Marie-Anne Auger-Rozenberg

Temporal and interspecific variation in rates of spread for insect species invading Europe during the last 200 years

Globalization is triggering an increase in the establishment of alien insects in Europe, with several species having substantial ecological and economic impacts. We investigated long-term changes in rates of species spread following establishment. We used the total area of countries invaded by 1171 insect species for which the date of first record in Europe is known, to estimate their current range radius (calculated as [invaded area]0.5/π). We estimated initial rates of radial spread and compared them among different groups of insects for all years (1800–2014) and for a subset of more recent decades (1950–2014). Accidentally introduced species spread faster than intentionally introduced species. Considering the whole period 1800–2014, spread patterns also differ between feeding guilds, with decreasing spread rates over residence time in herbivores but not in detritivores or parasitic species. These decreases for herbivorous species appeared mainly in those associated with herbaceous plants and crops rather than woody plants. Initial spread rate was significantly greater for species detected after 1990, roughly 3–4 times higher than for species that arrived earlier. We hypothesize that the political changes in Europe following the collapse of the Iron Curtain in 1989, and the further dismantling of customs checkpoints within an enlarged European Union (EU) have facilitated the faster spread of alien insect species. Also, the number of species first recorded in the Eastern Bloc of the politically-divided Europe before 1989 was lower than for the rest of Europe. A detailed analysis of six recent invaders indicated a dominant role of long-distance translocations related to human activities, especially with the plant trade, in determining rates of spread.

Complex patterns of global spread in invasive insects: eco-evolutionary and management consequences.

Abstract The advent of simple and affordable tools for molecular identification of novel insect invaders and assessment of population diversity has changed the face of invasion biology in recent years. The widespread application of these tools has brought with it an emerging understanding that patterns in biogeography, introduction history and subsequent movement and spread of many invasive alien insects are far more complex than previously thought. We reviewed the literature and found that for a number of invasive insects, there is strong and growing evidence that multiple introductions, complex global movement, and population admixture in the invaded range are commonplace. Additionally, historical paradigms related to species and strain identities and origins of common invaders are in many cases being challenged. This has major consequences for our understanding of basic biology and ecology of invasive insects and impacts quarantine, management and biocontrol programs. In addition, we found that founder effects rarely limit fitness in invasive insects and may benefit populations (by purging harmful alleles or increasing additive genetic variance). Also, while phenotypic plasticity appears important post-establishment, genetic diversity in invasive insects is often higher than expected and increases over time via multiple introductions. Further, connectivity among disjunct regions of global invasive ranges is generally far higher than expected and is often asymmetric, with some populations contributing disproportionately to global spread. We argue that the role of connectivity in driving the ecology and evolution of introduced species with multiple invasive ranges has been historically underestimated and that such species are often best understood in a global context.

Molecular phylogeny and evolution of host‐plant use in conifer seed chalcids in the genus Megastigmus (Hymenoptera: Torymidae)

Abstract.  Phylogenetic relationships amongst Megastigmus species (Chalcidoidea: Torymidae) associated with conifer seeds were inferred from DNA sequence data. Twenty‐nine species of seed chalcids were analysed using two different genes, cytochrome b (mitochondrial DNA) and the D2 domain of the 28S ribosomal DNA. Maximum‐parsimony and maximum‐likelihood analyses showed that taxa formed two monophyletic groups, one clade comprising all species associated with Cupressaceae and Taxodiaceae hosts with the exception of Chamaecyparis, and the other clade composed of species associated with Pinaceae. Species infesting Cupressaceae and Taxodiaceae seemed to be specialized to particular host genera or even to be species specific, which was consistent with a taxonomic radiation following initial host adaptation. By contrast, Megastigmus species associated with Pinaceae appeared capable of shifting onto different congeneric species or even onto a new host genus, with their evolution apparently less constrained by plant association. We hypothesized that the Megastigmus group associated with Pinaceae may have a much higher invasive potential than that related to Cupressaceae. The study also confirmed the presence of invasive Nearctic species in the Palaearctic, and demonstrated the existence of a cryptic species complex.

