Herbivory on the pedunculate oak along an urbanization gradient in Europe : Effects of impervious surface, local tree cover, and insect feeding guild

Urbanization is an important driver of the diversity and abundance of tree-associated insect herbivores, but its consequences for insect herbivory are poorly understood. A likely source of variability among studies is the insufficient consideration of intra-urban variability in forest cover. With the help of citizen scientists, we investigated the independent and interactive effects of local canopy cover and percentage of impervious surface on insect herbivory in the pedunculate oak (Quercus robur L.) throughout most of its geographic range in Europe. We found that the damage caused by chewing insect herbivores as well as the incidence of leaf-mining and gall-inducing herbivores consistently decreased with increasing impervious surface around focal oaks. Herbivory by chewing herbivores increased with increasing forest cover, regardless of impervious surface. In contrast, an increase in local canopy cover buffered the negative effect of impervious surface on leaf miners and strengthened its effect on gall inducers. These results show that-just like in non-urban areas-plant-herbivore interactions in cities are structured by a complex set of interacting factors. This highlights that local habitat characteristics within cities have the potential to attenuate or modify the effect of impervious surfaces on biotic interactions.Peer reviewe

Herbivory on the pedunculate oak along an urbanization gradient in Europe : Effects of impervious surface, local tree cover, and insect feeding guild

Urbanization is an important driver of the diversity and abundance of tree-associated insect herbivores, but its consequences for insect herbivory are poorly understood. A likely source of variability among studies is the insufficient consideration of intraurban variability in forest cover. With the help of citizen scientists, we investigated the independent and interactive effects of local canopy cover and percentage of impervious surface on insect herbivory in the pedunculate oak (Quercus robur L.) throughout most of its geographic range in Europe. We found that the damage caused by chewing insect herbivores as well as the incidence of leaf-mining and gall-inducing herbivores consistently decreased with increasing impervious surface around focal oaks. Herbivory by chewing herbivores increased with increasing forest cover, regardless of impervious surface. In contrast, an increase in local canopy cover buffered the negative effect of impervious surface on leaf miners and strengthened its effect on gall inducers. These results show that – just like in non-urban areas – plant-herbivore interactions in cities are structured by a complex set of interacting factors. This highlights that local habitat characteristics within cities have the potential to attenuate or modify the effect of impervious surfaces on biotic interactions.Agence Nationale de la Recherche, Grant/Award Number: ANR-10--LABX-45; Fondation BNP Paribas.info:eu-repo/semantics/publishedVersio

Herbivory on the pedunculate oak along an urbanization gradient in Europe: Effects of impervious surface, local tree cover, and insect feeding guild

Urbanization is an important driver of the diversity and abundance of tree-associated insect herbivores, but its consequences for insect herbivory are poorly understood. A likely source of variability among studies is the insufficient consideration of intra-urban variability in forest cover. With the help of citizen scientists, we investigated the independent and interactive effects of local canopy cover and percentage of impervious surface on insect herbivory in the pedunculate oak (Quercus robur L.) throughout most of its geographic range in Europe. We found that the damage caused by chewing insect herbivores as well as the incidence of leaf-mining and gall-inducing herbivores consistently decreased with increasing impervious surface around focal oaks. Herbivory by chewing herbivores increased with increasing forest cover, regardless of impervious surface. In contrast, an increase in local canopy cover buffered the negative effect of impervious surface on leaf miners and strengthened its effect on gall inducers. These results show that-just like in non-urban areas-plant-herbivore interactions in cities are structured by a complex set of interacting factors. This highlights that local habitat characteristics within cities have the potential to attenuate or modify the effect of impervious surfaces on biotic interactions

Tree biodiversity and forest resistance to biological invasions : application on chestnut and its exotic pest complex, chestnut blight (Cryponectria parasitica) and Asian chestnut gall wasp (Dryocosmus Kuriphilus)

