The Chalcidoidea bush of life: evolutionary history of a massive radiation of minute wasps.

Chalcidoidea are mostly parasitoid wasps that include as many as 500 000 estimated species. Capturing phylogenetic signal from such a massive radiation can be daunting. Chalcidoidea is an excellent example of a hyperdiverse group that has remained recalcitrant to phylogenetic resolution. We combined 1007 exons obtained with Anchored Hybrid Enrichment with 1048 ultra-conserved elements (UCEs) for 433 taxa including all extant families, >95% of all subfamilies, and 356 genera chosen to represent the vast diversity of the superfamily. Going back and forth between the molecular results and our collective knowledge of morphology and biology, we detected bias in the analyses that was driven by the saturation of nucleotide data. Our final results are based on a concatenated analysis of the least saturated exons and UCE datasets (2054 loci, 284 106 sites). Our analyses support an expected sister relationship with Mymarommatoidea. Seven previously recognized families were not monophyletic, so support for a new classification is discussed. Natural history in some cases would appear to be more informative than morphology, as illustrated by the elucidation of a clade of plant gall associates and a clade of taxa with planidial first-instar larvae. The phylogeny suggests a transition from smaller soft-bodied wasps to larger and more heavily sclerotized wasps, with egg parasitism as potentially ancestral for the entire superfamily. Deep divergences in Chalcidoidea coincide with an increase in insect families in the fossil record, and an early shift to phytophagy corresponds with the beginning of the "Angiosperm Terrestrial Revolution". Our dating analyses suggest a middle Jurassic origin of 174 Ma (167.3-180.5 Ma) and a crown age of 162.2 Ma (153.9-169.8 Ma) for Chalcidoidea. During the Cretaceous, Chalcidoidea may have undergone a rapid radiation in southern Gondwana with subsequent dispersals to the Northern Hemisphere. This scenario is discussed with regard to knowledge about the host taxa of chalcid wasps, their fossil record and Earth's palaeogeographic history.

Re-description and systematic re-appraisal of the genus Kokkocynips Pujade-Villar amp; Melika, (Hymenoptera: Cynipidae: Cynipini), including new combinations of Nearctic species and the description of a new species from Panama.

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.

Climate as a possible driver of gall morphology in the chestnut pest Dryocosmus kuriphilus across Spanish invaded areas.

The alien cynipid wasp Dryocosmus kuriphilus Yasumatsu, 1951 is a serious pest of chestnuts (Castanea spp.) in Japan, North America and Europe, causing fruit losses while inducing galls in buds. While D. kuriphilus galls have a recognizable and roughly invariable globular shape, their size varies, reaching up to 4 cm in diameter. Among other factors, such variation may depend on different climatic conditions in different attacked areas. Here, we sampled and measured 375 D. kuriphilus galls from 25 localities throughout the Iberian Peninsula, including 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 morphology. The analyses indicate that gall mass and volume follow a pattern that can be associated with a climatic cline. In particular, the Eurosiberian galls were smaller than the Mediterranean galls according to differences in climatic conditions. In the southern areas, the greater insolation regime does not allow the chestnut trees to be distributed at lower altitudes, but the high rainfall and humidity regime of the mountain enclaves allow their presence. These conditions of insolation and precipitation seem to influence the morphological characteristics of the galls of D. kuriphilus.

Description of three new species of oak gallwasps of the genus Amphibolips Reinhard from Mexico (Hymenoptera, Cynipidae).

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 & Castillejos-Lemus. 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.

Variation in parasitoidism of Protocalliphora azurea (Diptera: Calliphoridae) by Nasonia vitripennis (Hymenoptera: Pteromalidae) in Spain.

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.

Andricus Barriosi: a new species of oak gall wasp (Hymenoptera: Cynipidae: Cynipini) from Panama.

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.

Genomic dissection of an extended phenotype: Oak galling by a cynipid gall wasp.

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 innovations associated with cynipid gall induction.

Terminal-instar larval systematics and biology of west European species of Ormyridae associated with insect galls (Hymenoptera, Chalcidoidea).

