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.

Environmental factors driving the abundance of Philaenus spumarius in mesomediterranean habitats of Corsica (France).

Philaenus spumarius (Ps) is considered the main insect vector of the bacterium Xylella fastidiosa (Xf) in Europe. As such, it is a key actor of the Xf pathosystem on which surveillance and management strategies could be implemented. Although research effort has increased in the past years, the ecological factors shaping Ps abundance and distribution across landscapes are still poorly known in most regions of Europe. We selected 64 plots of 500m2 in Corsican semi-natural habitats in which we sampled nymphs and adults of Ps during three years. While local or surrounding vegetation structure (low or high scrubland) had little effect on Ps abundance, we highlighted a positive relationship between Ps abundance and the density of Cistus monspeliensis in the plots. We also found larger populations of Ps in cooler and moister plots. The pattern of host association highlighted here is unique, which calls for more studies on the ecology of Ps in Europe, to help designing surveillance and management strategy for Xf.

Vectors as Sentinels: Rising Temperatures Increase the Risk of Xylella fastidiosa Outbreaks.

Global change is expected to modify the threat posed by pathogens to plants. However, little is known regarding how a changing climate will influence the epidemiology of generalist vector-borne diseases. We developed a high-throughput screening method to test for the presence of a deadly plant pathogen, Xylella fastidiosa, in its insect vectors. Then, using data from a four-year survey in climatically distinct areas of Corsica (France), we demonstrated a positive correlation between the proportion of vectors positive to X. fastidiosa and temperature. Notably, a higher prevalence corresponded with milder winters. Our projections up to 2100 indicate an increased risk of outbreaks. While the proportion of vectors that carry the pathogen should increase, the climate conditions will remain suitable for the bacterium and its main vector, with possible range shifts towards a higher elevation. Besides calling for research efforts to limit the incidence of plant diseases in the temperate zone, this work reveals that recent molecular technologies could and should be used for massive screening of pathogens in vectors to scale-up surveillance and management efforts.

Ultra-Conserved Elements and morphology reciprocally illuminate conflicting phylogenetic hypotheses in Chalcididae (Hymenoptera, Chalcidoidea).

Recent technical advances combined with novel computational approaches have promised the acceleration of our understanding of the tree of life. However, when it comes to hyperdiverse and poorly known groups of invertebrates, studies are still scarce. As published phylogenies will be rarely challenged by future taxonomists, careful attention must be paid to potential analytical bias. We present the first molecular phylogenetic hypothesis for the family Chalcididae, a group of parasitoid wasps, with a representative sampling (144 ingroups and seven outgroups) that covers all described subfamilies and tribes, and 82% of the known genera. Analyses of 538 Ultra-Conserved Elements (UCEs) with supermatrix (RAxML and IQTREE) and gene tree reconciliation approaches (ASTRAL, ASTRID) resulted in highly supported topologies in overall agreement with morphology but reveal conflicting topologies for some of the deepest nodes. To resolve these conflicts, we explored the phylogenetic tree space with clustering and gene genealogy interrogation methods, analyzed marker and taxon properties that could bias inferences and performed a thorough morphological analysis (130 characters encoded for 40 taxa representative of the diversity). This joint analysis reveals that UCEs enable attainment of resolution between ancestry and convergent/divergent evolution when morphology is not informative enough, but also shows that a systematic exploration of bias with different analytical methods and a careful analysis of morphological features is required to prevent publication of artifactual results. We highlight a GC content bias for maximum-likelihood approaches, an artifactual mid-point rooting of the ASTRAL tree and a deleterious effect of high percentage of missing data (>85% missing UCEs) on gene tree reconciliation methods. Based on the results we propose a new classification of the family into eight subfamilies and ten tribes that lay the foundation for future studies on the evolutionary history of Chalcididae.

Citizen science and niche modeling to track and forecast the expansion of the brown marmorated stinkbug Halyomorpha halys (Stål, 1855).

