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The genus Plagiotrochus Mayr was originally described from the western Mediterranean region, which remains its known centre of diversity with 14 currently recognised species, all inducing galls on evergreen oaks (mainly Quercus section Ilex; one on section Cerris). Nine species have been described from Asian bioregions. While European Plagiotrochus is undoubtedly monophyletic, some recently described Asian Plagiotrochus show anomalous morphology and induce galls on a different oak lineage, Quercus section Cyclobalanopsis, suggesting their erroneous genus-level classification. We used molecular data to test generic limits of Plagiotrochus on a global scale, plus species limits of European species. We sequenced four genes (two mitochondrial, two nuclear) for 17 European Plagiotrochus taxa, four Asian Plagiotrochus species (one galling section Ilex, one galling section Cerris, two galling Cycloblanopsis oaks), and a selection of outgroups from the same major lineage within the Cynipini. We demonstrate that Plagiotrochus species galling section Ilex hosts are a primarily European, monophyletic radiation that also includes the Asian P. tarokoensis. In contrast, the section Cerris associated species P. kunugiphagus has been erroneously transferred to this genus, being most closely related to other Cerris-group gallers, while species galling Cyclobalanopsis oaks are also outside the limits of true Plagiotrochus. Molecular data match up some European taxa previously known only from a single generation and synonymise two asexual-only species. These data also confirm generation pairings previously based on experimental rearings, confirm the nomenclature of two clearly-defined species, and assist delimitation of two new species.

We provide the first description and analysis of species-rich communities of Fagaceae host plants, cynipid gall inducers and Hymenopteran parasitoids from two sites in western Sichuan, China. We use quantified interaction data to test the hypothesis that metabolically intimate cynipid-Fagaceae interactions are more specialised, resulting in more modular networks and stronger signatures of turnover than nestedness for beta diversity, than associations between parasitoids and Fagaceae. Rearing of nearly 27,000 cynipid gall specimens from 22 host plant species in Castanea, Castanopsis, Lithocarpus and Quercus (sections Cerris, Cyclobalanopsis, Ilex and Quercus) yielded 168 morphologically distinct cynipid gall morphotypes, and 3800 parasitoids in 67 morphospecies. The Sichuan parasitoid assemblage is similar in composition to cynipid-centred communities in the Western Palearctic and Nearctic. All of our predictions were confirmed. We discuss our findings in light of the processes thought to structure tritrophic interactions centred on gall inducing insect herbivores.

Co-evolutionary interactions between insect herbivores and their host plants underlie much contemporary biodiversity and are vital to assembly of natural ecosystems. Assemblages of galls on oaks induced by Cynipini wasps (Hymenoptera: Cynipidae) occur across much of the Northern Hemisphere, their diversity peaking in the Nearctic and on white oaks (Quercus section Quercus). However, the evolutionary history of the clade has been debated with respect to geographic origins, direction and timings of dispersal events, and shifts in host plant associations. We examined these questions using a global-scale, 6-gene phylogeny of 430 Cynipini species and a dataset of their associated host plants encompassing all eight sections within Quercus plus five Fagaceae genera. Likelihood-based ancestral state reconstructions demonstrate a Nearctic origin of the Cynipini followed by repeated colonisations of the Palearctic via both westwards and eastwards dispersal. These inferences are robust to bias in taxon sampling across continents and the inclusion of Protobalandricus as the sister lineage to Cynipini sensu stricto. Likewise, the association with white oaks is probably ancestral and has been retained by many Cynipini lineages. However, host shifts to other sections within Quercus and related Fagaceae genera are widely distributed across the cynipid phylogeny. They are associated with both global-scale range shifts and within-bioregion exploitation of alternative hosts, and their frequency typically correlates with host-plant relatedness. These findings highlight the evolutionary success of cynipids on white oak hosts and the connectedness of continental assemblages of gall wasps over evolutionary time.

Revealing processes that structure species interactions is central to understanding community assembly and dynamics. Species interact via their phenotypes, but identifying and quantifying the traits that structure species-specific interactions (links) can be challenging. Where these traits show phylogenetic signal, however, link properties are predictable using models that incorporate phylogenies in place of trait data. We analysed variation in link richness, frequency, and identity in a multi-site dataset of interactions between host oak cynipid galls and parasitoid natural enemies, using a Bayesian mixed modelling framework allowing concurrent fitting of terms for phylogenies of both trophic levels. In both link incidence (presence/absence) and link frequency datasets, we identified strong signatures of cophylogeny (related parasitoids attack related host galls) and patterns independent of either phylogeny. Our results are robust to simulations of substantially reduced sample completeness, and are consistent with the structuring of trophic interactions by a combination of phylogenetically conserved and convergently evolving traits at both trophic levels. We discuss our results in light of phenotypic traits thought to structure gall-parasitoid interactions and wider applications of this approach, including inference of underlying community assembly processes and prediction of economically important trophic interactions.

[Background]: A common strategy to limit the negative impact of biological invasions is biological control through the release of specialized alien natural enemies. However, biological control plans are not without risks, which include parasitism of native hosts and hybridization with related native species, particularly those that are potential natural enemies of the invasive species. Here, we evaluate these potential risks resulting from the introduction of the parasitoid wasp Torymus sinensis (Hymenoptera: Torymidae) in Europe to control the invasive Asian chestnut gall wasp Dryocosmus kuriphilus (Hymenoptera: Cynipidae)., [Results]: We found that, under laboratory conditions, the physiological host range of T. sinensis includes several native non-target species of Cynipidae, with oviposition observed on the galls of 8 of the 11 species tested. However, physiological host range of T. sinensis appears to be limited under field conditions, as we observed only one parasitized gall of Andricus curvator in the field. Regarding hybridization, inter-species mating between T. sinensis and its phylogenetically closest native Torymus species was not observed in the laboratory. Moreover, discordance between nuclear (ITS2) and mitochondrial (coxI) data does not support the presence of genetic introgression, suggesting that hybridization between T. sinensis and native Torymus species does not occur. In addition, we cite and discuss the unexpected presence of one individual of the related alien species Torymus beneficus in Spain., [Conclusion]: Our results suggest that T. sinensis may negatively impact several non-target species, highlighting the need for careful monitoring of the extent of such undesired behaviour and its effects on the native fauna.