A systematic study of the genus Ormyrus (Chalcidoidea, Ormyridae) was conducted based on the morphology and biology of the terminal-instar larvae of ten west European species that are parasitoids of gall wasps and gallflies of the families Cynipidae, Eurytomidae and Tephritidae. The first detailed descriptions are provided of the terminal-instar larvae of these ten species using SEM images to illustrate diagnostic characters with systematic values. A key is provided for the identification of ormyrid larvae associated with galls in Europe, which is based particularly on characters of the head, mouthparts and mandibles. Although only limited informative variation in body shape was found, the setation of the head provided several characters of potential taxonomic value. The larval biology of the ten ormyrid species inhabiting different galls is also summarised. Although Ormyrus larvae are usually solitary idiobiont ectoparasitoids of the host larva of various gall-inhabiting insects, evidence of secondary phytophagy was observed in some species.

Taxonomy and ecology of a new species of Corynura (Hymenoptera: Halictidae: Augochlorini) from Chile and Argentina.

We describe a new species of Corynura Spinola, a socially polymorphic genus within the bee tribe Augochlorini (Halictidae). We present a detailed description of both sexes of Corynura (Corynura) moscosensis n. sp. González-Vaquero, images of diagnostic morphological structures and antennal sensilla, floral associations, distributional data and notes on its nesting biology. Corynura moscosensis n. sp. is similar in appearance to C. (C.) chloromelas (Alfken), but it can be distinguished by the sculpturing of the mesoscutum and the hair types on the propodeum in the female, and by the genitalia and extent of the area exclusively covered by sensilla placoidea on the antennal flagellum of the male. A lectotype is designated for C. chloromelas. A Chilean Patagonian population of C. moscosensis n. sp. was observed to dig spatially clumped nests in an earth bank with a moderately hard and humid soil. The presence of multiple females within single nest tunnels is unusual and may preliminary suggest nest-sharing, but more data are necessary to define the social organization of this species. The species appears to be polylectic. Bee nests were attacked by satellite flies and possibly velvet ants.

Spectral cathodoluminescence analysis of hymenopteran mandibles with different levels of zinc enrichment in their teeth.

The inclusion of Zn in insect mandibles affects their hardness and is functional to their use during feeding or reproducing. However, little is known on the chemical/structural base of Zn enrichment. Here, we found that cathodoluminescence (CL) technique revealed two different types of CL spectra in the mandibles of Hymenoptera, depending on the Zn enrichment level assessed by Energy Dispersive X-ray Spectroscopy (EDS). Individuals having negligible traces to low % of Zn in their mandible teeth (≤3 wt%) presented a wide band of luminescence in the visible range which resembled those observed in the CC structures of graphite. This spectrum is probably characteristic for un-enriched cuticle, since it did not differ from those obtained from the Zn-lacking inner part of mandibles. Individuals with moderate to high % of Zn in their mandible teeth (≥7 wt%), instead, presented additional CL peaks in the ultraviolet range. Comparisons with different minerals of Zn suggest that these peaks could be related with OZnO bonds, with hydroxyl groups and with zinc-chlorine links (in agreement with Cl high levels detected by the EDS). Being a non-destructive technique, CL allows large comparative studies of the chemistry of metal-enriched insect cuticle even using unique specimens, such as those deposited in Natural History Museums.

Pheomelanin in the secondary sexual characters of male parasitoid wasps (Hymenoptera: Pteromalidae).

The occurrence and distribution of eumelanin and pheomelanin, the most prevalent biological pigments, has been rarely investigated in insects. Particularly yellowish to brownish body parts, which in many vertebrates are associated with pheomelanin, are visible in many insects but their chemical nature was rarely examined to a similar detail. Here, by using Dispersive Raman spectroscopy analysis, we found both eumelanin and pheomelanin in different body parts of male parasitoid wasps of three species of the genus Mesopolobus (Hymenoptera: Pteromalidae), which are known to have species-specific spots and coloured stripes on the legs and/or antennae which are displayed to females during courtship. We found a strong eumelanin signal in the antennal clava of all studied Mesopolobus species and in the circular black spot or callosity and the triangular black projection on the outer apical angle of the typically expanded middle tibia of Mesopolobus tibialis and Mesopolobus xanthocerus. Eumelanin was also the predominant pigment in the black thorax of Mesopolobus and other members of the family. Pheomelanin, on the other hand, was detected as predominant only in certain body parts of M. tibialis and M. xanthocerus, precisely in a very narrow, longitudinal brownish stripe on the middle femur and, only in M. tibialis, in a brownish oval-longitudinal stripe on the middle tibia. The two melanin types co-occurred in most pigmented areas, but more often one is clearly predominant relative to the other, according to the variation of Raman signal intensity of their signature peaks. A further tibial yellowish-orange stripe present in both these species did not include melanins of any type. Pheomelanin, could be more widespread than previously known in insects. A convergent evolution of melanin-based male sexual ornaments between vertebrates (e.g. bird feathers) and wasps can be suggested, opening to a new line of comparative evolutionary studies.