Halyomorpha halys (Stål, 1855), the Brown Marmorated StinkBug (BMSB) is a highly successful invasive species native to eastern Asia that managed to spread into North America and Europe in recent decades. We set up a citizen science survey to monitor BMSB expansion in France in 2012 and analyzed the data it yielded between 2012 and 2019 to examine the local expansion of the insect. These data were gathered with occurrences form various sources (GBIF, literature) to calibrate a species niche model and assess potential current BMSB range. We evaluated the potential changes to the BMSB range due to climate change by projecting the model according to 6 global circulation models (GCM) and the shared socio-economic pathways SSP245 in two time periods 2021-2040 and 2041-2060. Citizen science allowed to track BMSB expansion in France and provided information about its phenology and its habitat preferences. The model highlighted the potential for further range expansion in Europe and illustrated the impact of climate change. These results could help managing the current BMSB invasion and the framework of this survey could contribute to a better preparedness of phytosanitary authorities either for the BMSB or other invasive pests.

A crowdsourcing approach to track the expansion of the brown marmorated stinkbug Halyomorpha halys (Stål, 1855) in France.

BACKGROUND: Halyomorpha halys (Stål, 1855), the brown marmorated stinkbug (BMSB) is a highly successful invasive species, native to eastern Asia. It has managed to spread into North America and Europe in recent decades, causing severe damage to various crops. BMSB has been detected in Europe in 2004 and has since expanded in more than 20 countries from Sweden to Greece and Spain to Turkey, the South European Territory of Russia (Krasnodar region) and Abkhazia. In 2012, we set up a citizen science survey to monitor BMSB expansion in France. NEW INFORMATION: The present crowdsourcing survey was initiated in 2012 and provided a large number of occurrence points of BMSB. These data allowed to track the expansion of the species in France from 2012 to 2019 and brought information about its phenology and distribution in various habitats. The dataset comprises both valid and invalid sightings, thereby allowing us to examine changes in the quality of citizen reports during the course of the survey. Despite a large proportion of misidentifications, the survey provided a large number of valid occurrences. Furthermore, valuable information on hemipterans of Pentatomidae and Coreidae families entering habitations were also gathered. The dataset also illustrates that, although designed for a large public, the Agiir application was mostly used by urban dwellers with very few sightings stemming from professionals of agricultural sectors.

Ooctonus vulgatus (Hymenoptera, Mymaridae), a potential biocontrol agent to reduce populations of Philaenus spumarius (Hemiptera, Aphrophoridae) the main vector of Xylella fastidiosa in Europe.

As a vector of Xylella fastidiosa (Wells, 1987) in Europe, the meadow spittlebug Philaenus spumarius (Linnaeus, 1758) (Hemiptera, Aphrophoridae) is a species of major concern. Therefore, tools and agents to control this ubiquitous insect that develops and feeds on hundreds of plant species are wanted. We conducted a field survey of P. spumarius eggs in Corsica and provide a first report of Ooctonus vulgatus Haliday, 1833 (Hymenoptera, Mymaridae) as a potential biocontrol agent of P. spumarius in Europe. To allow species identification, we summarized the main characters distinguishing O. vulgatus from other European species of Ooctonus and generated COI DNA barcodes. Parasitism rates were variable in the four localities included in the survey but could reach 69% (for an average number of eggs that hatched per locality of 109). Based on the geographic occurrences of O. vulgatus obtained from the literature, we calibrated an ecological niche model to assess its potential distribution in the Holarctic. Obviously, several questions need to be addressed to determine whether O. vulgatus could become an effective biocontrol agent of P. spumarius in Europe. So far, O. vulgatus has been reared only from P. spumarius eggs, but its exact host-range should be evaluated to ensure efficiency and avoid non-target effect. The top-down impact of the parasitoid on vector populations should also be assessed on large data sets. Finally, the feasibility of mass rearing should be tested. We hope this report serves as a starting point to initiate research on this parasitoid wasp to assess whether it could contribute to reduce the spread and impact of X. fastidiosa in Europe.