The phylogeny of gall wasps (Cynipidae) and their parasitic relatives has attracted considerable attention in recent years. The family is now widely recognized to fall into twelve natural lineages, designated tribes, but the relationships among them have remained elusive. This has stymied any progress in understanding how cynipid gall inducers evolved from insect parasitoids, and what role inquilines (species that develop inside the galls of other species) might have played in this transition. A recent analysis of ultraconserved elements (UCEs) represents the first attempt at resolving these questions using phylogenomics. Here, we present the first analysis of gall-wasp relationships based on protein-coding sequences from genome and transcriptome assemblies. To address potential problems due to model misfit, we focus on models that accommodate site-specific amino-acid profiles and that are less sensitive to long-branch attraction than standard models. Our results show that the Cynipidae as previously circumscribed are not monophyletic. Specifically, the Paraulacini and a clade formed by Diplolepidini + Pediaspidini both fall outside of a core clade (Cynipidae sensu stricto), which is more closely related to Figitidae. This result is robust to the exclusion of long-branch taxa that could potentially mislead the analysis, and it is consistent with the previous UCE analysis. Given this, we propose that the Cynipidae be divided into three families: the Paraulacidae, Diplolepididae and Cynipidae (s. str.). Our results suggest that the Eschatocerini are the sister group of the remaining Cynipidae (s. str.). Within Cynipidae s. str., our results are consistent with the UCE analysis but place two additional tribes: (1) the Aylacini (s.str.), which are more closely related to the oak gall wasps (Cynipini) and some of their inquilines (Ceroptresini) than to the other herb gallers (Aulacideini and Phanacidini); and (2) the Qwaqwaiini, which are likely the sister group to Syner

Andricus dentimitratus (Rejtõ, 1887) and Andricus pictus (Hartig, 1856) are two European gall wasps (Hymenoptera, Cynipidae) that induce galls on species of Quercus. The distribution and ecological niches of these species have not been studied in detail, though they are known to have a different distribution pattern in the Iberian Peninsula in Europe. To investigate this difference and its potential relationship with climate and host species distribution, we analysed the potential distribution of both species in the Iberian Peninsula using six algorithms and a consensus model based on 600 iterations for each species. We compared the models obtained for each species with the distribution of their host Quercus species. The results show that A. dentimitratus and A. pictus have a complementary distribution delimited by the Ebro valley, with A. dentimitratus occurring northeast of the valley and A. pictus southwest. The observed distribution patterns might be due to differences in the climatic requirements of each species or to the distribution of their host species given that A. dentimitratus is specific to Q. humilis and Q. cerris (except in the northeastern Iberian Peninsula) and A. pictus, to marcescent Mediterranean oaks (Q. faginea and Q. pyrenaica) and Q. suber. We propose two hypotheses to explain the nonoverlapping distribution of the two gall wasp species in the Iberian Peninsula: in the first scenario, A. dentimitratus arrived to the to the Iberian Peninsula from the eastern Palearctic by way of Europe and A. pictus, from the north coast of Africa; in the second, their distribution is a result of their speciation in different glacial refugia: A. dentimitratus in the Italian Peninsula and A. pictus in the Iberian Peninsula.

The Asian chestnut gall wasp, Dryocosmus kuriphilus Yasumatsu, 1951 (Hymenoptera: Cynipidae), is a parthenogenetic species original from China that rapidly invaded Europe since its detection in 2002. The wasps lay eggs in the buds of chestnut trees (Castanea spp.) inducing galls on leaves, stipule, flowers and shoots. The consequences are significant economic damage due to a reduction of nuts and wood production, as well as a progressive decrease in tree vigour, which can make the tree susceptible to other secondary pests. We report here the results of a 4-year study of this species conducted in Galicia (NW of Iberian Peninsula), one of the most important areas for chestnut production in southern Europe. The insect was detected in 2014 but was apparently present at least two years earlier. It has since spread rapidly throughout the territory. It mainly dispersed naturally, although some long distance dispersal events influenced the rate of spread across the landscape. Population abundance varies throughout the territory, and we were able to establish hotspot and coldspot areas, mainly associated with maximum temperatures in the summer months. At a local scale, the trees attacked by D. kuriphilus and the number of galls on each tree were spatially clustered in the first years from the invasion. Tree height significantly predicted both variables, suggesting that gall wasps may use visual cues to locate suitable host trees, at least in the early stage of invasion. The species showed a rapid recruitment of native parasitoids: three years after detection we found 19 species of native parasitoids attacking D. kuriphilus in Galicia. Percent parasitism by native parasitoids decreased greatly with time since introduction. This was because galls became larger, more thickly walled, and with more individuals per gall as the abundance of D. kuriphilus increased. Per capita growth rate of D. kuriphilus was linearly density dependent with an apparent equilibrium of 5.2 galls/shoot