Revision of the Neotropical genus Eschatocerus Mayr (Hymenoptera, Cynipidae, Eschatocerini) with biological notes and the first description of the terminal larva.

The gall wasp genus Eschatocerus (Hymenoptera, Cynipidae, Eschatocerini), a cynipid genus of gall inducers on Prosopis and Acacia species (Fabaceae), endemic to southern South America, is revised. Complete descriptions of the external morphology of the genus and its three known species, illustrated with scanning electron photographs, are given for the first time, and an updated key for the identification of the species is provided. The biology of the species of Eschatocerus and their galls is described. Host plant associations are given, and the terminal larva of Eschatocerus niger is described for the first time. Preliminary notes on the inquiline and parasitoid community associated with the galls of Eschatocerus species are also given.

Phylogeny, evolution and classification of gall wasps: the plot thickens.

Gall wasps (Cynipidae) represent the most spectacular radiation of gall-inducing insects. In addition to true gall formers, gall wasps also include phytophagous inquilines, which live inside the galls induced by gall wasps or other insects. Here we present the first comprehensive molecular and total-evidence analyses of higher-level gall wasp relationships. We studied more than 100 taxa representing a rich selection of outgroups and the majority of described cynipid genera outside the diverse oak gall wasps (Cynipini), which were more sparsely sampled. About 5 kb of nucleotide data from one mitochondrial (COI) and four nuclear (28S, LWRh, EF1alpha F1, and EF1alpha F2) markers were analyzed separately and in combination with morphological and life-history data. According to previous morphology-based studies, gall wasps evolved in the Northern Hemisphere and were initially herb gallers. Inquilines originated once from gall inducers that lost the ability to initiate galls. Our results, albeit not conclusive, suggest a different scenario. The first gall wasps were more likely associated with woody host plants, and there must have been multiple origins of gall inducers, inquilines or both. One possibility is that gall inducers arose independently from inquilines in several lineages. Except for these surprising results, our analyses are largely consistent with previous studies. They confirm that gall wasps are conservative in their host-plant preferences, and that herb-galling lineages have radiated repeatedly onto the same set of unrelated host plants. We propose a revised classification of the family into twelve tribes, which are strongly supported as monophyletic across independent datasets. Four are new: Aulacideini, Phanacidini, Diastrophini and Ceroptresini. We present a key to the tribes and discuss their morphological and biological diversity. Until the relationships among the tribes are resolved, the origin and early evolution of gall wasps will remain elusive.

Comparative flight morphology in queens of invasive and native Patagonian bumblebees (Hymenoptera: Bombus).

Since its introduction in Chile, the European Bombus terrestris L. (Hymenoptera: Apidae) has progressively reduced the abundance of the native Patagonian bumblebee, Bombus dahlbomii Guérin. Because an important cause of successful invasion of a species may depend on a potentially advantageous phenotype, we studied morphologies related to flight performance (flight muscle ratio (FMR), wing loading (WL), excess power index (EPI, which integrates FMR and WL) and wing aspect ratio (AR)) in the queens of the two species. Previous empirical studies showed that greater FMR, AR and EPI, and lower WL increase flight performance. In the Patagonian Chilean fjord where the study was carried out, B. dahlbomii was 40% heavier than B. terrestris, a difference theoretically allowing the queens of the native species to take off with heavier loads, despite the fact that the two species have virtually identical FMRs. However, FMR negatively depended on body mass at the intra-specific level. The total wing area was 35% greater in B. dahlbomii, but the difference in forewing length was only of 16%. Once taken into account the effect of body size, WL, was significantly lower in B. terrestris. AR increased with body mass and did not differ between species. EPI was weakly but significantly higher in B. terrestris. Experiments formally linking such parameters with flight performance may help to explain the observed quick and wide spread of this alien species in Patagonia in the last few years.

The type specimens of sawflies (Hymenoptera: Symphyta) of the Museo Nacional de Ciencias Naturales, Madrid.