Competition between Exotic and Native Insects for Seed Resources in Trees of a Mediterranean Forest Ecosystem

The seeds of both cedar-of-Lebanon (Cedrus libani) and Cyprus cedar (Cedrus brevifolia) are attacked in their natural range by a specialised chalcid, Megastigmus schimitscheki. From 1995 to 1999, seeds were screened for insect damage in the main cedar plantations of southern France, as well as in the stands where cedar is mixed with firs (Abies spp.). X-rays were used to identify chalcid-infested seeds from which the insects were then reared. The surveys revealed the presence of M. schimitscheki in all the stands of Atlas cedar, Cedrus atlantica, planted at Mt Ventoux, southeastern France. The chalcid also infested seeds of an exotic fir, Abies pinsapo, planted in the same area. However, it has not yet reached the cedar plantations in southwestern France, where the seeds are colonised by a related exotic insect, Megastigmus pinsapinis, originating from North Africa. The latter species was common in cedar seeds at Mt Ventoux in the early 1990s but seems to have been supplanted by M. schimitscheki in the invasion zone. A native chalcid species, Megastigmus suspectus, was also shown to have shifted to a slight extent from a native fir, A. alba, onto cedar. The presence of three chalcid species competing for cedar seed resources may result in a substantial decline of the regeneration potential of that tree species. At Mt Ventoux, up to 92.6% of the cedar seeds were attacked, with 86.8% due to M. schimitscheki. The survey also revealed the widespread presence of another North American chalcid, Megastigmus rafni, in the fir stands of southern France.

Relationship of Scots pine clone characteristics and water stress to hatching and larval performance of the sawfly Diprion pini (Hymenoptera : Diprionidae)

Abstract Most Symphyta are phytophagous and lay their eggs in the host plant in contact with the plant tissue. We tested the hypothesis that Diprion pini L. egg hatching and larval survival vary between Scots pine clones with particular foliage characteristics. We also tested the influence of water limitation on hatching rates. Larval survival and hatching rate of D. pini eggs differed significantly among Pinus sylvestris L. genotypes and they were not correlated. We confirmed that the presence of taxifolin, a phenolic compound, and quantity of 3-carene were correlated with low larval survival. Foliage analysis showed an inverse relationship between hatching rate and dry matter content of foliage. The hatching rate on a tree varies, according to needle water content. This suggests that the success of egg development might be important for the population dynamics of D. pini.

Inferences on population history of a seed chalcid wasp: invasion success despite a severe founder effect from an unexpected source population

Most invasive species established in Europe originate from either Asia or North America, but little is currently known about the potential of the Anatolian Peninsula (Asia Minor) and/or the Near East to constitute invasion sources. Mediterranean forests are generally fragile ecosystems that can be threatened by invasive organisms coming from different regions of the Mediterranean Basin, but for which historical data are difficult to gather and the phylogeographic patterns are still poorly understood for most terrestrial organisms. In this study, we characterized the genetic structure of Megastigmus schimitscheki, an invasive seed‐feeding insect species originating from the Near East, and elucidated its invasion route in South‐eastern France in the mid 1990s. To disentangle the evolutionary history of this introduction, we gathered samples from the main native regions (Taurus Mountains in Turkey, Lebanon and Cyprus) and from the invaded region that we genotyped using five microsatellite markers and for which we sequenced the mitochondrial Cytochrome Oxidase I gene. We applied a set of population genetic statistics and methods, including approximate Bayesian computation. We proposed a detailed phylogeographic pattern for the Near East populations, and we unambiguously showed that the French invasive populations originated from Cyprus, although the available historical data strongly suggested that Turkey could be the most plausible source area. Interestingly, we could show that the introduced populations were founded from an extremely restricted number of individuals that realized a host switch from Cedrus brevifolia to C. atlantica. Evolutionary hypotheses are discussed to account for this unlikely scenario.

Seed wasp invasions promoted by unregulated seed trade affect vegetal and animal biodiversity