Les plantes sont au centre d’une grande diversité d’interactions biotiques entre organismes plus ou moins proches qui les exploitent en tant que ressources. L’objectif de cette thèse a été de comprendre comment les infections fongiques de la plante et la diversité des arbres en forêt modifient les interactions arbres-insectes. Nous avons tout d’abord effectué une méta-analyse pour poser le cadre théorique des effets indirects des infections fongiques sur les insectes herbivores associés aux mêmes plantes hôtes. L'effet de l’infection préalable des plantes par les champignons sur les préférences et performances des insectes s’avère généralement négatif. Cependant, la magnitude de cet effet délétère varie selon le mode de vie du champignon, la guilde trophique de l’insecte et la spatialité des interactions (interactions locales vs distantes). Nous avons ensuite analysé de façon empirique les interactions tripartites entre le châtaignier européen (Castanea sativa) et deux de ses bioagresseurs exotiques: le cynips (Dryocosmus kuriphilus), insecte galligène, et Cryphonectria parasitica, champignon pathogène responsable de la maladie du chancre. L'effet sur les taux d’infestation par le cynips de la composition spécifique en essences forestières des forêts de châtaigniers atteintes de chancre a été également étudié. Afin d'identifier les mécanismes sous-jacents aux effets de la diversité des forêts sur cet insecte invasif, les communautés d'insectes parasitoïdes et de champignons endophytes présents dans les galles ont été décrites. Les taux d’infection par le cynips étaient plus faibles dans les mélanges de châtaignier avec du chêne et du frêne que dans des parcelles de châtaignier monospécifiques ou dans les mélanges avec du pin. La composition des forêts influence aussi la composition des communautés de parasitoïdes associés aux galles du cynips mais pas leur abondance, richesse ou diversité. Les communautés de champignons endophytes des galles, étudiées par des méthodes de séquençage de nouvelle génération, sont indépendantes de la composition forestière. Par contre, celles présentes dans les galles différent fortement de celles des tissus foliaires adjacents. Nous avons ainsi apporté de nouvelles preuves que la diversité des plantes et les champignons pathogènes sont des facteurs clés déterminant les interactions plantes-insectes. Etudier comment les plantes interagissent avec leurs insectes et champignons associés, et les mécanismes sous-jacents à l’effet de la diversité des plantes sur ces interactions, doit permettre de mieux comprendre les relations entre diversité et fonctionnement des écosystèmes et de proposer des applications pour la gestion des bio-agresseurs forestiers natifs et exotiques.Plants are the playground of a large diversity of biotic interactions between related and unrelated organisms exploiting them as common resources. The aim of this thesis was to understand how plant-insect interactions vary with fungal infection of their host plant and plant diversity. I first performed a meta-analysis to provide a theoretical background for plant-mediated effects of fungal infection on herbivorous insects. Overall, I found a negative plant-mediated effect of fungi on both insect preference and performance. However, this effect varied according to fungus lifestyle, insect feeding guild and spatial location of the interactions (local vs distant). Then I experimentally tested plant-fungus-insect tripartite interactions in the particular case of exotic bio-aggressors of the European chestnut (Castanea sativa): the Asian chestnut Gall Wasp (ACGW, Dryocosmus kuriphilus), and the fungal pathogen Cryphonectria parasitica, the causal agent of chestnut blight. I performed an observational study, in natural chestnut forest stands in Italy, where I tested how ACGW infestation rates vary with the tree species composition. I also investigated the mechanisms underlying plant diversity effects on the invasive pest, with a particular focus on its natural enemies such as insect parasitoids and endophytic fungi. ACGW infestation rates was lower in oak and ash chestnut mixtures compared to monocultures or pine-chestnut mixtures. Plot composition also influenced ACGW parasitoid community composition but not their abundances, diversity or richness. Endophytic communities of galls, described by using next generation sequencing methods, did not vary with plot composition. However, they strongly differed from surrounding leaf tissues. We thus provided evidence that plant diversity and fungal pathogens are key drivers of plant-insect interactions. Understanding how plants interact with associated insects and fungi, and mechanisms underlying plant diversity effect on these interactions, will improve our knowledge on diversity-ecosystem functioning relationships and will have practical applications for the management of native and exotic forest pests

Tree biodiversity and forest resistance to biological invasions : application on chestnut and its exotic pest complex, chestnut blight (Cryponectria parasitica) and Asian chestnut gall wasp (Dryocosmus Kuriphilus)