The type specimens of sawflies (Hymenoptera: Symphyta) housed in the Museo Nacional de Ciencias Naturales, Madrid, were examined. Lectotypes are designated and illustrated for the following 32 nominal taxa (preserved in the MNCN collection if not stated otherwise): Tenthredo acutiscutis Konow, 1908; Tenthredo aericeps Konow, 1907; Allantus albipectus Konow, 1907; Athalia bolivari Dusmet, 1896; Tristactus punctatus var. candidatus Konow, 1899; Tenthredo capistrata Konow, 1907; Megalodontes capitalatus Konow 1904 (coll. SDEI); Tenthredo casta Konow, 1908; Clydostomus cestatus Konow, 1908; Miocephala chalybea Konow, 1907 (coll. SDEI); Peus cupreiceps Konow, 1907; Metallopeus cupreolus Malaise, 1945 (coll. NHRS); Allantus dusmeti Konow, 1894 (coll. SDEI); Megalodontes dusmeti Enslin, 1914 (coll. ZSM); Megalodontes escalerai Konow, 1899; Tenthredo flavitarsis Konow, 1908; Sciopteryx galerita Konow, 1907; Tenthredo habenata Konow, 1907; Allantus inguinalis Konow, 1908; Clydostomus merceti Konow, 1908; Megalodontes merceti Konow 1904 (coll. SDEI); Tenthredo mordax Konow, 1908; Megalodontes mundus Konow, 1904; Tenthredo nimbata Konow, 1906; Tenthredo oculissima Konow, 1907; Peus pannulosus Konow, 1907; Tenthredo podagrica Konow, 1907; Arge segmentaria var. rufiventris Konow, 1899; Tenthredo rugiceps Konow, 1908; Tenthredo segrega Konow, 1908; Peus splendidus Konow 1907; Tenthredo suta Konow, 1906. Peus cupreiceps Konow, 1907, is considered to be a valid species. New synonymy is proposed for Tenthredo (Metallopeus) cupreiceps (Konow, 1907), comb. nov., spec. rev. (=Metallopeus cupreolus Malaise, 1945, syn. nov.; =Metallopeus inermis Malaise, 1945, syn. nov.). 

Barucynips panamensis , a new genus and species of oak gallwasps (Hymenoptera, Cynipidae, Cynipini) from Panama, and description of one new species of Coffeikokkos.

Barucynips panamensis Medianero & Nieves-Aldrey, a new genus and species of oak gallwasps (Hymenoptera: Cynipidae: Cynipini), is described from adults reared from galls on Quercus bumelioides in Panama. The new genus is taxonomically close to the recently described Coffeikokkos from Costa Rica, but differs from it and all of the described genera of Cynipini, by the shape and setation of the projecting part of the ventral spine of the hypopygium and by the sculpture of the propodeum. A new species of Coffeikokkos is also described from the same area, the Volcán Barú in Panama. Diagnostic characters, gall description, distribution, and biological data of the new genus and the two new species are given. The new genus is the first genus of oak gallwasps of the tribe Cynipini described in Panama.

Catalogue of parasitoids and inquilines in cynipid oak galls in the West Palaearctic.

A quantitative catalogue of the parasitoids (almost exclusively Chalcidoidea) and inquiline Cynipidae recorded in the western Palaearctic from galls induced on Quercus by Cynipidae (Cynipini) is presented. Quantitative and national data are included with bibliographic references to almost all records published in 2011 and earlier. The catalogue is followed by two checklists, firstly one of the Chalcidoidea with numbers of each species recorded from each type of host gall (galls of the sexual and asexual generations of the host gall wasps are listed separately), and secondly one of inquiline Cynipidae with host galls. Compared to non-oak gall wasps, the Cynipini support a much larger parasitoid and especially inquiline fauna, and this fauna is very largely restricted at the species level to Cynipini galls. About one hundred chalcidoid species are recorded from galls of Cynipini, distributed over six families: Pteromalidae and Eulophidae (29 species each), Torymi-dae (21 species), Eurytomidae (10 species), Eupelmidae (8 species) and Ormyridae (at least 2 species). Polyphagy is usual in the chalcidoid parasitoids, most species having a broad host gall range, but quantitatively the fauna of each type of oak gall is rather characteristic and is strongly influenced by gall morphology, situation on the tree, season of growth and host tree species. These and other extrinsic factors restrict the full exploitation of the chalcidoids' potential host gall range.

Reconstructing community assembly in time and space reveals enemy escape in a Western Palearctic insect community.