Cone and seed insects are considered the most important predators of tree seeds during the pre-dispersal phase of development. Among them, exotic seed chalcids in the genus Megastigmus invaded Europe as a result of the rapidly-increasing and mostly unregulated seed trade for afforestation and ornamental plantations. Unlike their economic impact in seed orchards, until recently, little attention was paid to the ecological impact of these insects. In the present study, selected case studies of alien Megastigmus spp. were considered to assess their specific impact on the potential of natural regeneration of native woody plants and on the native entomofauna competing for seed resource. We re-analyzed data from former studies that did not focus on these ecological interactions and, here, present previously unpublished results. Seeds of Douglas-fir, true cedars, true firs and wild roses were sampled all over Europe, and the relative importance of the native and invasive chalcid species was assessed as well as their specific impact on seed yield. In most cases, the recent arrival of alien chalcids resulted in a significant decrease in the regeneration potential of the host trees. In the absence of competitors, alien chalcids occupied the entire seed niche in Douglas-fir, but their impact tended to decrease after the arrival of invasive seed bugs. In firs, alien chalcids tended to displace the native chalcids, but not in wild roses and cedars, where their damage was increasing. Different biological traits that might explain invasive success of alien chalcids are discussed. However, no general invasive patterns seem to exist.

Epidemiology of asexuality induced by the endosymbiotic Wolbachia across phytophagous wasp species: host plant specialization matters

Among eukaryotes, sexual reproduction is by far the most predominant mode of reproduction. However, some systems maintaining sexuality appear particularly labile and raise intriguing questions on the evolutionary routes to asexuality. Thelytokous parthenogenesis is a form of spontaneous loss of sexuality leading to strong distortion of sex ratio towards females and resulting from mutation, hybridization or infection by bacterial endosymbionts. We investigated whether ecological specialization is a likely mechanism of spread of thelytoky within insect communities. Focusing on the highly specialized genus Megastigmus (Hymenoptera: Torymidae), we first performed a large literature survey to examine the distribution of thelytoky in these wasps across their respective obligate host plant families. Second, we tested for thelytoky caused by endosymbionts by screening in 15 arrhenotokous and 10 thelytokous species for Wolbachia, Cardinium, Arsenophonus and Rickettsia endosymbionts and by performing antibiotic treatments. Finally, we performed phylogenetic reconstructions using multilocus sequence typing (MLST) to examine the evolution of endosymbiont‐mediated thelytoky in Megastigmus and its possible connections to host plant specialization. We demonstrate that thelytoky evolved from ancestral arrhenotoky through the horizontal transmission and the fixation of the parthenogenesis‐inducing Wolbachia. We find that ecological specialization in Wolbachias hosts was probably a critical driving force for Wolbachia infection and spread of thelytoky, but also a constraint. Our work further reinforces the hypothesis that community structure of insects is a major driver of the epidemiology of endosymbionts and that competitive interactions among closely related species may facilitate their horizontal transmission.

Modelling the impact of an invasive insect via reaction-diffusion.

An exotic, specialist seed chalcid, Megastigmus schimitscheki, has been introduced along with its cedar host seeds from Turkey to southeastern France during the early 1990s. It is now expanding in plantations of Atlas Cedar (Cedrus atlantica). We propose a model to predict the expansion and impact of this insect. This model couples a time-discrete equation for the ovo-larval stage with a two-dimensional reaction-diffusion equation for the adult stage, through a formula linking the solution of the reaction-diffusion equation to a seed attack rate. Two main diffusion operators, of Fokker-Planck and Fickian types, are tested. We show that taking account of the dependence of the insect mobility with respect to spatial heterogeneity, and choosing the appropriate diffusion operator, are critical factors for obtaining good predictions.

Potential invasion of China by exotic insect pests associated with tree seeds

A total of 39 insect species, mostly seed chalcids in the genus Megastigmus (Hymenoptera), but also midges (Diptera), are listed as potential seed-borne invaders of Chinese conifers. Although the number of native seed insects per conifer genus does not differ between China and other biogeographical regions, there are significantly fewer seed insects associated with each conifer genus in China than potential invaders. The eventual success of the invaders is likely to depend on the presence of native Chinese conifers that are congeneric with the original host, or on the presence of the original host as an exotic. When a substantial entomofauna is already associated with cones, competition for seed resources may limit the potential impact of invaders because seed insects are usually the last organisms to colonize the cone. A survey of 26 fir species, both native and introduced to Europe, showed that overall seed infestation by five species of exotic chalcids is negatively correlated to levels of damage by native insects, except on the original hosts of the chalcids. Similar patterns are hypothesized for native firs, spruces, Douglas firs, and larches in China. Uncontrolled importation of seeds and nuts of broad-leaved trees could also facilitate the introduction of seed chalcids, seed bruchids, tortricid moths and nut weevils into China. Only six species of seed chalcids are present in China, out of the 72 known to attack broad-leaved seeds over the world.