A new genus of herb gall wasps, Fumariphilus Nieves-Aldrey gen. nov., (Hymenoptera, Cynipidae, Aulacideini), is described. Previous molecular phylogenetic analyses, and resulting systematic changes at the tribe level, have shown that the related species Aylax hypecoi Trotter and Neaylax versicolor Nieves-Aldrey have been incorrectly placed within their respective genera. In order to reflect their monophyletic lineage and to accommodate these species within the tribe Aulacideini, it is necessary to erect a new genus. The two species, Fumariphilus versicolor (Nieves-Aldrey, 1985) n. comb. and Fumariphilus hypecoi (Trotter, 1913) n. comb. placed in the new genus induce galls in fruits of species of, respectively, Hypecoum L. (Papaveraceae: Fumarioideae: Hypecoeae) and Fumaria L. (Papaveraceae: Fumarioideae:Fumarieae), reflecting a host-plant association that is unique among all the cynipids. Diagnostic morphological features of both the adult and the larvae and an identification key to the species of the new genus are provided. Adults of Fumariphilus versicolor n. comb., are also re-described and, for both species of Fumariphilus, the terminal instar larva is described and illustrated with SEM images. Following the erection of the new genus Fumariphilus, Aylax Hartig, sensu NievesAldrey, 1994 is now sufficiently delimited within the tribe Aylacini sensu stricto Ashmead, 1903, which comprises species associated with only plants of the genus Papaver (Papaveraceae). Likewise, Neaylax Nieves-Aldrey, 1994, which now includes only the gall wasp species that associate with species of Salvia (Lamiaceae), is sufficiently delimited within the tribe Aulacideini Nieves-Aldrey, Nylander & Ronquist, 2015. Also provided in this work is an updated and illustrated identification key to the genera of the Aulacideini.

Nothofagus obliqua (Nothofagaceae) is a native tree to Chile, which hosts at least seven gall morphotypes, two of them induced by Hymenoptera, Pteromalidae. The life cycle of these pteromalids, particularly the feeding activity of their larvae, has a high impact on host plant tissues toward new shapes and functions, leading the gall formation. The Pteromalidae genus are mostly parasitoids of other insects, rarely inducing galls. This rare phenomenon is herein addressed in two models of study, the galls induced by Espinosa nothofagi and Plastobelyta gallicola on N. obliqua in Chilean flora. We assume that the two pteromalids have a similar impact over their host plant tissues, inducing similar galls regarding their anatomical and histochemical profiles. Our results confirm that E. nothofagi induces a clavate bud gall (CG), but P. gallicola induces a stem globoid gall (GG) herein describeb for the first time. Our premise is not supported, because the CG follows the structural and histochemical patterns described for hymenopteran galls, but the GG is peculiarly different. The CG has concentric layers of specialized tissues around the larval chamber, where lipids, proteins, and starch accumulate, while starch, alkaloids, proanthocyanidins, and lignins accumulate in the outer tissue compartment. The GG is simpler than the CG on the anatomical basis as it does not develop specialized tissue layers, and in addition, it does not accumulate starch and proanthocyanidins in any tissue layers. The distinct anatomical and histochemical profiles of the two galls indicate the specific impact of E. nothofagi and P. gallicola in the inner tissue layers. The similarities of the outer tissue layers between the two galls relate to the host stem morphogenetical constraints and lower plasticity.

Hymenoptera evolved structures on the legs which are able to remove particles from their antennae through grooming behaviour. The antenna cleaner (strigil) consists of an apical and modified protibial spur (calcar, composed of a trunk and a velum) and a modified basitarsus including a fine comb made up of setae and a notched inner surface. In “Terebrantia”, the non-aculeate Apocrita, large comparative studies of strigil are scarce, especially within lineages. Here, we studied in detail this structure in Cynipoidea, a group of wasps including parasitoids (Ibaliidae, Liopteridae and Figitidae), gall-inducers (Cynipidae) and gall-inquilines (Cynipidae), through a SEM analysis. We found some traits quite conserved across species and lineages. For example, the shape of protibia is almost invariably broadening towards apex, and one single, straight rather than curved, dorso-apical socketed spur on apical margin of protibia occurs in almost all species. Other characters roughly differentiate families. For example, thick and long setae on the protibia are arranged in one row especially in Figitidae, Ibaliidae and Liopteridae, while they more often occur in more than one rows in Cynipidae, which was the family with the shortest calcar. Figitidae have shorter basitarsal notch, but had longer and denser setae on the notch, compared with Cynipidae and Ibaliidae. Further characters were extremely variable across and within lineages. The observed morphological variation did not seem to reflect the phylogeny of Cynipoidea, and a role of life-history traits on such variation was not suggested, at least with the methodological approach used here.

The host range of parasitoids varies greatly among species: Some only parasitize one to a few hosts (specialists), while others parasitize multiple species or a va-riety of host types (generalists). The direction of most host range shifts in parasi-toid groups, that is from generalist to specialist or, alternatively, from specialist to generalist, is unknown. To explore the origin of host range shifts, we studied a clade within the genus Torymus(Hymenoptera, Chalcidoidea) that includes both generalist and specialist parasitoids of Cynipidae (Hymenoptera). We ana-lysed the phylogenetic relationships of the species of Torymuson the basis of two gene fragments (cox1and ITS2) of 246 specimens and performed an ancestral state reconstruction of the specialist/generalist trait. Our results revealed the fol-lowing: (a) The ancestral state of this group of Torymusis specialist, with the generalist state evolving through a loss of specialization. (b) The species Torymus cyaneusand Torymus flavipesboth have a strong genetic structure, suggesting the existence of different biological identities. (c) There has been a host plant shift in the lineage(s) leading to Torymus rubiand Torymus bedeguarisfrom galls on Quercusto those on Rosaceae. (d) The alien species Torymus sinensisand the native European species Torymus notatusare phylogenetically closely related. (e) Speciation within Torymuswas likely associated with the diversification of their cynipid hosts, which itself was driven by the dramatic changes in climate and vegetation that occurred during the Miocene.

Paraulax Kieffer and Cecinothofagus Nieves-Aldrey & Liljeblad (Cynipidae: Paraulacini) were long supposed to be gall-makers on southern beeches (Nothofagus, Nothofagaceae). Dissections of galls on Nothofagus Blume, suggested that Cecinothofagus could be instead either endoparasitoid or inquiline of Aditrochus larva (Chalcidoidea). We sequenced the universal COI barcode and Ultra-Conserved Elements (UCEs) from young larvae of Aditrochus collected from galls on Nothofagus and highlighted that one of them also contained DNA from Cecinothofagus ibarrai Nieves-Aldrey & Liljeblad. So far, when galls attributed to Aditrochus were dissected in early development stages they all contained only a single larva and no remains of other larvae. Conversely, when Cecinothofagus ibarrai was reared from galls on Nothofagus, remains of the host larva were observed inside the larval chamber. Altogether, biological observations and molecular results suggest that Cecinothofagus ibarrai is likely an endoparasitoid of Adritrochus. This result confirms the tribe Paraulacini as being entomophagous and supports the hypothesis of an ancestral parasitoid lifestyle for Cynipoidea.

