Approximate Bayesian computation (ABC) is a powerful and widely used approach in inference of population history. However, the computational effort required to discriminate among alternative ...historical scenarios often limits the set that is compared to those considered more likely a priori. While often justifiable, this approach will fail to consider unexpected but well‐supported population histories. We used a hierarchical tournament approach, in which subsets of scenarios are compared in a first round of ABC analyses and the winners are compared in a second analysis, to reconstruct the population history of an oak gall wasp, Synergus umbraculus (Hymenoptera, Cynipidae) across the Western Palaearctic. We used 4,233 bp of sequence data across seven loci to explore the relationships between four putative Pleistocene refuge populations in Iberia, Italy, the Balkans and Western Asia. We compared support for 148 alternative scenarios in eight pools, each pool comprising all possible rearrangements of four populations over a given topology of relationships, with or without founding of one population by admixture and with or without an unsampled “ghost” population. We found very little support for the directional “out of the east” scenario previously inferred for other gall wasp community members. Instead, the best‐supported models identified Iberia as the first‐regional population to diverge from the others in the late Pleistocene, followed by divergence between the Balkans and Western Asia, and founding of the Italian population through late Pleistocene admixture from Iberia and the Balkans. We compare these results with what is known for other members of the oak gall community, and consider the strengths and weaknesses of using a tournament approach to explore phylogeographic model space.
Oak galls are spectacular extended phenotypes of gallwasp genes in host oak tissues and have evolved complex morphologies that serve, in part, to exclude parasitoid natural enemies.Parasitoids and ...their insect herbivore hosts have coevolved to produce diverse communities comprising about a third of all animal species. The factors structuring these communities, however, remain poorly understood. An emerging theme in community ecology is the need to consider the effects of host traits, shaped by both natural selection and phylogenetic history, on associated communities of natural enemies. Here we examine the impact of host traits and phylogenetic relatedness on 48 ecologically closed and species-rich communities of parasitoids attacking gall-inducing wasps on oaks. Gallwasps induce the development of spectacular and structurally complex galls whose species- and generation-specific morphologies are the extended phenotypes of gallwasp genes. All the associated natural enemies attack their concealed hosts through gall tissues, and several structural gall traits have been shown to enhance defence against parasitoid attack. Here we explore the significance of these and other host traits in predicting variation in parasitoid community structure across gallwasp species. In particular, we test the "Enemy Hypothesis," which predicts that galls with similar morphology will exclude similar sets of parasitoids and therefore have similar parasitoid communities. Having controlled for phylogenetic patterning in host traits and communities, we found significant correlations between parasitoid community structure and several gall structural traits (toughness, hairiness, stickiness), supporting the Enemy Hypothesis. Parasitoid community structure was also consistently predicted by components of the hosts' spatiotemporal niche, particularly host oak taxonomy and gall location (e.g., leaf versus bud versus seed). The combined explanatory power of structural and spatiotemporal traits on community structure can be high, reaching 62% in one analysis. The observed patterns derive mainly from partial niche specialisation of highly generalist parasitoids with broad host ranges (>20 hosts), rather than strict separation of enemies with narrower host ranges, and so may contribute to maintenance of the richness of generalist parasitoids in gallwasp communities. Though evolutionary escape from parasitoids might most effectively be achieved via changes in host oak taxon, extreme conservatism in this trait for gallwasps suggests that selection is more likely to have acted on gall morphology and location. Any escape from parasitoids associated with evolutionary shifts in these traits has probably only been transient, however, due to subsequent recruitment of parasitoid species already attacking other host galls with similar trait combinations.
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DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
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•A review of Synergus species from mainland China is carried out for the first time.•Three new species are described, and some are redescribed.•A key to Eastern Palaearctic and ...Oriental species is provided.•East and Southeast Asia harbors a great diversity of Fagaceae trees.•Further studies will discover more new taxa from these regions.
