Primary and secondary host selection by Ips typographus depends on Norway spruce crown characteristics and phenolic-based defenses.

Climate change is expected to intensify bark beetle population outbreaks in forests globally, affecting biodiversity and trajectories of change. Aspects of individual tree resistance remain poorly quantified, particularly with regard to the role of phenolic compounds, hindering robust predictions of forest response to future conditions. In 2003, we conducted a mechanical wounding experiment in a Norway spruce forest that coincided with an outbreak of the bark beetle, Ips typographus. We collected phloem samples from 97 trees and monitored tree survival for 5 months. Using high-performance liquid chromatography, we quantified induced changes in the concentrations of phenolics. Classification and regression tools were used to evaluate relationships between phenolic production and bark beetle resistance, in the context of other survival factors. The proximity of beetle source populations was a principal determinant of survival. Proxy measures of tree vigor, such as crown defoliation, mediated tree resistance. Controlling for these factors, synthesis of catechin was found to exponentially increase tree survival probability. However, even resistant trees were susceptible in late season due to high insect population growth. Our results show that incorporating trait-mediated effects improves predictions of survival. Using an integrated analytical approach, we demonstrate that phenolics play a direct role in tree defense to herbivory.

Characterization of antibiosis and antixenosis to the woolly poplar aphid (Hemiptera: Aphididae) in the bark of different poplar genotypes.

The woolly poplar aphid, Phloeomyzus passerinii (Signoret) (Hemiptera: Aphididae), is a major pest of poplar plantations in the Mediterranean basin and the Near East. Aphids colonize poplar trunks and feed upon the cortical parenchyma. Despite the economic importance of poplar, little is known about the mechanisms involved in poplar resistance to this pest. However, Populus x canadensis Moench genotypes show various levels of resistance to P. passerinii. This study has investigated the type of poplar resistance (antibiosis or antixenosis) by assessing aphid settlement, physiology (survival, development, and reproduction), and stylet penetration behavior (electrical penetration graph) on three P. x canadensis genotypes; '1214' (susceptible), 'Brenta' (resistant), and '145/51' (intermediate). Because settlement was reduced, the highly resistant genotype Brenta exhibited surface antixenosis. In addition, nymphal survival was null on Brenta, and twice less adult aphid initiated a sustained intracellular phase in the cortical parenchyma of that genotype compared with the other two genotypes. Thus, Brenta also showed parenchyma-located antixenosis coupled with antibiosis characteristic. In contrast, P. passerinii had no difficulty to initiate a sustained ingestion in the cortical parenchyma of the intermediate genotype 145/51, but decreased fecundity and lower intrinsic rate of natural increase were clear expressions of antibiosis.

Exceptional plant penetration and feeding upon cortical parenchyma cells by the woolly poplar aphid.

Forty percent of aphids live wholly or partly on trees, most species being associated with leaves or petioles. Species able to exploit woody parts have either specific adaptations, such as extra long stylets that allow them to reach the phloem, or the ability to induce galls. The woolly poplar aphid, Phloeomyzus passerinii (Signoret) (Hemiptera: Aphididae), colonizes the trunks and base of the lower branches of mature poplars and causes cortical necrosis leading to the death of trees where infestation is heavy. Very little is known about the mode of feeding of P. passerinii. This study looked at the feeding behavior of P. passerinii on stem-cuttings of Populus x canadensis Moench using: (i) histological analyses of the feeding site and stylet pathway and (ii) electrical penetration graphs (EPG, DC) based on parthenogenetic apterous females on woody tissues. The histological and EPG results showed that stylets of P. passerinii penetrated into the plant tissues following a straight unbranched extracellular and intracellular pathway to reach the cortical parenchyma. Compared to EPGs for phloem sap feeding aphids, there were differences in the waveforms A and C whereas a new waveform Icp was described. Based on histological analyses and previous descriptions of EPG waveforms, correlations with the stylet tip position and aphid activities within bark tissues are discussed. A pathway and a sustained intracellular phase were distinguished, both occurring in the cortical parenchyma cells. The bark aphid feeding mode is discussed in relation to the damage caused and in terms of changes in the aphid's diet.

Novel insect-tree associations resulting from accidental and intentional biological 'invasions': a meta-analysis of effects on insect fitness.