A new genus, Prokius Nieves Aldrey, Medianero & Nicholls, gen. nov., and two new species of oak gall wasps (Hymenoptera: Cynipidae: Cynipini), Prokius cambrai Medianero & Nieves-Aldrey sp. nov. and Prokius lisethiae Medianero & Nieves-Aldrey sp. nov., are described from adults reared from galls on Quercus bumelioides Liebm (Fagaceae, sect. Quercus, white oaks) collected in Panama. The new genus is phylogenetically and morphologically close to Dros Kinsey and forms part of a large clade that includes species from several other genera that appear to require revision, including Andricus Hartig and Phylloteras Ashmead. Molecular and morphological data, diagnostic characters, gall descriptions, distribution and biological data of the new genus and the new species are given. This new genus represents the fourth recently described genus of Cynipidae endemic to the Neotropical region.

The monophyly and taxonomic validity of some currently accepted genera of gall wasps in the Cynipini (Hymenoptera: Cynipidae) are being challenged by recent systematic studies. Here we used morphological and molecular data to re-describe and revise the taxonomic limits of the monotypic genus Kokkocynips Pujade-Villar & Melika, previously recorded only from Mexico. We describe a new species from Panama, Kokkocynips panamensis Medianero & Nieves-Aldrey, reared from galls on Quercus salicifolia Neé, and add new records from Mexico for the type species of the genus K. doctorrosae Pujade-Villar. Six Nearctic species, Dryocosmus rileyi (Ashmead, 1896), D. imbricariae (Ashmead, 1896), D. coxii (Basset, 1881), D. deciduus (Beutenmueller, 1913), Callirhytis difficilis (Ashmead, 1887) and C. attractans (Kinsey, 1922) are transferred to Kokkocynips. Species of Kokkocynips are associated only with red oaks (Quercus section Lobatae (Fagaceae)) and are distributed in the Nearctic and Neotropical regions, from Canada through Mexico and Panama. Taxonomic limits of Kokkocynips are discussed in light of the molecular phylogenetic relationships of the studied species. Diagnostic characters, gall description, distribution, and biological data of Kokkocynips species are given, including a key for the identification of the asexual generations of seven species.

The monophyly and taxonomic validity of some currently accepted genera of gall wasps in the Cynipini (Hymenoptera: Cynipidae) are being challenged by recent systematic studies. Here we used morphological and molecular data to redescribe and revise the taxonomic limits of the monotypic genus Kokkocynips Pujade-Villar & Melika, previously recorded only from Mexico. We describe a new species from Panama, Kokkocynips panamensis Medianero & Nieves-Aldrey, reared from galls on Quercus salicifolia Neé, and add new records from Mexico for the type species of the genus K. doctorrosae Pujade-Villar. Six Nearctic species, Dryocosmus rileyi (Ashmead, 1896), D. imbricariae (Ashmead, 1896), D. coxii (Basset, 1881), D. deciduus (Beutenmueller, 1913), Callirhytis difficilis (Ashmead, 1887) and C. attractans (Kinsey, 1922) are transferred to Kokkocynips. Species of Kokkocynips are associated only with red oaks (Quercus section Lobatae (Fagaceae)) and are distributed in the Nearctic and Neotropical regions, from Canada through Mexico and Panama. Taxonomic limits of Kokkocynips are discussed in light of the molecular phylogenetic relationships of the studied species. Diagnostic characters, gall description, distribution, and biological data of Kokkocynips species are given, including a key for the identification of the asexual generations of seven species.

1. Parasites are one of the main actors in host–parasite interactions. Still,their role as a prey and the related consequences for such interactions and in otherrespects, such as food webs, are frequently overlooked., 2. This paper analyses predation pressure on a ubiquitous avian ectoparasitic fly,Carnus hemapterus, identifies their main natural enemies and quantifies their relativeeffect on the abundance of the parasite. Also, the effect of nest-site type on their mainenemies’ predation pressure was analysed., 3. Several ant species were found in the nest boxes of the host species, the Europeanroller (Coracias garrulus), during the breeding season and preyed upon adult and larvalstages of Carnus., 4. Ants were also the putative predators of carnid pupae after the breeding season, whensignificant reduction (on average, by half) in the abundance of carnid pupae occurred in75% of nest boxes within few months., 5. Carnid pupae are also reported here, for the first time, to be parasitised by theparasitoid waspChartocerus conjugalis, whose prevalence was around 21%., 6. Nest-site type had no clear effect on the predation rate of carnid pupae after thebreeding season., 7. It was concluded that predation is an important factor regulating the abundance ofectoparasites, and thus, it may influence the outcome of host–parasite relationships.

Dryocosmus kuriphilus (Hymenoptera: Cynipidae) is a global invasive gall wasp and a pest of chestnuts (Castanea spp.). A study of the Chalcidoidea parasitoid community of D. kuriphilus was undertaken over two years, from March 2017 to March 2019, at 15 sites in south and northwest Spain (Málaga and Galicia regions). More than 18,000 galls were collected, and 1153 parasitoids belonging to 22 species of seven chalcidoid families, plus two individuals of an inquiline Cynipidae, Synergus facialis, emerged. Richness was higher in the Málaga region, with 20 species, while 17 parasitoids and one inquiline were identified in Galicia. The parasitism rate of native chalcid parasitoid species in both regions was low. Eupelmus urozonus and Mesopolobus lichtensteini were the most abundant native species. Mesopolobus tibialis was a dominant species in south Spain, while Ormyrus pomaceus was a dominant species in northwest Spain. Our results revealed the existence of a sub-community of univoltine, probably host specialized, parasitoids in south Spain, which overwinter in galls, exhibiting a similar life cycle to Torymus sinensis. These species were Torymus notatus, Aulogymnus bicolor, Aulogymnus obscuripes and Aulogymnus balani. Data on the recovery of T. sinensis after release in the south Spain region show it to be well established, but its numbers are still low in northwest Spain.