How geographically widespread biological communities assemble remains a major question in ecology. Do parallel population histories allow sustained interactions (such as host-parasite or plant-pollinator) among species, or do discordant histories necessarily interrupt them? Though few empirical data exist, these issues are central to our understanding of multispecies evolutionary dynamics. Here we use hierarchical approximate Bayesian analysis of DNA sequence data for 12 herbivores and 19 parasitoids to reconstruct the assembly of an insect community spanning the Western Palearctic and assess the support for alternative host tracking and ecological sorting hypotheses. We show that assembly occurred primarily by delayed host tracking from a shared eastern origin. Herbivores escaped their enemies for millennia before parasitoid pursuit restored initial associations, with generalist parasitoids no better able to track their hosts than specialists. In contrast, ecological sorting played only a minor role. Substantial turnover in host-parasitoid associations means that coevolution must have been diffuse, probably contributing to the parasitoid generalism seen in this and similar systems. Reintegration of parasitoids after host escape shows these communities to have been unsaturated throughout their history, arguing against major roles for parasitoid niche evolution or competition during community assembly.

Community impacts of anthropogenic disturbance: natural enemies exploit multiple routes in pursuit of invading herbivore hosts.

BACKGROUND: Biological invasions provide a window on the process of community assembly. In particular, tracking natural enemy recruitment to invading hosts can reveal the relative roles of co-evolution (including local adaptation) and ecological sorting. We use molecular data to examine colonisation of northern Europe by the parasitoid Megastigmus stigmatizans following invasions of its herbivorous oak gallwasp hosts from the Balkans. Local host adaptation predicts that invading gallwasp populations will have been tracked primarily by sympatric Balkan populations of M. stigmatizans (Host Pursuit Hypothesis). Alternatively, ecological sorting allows parasitoid recruitment from geographically distinct populations with no recent experience of the invading hosts (Host Shift Hypothesis). Finally, we test for long-term persistence of parasitoids introduced via human trade of their hosts' galls (Introduction Hypothesis). RESULTS: Polymorphism diagnostic of different southern refugial regions was present in both mitochondrial and nuclear microsatellite markers, allowing us to identify the origins of northern European invaded range M. stigmatizans populations. As with their hosts, some invaded range populations showed genetic variation diagnostic of Balkan sources, supporting the Host Pursuit Hypothesis. In contrast, other invading populations had an Iberian origin, unlike their hosts in northern Europe, supporting the Host Shift Hypothesis. Finally, both British and Italian M. stigmatizans populations show signatures compatible with the Introduction Hypothesis from eastern Mediterranean sources. CONCLUSIONS: These data reveal the continental scale of multi-trophic impacts of anthropogenic disturbance and highlight the fact that herbivores and their natural enemies may face very different constraints on range expansion. The ability of natural enemies to exploit ecologically-similar hosts with which they have had no historical association supports a major role for ecological sorting processes in the recent assembly of these communities. The multitude of origins of invading natural enemy populations in this study emphasises the diversity of mechanisms requiring consideration when predicting consequences of other biological invasions or biological control introductions.

Concordant phylogeography and cryptic speciation in two Western Palaearctic oak gall parasitoid species complexes.

Little is known about the evolutionary history of most complex multi-trophic insect communities. Widespread species from different trophic levels might evolve in parallel, showing similar spatial patterns and either congruent temporal patterns (Contemporary Host-tracking) or later divergence in higher trophic levels (Delayed Host-tracking). Alternatively, host shifts by natural enemies among communities centred on different host resources could disrupt any common community phylogeographic pattern. We examined these alternative models using two Megastigmus parasitoid morphospecies associated with oak cynipid galls sampled throughout their Western Palaearctic distributions. Based on existing host cynipid data, a parallel evolution model predicts that eastern regions of the Western Palaearctic should contain ancestral populations with range expansions across Europe about 1.6 million years ago and deeper species-level divergence at both 8-9 and 4-5 million years ago. Sequence data from mitochondrial cytochrome b and multiple nuclear genes showed similar phylogenetic patterns and revealed cryptic genetic species within both morphospecies, indicating greater diversity in these communities than previously thought. Phylogeographic divergence was apparent in most cryptic species between relatively stable, diverse, putatively ancestral populations in Asia Minor and the Middle East, and genetically depauperate, rapidly expanding populations in Europe, paralleling patterns in host gallwasp species. Mitochondrial and nuclear data also suggested that Europe may have been colonized multiple times from eastern source populations since the late Miocene. Temporal patterns of lineage divergence were congruent within and across trophic levels, supporting the Contemporary Host-tracking Hypothesis for community evolution.