The translocation of species beyond their native range is a major threat to biodiversity. Invasions by tree-feeding insects attacking native trees and the colonization of introduced trees by native insects result in new insect-tree relationships. To date there is uncertainty about the key factors that influence the outcome of these novel interactions. We report the results of a meta-analysis of 346 pairwise comparisons of forest insect fitness on novel and ancient host tree species from 31 publications. Host specificity of insects and phylogenetic relatedness between ancient and novel host trees emerged as key factors influencing insect fitness. Overall, fitness was significantly lower on novel host species than on ancient hosts. However, in some cases, fitness increased on novel hosts, mainly in polyphagous insects or when close relatives of ancient host trees were colonized. Our synthesis enables greatly improved impact prediction and risk assessment of biological invasions.

Seasonal water stress and the resistance of Pinus yunnanensis to a bark-beetle-associated fungus.

We examined the influence of seasonal water stress on the resistance of Pinus yunnanensis (Franch.) to inoculation with Leptographium yunnanense, a pathogenic fungus associated with the aggressive bark beetle, Tomicus n. sp. Experiments took place between October 1997 and November 1999 in two plots located at the top and at the foot of a hill in Shaogiu, China, a region characterized by dry winters and wet summers. Following isolated and mass fungal inoculations, we observed the reaction zone length, fungal growth in the phloem, and the occlusion, blue-staining and specific hydraulic conductivity of the sapwood. Measurements of soil and needle water contents and predawn needle water potentials confirmed that trees were subject to mild water stress during winter, especially at the drier hilltop site. Measures of tree resistance to fungal infection of phloem and sapwood were congruent and indicated that trees were most susceptible to inoculation during the wet summer, especially at the lower-elevation plot. Specific hydraulic conductivity decreased after inoculation in summer. The results indicate that mild seasonal water stress is not likely responsible for the recent extensive damage to young P. yunnanensis stands by Tomicus n. sp. in the vicinity of our study plots. Rather, the results suggest that mild water stress enhances tree resistance to fungal pathogens associated with Tomicus n. sp.

[Comparison of coleopteran and spider communities between a reforested area and a still steppe area in a pre-Saharan region of Algeria]. / Comparaison des peuplements de Coléoptères et d'Araignées en zone reboisée et en zone steppique dans une région présaharienne d'Algérie.

In the pre-Saharan region of Djelfa (Algeria), the coleopteran and spider communities are compared between two neighbouring ecosystems, the one kept as a steppe, the other one reforested with Aleppo pine. In three plots in each area, pitfall (Barber) traps were sampled monthly during one year and arthropods were identified and counted. Two thousand five hundred and fifty-seven individuals, distributed into 95 species, were collected, belonging to four coleopteran families (Carabidae, Scarabaeidae, Tenebrionidae, and Curculionidae), and 14 spider families. The Tenebrionidae (43% of the species and 87% of the individuals) were clearly the most represented group of insects, numerically because of the considerable abundance of Pimelia mauritanica in the reforested area, where this species accounted for half the total number of catches. Among the spiders, the Gnaphosidae represented 40% of the individuals and one-third of the species. The total species richness of the two areas was the same, but, considering the total number of collected species, individual abundance and diversity indexes (Shannon, Simpson) were lower in the reforested zone. However, these parameters became fully comparable when excluding P. mauritanica from the calculations. Without P. mauritanica, numbers and species richness of Tenebrionidae were similar in both areas. Curculionidae had higher populations and higher species diversity in the steppe area, while Scarabeidae were equally abundant in the two ecosystems types. The number, the species richness, and the diversity indexes of Carabidae were much lower in the reforested area than in the steppe one. The opposite was observed with spiders. Nevertheless, more or less important variations in the number of individuals, in one or another direction depending on the species, existed among each taxonomic group. It is concluded that reforestation has not changed biological diversity, but has induced dramatic modifications in the organisation and functioning of the arthropod communities.

High intercontinental migration rates and population admixture in the sapstain fungus Ophiostoma ips.

Ophiostoma ips is a common fungal associate of various conifer-infesting bark beetles in their native ranges and has been introduced into non-native pine plantations in the Southern Hemisphere. In this study, we used 10 microsatellite markers to investigate the population biology of O. ips in native (Cuba, France, Morocco and USA) and non-native (Australia, Chile and South Africa) areas to characterize host specificity, reproductive behaviour, and the potential origin as well as patterns of spread of the fungus and its insect vectors. The markers resolved a total of 41 alleles and 75 haplotypes. Higher genetic diversity was found in the native populations than in the introduced populations. Based on the origin of the insect vectors, the populations of O. ips in Australia would be expected to reflect a North American origin, and those in Chile and South Africa to reflect a European origin. However, most alleles observed in the native European population were also found in the native North American population; only the allele frequencies among the populations varied. This admixture made it impossible to confirm the origin of the introduced Southern Hemisphere (SH) populations of O. ips. There was also no evidence for specificity of the fungus to particular bark beetle vectors or hosts. Although O. ips is thought to be mainly self-fertilizing, evidence for recombination was found in the four native populations surveyed. The higher genetic diversity in the North American than in the European population suggests that North America could be the possible source region of O. ips.