The success of invasive species largelydepends on the resistance offered by the native communities. Dryocosmus kuriphilusis a cynipidnative from China that is a serious pest of chestnuts worldwide. This species recruits natural enemies in theareas of introduction; nevertheless, their role inregulating pest population is frequently questioned, although the causes are not clear. We sampled parasitoids from 12,525 galls collected in 192 sitesin four geographic areas differing in climate andresidence time of the wasp of Galicia (NW Iberian Peninsula). We also dissected 2994 galls to assess the effects of gall characteristics on parasitism level. We found a rich community of native enemies thatcontributed to biotic resistance against the invasive:19 species of native parasitoids were feeding on thewasp, with mortalities ranging from 14 to 37%.Rarefaction analyses indicated that this species listaccounts for most of the native enemies that attack D.kuriphilusin inland Galicia. However, percent parasitism of D. kuriphilusby native parasitoids decreasedgreatly with time since introduction. This was becausegalls became larger, more thickly walled, and withmore D. kuriphilusper gall as the abundance of D. kuriphilusincreased. As the galls became larger andmore populated, a declining fraction of the D. kuriphiluswithin were parasitized by native enemies (decrease from [25% parasitism to\2% as gall sizeincreased from 0.2 to 3 cm3). Consequently, native parasitoids may play a role in slowing the invasive atlow densities, but this effect becomes increasingin consequential in highly invaded areas.

The alien cynipid waspDryocosmus kuriphilusYasumatsu, 1951 is a serious pest of chestnuts(Castaneaspp.) in Japan, North America and Europe, causing fruit losses while inducing gallsin buds. WhileD. kuriphilusgalls have a recognizable and roughly invariable globular shape,their size varies, reaching up to 4 cm in diameter. Among other factors, such variation maydepend on different climatic conditions in different attacked areas. Here, we sampled andmeasured 375D. kuriphilusgalls from 25 localities throughout the Iberian Peninsula, includ-ing both cold and rainy northern (Eurosiberian) areas and warm and dry central-southern(Mediterranean) areas, to test the effects of climate and geographical location on gall morph-ology. The analyses indicate that gall mass and volume follow a pattern that can be associatedwith a climatic cline. In particular, the Eurosiberian galls were smaller than the Mediterraneangalls according to differences in climatic conditions. In the southern areas, the greaterinsolation regime does not allow the chestnut trees to be distributed at lower altitudes, butthe high rainfall and humidity regime of the mountain enclaves allow their presence. Theseconditions of insolation and precipitation seem to influence the morphological characteristicsof the galls ofD. kuriphilus.

Diplolepis (Hymenoptera: Cynipidae) are gall wasps that induce conspicuous galls on Rosa spp. (Rosaceae). These species are distributed globally and in Europe some are especially common and are founder organisms of biological communities composed of different insects. However, the ecological niches of these species have not been studied in detail. We modelled the potential distributions of these species using the locations of the galls of the four most abundant species of Diplolepis on the Iberian Peninsula (Diplolepis mayri, Diplolepis rosae, Diplolepis eglanteriae and Diplolepis nervosa, the galls of latter two are indistinguishable) using four different algorithms and identified the resulting consensus for the species. We compared the potential distributions of these species, considering their spatial complementarity and the distributions of their host plants. We found that D. mayri and D. eglanteriae/nervosa have complementary distributions on the Iberian Peninsula. The former species is found in the Mediterranean region, while D. eglanteriae and D. nervosa are distributed mainly in the Eurosiberian region. Diplolepis rosae has the widest distribution on the Iberian Peninsula. Our models constitute the first effort to identify suitable areas for species of Diplolepis species on the Iberian Peninsula and could be useful for understanding the evolutionary ecology of these species throughout their distribution in the western Palearctic.

[EN] A revised checklist of eight of the twelve known palaearctic species in the subgenus Macroneura of Eupelmus occurring in the Iberian Peninsula, Balearic and Macaronesian islands (Azores, Balearics, Canaries, Madeira) is presented. Records of the distribution in the region of all the revised species are given with new biological notes, including host insects and plants. Eupelmus (Macroneura) seculatus is a new record for the Canary Islands and Eupelmus(Macroneura) vesicularis is recorded for the first time in Portugal. Additionally, a study of Malaise trap captures of species of Macroneura in an arid habitat in south-east Spain is presented and discussed. 358 individuals belonging to six species were collected and identified. Eupelmus (Macroneura)muellneri comprised 43.5% of the Macroneura and was the most abundant species followed by E. (M.) barai (25.4%)., [ES] Se presenta la revision y una lista anotada de ocho de las doce especies paleárticas de Eupelmus (Macro-neura) que se conocen de la Península Ibérica, Islas Baleares y Macaronesia (Canarias, Madeira y Azores). Se aportan para cada especie las citas en cada ámbito geográfico, incluyendo nuevos datos biológicos, como los de sus insectos y plantas hospedadoras. Eupelmus (Macroneura) seculatus es nueva cita para las Islas Canarias y Eupelmus(Macroneura) vesicularis se cita por primera vez de Portugal. Como complemento a esta revisión, se presentan los resultados de las capturas de especies de Macroneura efectuadas con una trampa Malaise que se mantuvo operativa en primavera y verano de 2013 en una zona árida del sureste español (Mojácar, Almería). Se capturaron 358 ejemplares de seis especies, siendo Eupelmus (Macroneura)muellneri (43,5% del total) la especie más abundante en las muestras, seguida de E. (M.)barai (25,4%).