The Eastern Palaearctic and Oriental fauna of gall wasp inquilines (Hymenoptera: Cynipidae) has been historically understudied. Previously, ten species of Synergus Hartig (Hymenoptera: Cynipidae: Synergini), the most speciose genus of inquilines associated with Fagaceae galls worldwide, were known to occur in mainland China despite this region harbors a great number of potential fagaceous tree hosts. After the examination of all the Synergus specimens deposited at the Parasitic Hymenoptera Collection of the Agriculture and Forestry University of Zhejiang (ZAFU, China), we provide a revision of this genus in mainland China. Three new species (S. drouarti Pujade-Villar, sp. nov., S. ponsatiae Lobato-Vila & Pujade-Villar, sp. nov. and S. rovirae Lobato-Vila & Pujade-Villar, sp. nov.) are described and illustrated. Four species (S. areolatus (Weld, 1926) comb. nov.; S. formosanus Schwéger & Melika, 2015, S. gifuensis Ashmead, 1904 and S. ishikarii Melika & Schwéger, 2015) are new records for this area. Saphonecrus reticulatus Pujade-Villar, Wang & Guo, 2014 is proposed as syn. nov. of Synergus castaneus Pujade-Villar, Bernardo & Viggiani, 2013, and Synergus changtitangi Melika & Schwéger, 2015 as syn. nov. of S. deqingensis Pujade-Villar, Wang & Chen, 2015. We provide new biological and distributional data for all the species as well as redescriptions, pictures and other corrections for those which were originally poorly characterized. Lastly, we provide an updated key to the Eastern Palaearctic and Oriental Synergus species, with a table summarizing all of them.
Diversification of insect herbivores is often associated with coevolution between plant toxins and insect countermeasures, resulting in a specificity that restricts host plant shifts. Gall inducers, ...however, bypass plant toxins and the factors influencing host plant associations in these specialized herbivores remain unclear. We reconstructed the evolution of host plant associations in Western Palaearctic oak gallwasps (Cynipidae: Cynipini), a species-rich lineage of specialist herbivores on oak (Quercus). (1) Bayesian analyses of sequence data for three genes revealed extreme host plant conservatism, with inferred shifts between major oak lineages (sections Cerris and Quercus) closely matching the minimum required to explain observed diversity. It thus appears that the coevolutionary demands of gall induction constrain host plant shifts, both in cases of mutualism (e.g., fig wasps, yucca moths) and parasitism (oak gallwasps). (2) Shifts between oak sections occurred independently in sexual and asexual generations of the gallwasp lifecycle, implying that these can evolve independently. (3) Western Palaearctic gallwasps associated with sections Cerris and Quercus diverged at least 20 million years ago (mya), prior to the arrival of oaks in the Western Palaearctic from Asia 5–7 mya. This implies an Asian origin for Western Palaearctic gallwasps, with independent westwards range expansion by multiple lineages.
Native parasitoids may play an important role in biological control. They may either support or hinder the effectiveness of introduced nonnative parasitoids released for pest control purposes. ...Results of a three-year survey (2011–2013) of the Asian chestnut gall wasp (ACGW) Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae) populations and on parasitism rates by native indigenous parasitoids (a complex of chalcidoid hymenopterans) in Italian chestnut forests are given. Changes in D. kuriphilus gall size and phenology were observed through the three years of study. A total of 13 species of native parasitoids were recorded, accounting for fluctuating parasitism rates. This variability in parasitism rates over the three years was mainly due to the effect of Torymus flavipes (Walker) (Hymenoptera: Torymidae), which in 2011 accounted for 75% of all parasitoid specimens yet decreased drastically in the following years. This strong fluctuation may be related to climatic conditions. Besides, our data verified that parasitoids do not choose host galls based on their size, though when they do parasitize smaller ones, they exploit them better. Consequently, ACGWs have higher chances of surviving parasitism if they are inside larger galls.
Phyllonorycter issikii (Lepidoptera: Gracillariidae) has recently spread over a considerable part of the Eurasian continent. So far, in Europe, unlike in Russia, it has not significantly affected its ...food plant (Tilia cordata), but its effect may increase over time. The regulating potential of associated parasitoid natural enemies is still low, which probably allows further leaf miner range expansion and population growth. This species is potentially capable of inflicting considerable damage on its host. Therefore, it is important to study its natural enemies, especially their oviposition behaviour and biology. In this paper, we summarize the results of rearing experiments throughout the area it has recently occupied, including our own study in Hungary. Here we list 46 species of parasitoids recorded from P. issikii. None of these species are likely to be able regulate the abundance of P. issikii. Further studies are needed to determine their effect on the abundance of this host.