Host selection in Tomicus piniperda L.: composition of monoterpene hydrocarbons in relation to attack frequency in the shoot feeding phase.

The aim of this study was to investigate the host selection capacity of the pine shoot beetle, Tomicus piniperda, in the shoot-feeding phase and analyze the chiral and non-chiral host volatiles by means of GC-MS and 2D-GC in five Pinus species originating from France (Pinus sylvestris, P. halepensis, P. nigra laricio, P. pinaster maritima, P. pinaster mesogeensis). Dominating monoterpenes were (-)-alpha-pinene, (+)-alpha-pinene, (-)-beta-pinene and (+)-3-carene. The amounts of the enantiomers varied considerably within and among the species. In a principal component analysis-plot, based on the absolute amounts of 18 monoterpene hydrocarbons, separation of the pine species into two groups was obtained. P. halepensis and P. sylvestris were grouped according to the amount of (+)-alpha-pinene and (+)-3-carene, while P. nigra laricio, P. pinaster maritima and P. pinaster mesogeensis were grouped according to (-)-alpha-pinene and (-)-beta-pinene. P. nigra laricio was the species most attacked and P, halepensis the one least attacked by T. piniperda.

Intra- and inter-provenance variability in phloem phenols of Picea abies and relationship to a bark beetle-associated fungus.

One hundred Norway spruce (Picea abies (L.) Karst.) clones (three ramets per clone) were analyzed for phloem phenol composition and concentration before and 10 days after wound inoculation with sterile malt agar. Fifty clones (Experiment 1) belonged to the same provenance, whereas the remaining clones (Experiment 2) belonged to five provenances from three geographic areas. In Experiment 2, two additional ramets from the same clones were mass inoculated with Ceratocystis polonica (Siem.) C. Moreau to quantify the resistance of each clone. Tree response to wound inoculations was characterized by increased catechin concentration in both experiments, accompanied by increases in astringin and decreases in piceid in Experiment 1. In both experiments, we observed a diverse group of phenolic compounds whose concentrations increased (catechin, astringin) or did not vary (taxifolin glucoside) in response to wound inoculations, whereas concentrations of a homogeneous group of stilbene compounds decreased (piceid) or did not vary (isorhapontin, unidentified stilbene). In Experiment 2, provenances from the alpine and Hercynian-Carpatic areas differed from provenances from the Baltic area with respect to the relative importance of these two groups of compounds, further indicating that the two groupings of phenolic compounds structure the Norway spruce populations. Eighty days after mass inoculation, the percentage of healthy sapwood, which was taken as a measure of tree resistance, indicated that clones from the Baltic area were less resistant to mass inoculations than clones from the alpine and Hercynian-Carpatic areas. We conclude that the degree of resistance of Norway spruce trees to mass inoculations with a bark beetle-associated fungus can be predicted based on the diversity of constitutive phloem phenols and the ability to induce phenol synthesis in response to wounding.

Population genetic structure of Tomicus piniperda L. (Curculionidae: Scolytinae) on different pine species and validation of T. destruens (Woll.).

Genetic diversity and population structure of Tomicus piniperda was assessed using mitochondrial sequences on 16 populations sampled on 6 pine species in France. Amplifications of Internal transcribed space 1 (ITS1) were also performed. Our goals were to determine the taxonomic status of the Mediterranean ecotype T. piniperda destruens, and to test for host plant or geographical isolation effect on population genetic structure. We showed that T. piniperda clusters in two mtDNA haplotypic groups. Clade A corresponds to insects sampled in continental France on Pinus sylvestris, P. pinaster and P. uncinata, whereas clade B gathers the individuals sampled in Corsica on P. pinaster and P. radiata and in continental France on P. pinea and P. halepensis. Insects belonging to clade A and clade B also consistently differ in the length of ITS1. Individuals belonging to both clades were found once in sympatry on P. pinaster. Genetic distances between clades are similar to those measured between distinct species of Tomicus. We concluded that clade B actually corresponds to the destruens ecotype and forms a good species, T. destruens. Analyses of molecular variance ( amova ) were conducted separately on T. destruens and T. piniperda to test for an effect of either geographical isolation or host species. Interestingly, the effect of host plant was significant for T. piniperda only, while the effect of geographical isolation was not. Pine species therefore seems to act as a significant barrier to gene flow, even if host race formation is not observed. These results still need to be confirmed by nuclear markers.