In several groups of insects, body structures related to feeding and oviposition are known to have a hardened cuticle by incorporation of transition metals. However, a functional link between metal enrichment and ecological pressures (i.e., adaptation) has been only rarely shown, opening the possibility that in some lineages, the evolutionary history may account for most of the observed variation (i.e., phylogenetic constraint). Here, we addressed this question in the hyper-diverse Hymenoptera (bees, wasps, ants, and sawflies), in which Zn and/or Mn have been found enriching the mandibles of a number of species. Across 87 species spanning most of the extant superfamilies, we found Zn enrichment to be widespread (57 species). Although lacking in the most primitive “Symphyta”, our ancestral state reconstruction was not conclusive in determining whether Zn enrichment was a derived state for the complete order, but it was clearly the ancestral state for the Apocrita, where it was lost in few lineages, notably in Aculeata (where it was then reacquired at least three times). Mn, on the other hand, occurred very rarely in mandibles (10 species). Our comparative analysis revealed a strong phylogenetic effect explaining most Zn % and Mn % variation in mandibles. Additionally, species with herbivorous larvae were less prone to have Zn (but not Mn) than those with carnivorous larvae, although a causal effect of diet on this trait was unlikely. Furthermore, species emerging from concealed vs. unconcealed development sites have similar likelihood to have metal-enriched mandibles. Evolutionary history seems to constrain metal enrichment in hymenopteran mandibles, yet the few observed losses and regains of this trait during evolution claim for deeper investigations on the role of alternative, here untested, ecological pressures.

Mating behaviour in insects is largely mediated by chemical signals emitted by one sex and recognized by the other (sex pheromones). A particular type of integumentary gland produces a sex pheromone in the antennae of male Cynipoidea (Hymenoptera), a rich and diverse wasp group consisting of species forming galls on plants, species which act as inquilines or parasitoids of these galls, and non-gall-associated parasitoids. This gland is connected to the surface by a modified structure which releases and spreads the chemicals on females (release and spread structure (RSS)). Here, we performed a scanning electron microscope investigation to evaluate the diversity of RSS in detail through an analysis of 49 species spanning almost all extant lineages, with special emphasis on the family Cynipidae (gall-wasps). Up to three strongly modified flagellomeres harboured the RSS in Cynipoidea, more often the first (proximal) flagellomere (F). The Fwas in most cases cylindrical or slightly flattened laterally, in this case often with a longitudinal ridge present, with few cases of a distinctly excavated (only in Cynipidae and the basal parasitoid family Ibaliidae) or strongly distally inflated (only in the cynipid gall-inquiline tribe Synergini) shape. The RSS ranged from long and narrow to short and wide. Pores of the RSS, more often scattered than closely spaced, were well visible in most of species. The observed conspicuous variability in RSS morphology seemed very weakly associated both with phylogenetic relationships among lineages and with life-history (galler, gall-associated inquilines or parasitoids, and non-gall-associated parasitoids). New studies are necessary to shed light on the evolution of this structure.

The known species richness of oak gallwasps in Asia has increased tremendously in the past decade. However, the vast majority of taxa have been described from the east coast of Asia, and knowledge of oak gallwasps from Central Asia is still scant. Here we use molecular and morphological characters to describe a new genus of cynipid oak gallwasp, Heocynips Fang, Nieves-Aldrey, and Melika, new genus (Hymenoptera: Cynipidae: Cynipini), and a new species Heocynips furvoaurantius Tang, Nicholls, and Stone, new species from Emeishan, Sichuan Province, China. We present diagnostic characters for the new genus, including adult morphology and DNA sequence data, and provide biological information on host plant association, gall morphology and developmental phenology for the new species.

Temporal comparisons of a site's biodiversity depend on the availability of at least two asynchronous data sets on a bioindicator group. The Sierra de Guadarrama, a mountain range in central Spain has a high biodiversity and representative sites such as El Ventorrillo Biological Station (Madrid province) play an important role in research and monitoring of the Sierra biodiversity. Although unevenly and randomly, hoverflies of the Sierra de Guadarrama have been surveyed for decades, but never at El Ventorrillo. This important group of pollinators, aphid predators and bioindicators was sampled with a Malaise trap at El Ventorrillo from June 1989 to June 1990. A total of 51 species were identified (20 spp. of Eristalinae and 31 spp. of Syrphinae), 22 of which were new to the Sierra de Guadarrama and 17 to the Madrid province. The genera Brachypalpus and Didea were also new to the Madrid province. The Sierra de Guadarrama now has 126 species recorded and the Madrid province 150. Within the Sierra de Guadarrama, El Ventorrillo becomes the second locality in number of hoverfly species recorded after El Escorial (64 spp.). In the Malaise trap catch, hoverflies with zoophagous larvae had a higher representation than those of other guilds, both in species richness and abundance. The results provided here for El Ventorrillo represent a baseline for future biodiversity studies at this site with a similar experimental design, for example, to detect changes in hoverfly diversities along time.

Aim of study: To update the distribution of Dryocosmus kuriphilus, the Asian chestnut gall wasp, focusing on the central area of the country (Sistema Central). Gall samplings of these areas were carried to obtain the first records of parasitoids on D. kuriphilus in this area. Area of study: Spain and, especially, the Sistema Central. Material and methods: Georeferenced new records were used to produce a map with the updated distribution of D. kuriphilus as of 2019. Galls were collected and stored in emergence boxes. Parasitoids that emerge from these galls were collected and identified at the most detailed taxonomic level. Main results: The alien species D. kuriphilus was found in the Spanish Sistema Central. Infestation focal points were detected in the Valle del Jerte (South of Sierra de Gredos), Sierra de Francia and in the mountains southwest of Madrid. In all detected focal points of infes-tation, the available evidence indicates that introduction was caused by the accidental mobilization of infested chestnut material. Alongside other parasitoids recruited by D. kuriphilus, we found individuals of Torymus sinensis, constituting the first records of this foreign species in the Sistema Central. Research highlights: We updated the previously published distribution of D. kuriphilus in Spain, an alien species and pest of chestnut forests and orchards. We also reported the first list of recruited parasitoids by D. kuriphilus and, more specifically, the first records of T. sinensis in the Sistema Central.