Live oak (Quercus virginiana Mill.) trees are hosts to a complex of gall making arthropods. However, the bullet galls produced by the asexual generation of the cynipid Disholcaspis quercusvirens ...(Ashmead) can esthetically and physically damage nursery and street trees, and thus reduce tree value. We sought to describe the unknown sexual generation of D. quercusvirens, describe the development of galls from both generations, record adult cynipid and parasitoid activity periods, and evaluate the efficacy of several insecticides to suppress the gall makers and prevent additional gall formation. The oviposition period for asexual females occurred from late November to January in both years of the caging study. Eggs laid into dormant buds resulted in small bud galls in which the sexual generation developed for 4–5 mo. Sexual adults emerged and laid eggs in young elongating shoots in April. Bullet galls began protruding from branches in June, and asexual wasps emerged 5–7 mo later. Cynipids that emerged from the bullet (asexual generation) and bud (sexual generation) galls were genetically identical. Both generations were heavily parasitized. Targeting asexual females with an early December treatment of bifenthrin or acephate significantly reduced the number of bud galls, but control did not extend to the next generation of bullet galls, possibly because of reinvasion from neighboring infested trees.
A new genus of oak gallwasp, Coffeikokkos Pujade-Villar & Melika, gen. n., is described from Costa Rica. Diagnostic characters and generic limits of the new genus are discussed in detail. The new ...genus includes Coffeikokkos copeyensis Pujade-Villar & Melika, sp. n., which induces galls on stems of Quercus bumelioides, an endemic oak to Costa Rica, Honduras and Panama. The new species and galls are described and illustrated.
Background and Purpose: Dryocosmus kuriphilus is a globally invasive insect pest, spreading very quickly in new habitats and making serious damage to sweet chestnut forests in Croatia and in several ...other European countries. Indigenous parasitoid species trophically associated with oak gallwasps have adapted to this new host but cannot effectively regulate its population density. Classical biological control using parasitoid Torymus sinensis has been proven to be the only effective method of controlling the populations of D. kuriphilus and has been successfully applied in Japan, South Korea, the USA and Italy. The aim of this review paper is to provide overview and up-to date knowledge about biological control of D. kurphilus and to describe first steps of introduction of T. sinensis to sweet chestnut forests in Croatia.
Conclusions and Future Prospects: Results presented in this paper show adapted biology and behavioural traits of T. sinensis to its host D. kuriphilus. The history and results of introductions of T. sinensis to Japan, the USA, Italy, France and Hungary are shown. The first report of release of T. sinensis to sweet chestnut forests in Croatia is given with discussion on native parasitoids attacking D. kuriphilus. Possible negative effects of T. sinensis on native parasitoid fauna and risks that could influence the successful establishment of T. sinensis in Croatia are discussed. Previous experiences have shown that T. sinensis can successfully control the population density of D. kuriphilus, slowing down the spread and mitigating negative impact of this invasive chestnut pest and keeping the damage of D. kuriphilus at acceptable level. High specificity of T. sinensis suggests that it has limited potential of exploiting native hosts but further detailed monitoring of native parasitoid and possible interactions with introduced T. sinensis is strongly suggested.
Oak gallwasps (Hymenoptera, Cynipidae, Cynipini) are one of seven major animal taxa that commonly reproduce by cyclical parthenogenesis (CP). A major question in research on CP taxa is the frequency ...with which lineages lose their sexual generations, and diversify as purely asexual radiations. Most oak gallwasp species are only known from an asexual generation, and secondary loss of sex has been conclusively demonstrated in several species, particularly members of the holarctic genus Andricus. This raises the possibility of widespread secondary loss of sex in the Cynipini, and of diversification within purely parthenogenetic lineages. We use two approaches based on analyses of allele frequency data to test for cryptic sexual generations in eight apparently asexual European species distributed through a major western palaearctic lineage of the gallwasp genus Andricus. All species showing adequate levels of polymorphism (7/8) showed signatures of sex compatible with cyclical parthenogenesis. We also use DNA sequence data to test the hypothesis that ignorance of these sexual generations (despite extensive study on this group) results from failure to discriminate among known but morphologically indistinguishable sexual generations. This hypothesis is supported: 35 sequences attributed by leading cynipid taxonomists to a single sexual adult morphospecies, Andricus burgundus, were found to represent the sexual generations of at least six Andricus species. We confirm cryptic sexual generations in a total of 11 Andricus species, suggesting that secondary loss of sex is rare in Andricus.