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• Our aim is to present why the hypothesis, that Ophiostomatoid fungi play an important role in the establishment of most bark beetle species on living conifers, is valuable.• After summarizing knowledge about the relationships of bark beetles with conifers and fungi, we conclude that controversy results from misinterpretations when using fungal pathogenicity to demonstrate the role of Ophiostomatoid fungi in beetle establishment on host trees.• We demonstrate that fungal pathogenicity is not the right parameter to appreciate the role of fungus in beetle establishment on host trees. We argue that artificial low density inoculations that allow the appreciation of fungus ability to stimulate tree defenses and thus to help beetles in overcoming tree resistance must be used in complement to mass inoculations. In both cases, results must be expressed in terms of tree defense stimulation rather than in terms of tree killing.Fungal species stimulating tree defenses are generally not those that grow the best in the sapwood.We argue that beetle development in the phloem, fungal invasion of the sapwood and phloem, and tree death, occur after tree defenses are exhausted, and that any fungus present in the beetle gallery could thus potentially invade the sapwood after defense exhaustion.• We conclude that stimulation of the tree defense reactions in both the phloem and the superficial sapwood is a real benefit brought by fungi to the beetles during the first phase of establishment (overcoming tree resistance).• Considering the origin of the bark beetle fungus associations attacking living trees and their general functioning based on stimulation of tree defenses, we develop three hypotheses:any beetle species would be helped in its establishment in a given tree species by developing an association, even loosely, with a fungus species belonging to the Ophiostomatoid flora of that tree species;the necessity of a considerably low level of tree resistance for fungus extension into the tree is the selection pressure that has led fungi to develop their intrinsic ability to stimulate tree defenses, through their ability to grow into the phloem. This association can be completed by antagonistic fungal species controlling extension of the previous fungal species in the tree tissues;Beetle species using the strategy of overcoming tree resistance are associated with a fungal complex, of which species could assume three roles regarding relationships between beetles and trees: 1- to stimulate tree defenses in the phloem and superficial sapwood, 2- to grow into the sapwood after tree resistance is overcome, and 3- to control phloem extension of the first other two categories. Bringing nutrients to the beetle progeny can be a fourth role.• We propose that bark beetle — Ophiostomatoid associations can be categorized, based on associations’ frequency and complexity while taking into account beetle aggressiveness. We show that a close correspondence exists between beetles’ aggressiveness and the ability of their main associated fungal species to stimulate the defenses of their host tree.• We conclude with suggesting that most sapwood invading fungi might be “cheaters” which have taken advantage of the efficiency of the relationship between beetles and fungi that stimulate tree defenses.

Key messageFor an optimal deployment of poplar resistance to the gall-inducing aphid Phloeomyzus passerinii, a laboratory susceptibility assay has been developed. The nature of aphid–tree interactions during compatible and incompatible situations has been studied in detail. This should help at identifying specific resistance markers and at testing effects of site conditions on tree–pest interactions.ContextP. passerinii is a major pest of poplar plantations in Europe, and the plantation of resistant poplar genotypes is regarded as the best long-term management strategy for this pest. This requires a sound knowledge of the interactions among the pest, its host and their environment.AimsHere, we review the recent advances aiming at developing an optimal deployment of host resistance versus P. passerinii.ResultsInvestigations on aphid-host interactions demonstrated that P. passerinii induces pseudogalls within the bark of susceptible hosts. This results in a reduction of starch bark content during aphid outbreaks, which could be involved in tree death. The constitutive level of starch in the bark could be related to the tolerance level of trees. A susceptibility test has been designed for poplar genotypes, discriminating three categories of susceptibility depending on tree’s ability to totally or partially inhibit pseudogall induction. The test still has several limitations however. It neither takes into account the large level of individual genetic diversity of the aphid in France, nor the environmental modulation of tree resistance and tolerance, while water deficit and fertilization could potentially affect these parameters.ConclusionThe hypotheses drawn regarding the processes leading to tree death or resistance should help at identifying resistance markers, and at testing effects of site conditions on tree–pest interactions.

ContextThe bark beetle, Pityogenes chalcographus, is one of the main pests in Picea abies stands, and it has also been found on other European Pinaceae species. With massive introductions of North American conifer species into European forests, this insect has extended its host range to exotic Pinaceae species.AimThis study assessed whether a wider host range has influenced the genetic structure of P. chalcographus and has led to host specialization.MethodsInsects were collected from two different regions of France, where eight native and exotic conifer species coexist and were analyzed using mitochondrial and nuclear genetic markers.ResultsConsiderable haplotypic diversity was observed within the regions and within host species from where P. chalcographus populations were collected. No genetic differentiation, especially with respect to host species associations, could be detected. Moreover, no relationship could be established between closely related P. chalcographus haplotypes and taxonomically related conifer species.ConclusionThe capacity of P. chalcographus for host shifting and dispersal may have played a key role in the rapid extension of its host range. These findings are important for pest management in forests and health and phytosanitary measures in the timber trade, especially for risk assessment in mixed coniferous forests including tree species of major economic importance.

•The study aimed at clarifying basic life traits (fecundity, mortality and development) of Monochamus galloprovincialis, the vector of Bursaphelenchus xylophilus, the causative agent of the Pine Wilt Disease, in Portugal.• Mating and oviposition experiments were conducted on P. sylvestris logs under laboratory conditions during two years. Larval development was followed outdoors.• Fecundity was high, with 138.2 eggs per female.• For the first time, the number of larval instars (4) in M. galloprovincialis was recognized and instars were described.• Informations are provided on the shape and size of the larval galleries. Boring of galleries in sapwood and heartwood started at the third instar. All instars could overwinter but adults emerged simultaneously resulting in a univoltine life cycle for the majority of the individuals.• An obligatory diapause in the forth instar was assumed. However, 8.1% of the insects had a two year development.• Important differences in mortality were noted between the two experimental years, which could be explained by differences on the size of the breeding logs.• There is a high biological proximity between M. galloprovincialis populations in France and those in Portugal and Northern Europe. However, a higher fecundity and longevity was observed in France, as well as a higher percentage of insects with a two year development, compared with portuguese populations.