Three new species of oak gall wasps of the genus Amphibolips Reinhard, 1865 (Hymenoptera: Cynipidae: Cynipini) are described from Mexico: Amphibolips magnigalla Nieves-Aldrey & Castillejos-Lemus, Amphibolips kinseyi Nieves-Aldrey & Castillejos-Lemus and Amphibolips nigrialatus Nieves-Aldrey & CastillejosLemus. The specimens of the first two species were representative of sexual generations and come from the State of Oaxaca, while only a female, collected in the State of Veracruz, is described for A. nigrialatus. The new species induces galls on Quercus zempoaltepecana and Q. sapotifolia (Fagaceae, section Lobatae, red oaks). Descriptions of the diagnostic morphological characteristics of the three species and a key for their identification are provided. The taxonomic relationships of the new species with other species of Amphibolips are discussed; the three new species are closely allied amongst themselves and are related to A. dampfi Kinsey, 1937. With the three newly-described species, the number of Amphibolips in Mexico is increased to 23.

Parasitoid wasps may act as hyperparasites and sometimes regulate the populations of their hosts by a top-down dynamic. Nasonia vitripennis (Walker, 1836) is a generalist gregarious parasitoid that parasitizes several host flies, including the blowfly Protocalliphora Hough, 1899 (Diptera, Calliphoridae), which in turn parasitizes bird nestlings. Nonetheless, the ecological factors underlying N. vitripennis prevalence and parasitoidism intensity on its hosts in natural populations are poorly understood. We have studied the prevalence of N. vitripennis in Protocalliphora azurea (Fallén, 1817) puparia parasitizing wild populations of pied flycatcher (Ficedula hypoleuca) and blue tit (Cyanistes caeruleus) birds in two Mediterranean areas in central and southern Spain. We found some evidence that the prevalence of N. vitripennis was higher in moist habitats in southern Spain. A host-dependent effect was found, since the greater the number of P. azurea puparia, the greater the probability and rate of parasitoidism by the wasp. Our results also suggest that N. vitripennis parasitizes more P. azurea puparia in blue tit nests than in pied flycatcher nests as a consequence of a higher load of these flies in the former. Based on the high prevalence of N. vitripennis in P. azurea puparia in nature, we propose that this wasp may regulate blowfly populations, with possible positive effects on the reproduction of both bird species

Galls are plant tissues whose development is induced by another organism for the inducer's benefit. 30,000 arthropod species induce galls, and in most cases the inducing effectors and target plant systems are unknown. Cynipid gall wasps are a speciose monophyletic radiation that induce structurally complex galls on oaks and other plants. We used a model system comprising the gall wasp Biorhiza pallida and the oak Quercus robur to characterise inducer and host plant gene expression at defined stages through the development of galled and ungalled plant tissues, and tested alternative hypotheses for the origin and type of galling effectors and plant metabolic pathways involved. Oak gene expression patterns diverged markedly during development of galled and normal buds. Young galls showed elevated expression of oak genes similar to legume root nodule Nod factor-induced early nodulin (ENOD) genes and developmental parallels with oak buds. In contrast, mature galls showed substantially different patterns of gene expression to mature leaves. While most oak transcripts could be functionally annotated, many gall wasp transcripts of interest were novel. We found no evidence in the gall wasp for involvement of third-party symbionts in gall induction, for effector delivery using virus-like-particles, or for gallwasp expression of genes coding for plant hormones. Many differentially and highly expressed genes in young larvae encoded secretory peptides, which we hypothesise are effector proteins exported to plant tissues. Specifically, we propose that host arabinogalactan proteins and gall wasp chitinases interact in young galls to generate a somatic embryogenesis-like process in oak tissues surrounding the gall wasp larvae. Gall wasp larvae also expressed genes encoding multiple plant cell wall degrading enzymes (PCWDEs). These have functional orthologues in other gall inducing cynipids but not in figitid parasitoid sister groups, suggesting that they may be evolutionary innovat, [Author summary] Plant galls are induced by organisms that manipulate host plant development to produce novel structures. The organisms involved range from mutualistic (such as nitrogen fixing bacteria) to parasitic. In the case of parasites, the gall benefits only the gall-inducing partner. A wide range of organisms can induce galls, but the processes involved are understood only for some bacterial and fungal galls. Cynipid gall wasps induce diverse and structurally complex galls, particularly on oaks (Quercus). We used transcriptome and genome sequencing for one gall wasp and its host oak to identify genes active in gall development. On the plant side, when compared to normally developing bud tissues, young gall tissues showed elevated expression of loci similar to those found in nitrogen-fixing root nodules of leguminous plants. On the wasp side, we found no evidence for involvement of viruses or microorganisms carried by the insects in gall induction or delivery of inducing stimuli. We found that gall wasps express many genes whose products may be secreted to the host, including enzymes that degrade plant cell walls. Genome comparisons between galling and non-galling relatives showed cell wall-degrading enzymes are restricted to gall inducers, and hence potentially key components of a gall inducing lifestyle.