Les plantes sont au centre d’une grande diversité d’interactions biotiques entre organismes plus ou moins proches qui les exploitent en tant que ressources. L’objectif de cette thèse a été de comprendre comment les infections fongiques de la plante et la diversité des arbres en forêt modifient les interactions arbres-insectes. Nous avons tout d’abord effectué une méta-analyse pour poser le cadre théorique des effets indirects des infections fongiques sur les insectes herbivores associés aux mêmes plantes hôtes. L'effet de l’infection préalable des plantes par les champignons sur les préférences et performances des insectes s’avère généralement négatif. Cependant, la magnitude de cet effet délétère varie selon le mode de vie du champignon, la guilde trophique de l’insecte et la spatialité des interactions (interactions locales vs distantes). Nous avons ensuite analysé de façon empirique les interactions tripartites entre le châtaignier européen (Castanea sativa) et deux de ses bioagresseurs exotiques: le cynips (Dryocosmus kuriphilus), insecte galligène, et Cryphonectria parasitica, champignon pathogène responsable de la maladie du chancre. L'effet sur les taux d’infestation par le cynips de la composition spécifique en essences forestières des forêts de châtaigniers atteintes de chancre a été également étudié. Afin d'identifier les mécanismes sous-jacents aux effets de la diversité des forêts sur cet insecte invasif, les communautés d'insectes parasitoïdes et de champignons endophytes présents dans les galles ont été décrites. Les taux d’infection par le cynips étaient plus faibles dans les mélanges de châtaignier avec du chêne et du frêne que dans des parcelles de châtaignier monospécifiques ou dans les mélanges avec du pin. La composition des forêts influence aussi la composition des communautés de parasitoïdes associés aux galles du cynips mais pas leur abondance, richesse ou diversité. Les communautés de champignons endophytes des galles, étudiées par des méthodes de séquençage de nouvelle génération, sont indépendantes de la composition forestière. Par contre, celles présentes dans les galles différent fortement de celles des tissus foliaires adjacents. Nous avons ainsi apporté de nouvelles preuves que la diversité des plantes et les champignons pathogènes sont des facteurs clés déterminant les interactions plantes-insectes. Etudier comment les plantes interagissent avec leurs insectes et champignons associés, et les mécanismes sous-jacents à l’effet de la diversité des plantes sur ces interactions, doit permettre de mieux comprendre les relations entre diversité et fonctionnement des écosystèmes et de proposer des applications pour la gestion des bio-agresseurs forestiers natifs et exotiques.Plants are the playground of a large diversity of biotic interactions between related and unrelated organisms exploiting them as common resources. The aim of this thesis was to understand how plant-insect interactions vary with fungal infection of their host plant and plant diversity. I first performed a meta-analysis to provide a theoretical background for plant-mediated effects of fungal infection on herbivorous insects. Overall, I found a negative plant-mediated effect of fungi on both insect preference and performance. However, this effect varied according to fungus lifestyle, insect feeding guild and spatial location of the interactions (local vs distant). Then I experimentally tested plant-fungus-insect tripartite interactions in the particular case of exotic bio-aggressors of the European chestnut (Castanea sativa): the Asian chestnut Gall Wasp (ACGW, Dryocosmus kuriphilus), and the fungal pathogen Cryphonectria parasitica, the causal agent of chestnut blight. I performed an observational study, in natural chestnut forest stands in Italy, where I tested how ACGW infestation rates vary with the tree species composition. I also investigated the mechanisms underlying plant diversity effects on the invasive pest, with a particular focus on its natural enemies such as insect parasitoids and endophytic fungi. ACGW infestation rates was lower in oak and ash chestnut mixtures compared to monocultures or pine-chestnut mixtures. Plot composition also influenced ACGW parasitoid community composition but not their abundances, diversity or richness. Endophytic communities of galls, described by using next generation sequencing methods, did not vary with plot composition. However, they strongly differed from surrounding leaf tissues. We thus provided evidence that plant diversity and fungal pathogens are key drivers of plant-insect interactions. Understanding how plants interact with associated insects and fungi, and mechanisms underlying plant diversity effect on these interactions, will improve our knowledge on diversity-ecosystem functioning relationships and will have practical applications for the management of native and exotic forest pests

Diversité des arbres et résistance des forêts aux invasions biologiques : application au chataignier et son complexe de bioagresseurs exotiques, chancre (Cryphonectria parasitica) et cynips (Dryocosmus Kuriphilus)

