gall wasp communities and their intricately associated parasitoids and inquilines. An understanding of these communities may provide evidence of native parasitoid species with the potential to attack the invading gall wasp. Importantly, an understanding of these associations will also allow for effective monitoring of potential competition between the introduced parasitoid and native gall wasp parasitoids, which could occur if the host range of the introduced parasitoid is not limited to the invasive gall wasp species. Any such competition could be detrimental to native gall wasp communities. In Italy, damage to sweet chestnuts resulting from gall wasp attack was severe so a quick decision was made to release T. sinensis in an effort to reduce the economic and environmental losses associated with the pest, with the first successful release conducted in 2005 (Quacchia et al., 2008). At that point in time, although T. sinensis was thought to be host specific, its host range had not been thoroughly investigated (Quacchia et al., 2008); monitoring for host range has instead been mainly post-release. Parasitism levels in oak galls 4 Concerns have been previously raised over the continued use of T. sinensis as a biological control agent of Oriental chestnut gall wasp in Europe due to potential adverse effects, including the lack of information on its host range, potential hybridisation with native Torymus species, and the lack of evaluations on the direct and indirect non-target effects on the ecosytem (EFSA, 2010; Aebi et al., 2011; Gibbs et al., 2011). The European Food Safety Authority (EFSA, 2010) identified an undetermined risk that T. sinensis could parasitise native oak gall wasps and concluded that information on this risk was crucial before release occurred. Oak gall wasps have a very rich parasitoid community associated with them. This review identifies the oak gall wasp species present in the UK, provides detailed information on the life histories of these gall wasps, and details of their parasitoid wasp and inquiline associates. This review therefore aims to provide baseline information for these communites, which in conjunction with sampling in areas where D. kuriphilus is present (C. Malumphy, pers. comm.), will help to achieve effective monitoring for non-target effects and potential competition between T. sinensis and the native parasitoids of oak gall wasps, should the exotic parasitoid be released (or indeed, arrive naturally). Parasitism levels in oak galls 5 Chapter 2. Cynipid oak galls of Great Britain In 2010, EFSA concluded that there was undetermined risk (due to lack of investigation and evaluation) in relation to the potential ability of T. sinensis to parasitise native oak gall wasps (non-target effects); the EFSA panel also stated that gathering this information was crucial prior to release. This chapter therefore focuses on providing information about oak gall wasp species and their associated communities. Oak gall wasps (Hymenoptera: Cynipidae) are typically characterised by their complex cyclically parthenogenetic life cycles (heterogony), alternating between sexual and
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