Studies of insect host-parasitoid relationships are often confounded by the difficulties associated with species delimitation in taxonomically challenging groups. Eurytomidae (Hymenoptera) are common parasitoids associated with galls induced by Cynipidae (Hymenoptera) and are difficult to identify due to their small size, morphological conservatism, and unreliable published host records. This study tests the species limits of eurytomids associated with galls induced by Diplolepis Geoffroy (Hymenoptera: Cynipidae) in Canada using an integrative taxonomy approach including adult morphology, the mitochondrial gene cytochrome c oxidase I, host records, and geographical range. Incongruences between morphological and molecular data were found within the Eurytoma discordans Bugbee complex, as Eurytoma discordans, Eurytoma acuta Bugbee, and Eurytoma calcarea Bugbee were shown to be new synonyms. The results also revealed the presence of cryptic species within Eurytoma spongiosa Bugbee. Furthermore, issues that have impeded ecological and biological studies of eurytomids associated with rose galls such as host specificity and sex association were resolved using DNA barcodes, providing new insights into the evolutionary history of this difficult group.
Targeted enrichment of ultraconserved elements (UCEs) has emerged as a promising tool for inferring evolutionary history in many taxa, with utility ranging from phylogenetic and biogeographic questions at deep time scales to population level studies at shallow time scales. However, the methodology can be daunting for beginners. Our goal is to introduce UCE phylogenomics to a wider audience by summarizing recent advances in arthropod research, and to familiarize readers with background theory and steps involved. We define terminology used in association with the UCE approach, evaluate current laboratory and bioinformatic methods and limitations, and, finally, provide a roadmap of steps in the UCE pipeline to assist phylogeneticists in making informed decisions as they employ this powerful tool. By facilitating increased adoption of UCEs in phylogenomics studies that deepen our comprehension of the function of these markers across widely divergent taxa, we aim to ultimately improve understanding of the arthropod tree of life.
Cynipid gall wasps play an important role in structuring oak arthropod communities. Wasps in the Cynipini tribe typically lay their eggs in oaks (Quercus L.), and induce the formation of a ‘gall’, which is a tumor-like growth of plant material that surrounds the developing wasp. As the wasp develops, the cynipid and its gall are attacked by a diverse community of natural enemies, including parasitoids, hyperparasitoids, and inquilines. Determining what structures these species-rich natural enemy communities across cynipid gall wasp species is a major question in gall wasp biology. Additionally, gall wasps are ecosystem engineers, as the abandoned gall is used by other invertebrates. The gall-associated insect communities residing on live oaks (Quercus geminata Small and Quercus virginiana Mill.) are emerging as a model system for answering ecological and evolutionary questions ranging from community ecology to the evolution of new species. Documenting the arthropods associated with cynipids in this system will expand our understanding of the mechanisms influencing eco-evolutionary processes, record underexplored axes of biodiversity, and facilitate future work. Here, we present the community of natural enemies and other associates of the asexual generation of the crypt gall wasp, Bassettia pallida Ashmead. We compare the composition of this community to communities recently documented from two other cynipid gall wasps specializing on live oaks along the U.S. Gulf coast, Disholcaspis quercusvirens Ashmead and Belonocnema treatae Mayr. B. pallida and their galls support a diverse arthropod community, including over 25 parasitoids, inquilines, and other associated arthropods spanning 5 orders and 16 families.
Gall wasps (Hymenoptera: Cynipidae) specializing on live oaks in the genus Quercus (subsection Virentes) are a relatively diverse and well‐studied community with 14 species described to date, albeit with incomplete information on their biology, life history and genetic structure. Incorporating an integrative taxonomic approach, we combine morphology, phenology, behaviour, genetics and genomics to describe a new species, Neuroterus valhalla sp. nov. The alternating generations of this species induce galls on the catkins and stem nodes of Quercus virginiana and Quercus geminata in the southern United States. We describe both generations in the species' life cycle, and primarily use samples from a population in the centre of Houston, Texas, thus serving as an example of the undescribed biodiversity still present in well‐travelled urban centres. In parallel, we present a draft assembly of the N. valhalla genome providing a direct link between the type specimen and reference genome. The genome of N. valhalla is the smallest reported to date within the tribe Cynipini, providing an important comparative contrast to the otherwise large genome size of cynipids. While relatively small, the genome was found to be composed of >64% repetitive elements, including 43% unclassified repeats and 11% retrotransposons. A preliminary ab initio and homology‐based annotation revealed 32,005 genes, and a subsequent orthogroup analysis grouped 18,044 of these to 8186 orthogroups, with some evidence for high levels of gene duplications within Cynipidae. A mitochondrial barcode phylogeny linked each generation of the new species and a phylogenomic ultraconserved element (UCEs) phylogeny indicates that the new species groups with other Nearctic Neuroterus. However, both phylogenies present the genus Neuroterus in North America as polyphyletic.
Quantifying the frequency of shifts to new host plants within diverse clades of specialist herbivorous insects is critically important to understand whether and how host shifts contribute to the origin of species. Oak gall wasps (Hymenoptera: Cynipidae: Cynipini) comprise a tribe of ~1000 species of phytophagous insects that induce gall formation on various organs of trees in the family Fagacae, primarily the oaks (genus Quercus; ~435 sp). The association of oak gall wasps with oaks is ancient (~50 my), and most oak species are galled by one or more gall wasp species. Despite the diversity of both gall wasp species and their plant associations, previous phylogenetic work has not identified a strong signal of host plant shifting among oak gall wasps. However, most emphasis has been on the Western Palearctic and not the Nearctic where both oaks and oak gall wasps are considerably more species rich and where oaks are more phylogenetically diverse. We collected 86 species of Nearctic oak gall wasps from 10 of the 14 major clades of Nearctic oaks and sequenced >1000 Ultra Conserved Elements (UCEs) and flanking sequences to infer wasp phylogenies. We assessed the relationships of Nearctic gall wasps to one another and, by leveraging previously published UCE data, to the Palearctic fauna. We then used phylogenies to infer historical patterns of shifts among host tree species and tree organs. Our results indicate that oak gall wasps have moved between the Palearctic and Nearctic at least four times, that some Palearctic clades have their proximate origin in the Nearctic, and that gall wasps have shifted within and between oak tree sections, subsections, and organs considerably more often than the analysis of previous data have suggested. Given that host shifts have been demonstrated to drive reproductive isolation between host-associated populations in other phytophagous insects, our analyses of Nearctic gall wasps suggest that host shifts are key drivers of speciation in this clade, especially in hotspots of oak diversity. Though formal assessment of this hypothesis requires further study, two putatively oligophagous gall wasp species in our dataset show signals of host-associated genetic differentiation unconfounded by geographic distance, suggestive of barriers to gene flow associated with the use of alternative host plants.
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