Plants are the playground of a large diversity of biotic interactions between related and unrelated organisms exploiting them as common resources. The aim of this thesis was to understand how plant-insect interactions vary with fungal infection of their host plant and plant diversity. I first performed a meta-analysis to provide a theoretical background for plant-mediated effects of fungal infection on herbivorous insects. Overall, I found a negative plant-mediated effect of fungi on both insect preference and performance. However, this effect varied according to fungus lifestyle, insect feeding guild and spatial location of the interactions (local vs distant). Then I experimentally tested plant-fungus-insect tripartite interactions in the particular case of exotic bio-aggressors of the European chestnut (Castanea sativa): the Asian chestnut Gall Wasp (ACGW, Dryocosmus kuriphilus), and the fungal pathogen Cryphonectria parasitica, the causal agent of chestnut blight. I performed an observational study, in natural chestnut forest stands in Italy, where I tested how ACGW infestation rates vary with the tree species composition. I also investigated the mechanisms underlying plant diversity effects on the invasive pest, with a particular focus on its natural enemies such as insect parasitoids and endophytic fungi. ACGW infestation rates was lower in oak and ash chestnut mixtures compared to monocultures or pine-chestnut mixtures. Plot composition also influenced ACGW parasitoid community composition but not their abundances, diversity or richness. Endophytic communities of galls, described by using next generation sequencing methods, did not vary with plot composition. However, they strongly differed from surrounding leaf tissues. We thus provided evidence that plant diversity and fungal pathogens are key drivers of plant-insect interactions. Understanding how plants interact with associated insects and fungi, and mechanisms underlying plant diversity effect on these interactions, will improve our knowledge on diversity-ecosystem functioning relationships and will have practical applications for the management of native and exotic forest pests.Les plantes sont au centre d’une grande diversité d’interactions biotiques entre organismes plus ou moins proches qui les exploitent en tant que ressources. L’objectif de cette thèse a été de comprendre comment les infections fongiques de la plante et la diversité des arbres en forêt modifient les interactions arbres-insectes. Nous avons tout d’abord effectué une méta-analyse pour poser le cadre théorique des effets indirects des infections fongiques sur les insectes herbivores associés aux mêmes plantes hôtes. L'effet de l’infection préalable des plantes par les champignons sur les préférences et performances des insectes s’avère généralement négatif. Cependant, la magnitude de cet effet délétère varie selon le mode de vie du champignon, la guilde trophique de l’insecte et la spatialité des interactions (interactions locales vs distantes). Nous avons ensuite analysé de façon empirique les interactions tripartites entre le châtaignier européen (Castanea sativa) et deux de ses bioagresseurs exotiques: le cynips (Dryocosmus kuriphilus), insecte galligène, et Cryphonectria parasitica, champignon pathogène responsable de la maladie du chancre. L'effet sur les taux d’infestation par le cynips de la composition spécifique en essences forestières des forêts de châtaigniers atteintes de chancre a été également étudié. Afin d'identifier les mécanismes sous-jacents aux effets de la diversité des forêts sur cet insecte invasif, les communautés d'insectes parasitoïdes et de champignons endophytes présents dans les galles ont été décrites. Les taux d’infection par le cynips étaient plus faibles dans les mélanges de châtaignier avec du chêne et du frêne que dans des parcelles de châtaignier monospécifiques ou dans les mélanges avec du pin. La composition des forêts influence aussi la composition des communautés de parasitoïdes associés aux galles du cynips mais pas leur abondance, richesse ou diversité. Les communautés de champignons endophytes des galles, étudiées par des méthodes de séquençage de nouvelle génération, sont indépendantes de la composition forestière. Par contre, celles présentes dans les galles différent fortement de celles des tissus foliaires adjacents. Nous avons ainsi apporté de nouvelles preuves que la diversité des plantes et les champignons pathogènes sont des facteurs clés déterminant les interactions plantes-insectes. Etudier comment les plantes interagissent avec leurs insectes et champignons associés, et les mécanismes sous-jacents à l’effet de la diversité des plantes sur ces interactions, doit permettre de mieux comprendre les relations entre diversité et fonctionnement des écosystèmes et de proposer des applications pour la gestion des bio-agresseurs forestiers natifs et exotiques