[EN] The diversity of insect antennal structures involved in communication is still poorly known because of the limited number of comparative studies and as such studies often exclusively focus on one sex. Within Cynipoidea, a recent study on female gall-wasps (Hymenoptera: Cynipidae) and some of their non-gall associated relatives (Ibaliidae and Figitidae) showed a great diversity of the antennal sensillar equipment both between and within lineages. In order to better understand possible patterns of this diversity, we here present a study in males. By Scanning Electron Microscopy analysis of 45 species of Cynipoidea (30 of them gall-wasps) from most known lineages, we found that the male antennal flagellum is generally filiform and bears overall eight types of sensilla: s. placoidea (SP), s. coeloconica (SCo-A), s. campaniformia (SCa), s. basiconica (SB) and s. trichoidea (ST-A, ST-B, ST-C, ST-D) (5e8 types per species). The number, size and arrangement of sensilla greatly varied among and within cynipoid lineages, with only a partial effect of phylogeny on this variation. By using data of 27 species for which both sexes were analysed, we found that males generally possess a lower number of sensillar types than females in gallinducers and a greater number of sensillar types than females in non-gall-inducers. Sexes shared 40 e100% of sensillar types independently from their biology. Males seem to possess an overall greater number of SP than females, with a weak tendency of such difference to be larger in gall-inducers. A possible hypothesis is that gall-inducing female cynipids may have evolved a richer sensillar equipment in response to female-only activities (e.g. plant host recognition), while cynipid males may have evolved higher density of sensilla devoted to mate recognition.

[EN] A new species of Andricus Hartig 1840 (Hymenoptera: Cynipidae: Cynipini) is described from Panama: Andricus barriosi sp. nov. The new species induces galls on Quercus bumelioides Liebm. and Q. insignis M. Martens & Galeotti (Fagaceae, sect. Quercus) and is the first species of the genus Andricus ‘sensu lato’ recorded from Panama. Andricus barriosi is part of a species complex that includes A. nievesaldreyi Pujade-Villar, A. georgei Pujade-Villar and A. maesi Pujade-Villar, which differ from the other species of the genus Andricus by having a characteristic striate-reticulate sculpture on the second metasomal tergum that is typically smooth in the other Andricus species. The diagnostic characters, gall description, distribution and biological data of the new species are given, and a key for the differentiation of the new species from related species is also provided.

[Aim of study]: The globally invasive gall wasp, Dryocosmus kuriphilus Yasumatsu, 1951 (Cynipidae: Cynipini), reached Spain seven years ago and is already regarded as an important pest of chestnuts (Castanea spp.) in this country as well as worldwide. In this paper, we present comprehensive data on the establishment in Spain of Torymus sinensis Kamijo, 1982 (Chalcidoidea: Torymidae), an effective non-native natural enemy of this pest, as a result of both natural dispersal and settlement after controlled releases since 2015., [Area of study]: Sites of the Spanish autonomous communities of Galicia, Asturias, Basque Country, Catalonia, Andalusia and Madrid where D. kuriphilus is present., [Material and methods]: To study the natural dispersal of T. sinensis from France, we selected two sampling sites in Catalonia, six in the Basque Country and two in Navarra known for their heavy Asian chestnut gall wasp (ACGW) infestation; to study T. sinensis establishment after authorized controlled releases by the concerned authorities, the field samplings were done mainly in Galicia (35 sites) and Andalusia (8 sites). Additionally an experimental release study was made in Madrid. [Main Results]: Our results showed that T. sinensis has spread throughout Spain by natural dispersal across the French border and now occurs in Catalonia (two sites), the Basque Country (three sites) and Navarra (one site) but not in the neighbouring region of Cantabria. The percentage of parasitism by T. sinensis on D. kuriphilus is higher in the Basque Country sites, which are close to the French border, thus indicating that its establishment in these localities is not recent. After controlled releases, T. sinensis has been successfully established in five release sites in Andalusia (Valle del Genal and Sierra Blanca, Málaga Province) and one release site in Madrid. However, in the region of Galicia (NW Spain), where the number of authorized releases has been higher, the establishment of T. sinensis still appears to be very low., [Research highlights]: Established populations of T. sinensis may exert a positive buffer against D. kuriphilus-driven chestnut infestation in Spain, similar to what is observed in other invaded European countries.

[EN] The Asian chestnut gall wasp Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae) is a global pest of chestnuts (Castanea). After being reported by the first time from Spain in 2012, the species has rapidly spread over the distribution area of wild and cultivated chestnuts in the Iberian Peninsula. Since the invasion D. kuriphilus has recruited de novo a rich community of hymenopterous parasitoids as well as other arthropods secondarily associated with the galls. Here we describe a new species of Parochthiphila (Diptera Chamaemyiidae), one of the dipteran species that make up the secondary fauna of arthropods associated with galls of the chestnut gall wasp. The new species was collected in the Sierra de las Nieves Natural Park, an important protected area in Andalusia (southern Spain). The new species is related to P. stackelbergi and the two differ by morphological external and genital characters., [ES] La avispa oriental asiática Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae) es una plaga global de los castaños (Castanea). Después de que se encontrara por primera vez en España en el año 2012, la especie se ha extendido rápidamente por la mayor parte del área de distribución de los castaños silvestres y cultivados de la Península Ibérica. Desde el comienzo de la invasión D. kuriphilus ha reclutado de novo una rica comunidad de himenópteros parasitoides, así como de otros artrópodos secundariamente asociados a sus agallas. Aquí describimos una especie nueva de Parochthiphila (Diptera Chamaemyiidae), una de las especies de dípteros que integran la comunidad secundaria de artrópodos en las agallas de la avispilla del castaño. La especie nueva se encontró en el Parque Natural de la Sierra de las Nieves, una importante área natural de Andalucía (sur de España). La especie nueva está relacionada con P. stackelbergi y las dos difieren por caracteres morfológicos externos y genitales.

Las avispas inductoras de agallas o cinípidos integran una de las relaciones planta animal más complejas y fascinantes por su capacidad de inducir estructuras morfológicas anormales en las plantas, cuya formación está controlada por el insecto y representan una extensión de su propio fenotipo. En esta charla haremos un recorrido por más de 40 años de estudio de avispas de las agallas en el mundo, con especial énfasis en las expediciones realizadas a Chile, Panamá, Méjico y más recientemente a las montañas de Sichuán (China)