As a consequence of the recent introduction of the pine wood nematode Bursaphelenchus xylophilus in Portugal, nematodes of the genus Bursaphelenchus were looked for in various French pine forests, in trees attacked by Monochamus galloprovincialis, the vector insect of B. xylophilus, and in the insects themselves. Trap trees were felled in 12 localities distributed all over the country. Nematodes were extracted from transversal stem discs; insects emerging from the trap trees were studied. B. hellenicus, B. leoni, B. mucronatus and B. sexdentati were isolated, but not B. xylophilus. The presence of B. mucronatus and the absence of B. xylophilus were confirmed by molecular markers. B. mucronatus was isolated from several regions with an average prevalence of 19%. The infestation of M. galloprovincialis by B. mucronatus reached 26.7%. The wide distribution of B. mucronatus in France could have an effect on the extension of B. xylophilus in a case of an introduction.

Ce livre présente l'état actuel des recherches entomologiques dans les divers écosystèmes forestiers méditerranéens du monde. Unique en son genre, il comble une lacune importante dans le domaine de l'entomologie forestière et méditerranéenne. Il comprend 4 parties principales : bilans des travaux de recherches ; biodiversité entomologique ; cycles biologiques et relations avec les arbres hôtes ; surveillance et contrôle des populations d'insectes. Chercheurs, enseignants et étudiants du supérieur ainsi que gestionnaires et décideurs forestiers y trouveront matière à réflexion et de nombreuses informations actualisées.

Two Norway spruce (Picea abies Karst.) clones, one resistant and the other susceptible to mass inoculation with Ophiostoma polonicum Siem., were compared with regard to their phenolic compositions and chalcone synthase (CHS) and synthase activities of their phloem before and at 6 and 12 d after artificial inoculation with sterile malt agar or O. polonicum. In unwounded phloem, the resistant clone differed from the susceptible clone by the presence of taxifolin glycoside, lower concentrations of stilbene glycosides, and higher CHS activity. After inoculation, (+)-catechin concentration and CHS activity dramatically increased around the wound, particularly in the resistant clone. Stilbene synthase activity also increased, but more slowly and to a lower level, whereas the concentrations of stilbenes remained stable. Tanning ability decreased in the susceptible clone, whereas it remained stable in the resistant one. It is proposed that the induced phenolic response of Norway spruce phloem consists of an activation of the phenolic pathway, finally leading to tannins and insoluble polymers. It is suggested that resistance to O. polonicum depends on the ability of the tree to easily activate the flavonoid pathway.

The S-adenosyl-L-methionine:pinosylvin-O-methyltransferase $(PMT)^{2}$ gene was sequenced from Scots pine (Pinus sylvestris). The open reading frame is arranged in two exons spaced by one 102-bp intron. Promoter regulatory elements such as two \"CAAT\" boxes and one \"TATA\" box were identified. Several cis-regulatory elements were recognized: stress-responsive elements (Myb-responsive elements) as well as G, H, and GC boxes. Moreover, elicitor-responsive elements (W boxes) and a sequence resembling the simian virus 40 enhancer core were found. In phloem and needles of control trees, the transcripts of stilbene synthase (STS) and PMT were hardly detectable. Increased ozone fumigation up to 0.3 μL L-1 enhanced the transcript level of STS and PMT in needles but not in healthy phloem. Wounding, e.g. mock inoculation, of stem-phloem was characterized by a transient increase in STS and PMT transcripts, which was more pronounced in the case of fungal inoculation. Combination of fungal-challenge or mock treatment with ozone resulted in a positive interaction at 0.3 μL L-1. Scots pine stilbene formation appeared to be induced via STS and PMT gene expression upon ozone and fungal stress as well as wounding. The broad stress-responsiveness is in agreement with the range of various cis-acting elements detected in the STS and PMT promoters.

Phloeomyzus passerinii can induce a pseudogall within the bark of susceptible poplar genotypes, while in resistant genotypes the induction seems to be impeded by lignification processes. Phloeomyzus passerinii is a major pest of poplar stands in Europe, North Africa and the Near East. This aphid feeds in the cortical parenchyma of different poplar species and hybrids, and can affect their growth and survival through unknown mechanisms. In some genotypes, however, resistance prevents either the settlement or the development of aphid colonies. For a better understanding of tree reactions to aphid probing, we compared the anatomical and biochemical modifications undergone within the bark of stem cuttings, after different delays of either aphid colonization or mechanical wounding. To assess how resistance may modulate tree reactions, the comparison was performed using three poplar genotypes exhibiting different resistance levels. In these three genotypes, mechanical wounding induced a similar, localized, wound periderm. In contrast, aphid colonization triggered more extended reactions, which differed among genotypes. In the susceptible genotype, aphids induced a reaction tissue, characterized after a month by thin-walled hypertrophied cells and a depletion of soluble phenolic compounds and starch. Anatomical features of this reaction tissue suggest that the aphid initiates a pseudogall in the cortical tissues of its susceptible host. In the resistant genotypes, however, the differentiation of the reaction tissue was totally or partially inhibited, probably because of extended lignification processes. The implications of a pseudogall induction on susceptible hosts' physiology, and the impact of lignification on aphid development and behavior, are discussed.