Tree biodiversity and forest resistance to biological invasions : application on chestnut and its exotic pest complex, chestnut blight (Cryponectria parasitica) and Asian chestnut gall wasp (Dryocosmus Kuriphilus)

Les plantes sont au centre d’une grande diversité d’interactions biotiques entre organismes plus ou moins proches qui les exploitent en tant que ressources. L’objectif de cette thèse a été de comprendre comment les infections fongiques de la plante et la diversité des arbres en forêt modifient les interactions arbres-insectes. Nous avons tout d’abord effectué une méta-analyse pour poser le cadre théorique des effets indirects des infections fongiques sur les insectes herbivores associés aux mêmes plantes hôtes. L'effet de l’infection préalable des plantes par les champignons sur les préférences et performances des insectes s’avère généralement négatif. Cependant, la magnitude de cet effet délétère varie selon le mode de vie du champignon, la guilde trophique de l’insecte et la spatialité des interactions (interactions locales vs distantes). Nous avons ensuite analysé de façon empirique les interactions tripartites entre le châtaignier européen (Castanea sativa) et deux de ses bioagresseurs exotiques: le cynips (Dryocosmus kuriphilus), insecte galligène, et Cryphonectria parasitica, champignon pathogène responsable de la maladie du chancre. L'effet sur les taux d’infestation par le cynips de la composition spécifique en essences forestières des forêts de châtaigniers atteintes de chancre a été également étudié. Afin d'identifier les mécanismes sous-jacents aux effets de la diversité des forêts sur cet insecte invasif, les communautés d'insectes parasitoïdes et de champignons endophytes présents dans les galles ont été décrites. Les taux d’infection par le cynips étaient plus faibles dans les mélanges de châtaignier avec du chêne et du frêne que dans des parcelles de châtaignier monospécifiques ou dans les mélanges avec du pin. La composition des forêts influence aussi la composition des communautés de parasitoïdes associés aux galles du cynips mais pas leur abondance, richesse ou diversité. Les communautés de champignons endophytes des galles, étudiées par des méthodes de séquençage de nouvelle génération, sont indépendantes de la composition forestière. Par contre, celles présentes dans les galles différent fortement de celles des tissus foliaires adjacents. Nous avons ainsi apporté de nouvelles preuves que la diversité des plantes et les champignons pathogènes sont des facteurs clés déterminant les interactions plantes-insectes. Etudier comment les plantes interagissent avec leurs insectes et champignons associés, et les mécanismes sous-jacents à l’effet de la diversité des plantes sur ces interactions, doit permettre de mieux comprendre les relations entre diversité et fonctionnement des écosystèmes et de proposer des applications pour la gestion des bio-agresseurs forestiers natifs et exotiques.Plants are the playground of a large diversity of biotic interactions between related and unrelated organisms exploiting them as common resources. The aim of this thesis was to understand how plant-insect interactions vary with fungal infection of their host plant and plant diversity. I first performed a meta-analysis to provide a theoretical background for plant-mediated effects of fungal infection on herbivorous insects. Overall, I found a negative plant-mediated effect of fungi on both insect preference and performance. However, this effect varied according to fungus lifestyle, insect feeding guild and spatial location of the interactions (local vs distant). Then I experimentally tested plant-fungus-insect tripartite interactions in the particular case of exotic bio-aggressors of the European chestnut (Castanea sativa): the Asian chestnut Gall Wasp (ACGW, Dryocosmus kuriphilus), and the fungal pathogen Cryphonectria parasitica, the causal agent of chestnut blight. I performed an observational study, in natural chestnut forest stands in Italy, where I tested how ACGW infestation rates vary with the tree species composition. I also investigated the mechanisms underlying plant diversity effects on the invasive pest, with a particular focus on its natural enemies such as insect parasitoids and endophytic fungi. ACGW infestation rates was lower in oak and ash chestnut mixtures compared to monocultures or pine-chestnut mixtures. Plot composition also influenced ACGW parasitoid community composition but not their abundances, diversity or richness. Endophytic communities of galls, described by using next generation sequencing methods, did not vary with plot composition. However, they strongly differed from surrounding leaf tissues. We thus provided evidence that plant diversity and fungal pathogens are key drivers of plant-insect interactions. Understanding how plants interact with associated insects and fungi, and mechanisms underlying plant diversity effect on these interactions, will improve our knowledge on diversity-ecosystem functioning relationships and will have practical applications for the management of native and exotic forest pests

Citizen science and monitoring forest pests

Purpose of the Review One of the major threats to tree health, and hence the resilience of forests and their provision of ecosystem services, is new and emerging pests. Therefore, forest health monitoring is of major importance to detect invasive, emerging and native pest outbreaks. This is usually done by foresters and forest health experts, but can also be complemented by citizen scientists. Here, we review the use of citizen science for detection and monitoring, as well as for hypothesis-driven research and evaluation of control measures as part of forest pest surveillance and research. We then examine its limitations and opportunities and make recommendations on the use of citizen science for forest pest monitoring. Recent Findings The main opportunities of citizen scientists for forest health are early warning, early detection of new pests, monitoring of impact of outbreaks and scientific research. Each domain has its own limitations, opportunities and recommendations to follow, as well as their own public engagement strategies. The development of new technologies provides many opportunities to involve citizen scientists in forest pest monitoring. To enhance the benefits of citizen scientists’ inclusion in monitoring, it is important that they are involved in the cocreation of activities. Summary Future monitoring and research may benefit from tailor-made citizen science projects to facilitate successful monitoring by citizen scientists and expand their practice to countries where the forest health sector is less developed. In this sense, citizen scientists can help understand and detect outbreaks of new pests and avoid problems in the future

Fungal endophyte communities differ between chestnut galls and surrounding foliar tissues

Foliar endophytic fungi are present in almost all vascular plants. The composition of endophyte communities varies among plant individuals. Likely, but understudied, sources of this variation are the species composition of the plant community and initial attacks by insect herbivores. We addressed these issues by characterizing fungal endophyte communities on leaves of chestnut (Castanea saliva) grown in pure vs. mixed stands. We used ITS metabarcoding methods to identify endophytic fungi associated with galls caused by the invasive gall wasp, Dryocosmus kuriphilus, and with surrounding chestnut leaf tissues. We found 1378 different OTUs. The richness, diversity and composition of endophyte communities differed between galls and surrounding leaf tissues but were independent of forest stand composition. Fungal endophyte richness was lower in galls than in surrounding leaf tissues. Most differences in the composition of fungal endophyte communities between galls and foliar tissues were due to OTU turnover. These results suggest that insect-induced galls provide a particular habitat condition for endophytic microorganisms, regardless of forest species composition. A better understanding of endophyte biology is important to improve their use as biocontrol agents of galling insects

The effect of community-wide phytochemical diversity on herbivory reverses from low to high elevation

1. Theory predicts that a large fraction of phytochemical diversity—the richness of individual chemical compounds produced by plants—governs the complexity of interactions between plants and their herbivores. While the effect of specific classes of chemical compounds on plant resistance against herbivores has been largely documented, the effect of community-level variation in phytochemical diversity on plant–herbivore interactions has so far received minimal consideration. 2. We hypothesized that plant communities bearing on average higher levels of phytochemical diversity should sustain lower herbivory rates, overall. Yet, the magnitude of this effect could vary across different environmental conditions, potentially because of climate-mediated effects on phytochemical production and changes in herbivore community richness and composition. 3. To address these hypotheses, we used previous knowledge of species-level phytochemical make-up for more than 400 plant species of the Swiss Alps. Using common garden experiments, we estimated season-wide herbivore damage on low (average 3,500 unique molecules) and high (average 4,500 unique molecules) phytochemical diversity plant communities that were planted in the colline, mountain and alpine vegetation sites along two elevation transects in the Alps. 4. We found that high phytochemical diversity plant communities showed reduced levels of herbivore damage in the colline (low elevation) sites, but this pattern reversed in the alpine (high elevation) sites. Our results suggest that the outcome of phytochemical diversity on plant–herbivore interactions depends on the characteristics of the local herbivore communities, together with trade-offs between chemical defences and other plant traits (i.e. physical defences and plant palatability). 5. Synthesis. Phytochemical diversity is a key component of functional diversity, influencing community composition and dynamics. We show that the effect of phytochemical diversity on herbivory is environmental-dependent, generating ecological switches when moving from low to high elevation. Through upward movement of plants under climate change, phytochemical community structure will be likely modified, ultimately disrupting local community assembly processes.ISSN:0022-047